Summary1. An increasing number of studies are examining the distribution and congruence of ecosystem services, often with the goal of identifying areas that will provide multiple ecosystem service 'hotspots'. However, there is a paucity of data on most ecosystem services, so proxies (e.g. estimates of a service for a particular land cover type) are frequently used to map their distribution. To date, there has been little attempt to quantify the effects of using proxies on distribution maps of ecosystem services, despite the potentially large errors associated with such data sets. 2. Here, we provide the first study examining the effects of using proxies on ecosystem service maps and the degree of spatial congruence of these maps with primary data, using England as a case study. 3. We show that land cover based proxies provide a poor fit to primary data surfaces for biodiversity, recreation and carbon storage, and that correlations between ecosystem services change depending on whether primary or proxy data are used for the analyses. 4. The poor fit of proxies to primary data was also evident when we selected hotspots of single ecosystem services, and consistency between raw and modelled surfaces was extremely low when considering the locations that were coincident hotspots for multiple services. 5. Synthesis and applications. Proxies may be suitable for identifying broad-scale trends in ecosystem services, but even relatively good proxies are likely to be unsuitable for identifying hotspots or priority areas for multiple services.

Summary 1.Ecosystems support biodiversity and also provide goods and services that are beneficial to humans. The extent to which the locations that are most valuable for ecosystem services coincide with those that support the most biodiversity is of critical importance when designing conservation and land management strategies. There are, however, few studies on which to base any kind of conclusion about possible spatial patterns of association between ecosystem services and biodiversity. Moreover, little is known about the sensitivity of the conclusions to the quality of the data available, or to the choice and size of the region used for analysis. 2. Here, we first present national-scale estimates of the spatial covariance in areas important for ecosystem services and biodiversity (richness of species of conservation concern), using Britain as a case study. We then explore how these associations are sensitive to the spatial resolution of the available data, the spatial extent of our study region and to regional variation across the study area. 3. Our analyses reveal a mixture of negative and positive associations. In particular, the regionalization analysis shows that one can arrive at diametrically opposing conclusions about relationships between ecosystem services and biodiversity by studying the same question within different areas, even within a moderately small island. 4. Synthesis and applications. In a policy context, the location-specific nature of relationships between ecosystem services and biodiversity underscores the importance of multi-scale environmental decision-making, so as to reflect both local conditions and broader-scale priorities. The results also suggest that efforts to establish general patterns of congruence in ecosystem services and biodiversity may offer a less constructive way forward than do more regional approaches.

Abstract. Pressure on ecosystems to provide various different and often conflicting services is immense and likely to increase. The impacts and success of conservation prioritization will be enhanced if the needs of competing land uses are recognized at the planning stage. We develop such methods and illustrate them with data about competing land uses in Great Britain, with the aim of developing a conservation priority ranking that balances between needs of biodiversity conservation, carbon storage, agricultural value, and urban development potential. While both carbon stocks and biodiversity are desirable features from the point of view of conservation, they compete with the needs of agriculture and urban development. In Britain the greatest conflicts exist between biodiversity and urban areas, while the largest carbon stocks occur mostly in Scotland in areas with low agricultural or urban pressure. In our application, we were able successfully to balance the spatial allocation of alternative land uses so that conflicts between them were much smaller than had they been developed separately. The proposed methods and software, Zonation, are applicable to structurally similar prioritization problems globally.

Many species are extending their leading‐edge (cool) range margins polewards in response to recent climate change. In the present study, we investigated range margin changes at the northern (cool) range margins of 1573 southerly‐distributed species from 21 animal groups in Great Britain over the past four decades of climate change, updating previous work. Depending on data availability, range margin changes were examined over two time intervals during the past four decades. For four groups (birds, butterflies, macromoths, and dragonflies and damselflies), there were sufficient data available to examine range margin changes over both time intervals. We found that most taxa shifted their northern range margins polewards and this finding was not greatly influenced by changes in recorder effort. The mean northwards range margin change in the first time interval was 23 km per decade (N = 13 taxonomic groups) and, in the second interval, was 18 km per decade (N = 16 taxonomic groups) during periods when the British climate warmed by 0.21 and 0.28 °C per decade, respectively. For the four taxa examined over both intervals, there was evidence for higher rate of range margin change in the more recent time interval in the two Lepidoptera groups. Our analyses confirm a continued range margin shift polewards in a wide range of taxonomic groups.

Many studies have demonstrated the selection of stubble fields by farmland birds in winter, but none have shown whether provisioning of this key habitat positively influences national population trends for widespread farmland birds. We use two complementary extensive bird surveys undertaken at the same localities in summer and winter and show that the area of stubble in winter attracts increased numbers of several bird species of conservation concern. Moreover, for several farmland specialists, the availability of stubble fields in winter positively influenced the 10 year breeding population trend (1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003) whereas hedgerow bird species were less affected. For skylarks and yellowhammers, initially negative trends showed recovery with 10-20 ha of stubble per 1 km square. Thus, agri-environment schemes that promote retention of over-winter stubbles will attract birds locally and are capable of reversing current population declines if stubbles are available in sufficient quantity.

