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In 1949-1951, ecologist Robert H. Whittaker sampled plant community composition at 470 sites in the Siskiyou Mountains (Oregon and California; also known as Klamath or Klamath-Siskiyou Mountains). His primary goal was to develop methods to quantify plant community variation across environmental gradients, following on his seminal work challenging communities as discrete entities. He selected the Siskiyous because of their diverse and endemic-rich flora, which he attributed to geological complexity and an ancient stable climate. He chose sites to span gradients of topography, elevation, geologic substrate, and distance from the coast. He used the frequencies of indicator species in his data to assign sampling locations to positions on the topographic gradient, nested within the elevational and substrate gradients. He originated in this study the concept of diversity partitioning, in which gamma diversity (species richness of a community) equals alpha diversity (species richness in homogeneous sites) times beta diversity (species turnover among sites along gradients). Diversity partitioning subsequently became highly influential and new developments on it continue. Whittaker published his Siskiyou work covering paleohistory, biogeography, floristics, vegetation, gradient analysis, and diversity partitioning in Ecological Monographs in 1960. Discussed in 2 pages of his 60-page monograph, diversity partitioning accounts for95% of its current4300 citations. In 2006, we retrieved Whittaker's Siskiyou data in hard copy from the Cornell University archives and entered them in a database. We used these data for multiple published analyses, including some based on (re)sampling the approximate locations of a subset of his sites. Because of the continued interest in diversity partitioning and in historic data sets, here we present his data, including 359 sampling locations and their descriptors and, for each sample, a list of species with their estimated percent cover (herbs and shrubs) and numbers by diameter at breast height (DBH) category (trees). Site descriptors include the approximate location (road, trail, or stream), elevation, topographic aspect, geologic substrate (serpentine, gabbro, or diorite), and dominant woody vegetation of each location. For 111 sites, including the small number chosen to represent the distance-to-coast gradient, we could not locate his data. There are no copyright restrictions and users of these data should cite this data paper in any publications that result from its use. The authors are available for consultations about and collaborations involving the data.

Understanding how biotic communities respond to landscape spatial structure is critically important for conservation management as natural habitats become increasingly fragmented. However, empirical studies of the effects of spatial structure on plant species richness have found inconsistent results, suggesting that more comprehensive approaches are needed. We asked how landscape structure affects total plant species richness and the richness of a guild of specialized plants in a multivariate context. We sampled herbaceous plant communities at 56 dolomite glades (insular, fire-adapted grasslands) across the Missouri Ozarks, USA, and used structural equation modeling (SEM) to analyze the relative importance of landscape structure, soil resource availability, and fire history for plant communities. We found that landscape spatial structure, defined as the area-weighted proximity of glade habitat surrounding study sites (proximity index), had a significant effect on total plant species richness, but only after we controlled for environmental covariates. Richness of specialist species, but not generalists, was positively related to landscape spatial structure. Our results highlight that local environmental filters must be considered to understand the influence of landscape structure on communities and that unique species guilds may respond differently to landscape structure than the community as a whole. These findings suggest that both local environment and landscape context should be considered when developing management strategies for species of conservation concern in fragmented habitats.

Despite broad recognition that habitat loss represents the greatest threat to the world's biodiyersity, a mechanistic understanding of how habitat loss and associated fragmentation affect ecological systems has proven remarkably challenging. The challenge stems from the multiple interdependent ways that landscapes change following fragmentation and the ensuing complex impacts on populations and communities of interacting species. We confronted these challenges by evaluating how fragmentation affects individual plants through interactions with animals, across five herbaceous species native to longleaf pine savannas. We created a replicated landscape experiment that provides controlled tests of three major fragmentation effects (patch isolation, patch shape [i.e., edge-to-area ratio], and distance to edge), established experimental founder populations of the five species to control for spatial distributions and densities of individual plants, and employed structural equation modeling to evaluate the effects of fragmentation on plant reproductive output and the degree to which these impacts are mediated through altered herbivory, pollination, or pre-dispersal seed predation. Across species, the most consistent response to fragmentation was a reduction in herbivory. Herbivory, however, had little impact.on plant reproductive output, and thus we found little evidence for any resulting benefit to plants in fragments. In contrast, fragmentation rarely impacted pollination or pre-dispersal seed predation, but both of these interactions had strong and consistent impacts on plant reproductive output. As a result, our models robustly predicted plant reproductive output (r2 = 0.52-0.70), yet due to the weak effects of fragmentation on pollination and pre-dispersal seed predation, coupled with the weak effect of herbivory on plant reproduction, the effects of fragmentation on reproductive output were generally small in magnitude and inconsistent. This work provides mechanistic insight into landscape-scale variation in plant reproductive success, the relative importance of plant-animal interactions for structuring these dynamics, and the nuanced nature of how habitat fragmentation can affect populations and communities of interacting species.

