Invasion in a diversity hotspot: exotic cover and native richness in the Californian serpentine flora

Exotic species have been observed to be more prevalent in sites where the richness of native species is highest, possibly reflecting variation among sites in resources, propagule supply, heterogeneity, or disturbance. However, such a pattern leaves unclear whether natives at species-rich sites are subject to especially severe impacts from exotics as a result. We considered this question using path models in which relationships between exotic cover and native richness were evaluated in the presence of correlated environmental factors. At 109 sites on serpentine soils across California, USA, exotic cover was positively correlated with total native herbaceous richness and was negatively correlated with the richness of both serpentine-endemic and rare native herbs. However, in path models that accounted for the influences of soil chemistry, disturbance, overstory cover, and regional rainfall and elevation, we found no indication that exotic cover reduced any component of native herb richness. Rather, our results indicated similarities and differences in the conditions favoring exotic, native, endemic, and rare species. Our results suggest that, in spite of some localized impacts, exotic species are not exerting a detectable overall effect on the community richness of the unique native flora of Californian serpentine.     

Human impacts, plant invasion, and imperiled plant species in California.

Invasive species are one of the fastest growing conservation problems. These species homogenize the world's flora and fauna, threaten rare and endemic species, and impose large economic costs. Here, we examine the distribution of 834 of the more than 1000 exotic plant taxa that have become established in California, USA. Total species richness increases with net primary productivity; however, the exotic flora is richest in low-lying coastal sites that harbor large numbers of imperiled species, while native diversity is highest in areas with high mean elevation. Weedy and invasive exotics are more tightly linked to the distribution of imperiled species than the overall pool of exotic species. Structural equation modeling suggests that while human activities, such as urbanization and agriculture, facilitate the initial invasion by exotic plants, exotics spread ahead of the front of human development into areas with high numbers of threatened native plants. The range sizes of exotic taxa are an order of magnitude smaller than for comparable native taxa. The current small range size of exotic species implies that California has a significant "invasion debt" that will be paid as exotic plants expand their range and spread throughout the state.     

Wildlife Association with Human-Altered Water Sources in Semiarid Vegetation Communities

Based on common use in wildlife management, we hypothesized that human-constructed water sources influence faunal communities detectably compared to similar habitats that lack water. We examined 20 wildlife water units and 20 paired comparison sites without water from April to August 1992 in semiarid southern New Mexico to assess animal species associations. We sampled sites by using small-mammal live traps, herpetofaunal and invertebrate pitfall arrays, and 30-minute time-area counts. We compared animal species richness and species concordance among water units (rain catchments, earthen tanks, and windmills) and comparison sites in three vegetation communities (mixed scrub, grassland, and pinyon-juniper). We detected 134 animal taxa during field sampling. Animal species richness did not differ between water units and comparison sites among vegetation communities. Amphibians were found only at water units but occur far from units during seasonal wet periods. Greater numbers of individual small mammals and herpetofauna at water units versus comparison sites likely related to debris and disturbed soil present near water units. Taxa detected at water units and comparison sites were 65% concordant overall; discordant taxa were those rarely detected. Our data implied that definitive effects of artificial water sources on native wildlife species were not detectable. Providing water sources may be a strategic management tool but must be viewed critically regarding effect on distribution of native, feral, and exotic animals. Water units should be developed only when and where clear objectives have been stated, natural water sources have been quantified, commitment exists to ensure continued function, and feral and exotic animals will not benefit to the detriment of native species.     

Modifying native and exotic species richness correlations: the influence of fire and seed addition.