There is good evidence that species' distributions are shifting poleward in response to climate change and wide interest in the magnitude of such responses for scientific and conservation purposes. It has been suggested from the directions of climatic changes that species' distribution shifts may not be simply poleward, but this has been rarely tested with observed data. Here, we apply a novel approach to measuring range shifts on axes ranging through 360°, to recent data on the distributions of 122 species of British breeding birds during 1988-1991 and 2008-2011. Although previously documented poleward range shifts have continued, with an average 13.5 km shift northward, our analysis indicates this is an underestimate because it ignores common and larger shifts that occurred along axes oriented to the north-west and north-east. Trailing edges contracted from a broad range of southerly directions. Importantly, these results are derived from systematically collected data so confounding observer-effort biases can be discounted. Analyses of climate for the same period show that whilst temperature trends should drive species along a north-north-westerly trajectory, directional responses to precipitation will depend on both the time of year that is important for determining a species' distribution, and the location of the range margin. Directions of species' range centroid shift were not correlated with spatial trends in any single climate variable. We conclude that range shifts of British birds are multidirectional, individualistic and probably determined by species-specific interactions of multiple climate factors. Climate change is predicted to lead to changes in community composition through variation in the rates that species' ranges shift; our results suggest communities could change further owing to constituent species shifting along different trajectories. We recommend more studies consider directionality in climate and range dynamics to produce more appropriate measures of observed and expected responses to climate change.

Climate change is leading to the development of land-based mitigation and adaptation strategies that are likely to have substantial impacts on global biodiversity. Of these, approaches to maintain carbon within existing natural ecosystems could have particularly large benefits for biodiversity. However, the geographical distributions of terrestrial carbon stocks and biodiversity differ. Using conservation planning analyses for the New World and Britain, we conclude that a carbon-only strategy would not be effective at conserving biodiversity, as have previous studies. Nonetheless, we find that a combined carbon-biodiversity strategy could simultaneously protect 90% of carbon stocks (relative to a carbon-only conservation strategy) and > 90% of the biodiversity (relative to a biodiversity-only strategy) in both regions. This combined approach encapsulates the principle of complementarity, whereby locations that contain different sets of species are prioritised, and hence disproportionately safeguard localised species that are not protected effectively by carbon-only strategies. It is efficient because localised species are concentrated into small parts of the terrestrial land surface, whereas carbon is somewhat more evenly distributed; and carbon stocks protected in one location are equivalent to those protected elsewhere. Efficient compromises can only be achieved when biodiversity and carbon are incorporated together within a spatial planning process.

The timing of migration is one of the key life-history parameters of migratory birds. It is expected to be under strong selection, to be sensitive to changing environmental conditions and to have implications for population dynamics. However, most phenological studies do not describe arrival and departure phenologies for a species in a way that is robust to potential biases, or that can be clearly related to breeding populations. This hampers our ability to understand more fully how climate change may affect species' migratory strategies, their life histories and ultimately their population dynamics. Using generalized additive models (GAMs) and extensive large-scale data collected in the UK over a 40-year period, we present standardized measures of migration phenology for common migratory birds, and examine how the phenology of bird migration has changed in the UK since the 1960s. Arrival dates for 11 of 14 common migrants became significantly earlier, with six species advancing their arrival by more than 10 days. These comprised two species, Blackcap Sylvia atricapilla and Chiffchaff Phylloscopus collybita, which winter closest to Britain in southern Europe and the arid northern zone of Africa, Common Redstart Phoenicurus phoenicurus, which winters in the arid zone, and three hirundines (Sand Martin Riparia riparia, House Martin Delichon urbicum and Barn Swallow Hirundo rustica), which winter in different parts of Africa. Concurrently, departure dates became significantly later for four of the 14 species and included species that winter in southern Europe (Blackcap and Chiffchaff) and in humid zones of Africa (Garden Warbler Sylvia borin and Whinchat Saxicola rubetra). Common Swift Apus apus was the exception in departing significantly earlier. The net result of earlier arrival and later departure for most species was that length of stay has become significantly longer for nine of the 14 species. Species that have advanced their timing of arrival showed the most positive trends in abundance, in accordance with previous studies. Related in part to earlier arrival and the relationship above, we also show that species extending their stay in Great Britain have shown the most positive trends. Further applications of our modelling approach will provide opportunities for more robust tests of relationships between phenological change and population dynamics than have been possible previously.