Habitat fragmentation can create significant impediments to dispersal. A technique to increase dispersal between otherwise isolated fragments is the use of corridors. Although previous studies have compared dispersal between connected fragments to dispersal between unconnected fragments, it remains unknown how dispersal between fragments connected by a corridor compares to dispersal in unfragmented landscapes. To assess the extent to which corridors can restore dispersal in fragmented landscapes to levels observed in unfragmented landscapes, we employed a stable-isotope marking technique to track seeds within four unfragmented landscapes and eight experimental landscapes with fragments connected by corridors. We studied two wind- and two bird-dispersed plant species, because previous community-based research showed that dispersal mode explains how connectivity effects vary among species. We constructed dispersal kernels for these species in unfragmented landscapes and connected fragments by marking seeds in the center of each landscape with 'IN and then recovering marked seeds in seed traps at distances up to 200 m. For the two wind-dispersed plants, seed dispersal kernels were similar in unfragmented landscapes and connected fragments. In contrast, dispersal kernels of bird-dispersed seeds were both affected by fragmentation and differed in the direction of the impact: Morella cerifera experienced more and Rhus copallina experienced less long-distance dispersal in unfragmented than in connected landscapes. These results show that corridors can facilitate dispersal probabilities comparable to those observed in unfragmented landscapes. Although dispersal mode may provide useful broad predictions, we acknowledge that similar species may respond uniquely due to factors such as seasonality and disperser behavior. Our results further indicate that prior work has likely underestimated dispersal distances of wind-dispersed plants and that factors altering long-distance dispersal may have a greater impact on the spread of species than previously thought.

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.

Experiments in ecology can have unintended side effects. Recently, it has been suggested that the act of visiting a plant, inherent to studying herbivory, may alter plant performance and interactions. To evaluate the generality of this inference, we examined plant performance and herbivory on 14 plant species in three geographic regions. Visitation did not significantly affect any of the variables that we measured, including leaf damage, height, biomass, or survivorship, for any species. However, rates of herbivory varied significantly among sites and regions. Thus, our data do not support the generality of visitation impacting estimates of herbivory. We propose that future studies of herbivory will gain more by evaluating spatial heterogeneity in interaction outcomes than by quantifying possible experimenter-caused variation.

Landscape corridors are commonly used to mitigate negative effects of habitat fragmentation, but concerns persist that they may facilitate the spread of invasive species. In a replicated landscape experiment of open habitat, we measured effects of corridors on the invasive fire ant, Solenopsis invicta, and native ants. Fire ants have two social forms: polygyne, which tend to disperse poorly but establish at high densities, and monogyne, which disperse widely but establish at lower densities. In landscapes dominated by polygyne fire ants, fire ant abundance was higher and native ant diversity was lower in habitat patches connected by corridors than in unconnected patches. Conversely, in landscapes dominated by monogyne fire ants, connectivity had no influence on fire ant abundance and native ant diversity. Polygyne fire ants dominated recently created landscapes, suggesting that these corridor effects may be transient. Our results suggest that corridors can facilitate invasion and they highlight the importance of considering species' traits when assessing corridor utility.

In his classic study in the Siskiyou Mountains (Oregon, USA), one of the most botanically rich forested regions in North America, R. H. Whittaker (1960) foreshadowed many modern ideas on the multivariate control of local species richness along environmental gradients related to productivity. Using a structural equation model to analyze his data, which were never previously statistically analyzed, we demonstrate that Whittaker was remarkably accurate in concluding that local herb richness in these late-seral forests is explained to a large extent by three major abiotic gradients (soils, topography, and elevation), and in turn, by the effects of these gradients on tree densities and the numbers of individual herbs. However, while Whittaker also clearly appreciated the significance of large-scale evolutionary and biogeographic influences on community composition, he did not fully articulate the more recent concept that variation in the species richness of local communities could be explained in part by variation in the sizes of regional species pools. Our model of his data is among the first to use estimates of regional species pool size to explain variation in local community richness along productivity-related gradients. We find that regional pool size, combined with a modest number of other interacting abiotic and biotic factors, explains most of the variation in local herb richness in the Siskiyou biodiversity hotspot.

Species with relatively narrow niches, such as plants restricted (endemic) to particular soils, may be especially vulnerable to extinction under a changing climate due to the enhanced difficulty they face in migrating to suitable new sites. To test for community-level effects of climate change, and to compare such effects in a highly endemic-rich flora on unproductive serpentine soils vs. the flora of normal (diorite) soils, in 2007 we resampled as closely as possible 108 sites originally studied by ecologist Robert H. Whittaker from 1949 to 1951 in the Siskiyou Mountains of southern Oregon, USA. We found sharp declines in herb cover and richness on both serpentine and diorite soils. Declines were strongest in species of northern biogeographic affinity, species endemic to the region (in serpentine communities only), and species endemic to serpentine soils. Consistent with climatic warming, herb communities have shifted from 1949-1951 to 2007 to more closely resemble communities found on xeric (warm, dry) south-facing slopes. The changes found in the Siskiyou herb flora suggest that biotas rich in narrowly distributed endemics may be particularly susceptible to the effects of a warming climate.