Space Use and Habitat Selection by Bobcats in the Fragmented Landscape of South-Central Iowa

Abstract: Historically, bobcats (Lynx rufus) were found throughout the Corn Belt region, but they nearly disappeared from this area due to habitat loss and unregulated harvest that occurred during the century after European settlement. Reports of bobcat occurrences have been increasing in Iowa, USA, and biologists would like to understand the mechanisms enabling bobcats to recolonize this fragmented agricultural landscape. We determined space use and habitat selection of bobcats by radiocollaring 68 bobcats in south-central Iowa during 2003–2006. We triangulated 12,966 locations and recovered an additional 1,399 3-dimensional locations from Global Positioning System collars. We used a fixed kernel estimator to calculate 95% utilization distributions (UDs) for home ranges and 50% UDs for cores. Annual home range area of males (x̄ = 58.6 km², 95% CI = 49.2–69.9) was nearly 3 times that of females (x̄=19.9 km², 95% CI = 17.0–23.3). Females used smaller home ranges during April-September when they were suspected to have kittens with them (x̄ = 16.8 km², 95% CI = 13.7–20.7), as compared to October-March (x̄ = 24.1 km², 95% CI = 19.0–30.7), whereas home ranges of males did not differ between seasons. Similarly, core area of males (x̄ = 7.7 km², 95% CI = 6.2–9.6) was larger than that of females (x̄ = 2.3 km², 95% CI = 1.9–2.7). Females used significantly smaller cores in April-September (x̄ = 1.8 km², 95% CI = 1.4–2.3) as compared to October-March (x̄ = 2.8 km², 95% CI = 2.2–3.7), whereas males did not. For both sexes, compositional analysis indicated that forest habitat was ranked higher than all other habitat classes at both the landscape and local scale. Standardized habitat selection ratios illustrate that female and male bobcats selected forest habitat about twice as frequently as any other habitat class, including grassland and Conservation Reserve Program land. Predictive models indicated that home range and core area was smaller in landscapes where perennial forest and grassland habitats were less fragmented. Predictive models indicated home ranges were more irregular in shape in landscapes where row crop patches were less aggregated within home ranges. Our results have practical implications for wildlife managers regarding expected bobcat habitat use and distribution as the species becomes more abundant in the agricultural landscape of the Midwest.     

Habitat Selection and Movement of Pond-Breeding Amphibians in Experimentally Fragmented Pine Forests

ABSTRACT Population-level responses of amphibians to forest management regimes are partly dictated by individual behavioral responses to habitat alteration. We examined the short-term (i.e., 24-hr) habitat choices and movement patterns of 3 amphibian species—southern leopard frogs (Rana sphenocephala), marbled salamanders (Ambystoma opacum), and southern toads (Bufo terrestris)—released on edges between forest habitats and recent clear-cuts in the Upper Coastal Plain of South Carolina, USA. We predicted that adult frogs and salamanders would preferentially select forest using environmental cues as indicators of habitat suitability. We also predicted that movement patterns would differ in clear-cuts relative to forests, resulting in lower habitat permeability of clear-cuts for some or all of the species. Using fluorescent powder tracking, we determined that marbled salamanders selected habitat at random, southern toads preferred clear-cuts, and southern leopard frogs initially selected clear-cuts but ultimately preferred forests. Frogs exhibited long-distance, directional movement with few turns. In contrast, toads exhibited wandering behavior and salamanders moved relatively short distances before locating cover. Southern toads and southern leopard frogs moved farther in forests, and all 3 species made more turns in clear-cuts than in forests. Habitat selection by southern toads did not vary according to body size, sex, or the environmental cues we measured. However, marbled salamanders were more likely to enter clear-cuts when soil moisture was high, and southern leopard frogs were more likely to enter clear-cuts when relative humidity and air temperature were higher in the clear-cut than in adjacent forest. Although we found evidence of reduced habitat permeability of clear-cuts for southern leopard frogs and southern toads, none of the species exhibited strong behavioral avoidance of the small (4-ha) clear-cuts in our study. Further studies of long-term habitat use and the potential physiological and other costs to individuals in altered forests are needed to understand the effects of forest management on population persistence. To reduce potentially detrimental effects of clear-cutting on amphibians in the Southeast, wildlife managers should consider the vagility and behavior of species of concern, especially in relation to the size of planned harvests adjacent to breeding sites.     

