Impacts of road deicing salt on the demography of vernal pool-breeding amphibians.

Deicing agents, primarily road salt, are applied to roads in 26 states in the United States and in a number of European countries, yet the scale of impacts of road salt on aquatic organisms remains largely under-studied. The issue is germane to amphibian conservation because both adult and larval amphibians are known to be particularly sensitive to changes in their osmolar environments. In this study, we combined survey, experimental, and demographic modeling approaches to evaluate the possible effects of road salt on two common vernal-pond-breeding amphibian species, the spotted salamander (Ambystoma maculatum) and the wood frog (Rana sylvatica). We found that in the Adirondack Mountain Region of New York (USA), road salt traveled up to 172 m from the highway into wetlands. Surveys showed that egg mass densities of spotted salamanders (A. maculatum) and wood frogs (R. sylvatica) were two times higher in forest pools than roadside pools, but this pattern was better explained by road proximity than by increased salinity. Experiments demonstrated that embryonic and larval survival were reduced at moderate (500 muS) and high conductivities (3000 muS) in A. maculatum and at high conductivities in R. sylvatica. Demographic models suggest that such egg and larval stage effects of salt may have important impacts on populations near roads, particularly in the case of A. maculatum, for which salt exposure may lead to local extinction. For both species, the effect of road salt was dependent upon the strength of larval density dependence and declined rapidly with distance from the roadside, with the greatest negative effects being limited to within 50 m. Based on this evidence, we argue that efforts to protect local populations of A. maculatum and R. sylvatica in roadside wetlands should, in part, be aimed at reducing application of road salt near wetlands with high conductivity levels.     

The Ecological Road-Effect Zone of a Massachusetts (U.S.A.) Suburban Highway

Abstract: Ecological flows and biological diversity trace broad patterns across the landscape, whereas transportation planning for human mobility traditionally focuses on a narrow strip close to a road or highway. To help close this gap we examined the "road-effect zone" over which significant ecological effects extend outward from a road. Nine ecological factors—involving wetlands, streams, road salt, exotic plants, moose, deer, amphibians, forest birds, and grassland birds—were measured or estimated near 25 km of a busy four-lane highway west of Boston, Massachusetts. The effects of all factors extended > 100 m from the road, and moose corridors, road avoidance by grassland birds, and perhaps road salt in a shallow reservoir extended outwards > 1 km. Most factors had effects at 2–5 specific locations, whereas traffic noise apparently exerted effects along most of the road length. Creating a map of these effects indicates that the road-effect zone averages approximately 600 m in width and is asymmetric, with convoluted boundaries and a few long fingers. We conclude that busy roads and nature reserves should be well separated, and that future transportation systems across landscapes can provide for ecological flows and biological diversity in addition to safe and efficient human mobility.     

Effects of Road Fencing on Population Persistence

Abstract:  Roads affect animal populations in three adverse ways. They act as barriers to movement, enhance mortality due to collisions with vehicles, and reduce the amount and quality of habitat. Putting fences along roads removes the problem of road mortality but increases the barrier effect. We studied this trade-off through a stochastic, spatially explicit, individual-based model of population dynamics. We investigated the conditions under which fences reduce the impact of roads on population persistence. Our results showed that a fence may or may not reduce the effect of the road on population persistence, depending on the degree of road avoidance by the animal and the probability that an animal that enters the road is killed by a vehicle. Our model predicted a lower value of traffic mortality below which a fence was always harmful and an upper value of traffic mortality above which a fence was always beneficial. Between these two values the suitability of fences depended on the degree of road avoidance. Fences were more likely to be beneficial the lower the degree of road avoidance and the higher the probability of an animal being killed on the road. We recommend the use of fences when traffic is so high that animals almost never succeed in their attempts to cross the road or the population of the species of concern is declining and high traffic mortality is known to contribute to the decline. We discourage the use of fences when population size is stable or increasing or if the animals need access to resources on both sides of the road, unless fences are used in combination with wildlife crossing structures. In many cases, the use of fences may be beneficial as an interim measure until more permanent measures are implemented.     

Are calcareous soils in uplands less prone to damage from road salting than acidic soils?