An important goal in restoration is to increase the richness of native species while reducing exotic species. However, native species richness is often positively correlated with exotic species richness. In a grassland-savanna system in Michigan (USA), we show that management that focuses on changing the nature of the exotic-native richness relationship can be used to restore native communities. Native and exotic species richnesses were positively correlated, likely due to a shared coupling with aboveground live biomass (a surrogate for productivity). The addition of native seed shifted the exotic-native richness relationship from a linear positive to a monotonic relationship: in areas of intermediate levels of exotic species richness, seed addition increased native diversity without an associated effect on exotic diversity, but in areas of high or low exotic richness, it did not affect native species richness. Prescribed burning broke the correlation between native and exotic richness with no consistent effect on the richness of either group. However, when burning was combined with native-seed addition, the relationship between native and exotic richness was maintained and was shifted upwards, enhancing native recruitment. Although aboveground productivity was strongly related to species richness across the landscape, changes in productivity did not drive these shifts.     

High richness and dense seeding enhance grassland restoration establishment but have little effect on drought response

Restorations commonly utilize seed addition to formerly arable lands where the development of native plant communities is severely dispersal limited. However, variation in seed addition practices may profoundly affect restoration outcomes. Theory and observations predict that species-rich seed mixtures and seeding at high densities should enhance native plant community establishment, minimize exotic species cover, and may promote resistance and resilience to, and recovery from, environmental perturbations. We studied the post-seeding establishment of native plant communities in large grassland restoration plots, which were sown at two densities crossed with two levels of species richness on formerly arable land in Nebraska, USA, and their responses to drought. To evaluate drought resistance, recovery, and resilience of restored plant communities, we erected rainfall manipulation structures and tracked the response of seeded species cover and total plant biomass during experimental drought relative to controls and in the post-drought growing season. High seed richness and high-density seeding treatments resulted in greater richness and cover of native, seeded species per 0.5 m2 compared to low-richness and low-density treatments. Cover differences in response to seed mixture richness were driven by native forbs. Richness and cover of exotic species were lowest in high-richness and high-density treatments. We found little evidence of differential drought resistance, recovery, and resilience among seeding treatments. Increases in exotic species across years were restricted to drought subplots, and were not affected by seeding treatments. Grassland restoration was generally enhanced and exotic cover reduced both by the use of high-richness seed mixtures and high-density seeding. Given the lack of restoration treatment effects on the resistance, recovery, or resilience of seeded species exposed to drought, and the increases in exotic species following drought, other forms of active management may be needed to produce restored plant communities that are robust to climate change.     

Provenance, life span, and phylogeny do not affect grass species' responses to nitrogen and phosphorus

Successful conservation management requires an understanding of how species respond to intervention. Native and exotic species may respond differently to management interventions due to differences arising directly from their origin (i.e., provenance) or indirectly due to biased representations of different life history types (e.g., annual vs. perennial life span) or phylogenetic lineages among provenance (i.e., native or exotic origin) groups. Thus, selection of a successful management regime requires knowledge of the life history and provenance-bias in the local flora and an understanding of the interplay between species characteristics across existing environmental gradients in the landscape. Here we tested whether provenance, phylogeny, and life span interact to determine species distributions along natural gradients of soil chemistry (e.g., soil nitrogen and phosphorus) in 10 upland prairie sites along a 600-km latitudinal transect running from southern Vancouver Island in British Columbia, Canada, to the Willamette Valley in Oregon, USA. We found that soil nitrate, phosphorus, and pH exerted strong control over community composition. However, species distributions along environmental gradients were unrelated to provenance, life span, or phylogenetic groupings. We then used a greenhouse experiment to more precisely measure the response of common grass species to nitrogen and phosphorus supply. As with the field data, species responses to nutrient additions did not vary as a function of provenance, life span, or phylogeny. Native and exotic species differed strongly in the relationship between greenhouse-measured tolerance of low nutrients and field abundance. Native species with the greatest ability to maintain biomass production at low nutrient supply rates were most abundant in field surveys, as predicted by resource competition theory. In contrast, there was no relationship between exotic-species biomass at low nutrient levels and field abundance. The implications of these findings for management of invasive species are substantial in that they overturn a general belief that reduction of nutrient supplies favors native species. The idiosyncratic nature of species response to nutrients in this study suggests that manipulation of nutrient supplies is unlikely to alter the overall balance between native and exotic species, although it may well be useful to control specific exotic species.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Effect of Invasive Plant Species on Temperate Wetland Plant Diversity

