in all months, and mean precipitation increased in most months (Fig. 2a). 68Spatial variability in climatic change (Fig. 2b,c), necessitates local matching of phenological 69 and climatic datasets rather than the use of regionally-averaged climate data (e.g. Central 70England Temperatures) or large-scale climatic indicators (e.g. North Atlantic Oscillation). 71We did not make the restrictive assumption that biological events would be related to annual CSP precip varied less among trophic levels than the upper limit (Fig. 3d,f) consumers were less than those for primary consumers (Fig. 5a). This occurred because, 195averaged across species, the opposing climate responses of primary producers and secondary 196consumers are more similar in magnitude than are those for primary consumers (Fig. 3), 197 effectively "cancelling each other out". Our models suggest greater average advances for 198 crustacea, fish and insects than for other groups, such as freshwater phytoplankton, birds and 199 mammals (Fig. 5b). However, response-variation is high for crustacea (Fig. 5b). not estimated for marine plankton data (see above), and so the second-phase LME models 441 were run twice: once to examine correlations with temperature and precipitation for all but 442 the marine plankton phenological series (9,800 series), and once to examine only correlations 443 with temperature for the whole data set (10,003 series).

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Background: In the Neolithic, domestic sheep migrated into Europe and subsequently spread in westerly and northwesterly directions. Reconstruction of these migrations and subsequent genetic events requires a more detailed characterization of the current phylogeographic differentiation. Results: We collected 50 K single nucleotide polymorphism (SNP) profiles of Balkan sheep that are currently found near the major Neolithic point of entry into Europe, and combined these data with published genotypes from southwest-Asian, Mediterranean, central-European and north-European sheep and from Asian and European mouflons. We detected clines, ancestral components and admixture by using variants of common analysis tools: geography-informative supervised principal component analysis (PCA), breed-specific admixture analysis, across-breed f 4 profiles and phylogenetic analysis of regional pools of breeds. The regional Balkan sheep populations exhibit considerable genetic overlap, but are clearly distinct from the breeds in surrounding regions. The Asian mouflon did not influence the differentiation of the European domestic sheep and is only distantly related to present-day sheep, including those from Iran where the mouflons were sampled. We demonstrate the occurrence, from southeast to northwest Europe, of a continuously increasing ancestral component of up to 20% contributed by the European mouflon, which is assumed to descend from the original Neolithic domesticates. The overall patterns indicate that the Balkan region and Italy served as post-domestication migration hubs, from which wool sheep reached Spain and north Italy with subsequent migrations northwards. The documented dispersal of Tarentine wool sheep during the Roman period may have been part of this process. Our results also reproduce the documented 18th century admixture of Spanish Merino sheep into several central-European breeds.

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