Previous studies of upland roadside soils in Cumbria, that would normally be naturally acidic, have highlighted that (a) runoff from roads subjected to long-term road salting can dramatically raise soil pH down slope in upland areas; (b) the soil pH increase dramatically changes N cycling in soils down slope, increasing mineralisation of organic matter, ammonification, ammonium leaching down slope and nitrification and nitrate leaching; (c) the increase in nitrification substantially increases nitrate leaching to down-slope rivers, and this is readily detectable in field studies; and (d) loss of soil organic matter over decades of salting is so great that organic matter is no longer substantially solubilised by high salt concentrations found in soil solution below road drains. This paper tests and supports the hypothesis that such effects are minimal for more calcareous soil ecosystems. It examines the soil and soil solution chemistry on another Cumbrian upland highway, the A686 near Leadgate, Alston. Sodium % of soil CEC values for soil transects affected by spray containing road salt are similar at both the A6 and A686 sites. However, spatial trends in calcium, magnesium, ammonium, and nitrate concentrations as well as pH differ, as a direct result of the higher weathering rate of parent material and possibly also the presence of limestone walls above both spray-affected and control transects at the A686 site.     

A comparison of spatial learning and memory capabilities in intertidal gobies

For the majority of animals, the ability to orient in familiar locations is a fundamental part of life, and spatial memory allows individuals to remember key locations such as food patches, shelter, mating sites or areas regularly occupied by predators. This study determined if gobies collected from rocky platforms and sandy beaches differ in their ability to learn and memorise the locations of tide pools in a simulated rocky intertidal zone. Intertidal rock pool gobies show acute homing abilities and, therefore, should be expected to display superior learning and memory capabilities. In contrast, it is unlikely that natural selection would favour similar learning skills in sandy shore fishes because they simply shift back and forth with the tides. The learning abilities of gobies were tested using small replica rock platforms, containing four tide pools that retained varying depths of water at simulated low tide. Gobies were categorised as having learnt the task if they were able to consistently locate the tide pool that retained the most water at simulated low tide as the pool with the most favourable conditions. Rock pool species were able to locate the deepest pool to wait out low tide for ~95 % of the trials, while species from sandy shores were found in the deepest pool ~10 % of trials. Despite repeated stranding, sandy shore fish continued to follow the tide out. We propose that rock pool species memorised the location of rock pools during simulated high tide enabling them to relocate the best refuge for low tide.     

Forest Roads as Partial Barriers to Terrestrial Salamander Movement

Abstract:  Roads can fragment animal populations by disrupting movement among formerly continuous habitats. Although models have demonstrated that disrupted movement can contribute to long-term extinction, there are few empirical data on the effects of roads on animal movement. We used displacement and homing experiments to determine whether forest roads are barriers to the movement of terrestrial salamanders. We displaced 1471 red-backed salamanders ( Plethodon cinereus ) across five forest roads and compared return rates to those of salamanders displaced equal distances toward the forest interior. Roads significantly reduced the return rate of salamanders, with a mean reduction of 51%. Steep roadside verges further reduced return rates, particularly for salamanders moving downhill across verges. The permeability of roads to salamander movement did not appear to be related to road surface type. Gravel roads had both the highest and lowest observed permeability with the two paved roads intermediate between these. We conclude that narrow forest roads are partial barriers to salamander movement and that steep roadside verges may exacerbate these effects.     

Predicting species distributions from samples collected along roadsides

Predictive models of species distributions are typically developed with data collected along roads. Roadside sampling may provide a biased (nonrandom) sample; however, it is currently unknown whether roadside sampling limits the accuracy of predictions generated by species distribution models. We tested whether roadside sampling affects the accuracy of predictions generated by species distribution models by using a prospective sampling strategy designed specifically to address this issue. We built models from roadside data and validated model predictions at paired locations on unpaved roads and 200 m away from roads (off road), spatially and temporally independent from the data used for model building. We predicted species distributions of 15 bird species on the basis of point-count data from a landbird monitoring program in Montana and Idaho (U.S.A.). We used hierarchical occupancy models to account for imperfect detection. We expected predictions of species distributions derived from roadside-sampling data would be less accurate when validated with data from off-road sampling than when it was validated with data from roadside sampling and that model accuracy would be differentially affected by whether species were generalists, associated with edges, or associated with interior forest. Model performance measures (kappa, area under the curve of a receiver operating characteristic plot, and true skill statistic) did not differ between model predictions of roadside and off-road distributions of species. Furthermore, performance measures did not differ among edge, generalist, and interior species, despite a difference in vegetation structure along roadsides and off road and that 2 of the 15 species were more likely to occur along roadsides. If the range of environmental gradients is surveyed in roadside-sampling efforts, our results suggest that surveys along unpaved roads can be a valuable, unbiased source of information for species distribution models.     

