Consistent response of bird populations to climate change on two continents.

Global climate change is a major threat to biodiversity. Large-scale analyses have generally focused on the impacts of climate change on the geographic ranges of species and on phenology, the timing of ecological phenomena. We used long-term monitoring of the abundance of breeding birds across Europe and the United States to produce, for both regions, composite population indices for two groups of species: those for which climate suitability has been either improving or declining since 1980. The ratio of these composite indices, the climate impact indicator (CII), reflects the divergent fates of species favored or disadvantaged by climate change. The trend in CII is positive and similar in the two regions. On both continents, interspecific and spatial variation in population abundance trends are well predicted by climate suitability trends.

Continent-scale global change attribution in European birds - combining annual and decadal time scales.

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes-based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long-term population growth rates documents simultaneous responses that can be attributed to both climate change and land-use change, including long-term increases in populations of hot-dwelling species and declines in long-distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long-term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high-resolution time series are available in large-scale biodiversity surveys.

No evidence of the effect of extreme weather events on annual occurrence of four groups of ectothermic species.

Weather extremes may have strong effects on biodiversity, as known from theoretical and modelling studies. Predicted negative effects of increased weather variation are found only for a few species, mostly plants and birds in empirical studies. Therefore, we investigated correlations between weather variability and patterns in occupancy, local colonisations and local extinctions (metapopulation metrics) across four groups of ectotherms: Odonata, Orthoptera, Lepidoptera, and Reptilia. We analysed data of 134 species on a 1×1 km-grid base, collected in the last 20 years from the Netherlands, combining standardised data and opportunistic data. We applied dynamic site-occupancy models and used the results as input for analyses of (i) trends in distribution patterns, (ii) the effect of temperature on colonisation and persistence probability, and (iii) the effect of years with extreme weather on all the three metapopulation metrics. All groups, except butterflies, showed more positive than negative trends in metapopulation metrics. We did not find evidence that the probability of colonisation or persistence increases with temperature nor that extreme weather events are reflected in higher extinction risks. We could not prove that weather extremes have visible and consistent negative effects on ectothermic species in temperate northern hemisphere. These findings do not confirm the general prediction that increased weather variability imperils biodiversity. We conclude that weather extremes might not be ecologically relevant for the majority of species. Populations might be buffered against weather variation (e.g. by habitat heterogeneity), or other factors might be masking the effects (e.g. availability and quality of habitat). Consequently, we postulate that weather extremes have less, or different, impact in real world metapopulations than theory and models suggest.

Predicting rarity and decline in animals, plants, and mushrooms based on species attributes and indicator groups.

In decisions on nature conservation measures, we depend largely on knowledge of the relationship between threats and environmental factors for a very limited number of species groups, with relevant environmental factors often being deduced from the relationship between threat and species traits. But can relationships between traits and levels of threats be identified across species from completely different taxonomic groups; and how accurately do well-known taxonomic groups indicate levels of threat in other species groups? To answer these questions, we first made a list of 152 species attributes of morphological and demographic traits and habitat requirements. Based on these attributes we then grew random forests of decision trees for 1183 species in the 18 different taxonomic groups for which we had Red Lists available in the Netherlands, using these to classify animals, plants, and mushrooms according to their rarity and decline. Finally, we grew random forests for four species groups often used as indicator groups to study how well the relationship between attribute and decline within these groups reflected that relationship within the larger taxonomic group to which these groups belong. Correct classification of rarity based on all attributes was as high as 88% in animals, 85% in plants, and 94% in mushrooms and correct classification of decline was 78% in animals, 69% in plants, and 70% in mushrooms. Vertebrates indicated decline in all animals well, as did birds for all vertebrates and vascular plants for all plants. However, butterflies poorly indicated decline in all insects. Random forests are a useful tool to relate rarity and decline to species attributes thereby making it possible to generalize rarity and decline to a wider set of species groups. Random forests can be used to estimate the level of threat to complete faunas and floras of countries or regions. In regions like the Netherlands, conservation policy based on attributes known to be relevant for the decline to birds, vertebrates or plants will probably also impact all aboveground terrestrial and freshwater macrofauna or macrophytes.

