Abstract
The effect of urban development on butterfly species' richness and species' incidence is tested for the Greater Manchester conurbation and two sample areas, mapped at finer scales, within the southern part of the conurbation. The tests include measures of bias for recording effort (number of visits). Species' richness is found to increase with percentage urban cover for Greater Manchester (tetrad scale) and decrease with urban cover for the two sample areas in South West Manchester (1 km scale) and the Mersey Valley (100 m scale). For Greater Manchester, the increase in species' richness with increased urban cover is largely explained by lower species' richness at higher altitude in the Pennines bounding the conurbation. For the two sample areas, decreasing species' richness associated with increasing urban cover corresponds with reductions in the areas of a number of semi-natural habitats, hostplants and nectar sources. Despite these statistically significant correlations, the impact of urban cover on species' richness is weak. The maximum loss rate identified anywhere within the region is 0.81 species per 10% change in urban cover for South West Manchester. This finding may reflect on the generally low species diversity of the region. However, these results could be influenced by recording and sampling artefacts, particularly the failure of mapping programmes to distinguish vagrant individuals from breeding populations and a bias of records to vagrants. This is supported by the higher correlations between species' incidence and nectar sources than between species' incidence and their hostplants. Adult butterflies are opportunistic nectar users and nectar sources are more widely spread and thus less influenced by urban development than are specific butterfly hostplants. The finding may also reflect on the capacity of most of the butterfly species to breed successfully on tiny areas of hostplant existing within extensively built-up areas. Moreover, the capacity of butterfly species to persist by using small fragments of hostplants would be enhanced by vagrancy. If this is indeed the case, it is a finding that would support the value of small patches in butterfly metapopulations, albeit ones comprising incomplete complements of resources required during the life cycle. The incidence of most species decreases with increase in urban cover. Multivariate analyses indicate that this is owing to corresponding declines in hostplant-habitats and nectar sources. Five species increase with urban cover, but none attain formal significance. Associations among hostplants and habitat variables in a principal components analysis suggest that, in three cases (Pieris brassicae, P. rapae, Celastrina argiolus), this is owing to increasing areas of their hostplants within urban environments.
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