Platinum pollution in road dusts,roadside soils,and tree barks in Seoul,Korea

This study presents the level of platinum in urban environment in and around Seoul, the capital city of Korea. Road dust, roadside soil, and tree bark samples were collected from the sites of various traffic volumes and from control sites in the suburbs. The above samples were analyzed for Pt by ICP-MS and other heavy metals by ICP-OES. Platinum levels in road dusts and roadside soils from Seoul were in the range of 3.8–444 ng/g (av. 115.0 ng/g) and 0.7–221 ng/g (av. 49.7 ng/g), respectively, whereas those in the suburbs were in the range of 2.3–5.2 ng/g (av. 3.9 ng/g) in road dusts and 0.4–5.1 ng/g (av. 2.4 ng/g) in roadside soils. The highest Pt levels in road dusts were found from major roads with high traffic volume. The remarkable difference in average Pt level between heavy traffic roads (av. 132.2 ng/g) and light traffic roads (av. 22.8 ng/g) reflects that an important source of Pt in roadside environment is automobile catalytic converter. High Pt level in road dust was found from the site of erratic stop–start driving condition, for example, 178 ng/g Pt in road dust around a vehicle crossing gate. Platinum level in tree barks ranged from 0.9 to 4.5 ng/g, which indicates the existence of Pt-containing particulate matter in the atmosphere. Road dusts with high Pt level were enriched in traffic-related heavy metals.

     

Influences of traffic on Pb,Cu and Zn concentrations in roadside soils of an urban park in Dublin,Ireland

Soils in the vicinity of roads are recipients of contaminants from traffic emissions. In order to obtain a better understanding of the impacts of traffic on soils, a total of 225 surface soil samples were collected from an urban park (Phoenix Park, Dublin, Ireland) in a grid system. Metal (Pb, Cu and Zn) concentrations were determined using a portable X-ray fluorescence analyzer. Strong spatial variations for the concentrations of Pb, Cu and Zn were observed. The spatial distribution maps created using geographical information system techniques revealed elevated metal concentrations close to the main traffic route in the park. The relationships between the accumulation of Pb, Cu and Zn in the roadside soils and the distance from the road were well fitted with an exponential model. Elevated metal concentrations from traffic pollution extended to a distance of approximately 40 m from the roadside. The results of this study provide useful information for the management of urban parks particularly in relation to policies aimed at reducing the impact of traffic related pollution on soils.     

Platinum pollution in road dusts, roadside soils, and tree barks in Seoul, Korea

This study presents the level of platinum in urban environment in and around Seoul, the capital city of Korea. Road dust, roadside soil, and tree bark samples were collected from the sites of various traffic volumes and from control sites in the suburbs. The above samples were analyzed for Pt by ICP-MS and other heavy metals by ICP-OES. Platinum levels in road dusts and roadside soils from Seoul were in the range of 3.8-444 ng/g (av. 115.0 ng/g) and 0.7-221 ng/g (av. 49.7 ng/g), respectively, whereas those in the suburbs were in the range of 2.3-5.2 ng/g (av. 3.9 ng/g) in road dusts and 0.4-5.1 ng/g (av. 2.4 ng/g) in roadside soils. The highest Pt levels in road dusts were found from major roads with high traffic volume. The remarkable difference in average Pt level between heavy traffic roads (av. 132.2 ng/g) and light traffic roads (av. 22.8 ng/g) reflects that an important source of Pt in roadside environment is automobile catalytic converter. High Pt level in road dust was found from the site of erratic stop-start driving condition, for example, 178 ng/g Pt in road dust around a vehicle crossing gate. Platinum level in tree barks ranged from 0.9 to 4.5 ng/g, which indicates the existence of Pt-containing particulate matter in the atmosphere. Road dusts with high Pt level were enriched in traffic-related heavy metals.     

An adaptive plan for prioritizing road sections for fencing to reduce animal mortality

Mortality of animals on roads is a critical threat to many wildlife populations and is poised to increase strongly because of ongoing and planned road construction. If these new roads cannot be avoided, effective mitigation measures will be necessary to stop biodiversity decline. Fencing along roads effectively reduces roadkill and is often used in combination with wildlife passages. Because fencing the entire road is not always possible due to financial constraints, high-frequency roadkill areas are often identified to inform the placement of fencing. We devised an adaptive fence-implementation plan to prioritize road sections for fencing. In this framework, areas along roads of high, moderate, and low levels of animal mortality (respectively, roadkill hotspots, warmspots, and coldspots) are identified at multiple scales (i.e., in circles of different diameters [200–2000 m] in which mortality frequency is measured). Fence deployment is based on the relationship between the amount of fencing being added to the road, starting with the strongest roadkill hotspots, and potential reduction in road mortality (displayed in mortality-reduction graphs). We applied our approach to empirical and simulated spatial patterns of wildlife–vehicle collisions. The scale used for analysis affected the number and spatial extent of roadkill hot-, warm-, and coldspots. At fine scales (e.g., 200 m), more hotspots were identified than at coarse scales (e.g., 2000 m), but combined the fine-scale hotspots covered less road and less fencing was needed to reduce road mortality. However, many short fences may be less effective in practice due to a fence-end effect (i.e., animals moving around the fence more easily), resulting in a trade-off between few long and many short fences, which we call the FLOMS (few-long-or-many-short) fences trade-off. Thresholds in the mortality-reduction graphs occurred for some roadkill patterns, but not for others. Thresholds may be useful to consider when determining road-mitigation targets. The existence of thresholds at multiple scales and the FLOMS trade-off have important implications for biodiversity conservation.     