The Interplay of Habitat and Seed Type on Scatterhoarding Behavior in a Fragmented Afromontane Forest Landscape

Scatterhoarding by rodents, whereby seeds are collected and stored for later consumption, can result in seed dispersal. Seeds may be covered in litter on the forest floor (cached) or buried. This is particularly so in the Neotropics for large, nutritious seeds, and where primary dispersers are rare or missing. In African forests, contemporary anthropogenic pressures such as hunting, forest degradation, and fragmentation are contributing toward major declines in large frugivores, yet the potential for scatterhoarding to mitigate this loss is largely unknown. In this study, we used thread‐marked seed to explore the balance between seed predation and dispersal by rodents in Afromontane forest. We studied two tree species in three habitats: (1) continuous forest; (2) continuous forest edge, and (3) small, degraded riparian forest patches. We found that seed removal rates were high and almost the same in all three habitats for both tree species, but that the predation/dispersal balance differed among habitats. In continuous forest, more seeds of each species were scatterhoarded than depredated, and rates of scatterhoarding differed between the two species. In all habitats, burying seeds up to 2 cm belowground was more common than caching. Distances seeds were moved was approximately five times greater in continuous forest than in forest edge or riparian patches. We found strong evidence to suggest that the African pouched rat, Cricetomys sp. nov was responsible for the scatterhoarding.     

Habitat Selection by the Spur-thighed Tortoise Testudo graeca in a Multisuccessional Landscape: Implications for Habitat Management

As a result of human activities, natural Mediterranean landscapes (including agro-ecosystems) are characterised by a mosaic-like structure with habitat-patches at different successional stages. These systems have high biodiversity levels and are home to a large number of species protected by European laws whose habitats should be adequately managed. In the present work, we study habitat use from an applied point of view in the spur-thighed tortoise Testudo graeca, an endangered reptile present in semi-arid Mediterranean agro-ecosystems. Results show that, at a landscape scale, the species selects simplified vegetation structures and includes in its home range re-colonisation shrubland and small non-irrigated fields. Within the home range, habitat selection patterns vary and areas with higher vegetation cover and complexity are selected. Detected patterns are discussed in terms of the ecological requirements of the species and with a hierarchical view of resources and conditions. The implications of our findings for habitat management aimed at the conservation of the species are also discussed.     

Pond-Breeding Amphibian Species Richness and Habitat Selection in a Beaver-Modified Landscape

ABSTRACT Beaver (Castor canadensis) activity creates wetland habitats with varying hydroperiods important in maintaining habitat diversity for pond-breeding amphibians with significantly different breeding habitat requirements. We documented pond-breeding amphibian assemblages in 71 freshwater wetlands in Acadia National Park, Maine, USA. Using 15 variables describing local pond conditions and wetland landscape characteristics, we developed a priori models to predict sites with high amphibian species richness and used model selection with Akaike's Information Criterion to judge the strength of evidence supporting each model. We developed single-species models to predict wood frog (Rana sylvatica), bullfrog (R. catesbeiana), and pickerel frog (R. palustris) breeding site selection. Sites with high species richness were best predicted by 1) connectivity of wetlands in the landscape through stream corridors and 2) wetland modification by beaver. Wood frog breeding habitat was best predicted by temporary hydroperiod, lack of fish, and absence of current beaver activity. Wood frog breeding was present in abandoned beaver wetlands nearly as often as in nonbeaver wetlands. Bullfrog breeding was limited to active beaver wetlands with fish and permanent water. Pickerel frog breeding sites were best predicted by connectivity through stream corridors within the landscape. As beavers have recolonized areas of their former range in North America, they have increased the number and diversity of available breeding sites in the landscape for pond-breeding amphibians. The resulting mosaic of active and abandoned beaver wetlands both supports rich amphibian assemblages and provides suitable breeding habitat for species with differing habitat requirements. Land managers should consider the potential benefits of minimal management of beavers in promoting and conserving amphibian and wetland diversity at a landscape scale.     

