Papers by Keith Summerville
The American Naturalist, 2003
Species diversity may be additively partitioned within and among samples (alpha and beta diversit... more 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.
Environmental Entomology, Nov 1, 2008
The response of forest insect communities to disturbances such as timber harvest will likely depe... more 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.
Oikos, 2002
AbstractA nested pattern occurs whenever the species observed in depauperate habitat patches are ... more 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.
Agricultural and Forest Entomology, 2015
Journal of The Society for Gynecologic Investigation, 1998
The Wilson Journal of Ornithology, 2009
The American Naturalist, 2003
Species diversity may be additively partitioned within and among samples (alpha and beta diversit... more 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.
Restoration Ecology, 2005
Ecological restoration is deemed important for the longterm conservation of biodiversity, but eco... more Ecological restoration is deemed important for the longterm conservation of biodiversity, but ecologists still lack an understanding of how habitat availability and habitat quality in a restored system interact to determine species diversity. This problem seems particularly apparent in Tallgrass Prairie and savanna ecoregions, where restored management units represent the majority of extant habitat. In this study, we tested three principal hypotheses, each stating that the diversity of Lepidoptera would be greater in (1) patches of savanna habitat that were larger; (2) patches that were of higher habitat quality; and (3) patches that had greater connectivity to management units of similar physiognomy. Lepidoptera were sampled in 2003 from 13 unmanaged woodland remnants within Neal Smith National Wildlife Refuge, a 2,292-ha prairie and savanna reconstruction project. We also measured 11 environmental variables within each site to assess variation in habitat quantity and quality. Principal components analysis (PCA) was used to identify major gradients of environmental variation among the 13 sites. Our PCA differentiated among woodlands along three environmental gradients, defined by (1) stand size, shape, topography, and oak dominance; (2) degree of disturbance; and (3) isolation. Total lepidopteran species richness, however, was only predicted by variation in the first principal component. Species richness of Lepidoptera known to be oak specialists was significantly affected by variation along all three PCA gradients. Surprisingly, more isolated woodland remnants contained a greater richness of oak feeders. Our results suggest that approaches to restoring oak savannas should emphasize aspects of both habitat quantity and quality. Beyond making individual management units larger, priority sites for restoration should possess a low importance of trees that are indicative of past habitat disturbance (e.g., Honey locust, White mulberry) even if canopy closure is substantial. Connectivity among restored habitats may benefit savanna moth communities only when habitat linkages contain a flora similar in composition to focal patches.
Restoration Ecology, 2007
Understanding the degree to which species assemblages naturally vary over time will be critically... more Understanding the degree to which species assemblages naturally vary over time will be critically important when assessing whether direct management effects or contingency is responsible for species gain or loss. In this study, we tested three predictions related to short-term variation in prairie moth communities: (1) communities would only exhibit significant temporal variation in newly restored sites (1-3 years old); (2) prairie size and age would positively influence community reassembly, with larger, older restorations sampling a greater proportion of the regional species pool; and (3) older restorations (7-10 years old) would have yet to converge on the community composition of prairie remnants. Moths were sampled from 13 Tallgrass prairie restorations and remnants in central Iowa in [2004][2005]. Repeated measures analysis of variance revealed significant effects of sampling year on moth species richness and abundance as well as on the richness of two functional groups, but difference among prairie types was only observed in 2005. Rarefaction analysis revealed that older restorations and prairie remnants supported higher species richness compared to recently planted sites, and nonmetric, multidimensional scaling ordination indicated that restorations older than 7 years were clearly converging on the species composition of remnants. These results suggest that moth communities in restorations and remnants are highly variable in time but that as restorations age, they appear to reaccumulate moth species found in prairie remnants. The long-term persistence of a particular species assemblage within a given site, however, might be a difficult endpoint to attain in central Iowa prairies because of significant annual variation in species occurrence.
Oikos, 2002
Minireviews provides an opportunity to summarize existing knowledge of selected ecological areas,... more Minireviews provides an opportunity to summarize existing knowledge of selected ecological areas, with special emphasis on current topics where rapid and significant advances are occurring. Reviews should be concise and not too wide-ranging. All key references should be cited. A summary is required. . 2002. The additive partitioning of species diversity: recent revival of an old idea. Oikos 99: 3 -9.
Oikos, 2002
2002. Does variation in patch use among butterfly species contribute to nestedness at fine spatia... more 2002. Does variation in patch use among butterfly species contribute to nestedness at fine spatial scales A 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 m 2 plots within an old-field (patch areas: 225, 180, 135, 90, and 45 m 2 ). 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 m 2 ) 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.
Natural Areas Journal, 2009
Insect Conservation and Diversity, 2013
Forest Ecology and Management, 2009
Environmental Entomology, 2008
Community assembly and the factors that inßuence it have long been a topic of interest to ecologi... more Community assembly and the factors that inßuence it have long been a topic of interest to ecologists, but theory has yet to produce unequivocal evidence that communities assemble in predictable ways. The goal of this study was to document the relationship between ant communities and environmental variation between four habitat types. To accomplish this, ant communities and 16 environmental variables were sampled across four different habitat types in the Black Belt Prairie and Flatwoods regions in Mississippi. Furthermore, ant species were placed into functional groups for an analysis of the relationship between the assembly of ecological communities and variation in ecosystem function. A total of 20,916 ants representing 68 species was collected across the four habitat types. Nonmetric multidimensional scaling and analysis of covariance analyses both revealed three distinct ant communities, which can be characterized by habitat type: pasture, prairie, and woodland. Principle components analysis (PCA) simpliÞed the 16 environmental variables into four principle components that explained 78% of the variation among sites. Results of multiple regression using the four PCA axes as predictor variables suggest that regional variation in soil structure, land cover type, and the presence of grazing have had major inßuences on ant community composition. Variation in ßora and habitat architecture had smaller but signiÞcant effects on ant species diversity and functional group composition. Our results imply that restoration of native ant communities in disturbed habitats must consider how current disturbance regimens likely interact with the presence of Solenopsis to lower ant biodiversity.
Environmental Entomology, 2008
The response of forest insect communities to disturbances such as timber harvest will likely depe... more 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.
Ecology Letters, 2000
There is a lack of quantitative syntheses of fragmentation effects across species and biogeograph... more There is a lack of quantitative syntheses of fragmentation effects across species and biogeographic regions, especially with respect to species life-history traits. We used data from 24 independent studies of butterflies and moths from a wide range of habitats and landscapes in Europe and North America to test whether traits associated with dispersal capacity, niche breadth and reproductive rate modify the effect of habitat fragmentation on species richness. Overall, species richness increased with habitat patch area and connectivity. Life-history traits improved the explanatory power of the statistical models considerably and modified the butterfly species-area relationship. Species with low mobility, a narrow feeding niche and low reproduction were most strongly affected by habitat loss. This demonstrates the importance of considering life-history traits in fragmentation studies and implies that both species richness and composition change in a predictable manner with habitat loss and fragmentation.
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Papers by Keith Summerville