Habitat quality and metapopulation e¡ects are the main hypotheses that currently explain the disproportionate decline of insects in cultivated Holarctic landscapes. The former assumes a degradation in habitat quality for insects within surviving ecosystems, the latter that too few, small or isolated islands of ecosystem remain in landscapes for populations to persist. These hypotheses are often treated as alternatives, and this can lead to serious con£ict in the interpretations of conservationists. We present the ¢rst empirical demonstration that habitat quality and site isolation are both important determinants of where populations persist in modern landscapes. We described the precise habitat requirements of Melitaea cinxia, Polyommatus bellargus and Thymelicus acteon, and quanti¢ed the variation in carrying capacity within each butter£y's niche. We then made detailed surveys to compare the distribution and density of every population of each species with the size, distance apart and quality of their speci¢c habitats in all their potential habitat patches in three UK landscapes. In each case, within-site variation in habitat quality explained which patches supported a species' population two to three times better than site isolation. Site area and occupancy were not correlated in any species. Instead of representing alternative paradigms, habitat quality and spatial e¡ects operate at di¡erent hierarchical levels within the same process: habitat quality is the missing third parameter in metapopulation dynamics, contributing more to species persistence, on the basis of these results, than site area or isolation. A reorientation in conservation priorities is recommended.

Summary 1.Many ecological studies and applications involve measuring the height of grassland swards. An evaluation was made of the practicality, accuracy and comparability of using the sward stick, drop disc and direct methods of measurement. Each method proved to be appropriate for measuring swards that contain a wide range of heights, each was quick to perform, and recorder effects were negligible. Yet each had strengths and weaknesses. 2. The sward stick gave the most variable results: it was considered the best method for recording the architecture of the sward surface, and hence invertebrate niches, but was poor for measuring short turf. The drop disc was the worst method for recording microheterogeneity in sward architecture and was completely unsuitable for measuring variation in short turf. But in medium-tall swards, it was considered to be the best method for measuring productivity, vertebrate herbivory and for large-scale monitoring of land managed for conservation and under agri-environment schemes. The direct method gave the most consistent and accurate results compared with an independent parameter, soil temperature. It was the only method suitable for measuring variation in short turf. 3. A serious problem exists when research results, recommendations and assays involve measurements made by more than one method. The sward stick consistently gave higher absolute values than either of the other methods and, apart from in short turf (for which it is unsuitable), the drop disc gave values that were 73% and up to 40% lower, respectively, than those obtained using the sward stick and direct methods. This can lead to major misapplications of ecological results and recommendations in conservation and agri-environmental projects.

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