Habitat Selection of Female Turkeys in a Managed Pine Landscape in Mississippi

Abstract: Intensive pine (Pinus spp.) management is a dominant form of forest management in the southeastern United States. Previous research has shown that managed pine forests provide suitable habitat for eastern wild turkeys (Meleagris gallopavo silvestris), but little research has examined seasonal habitat selection for female wild turkeys from a landscape perspective, particularly within managed pine landscapes. Therefore, we used a long-term (1986-1993) data set to describe seasonal habitat selection by female wild turkeys, using an information-theoretic approach from a landscape perspective, on an intensively managed pine landscape. Habitat use patterns during preincubation and autumn-winter were indicative of female wild turkeys moving between a bottomland hardwood-agricultural field complex during autumn-winter and upland managed pine stands during the remainder of the year. During spring and summer, female wild turkeys used landscapes primarily composed of intensively managed pine, including thinned and burned stands and roadsides. Our results confirm results of short-term, stand-based habitat analyses on our study area. We recommend variable fire return intervals of 3 to 7 years to improve habitat conditions for wild turkeys within intensively managed pine forests. Further research is needed to examine management actions, such as thinning, prescribed fire, and herbicide use, within the context of wild turkey use of intensively managed pine landscapes.     

Habitat Loss Reduces the Diversity of Frog Reproductive Modes in an Atlantic Forest Fragmented Landscape

Several amphibian species have ecological traits that can make them vulnerable to landscape changes, such as habitat preference and reproductive strategies. We evaluated how anuran species and their respective reproductive modes were distributed in an Atlantic Forest fragmented landscape, Southeastern Brazil. We sampled through visual encounter surveys three continuous forest sites, 12 forest fragments and five pasture areas (matrix) between July 2007 and March 2010. We recorded 50 anuran species with 15 reproductive modes, a third of them (33.3%) recorded only in continuous forest sites (modes 3, 8, 19, 25 and 36). These reproductive modes found only in continuous forest sites are typically forest‐associated, and seem to be more vulnerable to habitat loss. Additionally, we found a trend for larger fragments to harbor both higher number of types of reproductive sites and diversity of frog reproductive modes. Our findings suggests that fragment sizes can better predict the number of frog reproductive modes than the species richness in Atlantic Forest fragmented landscapes, and highlights the need to consider frog reproductive traits in future studies.     

Habitat Use and Selection by California Spotted Owls in a Postfire Landscape

ABSTRACT Forest fire is often considered a primary threat to California spotted owls (Strix occidentalis occidentalis) because fire has the potential to rapidly alter owl habitat. We examined effects of fire on 7 radiomarked California spotted owls from 4 territories by quantifying use of habitat for nesting, roosting, and foraging according to severity of burn in and near a 610-km²fire in the southern Sierra Nevada, California, USA, 4 years after fire. Three nests were located in mixed-conifer forests, 2 in areas of moderate-severity burn, and one in an area of low-severity burn, and one nest was located in an unburned area of mixed-conifer-hardwood forest. For roosting during the breeding season, spotted owls selected low-severity burned forest and avoided moderate- and high-severity burned areas; unburned forest was used in proportion with availability. Within 1 km of the center of their foraging areas, spotted owls selected all severities of burned forest and avoided unburned forest. Beyond 1.5 km, there were no discernable differences in use patterns among burn severities. Most owls foraged in high-severity burned forest more than in all other burn categories; high-severity burned forests had greater basal area of snags and higher shrub and herbaceous cover, parameters thought to be associated with increased abundance or accessibility of prey. We recommend that burned forests within 1.5 km of nests or roosts of California spotted owls not be salvage-logged until long-term effects of fire on spotted owls and their prey are understood more fully.     

Habitat Selection of a Large Carnivore along Human-Wildlife Boundaries in a Highly Modified Landscape

When large carnivores occupy peripheral human lands conflict with humans becomes inevitable, and the reduction of human-carnivore interactions must be the first consideration for those concerned with conflict mitigation. Studies designed to identify areas of high human-bear interaction are crucial for prioritizing management actions. Due to a surge in conflicts, against a background of social intolerance to wildlife and the prevalent use of lethal control throughout Japan, Asiatic black bears (Ursus thibetanus) are now threatened by high rates of mortality. There is an urgent need to reduce the frequency of human-bear encounters if bear populations are to be conserved. To this end, we estimated the habitats that relate to human-bear interactions by sex and season using resource selection functions (RSF). Significant seasonal differences in selection for and avoidance of areas by bears were estimated by distance-effect models with interaction terms of land cover and sex. Human-bear boundaries were delineated on the basis of defined bear-habitat edges in order to identify areas that are in most need of proactive management strategies. Asiatic black bears selected habitats in close proximity to forest edges, forest roads, rivers, and red pine and riparian forests during the peak conflict season and this was correctly predicted in our human-bear boundary maps. Our findings demonstrated that bears selected abandoned forests and agricultural lands, indicating that it should be possible to reduce animal use near human lands by restoring season-specific habitat in relatively remote areas. Habitat-based conflict mitigation may therefore provide a practical means of creating adequate separation between humans and these large carnivores.     