The Role of Roadsides in Plant Invasions: a Demographic Approach

Abstract:  Non-native plant species are common along roadsides, but presence does not necessarily indicate spread along the road axis. Roadsides may serve merely as habitat for a species spreading independently of roads. The potential conduit function of roads depends on the habitat specificity of the spreading species, its dispersal range relative to the spacing of roads in the landscape, and the relative importance of long- and short-range dispersal. We describe a demographic model of the road × species interaction and suggest methods of assessing conduit function in the field based on the model results. A species limited to roadside habitat will be constrained to spread along the road axis unless its long-range dispersal is sufficient to carry it across the intervening unfavorable area to another road. It will propagate along a road corridor at a rate determined by the scale of short-range dispersal. Effective management of an invasion requires distinguishing between the habitat and conduit functions, a distinction difficult to make with only snapshot data. Invasions can be reconstructed by several methods, but none is totally satisfactory. We suggest comparing stem distributions on transects parallel and perpendicular to the road axis, and beside the road, and away from it, with an idealized Gaussian curve. Such comparisons would allow discrimination between pattern determined by habitat suitability and pattern reflecting random and facilitated dispersal.     

Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil

Abstract: Roads through tropical forest create linear disturbances that have unknown consequences for forest birds. We studied how a narrow, rarely used road through otherwise undisturbed Amazonian forest affected the movements and area requirements of understory birds that form mixed-species flocks. Differences in road maintenance led to two distinct treatments along the same road. Trees along the "closed" road formed a partial canopy connecting the two sides of the road, although the roadway itself was kept open. The "open" road was regularly maintained, making a complete opening 10–30 m wide.We followed 15 flocks, 5 each in interior forest, along the open road, and along the closed road. These flocks were led by Thamnomanes antshrikes, and each flock had a discreet, permanent territory. Flock territory size (mean = 8.5 ha) did not differ among the three locations. The open road formed the territorial boundary for all five flocks, although birds moved within a few meters of the edge of the road. The closed road was less of a barrier: 2 of 5 flocks used both sides of the closed road. Playback experiments showed that flocks readily crossed the closed road to approach agonistic vocalizations. Along the open road, even though birds responded to playback by becoming agitated and moving to the extreme edge of the roadside vegetation, they were less likely to cross the road and did so only after a longer duration of playback. Our results suggest that flocks respond to a road as they would to a long linear gap. They use the vegetation along the edges of the road, but because they are unwilling to cross the open area, it becomes a flock territory boundary. Similarly, as in forest gaps, successional change along the closed road produced suitable habitat for flocks. Although this suggests that roads are a trivial problem, we caution that this result applies only to narrow roads that are not accompanied by deforestation or other disturbance.     

Platinum pollution in road dusts,roadside soils,and tree barks in Seoul,Korea

This study presents the level of platinum in urban environment in and around Seoul, the capital city of Korea. Road dust, roadside soil, and tree bark samples were collected from the sites of various traffic volumes and from control sites in the suburbs. The above samples were analyzed for Pt by ICP-MS and other heavy metals by ICP-OES. Platinum levels in road dusts and roadside soils from Seoul were in the range of 3.8–444 ng/g (av. 115.0 ng/g) and 0.7–221 ng/g (av. 49.7 ng/g), respectively, whereas those in the suburbs were in the range of 2.3–5.2 ng/g (av. 3.9 ng/g) in road dusts and 0.4–5.1 ng/g (av. 2.4 ng/g) in roadside soils. The highest Pt levels in road dusts were found from major roads with high traffic volume. The remarkable difference in average Pt level between heavy traffic roads (av. 132.2 ng/g) and light traffic roads (av. 22.8 ng/g) reflects that an important source of Pt in roadside environment is automobile catalytic converter. High Pt level in road dust was found from the site of erratic stop–start driving condition, for example, 178 ng/g Pt in road dust around a vehicle crossing gate. Platinum level in tree barks ranged from 0.9 to 4.5 ng/g, which indicates the existence of Pt-containing particulate matter in the atmosphere. Road dusts with high Pt level were enriched in traffic-related heavy metals.     