Abstract:  Invasive species are a major threat to global biodiversity and an important cause of biotic homogenization of ecosystems. Exotic plants have been identified as a particular concern because of the widely held belief that they competitively exclude native plant species. We examined the correlation between native and invasive species richness in 58 Ontario inland wetlands. The relationship between exotic and native species richness was positive even when we controlled for important covarying factors. In addition, we examined the relationship between the abundance of four native species (  Typha latifolia, T. angustifolia, Salix petiolaris, Nuphar variegatum ) and four invasive species (  Lythrum salicaria, Hydrocharis morsus-ranae, Phalaris arundinacea, Rhamnus frangula ) that often dominate temperate wetlands and native and rare native species richness. Exotic species were no more likely to dominate a wetland than native species, and the proportion of dominant exotic species that had a significant negative effect on the native plant community was the same as the proportion of native species with a significant negative effect. We conclude that the key to conservation of inland wetland biodiversity is to discourage the spread of community dominants, regardless of geographical origin.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Effects of Exotic Lonicera and Rhamnus on Songbird Nest Predation

Abstract: Habitat fragmentation and disturbance exacerbate the invasion of exotic plant species that, in turn, may attract nesting songbirds by providing a branch structure suitable for nest sites. We document that American Robin ( Turdus migratorius ) nests in two exotic plants, Lonicera maackii and Rhamnus cathartica , experienced higher predation than nests built in comparable native shrubs ( Crataegus, Viburnum ) and native tree species. This was due to a combination of lower nest height, the absence of sharp thorns on the exotic species, and perhaps a branch architecture that facilitated predator movement among the exotic species. In a more subtle interaction, nesting Wood Thrushes ( Hylocichla mustelina ) experienced apparent competition with robins for nest sites in Lonicera , and this interaction was further aggravated by an increased selectivity for Lonicera by nesting robins, possibly due to their early leaf flush and expansion. By documenting increased nest predation in songbirds nesting in exotic shrubs, our results suggest that restoring native plant communities may benefit the surrounding avian community.     

Abundance declines of a native forb have nonlinear impacts on grassland invasion resistance

The effects of declining plant biodiversity on ecosystem processes are well studied, with most investigations examining the role of species richness declines rather than declines of species abundance. Using grassland mesocosms, we examined how the abundance of a native, resident species, Hemizonia congesta (hayfield tarweed), affected exotic Centaurea solstitialis (yellow starthistle) invasion. We found that progressive H. congesta abundance declines had threshold effects on invasion resistance, with initial declines resulting in minor increases in invasion and subsequent declines leading to accelerating increases in invader performance. Reduced invasion resistance was explained by increased resource availability as H. congesta declined. We also found evidence that resident abundance might indirectly affect invasion by mediating invader impact on resident competitors; C. solstitialis disproportionately reduced H. congesta biomass in low-abundance rather than high-abundance populations. H. congesta's direct and indirect effects on invasion resistance illustrate that an individual species' declining abundance can have accelerating, deleterious effects on ecosystem functions of conservation value.     