Effects of Habitat on Competition Between Kit Foxes and Coyotes

Abstract: San Joaquin kit foxes (Vulpes macrotis mutica) are an endangered species with a narrow geographic range whose natural populations are limited by predation by coyotes (Canis latrans). In the warm, arid grassland and shrubland habitats where kit foxes occur, coyotes are more cover dependent than kit foxes, creating the possibility of habitat segregation. Effects of habitat variation on coyote and kit fox competition are unknown. We assessed exploitation and interference competition between coyotes and kit foxes in grassland and shrubland habitats to determine if such competition varies among habitats. With respect to exploitation competition, we evaluated habitat and spatial partitioning, diet, prey abundance, and survival for kit foxes and coyotes at the Lokern Natural Area in central California, USA, from January 2003 through June 2004. Kit foxes partitioned habitat, space, and diet with coyotes. Coyotes primarily used shrubland habitats whereas kit foxes selectively used burned grasslands. Kit foxes and coyotes had high dietary overlap with regards to items used, but proportional use of items differed between the 2 species. Kit foxes selected for Heermann's kangaroo rats (Dipodomys heermanni), which were closely tied to shrub habitats. With respect to interference competition, predation was the primary source of mortality for kit foxes, and survival of individual kit foxes was inversely related to proportion of shrub habitat within their home ranges. Our results suggest that a heterogeneous landscape may benefit kit foxes by providing habitat patches where predation risk may be lower.     

Habitat Selection by Prairie Dogs in a Disturbed Landscape at the Edge of Their Geographic Range

ABSTRACT Although habitat attributes of black-tailed prairie dog (Cynomys ludovicianus) colonies have been described for central and northern portions of the species' geographic range, little is known about these associations at the southern edge of this species' distribution. Because high-quality habitats are expected to be scarcer at the edge of the species' geographic range, different patterns of habitat selection might emerge in these populations. We analyzed habitat selection by black-tailed prairie dogs in a human-disturbed mosaic of desert grasslands and shrublands in northwestern Chihuahua, Mexico. We contrasted 151 used and 133 unused habitat units producing 11 case-control logistic regression models to explain site occupancy by prairie dogs with different combinations of environmental variables. Prairie dogs from Chihuahua occupy sites similar in most respects to sites in more northern regions, although these prairie dogs appear to be more tolerant of increased shrub density and reduced herbage cover. We found that site occupancy was best modeled by positive effects of soil moisture level, cover of forbs, cover of unpalatable vegetation, cover of bare ground, and amount of prairie-dog colony area within 1 km and by the inverse of altitude, shrub density, herbage height, and amount of hostile habitat within 1 km. The 2 most significant variables were herbage height and shrub density, which might reflect the prominent role that visibility plays in habitat selection by prairie dogs. In contrast, we found weak evidence that human features have significant impacts on site occupancy by prairie dogs. Our results support the prediction that environmental conditions of sites used by prairie dogs in edge regions partially differ from those observed in more northern latitudes. We suggest that reserve managers focus conservation efforts on areas with short vegetation, low density of shrubs, and high herbage cover, conditions that could be promoted by controlled burns, herbage mowing, and mechanical removal of shrubs.     

Population Ecology of the Eastern Box Turtle in a Fragmented Landscape

Abstract: In the mid-Atlantic region, urban sprawl and development have resulted in habitat alterations and fragmentation; however, the effects on eastern box turtle (Terrapene carolina carolina) populations are unclear. To investigate the status of eastern box turtle populations in a fragmented landscape, we used mark—recapture and radiotelemetry to estimate population density, sex ratio, age structure, and survival on 4 study areas with differing degrees of isolation and human disturbance in northern New Castle County, Delaware, USA. We estimated adult population densities ranging from 0.81 turtles/ha to 3.62 turtles/ha among our 4 study areas. Sex ratios were male-biased at 2 study areas and balanced at 2 study areas. Proportion of juveniles ranged from 0% to 31%. Estimated annual survival rate ranged from 0.813 to 0.977. Mortality of radiotagged and marked turtles was primarily due to natural causes, but mowing was the primary cause of human-induced mortality. We found evidence of population decline at one study area due to low survival and recruitment. Human disturbances, isolation, and habitat composition appear to have the greatest influence on the box turtle populations we studied. To minimize mortality from human disturbance, we suggest planting crops adjacent to forest habitat that require no mowing or mowing at a height ≥15 cm. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):745–753; 2008)     