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.

Global biodiversity: indicators of recent declines.

In 2002, world leaders committed, through the Convention on Biological Diversity, to achieve a significant reduction in the rate of biodiversity loss by 2010. We compiled 31 indicators to report on progress toward this target. Most indicators of the state of biodiversity (covering species' population trends, extinction risk, habitat extent and condition, and community composition) showed declines, with no significant recent reductions in rate, whereas indicators of pressures on biodiversity (including resource consumption, invasive alien species, nitrogen pollution, overexploitation, and climate change impacts) showed increases. Despite some local successes and increasing responses (including extent and biodiversity coverage of protected areas, sustainable forest management, policy responses to invasive alien species, and biodiversity-related aid), the rate of biodiversity loss does not appear to be slowing.

Avian population consequences of climate change are most severe for long-distance migrants in seasonal habitats.

One consequence of climate change is an increasing mismatch between timing of food requirements and food availability. Such a mismatch is primarily expected in avian long-distance migrants because of their complex annual cycle, and in habitats with a seasonal food peak. Here we show that insectivorous long-distance migrant species in The Netherlands declined strongly (1984-2004) in forests, a habitat characterized by a short spring food peak, but that they did not decline in less seasonal marshes. Also, within generalist long-distance migrant species, populations declined more strongly in forests than in marshes. Forest-inhabiting migrant species arriving latest in spring declined most sharply, probably because their mismatch with the peak in food supply is greatest. Residents and short-distance migrants had non-declining populations in both habitats, suggesting that habitat quality did not deteriorate. Habitat-related differences in trends were most probably caused by climate change because at a European scale, long-distance migrants in forests declined more severely in western Europe, where springs have become considerably warmer, when compared with northern Europe, where temperatures during spring arrival and breeding have increased less. Our results suggest that trophic mismatches may have become a major cause for population declines in long-distance migrants in highly seasonal habitats.

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.

An indicator of the impact of climatic change on European bird populations.

Rapid climatic change poses a threat to global biodiversity. There is extensive evidence that recent climatic change has affected animal and plant populations, but no indicators exist that summarise impacts over many species and large areas. We use data on long-term population trends of European birds to develop such an indicator. We find a significant relationship between interspecific variation in population trend and the change in potential range extent between the late 20(th) and late 21(st) centuries, forecasted by climatic envelope models. Our indicator measures divergence in population trend between bird species predicted by climatic envelope models to be favourably affected by climatic change and those adversely affected. The indicator shows a rapid increase in the past twenty years, coinciding with a period of rapid warming.

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.

Developing indicators for European birds.

The global pledge to deliver 'a significant reduction in the current rate of biodiversity loss by 2010' is echoed in a number of regional and national level targets. There is broad consensus, however, that in the absence of conservation action, biodiversity will continue to be lost at a rate unprecedented in the recent era. Remarkably, we lack a basic system to measure progress towards these targets and, in particular, we lack standard measures of biodiversity and procedures to construct and assess summary statistics. Here, we develop a simple classification of biodiversity indicators to assist their development and clarify purpose. We use European birds, as example taxa, to show how robust indicators can be constructed and how they can be interpreted. We have developed statistical methods to calculate supranational, multi-species indices using population data from national annual breeding bird surveys in Europe. Skilled volunteers using standardized field methods undertake data collection where methods and survey designs differ slightly across countries. Survey plots tend to be widely distributed at a national level, covering many bird species and habitats with reasonable representation. National species' indices are calculated using log-linear regression, which allows for plot turnover. Supranational species' indices are constructed by combining the national species' indices weighted by national population sizes of each species. Supranational, multi-species indicators are calculated by averaging the resulting indices. We show that common farmland birds in Europe have declined steeply over the last two decades, whereas woodland birds have not. Evidence elsewhere shows that the main driver of farmland bird declines is increased agricultural intensification. We argue that the farmland bird indicator is a useful surrogate for trends in other elements of biodiversity in this habitat.