Keith Summerville

Species diversity may be additively partitioned within and among samples (alpha and beta diversity) from hierarchically scaled studies to assess the proportion of the total diversity (gamma) found in different habitats, landscapes, or regions. We developed a statistical approach for testing null hypotheses that observed partitions of species richness or diversity indices differed from those expected by chance, and we illustrate these tests using data from a hierarchical study of forest-canopy beetles. Two null hypotheses were implemented using individual- and sample-based randomization tests to generate null distributions for alpha and beta components of diversity at multiple sampling scales. The two tests differed in their null distributions and power to detect statistically significant diversity components. Individual-based randomization was more powerful at all hierarchical levels and was sensitive to departures between observed and null partitions due to intraspecific aggregation of individuals. Sample-based randomization had less power but still may be useful for determining whether different habitats show a higher degree of differentiation in species diversity compared with random samples from the landscape. Null hypothesis tests provide a basis for inferences on partitions of species richness or diversity indices at multiple sampling levels, thereby increasing our understanding of how alpha and beta diversity change across spatial scales.

AbstractA nested pattern occurs whenever the species observed in depauperate habitat patches are a subset of those found in more species-rich patches. Ecologists have documented many instances of nestedness caused by population-level processes such as colonization and extinction at biogeographic scales. However, few researchers have examined whether nestedness may exist at fine scales due to the ways in which individual organisms discriminate among potential habitat patches. In 1999, we experimentally fragmented an old-field habitat into patches of varying size to test whether nestedness could exist on a fine spatial scale. Five treatments of differing patch size were replicated five times in a Latin square design by selectively mowing 15×15 m2 plots within an old-field (patch areas: 225, 180, 135, 90, and 45 m2). Specifically, we tested whether butterflies foraging within a network of patches differing in area conformed to a nested subset structure. We also classified species according to (1) their flight height while foraging (high or low), and (2) their adult habitat breadth (ubiquitous, general, or restricted) to determine whether nestedness could be explained by difference in species’ tendency to discriminate among patches differing in area. We found significant evidence that a community of foraging Lepidoptera conformed to a nested subset structure based on the difference between the observed nestedness within the butterfly community and the nestedness obtained from randomly generated species presence/absence matrices. Poisson regression analyses demonstrated that high-flying, habitat-restricted species avoided the smallest patches (90 and 45 m2) in favor of larger remnants, whereas low-flying, habitat generalists used all patch sizes. Thus, our study is one of the first to demonstrate that nestedness among species subsets can be observed at fine spatial scales (within a single 1.5 hectare field) and may be maintained by species behavioral differences: discriminating species (i.e. high-flying, habitat restricted) avoided the smallest patches, and less discriminating species (i.e. low-flying, ubiquitous) were distributed throughout the field without regard to patch size. Our results also suggest that nestedness should be viewed as yet another scalar pattern in ecology, generated by variation in patch use by individuals at fine-scales as well as the more traditionally invoked processes of extinction and colonization of species at broad-scales.

Species diversity may be additively partitioned within and among samples (alpha and beta diversity) from hierarchically scaled studies to assess the proportion of the total diversity (gamma) found in different habitats, landscapes, or regions. We developed a statistical approach for testing null hypotheses that observed partitions of species richness or diversity indices differed from those expected by chance, and we illustrate these tests using data from a hierarchical study of forest-canopy beetles. Two null hypotheses were implemented using individual- and sample-based randomization tests to generate null distributions for alpha and beta components of diversity at multiple sampling scales. The two tests differed in their null distributions and power to detect statistically significant diversity components. Individual-based randomization was more powerful at all hierarchical levels and was sensitive to departures between observed and null partitions due to intraspecific aggregation of individuals. Sample-based randomization had less power but still may be useful for determining whether different habitats show a higher degree of differentiation in species diversity compared with random samples from the landscape. Null hypothesis tests provide a basis for inferences on partitions of species richness or diversity indices at multiple sampling levels, thereby increasing our understanding of how alpha and beta diversity change across spatial scales.

The response of forest insect communities to disturbances such as timber harvest will likely depend on the underlying spatial structure of species assemblages before the disturbance occurs. Unfortunately, many studies of forest management implicitly assume homogeneity of community structure before harvest; postlogging communities are inferred to be a direct product of the imposed management. The goal of this study is to examine variation in the community structure of forest Lepidoptera using the pretimber harvest data on Lepidoptera from 20 forest sites within three watersheds at Morgan Monroe State Forest, IN. A total of 14,537 individuals representing 324 species of Lepidoptera were sampled from Morgan-Monroe State Forest in 2007. Sampling efficiency was not a function of management unit, and, surprisingly, we found little evidence that management units differed in overall community composition. Diversity partitioning suggested that > 99% of Simpson diversity (species dominance) was determined at the local scale, and each site contained the same 10 dominant taxa in rank order. Variation in species richness seemed to be more a problem of sampling bias than underlying differences in habitat preference by moth feeding guilds. Finally, Mantel tests suggested that forest moth communities at Morgan-Monroe are not spatially autocorrelated. The results here are encouraging because they strongly suggest that shifts in lepidopteran community structure should reflect the community response to disturbance rather than inherent spatial heterogeneity of species composition.

... Keith S. Summerville 1,* ,; Thomas O. Crist 2 ,; Jonathan K. Kahn 2,3 ,; Jon C. Gering 4. Article first published online: 12 DEC 2003. ... 3. Species richness and abundance of caterpillars were higher on oaks and maples than on American beech. ...

Restoration ecologists are increasingly turning to the development of trait-filter models, which predict how evolved traits limit species membership within assemblages depending on existing abiotic or biotic constraints, as a tool to explain how species move from a regional species pool into a restored community. Two often untested assumptions of these models, however, are that species traits can reliably predict species' broadscale distribution and that the effects of traits on community membership do not vary between restored and remnant habitats. The goals of this study were to determine whether combinations of ecological traits predispose moth species toward recolonization of restored prairies and to assess the degree to which restored prairies contain moth assemblages comparable with prairie remnants. In 2004, we collected 259 moth species from 13 tallgrass prairie remnants and restorations in central Iowa. Principal components analysis (PCA) was used to identify significant combinations of ecological traits that were shared by groups of moth species. Logistic regression was then employed to test for significant effects of the trait combinations on the frequency of prairie sites occupied by moth species. PCA partitioned moth traits into four axes that explained a total of 81.6% of the variance. Logistic regression detected significant effects for all four PCA axes on the fraction of sites occupied by moths. Species frequently filtered from the regional species pool into prairies were those that had long flight periods and were multivoltine, displayed a feeding preference for legumes but not other forb families, and were regionally abundant but relatively small in body size. Ordination revealed significant differences in moth communities among prairies, suggesting that species traits and habitat characteristics likely interact to create observed patterns of species recolonization of restorations. Thus, the optimal approach to restoring the lepidopteran fauna of tallgrass prairies may involve locating prairie plantings adjacent to habitat remnants.

... Temperate forests, however, are rich in moth species ( [Summerville et al., 2001] and [Summerville & Crist, 2002] ), and several studies suggest that certain moth assemblages show promise as indicators of overall lepidopteran biodiversity and forest composition ( [Holloway ...