Spatial Overlap Between People and Non-human Primates in a Fragmented Landscape

In western Uganda, the landscape surrounding Kibale National Park (KNP) contains households, trading centers, roads, fields, and forest fragments. The mosaic arrangement of these landscape features is thought to enhance human–primate interaction, leading to primate population declines and increased bi-directional disease transmission. Using a social–ecological systems research framework that captures the complexity of interaction among people, wildlife, and environment, we studied five forest fragments near KNP and conducted intensive on-the-ground mapping to identify locations of human–primate spatial overlap. Primate locations and human activities were distributed within, on the edges, and far beyond fragment borders. Analysis of shared spaces indicated that 5.5% of human space overlapped with primate spaces, while 69.5% of primate spaces overlapped with human spaces. Nearest neighbor analysis indicated that human activities were significantly spatially clustered within and around individual fragments, as were primate locations. Getis–Ord statistics revealed statistically significant “hotspots” of human activity and primate activity, but only one location where spatial overlap between humans and primates was statistically significant. Human activities associated with collecting fuelwood and other forest products were the primary drivers of human–primate overlap; however, primates also spent time outside of forest fragments in agricultural spaces. These results demonstrate that fragmented landscapes are not uniform with respect to human–primate overlap, and that the implications of human–primate interaction, such as primate population declines and possible cross-species disease transmission, are spatially aggregated.     

Genetic Connectivity of the Moth Pollinated Tree Glionnetia sericea in a Highly Fragmented Habitat

Long-distance gene flow is thought to be one prerequisite for the persistence of plant species in fragmented environments. Human influences have led to severe fragmentation of native habitats in the Seychelles islands, with many species surviving only in small and isolated populations. The endangered Seychelles endemic tree Glionnetia sericea is restricted to altitudes between 450 m and 900 m where the native forest vegetation has been largely lost and replaced with exotic invasives over the last 200 years. This study explores the genetic and ecological consequences of population fragmentation in this species by analysing patterns of genetic diversity in a sample of adults, juveniles and seeds, and by using controlled pollination experiments. Our results show no decrease in genetic diversity and no increase in genetic structuring from adult to juvenile cohorts. Despite significant inbreeding in some populations, there is no evidence of higher inbreeding in juvenile cohorts relative to adults. A Bayesian structure analysis and a tentative paternity analysis indicate extensive historical and contemporary gene flow among remnant populations. Pollination experiments and a paternity analysis show that Glionnetia sericea is self-compatible. Nevertheless, outcrossing is present with 7% of mating events resulting from pollen transfer between populations. Artificial pollination provided no evidence for pollen limitation in isolated populations. The highly mobile and specialized hawkmoth pollinators (Agrius convolvuli and Cenophodes tamsi; Sphingidae) appear to promote extensive gene flow, thus mitigating the potential negative ecological and genetic effects of habitat fragmentation in this species. We conclude that contemporary gene flow is sufficient to maintain genetic connectivity in this rare and restricted Seychelles endemic, in contrast to other island endemic tree species with limited contemporary gene flow.     

Habitat Deterioration, Habitat Destruction, and Metapopulation Persistence in a Heterogenous Landscape

Levins's unstructured metapopulation model predicts that the equilibrium fraction of empty habitat patches is a constant function of the fractionhof suitable patches in the landscape and that this constant equals the threshold value for metapopulation persistence. Levins's model thus suggests that the minimum amount of suitable habitat necessary for metapopulation persistence can be estimated from the fraction of empty patches at steady state. In this paper we construct several more realistic structured metapopulation models that include variation in patch quality and the rescue effect. These models predict both positive and negative correlations between the fractions of suitable patches and empty patches. The type of correlation depends in an intricate manner on the strength of the rescue effect and on the quality distribution of the patches to be destroyed. Empty patches can be considered as the resource limiting metapopulation growth. Our results demonstrate that the correlation between the fractions of suitable patches and empty patches is positive if and only if the average value of the resource decreases as the number of patches increases.     

