Habitat Associations and Community Structure of Birds in Shrubsteppe Environments

We studied the relations between bird distribution and abundance and habitat characteristics at a regional scale of investigation, using surveys conducted over three consecutive years on 14 plots at nine locations in the shrubsteppe of the northwestern Great Basin of North America. Bivariate and multivariate analyses revealed no large suites of bird species that were correlated in their distribution and abundance, and few associations existed between pairs of species, suggesting that bird populations in this system vary largely independently of one another. Both bivariate and multivariate correlational analyses between birds and habitat physiognomy indicated that the bird species that were widely distributed in this shrubsteppe system had few significant associations with habitat features, while species with more localized distributions did exhibit habitat affinities, most notably with a suite of characters associated with the occurrence of rocky outcrops. Bird species whose primary distributions and habitat affinities lie in grassland regions to the east demonstrated the greatest degree of correlation with features of habitat physiognomy in this shrubsteppe region, increasing in abundance as vegetation coverage and stature increased and horizontal heterogeneity of habitats decreased. Some bird species, however, exhibited no correlations with the habitat features we measured, and multivariate analyses comparing variation in the bird abundances with variation in features of habitat physiognomy accounted for <17% of the overall variation in the bird abundance matrix. Consideration of coverages of different species of shrubs, however, generally produced more significant correlations with variations in bird abundances, especially for the widespread shrubsteppe species. Attributes of avian community structure varied among the locations surveyed. Variations in the abundance of Brewer's Sparrows, the most abundant species at most sites, accounted for 86% of the variation in total avian density. Species diversity was negatively correlated with total density (presumably because of the overwhelming influence of one species on density) but positively correlated with species richness. Richness variations, in turn, were a consequence of variations in the abundances of several locally distributed shrubsteppe species or grassland species. Richness decreased with increasing horizontal habitat heterogeneity and general vegetation sparseness, but increased with increasing structural diversity of the habitat. Variations in avian community biomass were largely a function of abundances of the widespread or peripheral species; none of the local shrubsteppe forms that contributed so much to species richness was correlated with total biomass variations. We compare the findings of these regional—scale analyses with those of a continental—scale study that included a habitat spectrum ranging from shrubsteppe through tallgrass prairies. Bird species exhibited different patterns of habitat correlations on the two spatial scales. In particular, the characteristic shrubsteppe species showed strong correlations with features of habitat physiognomy in the continental analysis, but in the regional study such associations were generally lacking and these birds instead were correlated with the coverages of various shrub species. This suggests that at a large scale, between—habitat level of analysis these birds may respond to some elements of general habitat configuration, but their within—habitat responses may be more strongly associated with details of habitat floristics. These results complicate studies of avian community/habitat relationships: complete understanding of ecological patterns apparently requires knowledge of vegetational floristics as well as physiognomy; the response of birds to habitat characteristics, and the habitat features that are important, may differ at different scales of spatial resolution. At both spatial scales, however, a substantial portion of the variation in avian abundance remains unexplained after consideration of habitat features. Relatively few significant correlations thus emerge, increasing the probability that those that are revealed may well be spurious, and reinforcing the view that biotic interactions such as competition probably play a minor role in structuring these communities.