Native plant diversity resists invasion at both low and high resource levels

Human modification of the environment is causing both loss of species and changes in resource availability. While studies have examined how species loss at the local level can influence invasion resistance, interactions between species loss and other components of environmental change remain poorly studied. In particular, the manner in which native diversity interacts with resource availability to influence invasion resistance is not well understood. We created experimental plant assemblages that varied in native species (1-16 species) and/or functional richness (defined by rooting morphology and phenology; one to five functional groups). We crossed these diversity treatments with resource (water) addition to determine their interactive effects on invasion resistance to spotted knapweed (Centaurea maculosa), a potent exotic invader in the intermountain West of the United States. We also determined how native diversity and resource addition influenced plant-available soil nitrogen, soil moisture, and light. Assemblages with lower species and functional diversity were more heavily invaded than assemblages with greater species and functional diversity. In uninvaded assemblages, experimental addition of water increased soil moisture and plant-available nitrogen and decreased light availability. The availability of these resources generally declined with increasing native plant diversity. Although water addition increased susceptibility to invasion, it did not fundamentally change the negative relationship between diversity and invasibility. Thus, native diversity provided strong invasion resistance even under high resource availability. These results suggest that the effects of local diversity can remain robust despite enhanced resource levels that are predicted under scenarios of global change.     

Disturbance influences oyster community richness and evenness, but not diversity

Foundation species in space-limited systems can increase diversity by creating habitat, but they may also reduce diversity by excluding primary space competitors. These contrasting forces of increasing associate diversity and suppressing competitor diversity have rarely been examined experimentally with respect to disturbance. In a benthic marine community in central California, where native oysters are a foundation species, we tested how disturbance influenced overall species richness, evenness, and diversity. Surprisingly, overall diversity did not peak across a disturbance gradient because, as disturbance decreased, decreases in overall species evenness opposed increases in overall species richness. Decreasing disturbance intensity (high oyster abundance) led to increasing species richness of sessile and mobile species combined. This increase was due to the facilitation of secondary sessile and mobile species in the presence of oysters. In contrast, decreasing disturbance intensity and high oyster abundance decreased the evenness of sessile and mobile species. Three factors likely contributed to this decreased evenness: oysters reduced abundances of primary sessile species due to space competition; oysters supported more rare mobile species; and oysters disproportionately increased the relative abundance of a few common mobile species. Our results highlight the need for further studies on how disturbance can differentially affect the evenness and richness of different functional groups, and ultimately how these differences affect the relationship between overall diversity and ecosystem function.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

Evaluation of Central North American Prairie Management Based on Species Diversity, Life Form, and Individual Species Metrics

Abstract:  Reintroduction of fire and grazing, alone or in combination, has increasingly been recognized as central to the restoration of North American mixed-grass and tallgrass prairies. Although ecological studies of these systems are abundant, they have generally been observational, or if experimental, have focused on plant species diversity. Species diversity measures alone are not sufficient to inform management, which often has goals associated with life-form groups and individual species. We examined the effects of prescribed fire, light cattle grazing, and a combination of fire and grazing on three vegetation components: species diversity, groups of species categorized by life-form, and individual species. We evaluated how successful these three treatments were in achieving specific management goals for prairies in the Iowa Loess Hills (U.S.A.). The grazing treatment promoted the greatest overall species richness, whereas grazing and burning and grazing treatments resulted in the lowest cover by woody species. Burning alone best achieved the management goals of increasing the cover and diversity of native species and reducing exotic forb and (predominantly exotic) cool-season grass cover. Species-specific responses to treatments appeared idiosyncratic (i.e., within each treatment there existed a set of species attaining their highest frequency) and nearly half of uncommon species were present in only one treatment. Because all management goals were not achieved by any one treatment, we conclude that management in this region may need refining. We suggest that a mosaic of burning and grazing (alone and in combination) may provide the greatest landscape-level species richness; however, this strategy would also likely promote the persistence of exotic species. Our results support the need to consider multiple measures, including species-specific responses, when planning and evaluating management .     