The Nature of Seedling Assemblages in a Fragmented Tropical Landscape: Implications for Forest Regeneration

Whether aging forest fragments are able to recover original assemblages or progressively move toward impoverished successional stages remains as an open question. This study tested the hypothesis that seedling assemblages in forest fragments differ from those across mature forest stands and examined to what extent the uncovered patterns supported the notion that edge‐affected habitats tend to support impoverished tree assemblages dominated by pioneer species. We contrasted a series of small forest remnants (3–91 ha) to old‐growth stands located in the largest (ca 3500 ha) and best preserved forest remnant in northeastern Brazil and found that tree seedling assemblages inhabiting forest fragments exhibited reduced species richness (up to 50%) at different spatial scales in comparison to seedling assemblages in mature forest and adult assemblages in both fragments and mature forest stands. Moreover, ordination analyses clearly segregated fragment seedling assemblages in taxonomic/functional terms and segregation correlated to the richness of pioneer species. Seedlings of pioneer species and those bearing medium‐sized seeds (0.6–1.5 cm) increased in fragments, whereas large‐seeded species (1.5–3.0 cm) were reduced by more than a half. Such a multiple‐scale replacement of the old‐growth flora by pioneers was also confirmed by an indicator species analysis and the resulting pioneer indicator species. Our results suggest that small forest fragments support impoverished and distorted seedling assemblages. This floristic/functional drift implies that forest remnants or edge‐affected habitats tend to be dominated by a small set of pioneer tree species rather than supporting a substantial portion of the old‐growth flora as do mature forest stands.     

栖息地选择的理论与模型

栖息地选择理论和模型的发展经历了两个主要阶段:理想自由分布模型和空间直观的栖息地选择模型。随着对理论模型假设的放宽,近年来产生了越来越多的新模型。通过对栖息地选择过程的分析,提出了栖息地选择的几个关键环节:栖息地偏好、信息获取、行为决策及选择行为。在建立栖息地选择模型的各个关键环节上均存在大量有待解决的问题。目前对栖息地偏好的研究主要为相关分析,栖息地信息获取的过程仍然是一黑箱;对动物在栖息地选择过程中的行为决策以及对其生理状态的影响尚不了解,而解决这些问题需要生态学、生态学及认知学等多个领域的研究结果支持,也有待新的理论及方法加以充实,甚至还需要其他学科的介入。     

Restoration of a Native Grassland as Habitat for the Golden Sun Moth Synemon plana Walker (Lepidoptera; Castniidae) at Mount Piper, Australia

Synemon plana (golden sun moth) is a small diurnal moth from the family Castniidae, now listed as endangered in all regions of southeastern Australia in which it occurs. This study details an attempt to restore the native grassland habitat of S. plana at one site by increasing the cover of its probable food plant, Austrodanthonia eriantha (wallaby grass), to at least 40%, the average percentage cover determined from all currently inhabited sites. The percentage cover of A. eriantha increased when seedlings were planted, although there was no significant increase in cover by seeding caryopsides (dispersal units) or by weeding. However, combining weeding with planting seedlings, or seeding increased cover by 22% and 27% to 43% and 64%, respectively. Two hundred caryopsides per square meter (compared with only 49 seedlings/m²) were required to increase cover to at least 40% due to the low viability of seeds and low germination in the field. Therefore, to restore grasslands dominated by A. eriantha as suitable habitat for S. plana, seedlings should be planted and weeds removed.     

Inoculation with a Native Soil Community Advances Succession in a Grassland Restoration

Restoration on post‐agricultural land may be hindered by the degradation of the soil community, which has been shown to contribute to structuring plant communities and driving succession. Our experiment tested the effect of inoculation with remnant grassland whole soil with or without nurse plants on the survival and growth of uninoculated early and late successional plant species. In 2007 and 2008, we planted uninoculated early, mid, and late successional plant species 0.25–2 m away from a central point of inoculated nurse plants. We found a negative response to inoculation on early successional plants and a positive response to inoculation on mid to late successional plants. This work suggests that the restoration of the soil community is critical to establishing a late successional plant community and that the benefit of inoculated plants can spread to neighbors.