Large vertebrates are strong interactors in food webs, yet they were lost from most ecosystems after the dispersal of modern humans from Africa and Eurasia. We call for restoration of missing ecological functions and evolutionary potential of lost North American megafauna using extant conspecifics and related taxa. We refer to this restoration as Pleistocene rewilding; it is conceived as carefully managed ecosystem manipulations whereby costs and benefits are objectively addressed on a case-by-case and locality-by-locality basis. Pleistocene rewilding would deliberately promote large, long-lived species over pest and weed assemblages, facilitate the persistence and ecological effectiveness of megafauna on a global scale, and broaden the underlying premise of conservation from managing extinction to encompass restoring ecological and evolutionary processes. Pleistocene rewilding can begin immediately with species such as Bolson tortoises and feral horses and continue through the coming decades with elephants and Holarctic lions. Our exemplar taxa would contribute biological, economic, and cultural benefits to North America. Owners of large tracts of private land in the central and western United States could be the first to implement this restoration. Risks of Pleistocene rewilding include the possibility of altered disease ecology and associated human health implications, as well as unexpected ecological and sociopolitical consequences of reintroductions. Establishment of programs to monitor suites of species interactions and their consequences for biodiversity and ecosystem health will be a significant challenge. Secure fencing would be a major economic cost, and social challenges will include acceptance of predation as an overriding natural process and the incorporation of pre-Columbian ecological frameworks into conservation strategies.
Livestock have been excluded from a 3,160-ha range in southeastern Arizona since 1968. Compared to an adjacent continuously grazed area, in 1981-82 a protected upland site supported 45% more grass cover, a comparatively heterogeneous grass community, and 4 times as many shrubs. Grama grasses (Bouteloua spp.) were equally common in and outside the exclosure, while a variety of other species, especially plains lovegrass (Eragrostis intermedia) and Arizona cottontop (Trichachne cal~ornicum) were much more abundant on the protected site. The grazed area supported significantly higher numbers of birds In summer, while densities did not differ in winter. Rodents were significantly more abundant inside the protected area. Species of birds and rodents more common in the grazed area included those typical of more xeric lowland habitats and those preferring open ground for feeding. Species more common on the protected site were those which characterize semidesert or plains grasslands, and which prefer substantial grass or shrub cover. Grazing appeared to favor birds as a class over rodents.
In a 4—yr field experiment, we tested the hypotheses that insectivorous birds (1) controlled densities of herbivorous grasshoppers in an ungrazed semiarid grassland in southeastern Arizona, and (2) functioned as keystone predators, by limiting abundances of grasshoppers that otherwise might change vegetation cover and species composition, and/or by mediating the effects of otherwise competitively superior members of the grasshopper assemblage. We measured grasshopper densities and vegetation on 32 464—m2 grassland plots for 1 yr, then enclosed 16 of these plots with bird exclosures and continued data collection for 3 yr. Eight of the 16 experimental plots were further modified in the last 2 yr of the study by installing fine—mesh 1 m high barriers designed to retard grasshopper dispersal. Microclimates of caged plots differed only slightly from open plots. Lizards and rodents increased inside the exclosures, but they were removed and released elsewhere such that their average abundances did not differ among treatments. By the final year of the study, mean annual adult grasshopper density was >2.2 times higher on plots from which birds were excluded, and where grasshoppers were enclosed by dispersal barriers, than on unmanipulated control plots. Mean nymph density was >3.0 times higher in the same comparison. Grasshoppers were significantly more abundant in bird exclosures with insect dispersal barriers, indicating that experimental plots were dispersal sources rather than sinks. Seven of 12 common grasshopper species were more abundant inside the bird exclosures, while none was less abundant. Among the more abundant taxa, those responding most positively were grass feeders: Eritettix simplex, Opeia obscura, Paropomala wyomingensis, and Phoetaliotes nebrascensis. We found no evidence that grasshoppers competed with one another under increased densities inside the bird exclosures. Although the amount of insect herbivory was somewhat higher inside the bird exclosures, and was positively correlated with grasshopper density across all 32 plots (r = 0.87), overall vegetation cover and species composition did not differ among treatments by the end of the study. Dactylotum variegatum, an aposematic species apparently immune to avian predation, showed no significant responses to the experiment. Birds clearly limited grasshoppers in this grassland ecosystem, but they failed to qualify as keystone predators, at least in the short term, for two reasons: (1) in their absence, increased grasshopper densities had no appreciable impact on vegetation cover or species composition; and (2) there was no evidence that birds mediated competition among grasshoppers.
Species richness and evenness are components of biological diversity that may or may not be correlated with one another and with patterns of species abundance. We compared these attributes among flowering plants, grasshoppers, butterflies, lizards, summer birds, winter birds, and rodents across 48 plots in the grasslands and mesquite-oak savannas of southeastern Arizona. Species richness and evenness were uncorrelated or weakly negatively correlated for each taxonomic group, supporting the conclusion that richness alone is an incomplete measure of diversity. In each case, richness was positively correlated with one or more measures of abundance. By contrast, evenness usually was negatively correlated with the abundance variables, reflecting the fact that plots with high evenness generally were those where all species present were about equally uncommon. Therefore richness, but not evenness, usually was a positive predictor of places of conservation value, if these are defined as places where species of interest are especially abundant. Species diversity was more positively correlated with evenness than with richness among grasshoppers and flowering plants, in contrast to the other taxonomic groups, and the positive correlations between richness and abundance were comparatively weak for grasshoppers and plants as well. Both of these differences can be attributed to the fact that assemblages of plants and grasshoppers were numerically dominated by small subsets of common species (grasses and certain spur-throated grasshoppers) whose abundances differed greatly among plots in ways unrelated to species richness of the groups as a whole.
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