Consistency of effects of tropical‐forest disturbance on species composition and richness relative to use of indicator taxa

Lawton et al. (1998) found, in a highly cited study, that the species richness of 8 taxa each responds differently to anthropogenic disturbance in Cameroon forests. Recent developments in conservation science suggest that net number of species is an insensitive measure of change and that understanding which species are affected by disturbance is more important. It is also recognized that all disturbance types are not equal in their effect on species and that grouping species according to function rather than taxonomy is more informative of responses of biodiversity to change. In a reanalysis of most of the original Cameroon data set (canopy and ground ants, termites, canopy beetles, nematodes, and butterflies), we focused on changes in species and functional composition rather than richness and used a more inclusive measure of forest disturbance based on 4 component drivers of change: years since disturbance, tree cover, soil compaction, and degree of tree removal. Effects of disturbance on compositional change were largely concordant between taxa. Contrary to Lawton et al.’s findings, species richness for most groups did not decline with disturbance level, providing support for the view that trends in species richness at local scales do not reflect the resilience of ecosystems to disturbance. Disturbance affected species composition more strongly than species richness for butterflies, canopy beetles, and litter ants. For these groups, disturbance caused species replacements rather than just species loss. Only termites showed effects of disturbance on species richness but not composition, indicating species loss without replacement. Although disturbance generally caused changes in composition, the strength of this relationship depended on the disturbance driver. Butterflies, litter ants, and nematodes were correlated with amount of tree cover, canopy beetles were most strongly correlated with time since disturbance, and termites were most strongly correlated with degree of soil disturbance. There were moderately divergent responses to disturbance between functional feeding groups. Disturbance was most strongly correlated with compositional differences of herbivores within beetles and nematodes and humus feeders within termites. Our results suggest that consideration of the impact of different forms of disturbance on species and functional composition, rather than on net numbers of species, is important when assessing the impacts of disturbance on biodiversity.     

Shifts in southern Wisconsin forest canopy and understory richness, composition, and heterogeneity

We resurveyed the under- and overstory species composition of 94 upland forest stands in southern Wisconsin in 2002-2004 to assess shifts in canopy and understory richness, composition, and heterogeneity relative to the original surveys in 1949-1950. The canopy has shifted from mostly oaks (Quercus spp.) toward more mesic and shade-tolerant trees (primarily Acer spp.). Oak-dominated early-successional stands and those on coarse, nutrient-poor soils changed the most in canopy composition. Understories at most sites (80%) lost native species, with mean species density declining 25% at the 1-m2 scale and 23.1% at the 20-m2 scale. Woody species have increased 15% relative to herbaceous species in the understory despite declining in absolute abundance. Initial canopy composition, particularly the abundance of red oaks (Quercus rubra and Q. velutina), predicted understory changes better than the changes observed in the overstory. Overall rates of native species loss were greater in later-successional stands, a pattern driven by differential immigration rather than differential extirpation. However, understory species initially found in early-successional habitats declined the most, particularly remnant savanna taxa with narrow or thick leaves. These losses have yet to be offset by compensating increases in native shade-adapted species. Exotic species have proliferated in prevalence (from 13 to 76 stands) and relative abundance (from 1.2% to 8.4%), but these increases appear unrelated to the declines in native species richness and heterogeneity observed. Although canopy succession has clearly influenced shifts in understory composition and diversity, the magnitude of native species declines and failure to recruit more shade-adapted species suggest that other factors now act to limit the richness, heterogeneity, and composition of these communities.     

Spatial and temporal patterns of seed dispersal: an important determinant of grassland invasion.

We measured spatial and temporal patterns of seed dispersal and seedling recruitment for 58 species in a grassland community to test whether seed dispersal could predict patterns of invasion after disturbance. For the 12 most abundant grasses, recruitment of native species was dependent on the propagule supply of both native and exotic species. Variability in seed rain on small spatial (1-10 m) and temporal (within season) scales led to qualitative differences in the outcome of disturbance colonization such that native species dominated disturbances when exotic seed supply was low but failed to establish when exotic seed supply was high. Local dispersal and spatial heterogeneity in species composition promoted coexistence of native and exotic species by creating refuges from high exotic seed supply within native dominated patches. Despite this, copious exotic seed production strongly limited recruitment of native species in exotic dominated patches. Most grasslands in California are presently dominated by exotic species, suggesting that competition at the seedling stage is a major barrier to native species restoration.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.