Permeability of Roads to Movement of Scrubland Lizards and Small Mammals

A primary objective of road ecology is to understand and predict how roads affect connectivity of wildlife populations. Road avoidance behavior can fragment populations, whereas lack of road avoidance can result in high mortality due to wildlife‐vehicle collisions. Many small animal species focus their activities to particular microhabitats within their larger habitat. We sought to assess how different types of roads affect the movement of small vertebrates and to explore whether responses to roads may be predictable on the basis of animal life history or microhabitat preferences preferences. We tracked the movements of fluorescently marked animals at 24 sites distributed among 3 road types: low‐use dirt, low‐use secondary paved, and rural 2‐lane highway. Most data we collected were on the San Diego pocket mouse (Chaetodipus fallax), cactus mouse (Peromyscus eremicus), western fence lizard (Sceloporus occidentalis), orange‐throated whiptail (Aspidoscelis hyperythra), Dulzura kangaroo rat (Dipodomys simulans) (dirt, secondary paved), and deer mouse (Peromyscus maniculatus) (highway only). San Diego pocket mice and cactus mice moved onto dirt roads but not onto a low‐use paved road of similar width or onto the highway, indicating they avoid paved road substrate. Both lizard species moved onto the dirt and secondary paved roads but avoided the rural 2‐lane rural highway, indicating they may avoid noise, vibration, or visual disturbance from a steady flow of traffic. Kangaroo rats did not avoid the dirt or secondary paved roads. Overall, dirt and secondary roads were more permeable to species that prefer to forage or bask in open areas of their habitat, rather than under the cover of rocks or shrubs. However, all study species avoided the rural 2‐lane highway. Our results suggest that microhabitat use preferences and road substrate help predict species responses to low‐use roads, but roads with heavy traffic may deter movement of a much wider range of small animal species.     

Covariates affecting spatial variability in bison travel behavior in Yellowstone National Park.

Understanding mechanisms influencing the movement paths of animals is essential for comprehending behavior and accurately predicting use of travel corridors. In Yellowstone National Park (USA), the effects of roads and winter road grooming on bison (Bison bison) travel routes and spatial dynamics have been debated for more than a decade. However, no rigorous studies have been conducted on bison spatial movement patterns. We collected 121 380 locations from 14 female bison with GPS collars in central Yellowstone to examine how topography, habitat type, roads, and elevation affected the probability of bison travel year-round. We also conducted daily winter bison road use surveys (2003-2005) to quantify how topography and habitat type influenced spatial variability in the amount of bison road travel. Using model comparison techniques, we found the probability of bison travel and spatial distribution of travel locations were affected by multiple topographic and habitat type attributes including slope, landscape roughness, habitat type, elevation, and distances to streams, foraging areas, forested habitats, and roads. Streams were the most influential natural landscape feature affecting bison travel, and results suggest the bison travel network throughout central Yellowstone is spatially defined largely by the presence of streams that connect foraging areas. Also, the probability of bison travel was higher in regions of variable topography that constrain movements, such as in canyons. Pronounced travel corridors existed both in close association with roads and distant from any roads, and results indicate that roads may facilitate bison travel in certain areas. However, our findings suggest that many road segments used as travel corridors are overlaid upon natural travel pathways because road segments receiving high amounts of bison travel had similar landscape features as natural travel corridors. We suggest that most spatial patterns in bison road travel are a manifestation of general spatial travel trends. Our research offers novel insights into bison spatial dynamics and provides conceptual and analytical frameworks for examining movement patterns of other species.     

Estimate of the Area Affected Ecologically by the Road System in the United States

Abstract: In view of an extensive road system, abundant and rapidly growing vehicular traffic, and a scattered literature indicating that some ecological effects of roads extend outward for > 100 m, it seems likely that the cumulative ecological effect of the road system in the United States is considerable. Two recent studies in The Netherlands and Massachusetts ( U.S.A.) evaluated several ecological effects of roads, including traffic noise effects, and provide quantitative evidence for a definable "road-effect zone." Based on the approximate width of this asymmetric convoluted zone, I estimate that about one-fifth of the U.S. land area is directly affected ecologically by the system of public roads. I identify a series of assumptions and variables suggesting that over time this preliminary estimate is more likely to rise than drop. Several transportation planning and policy recommendations, ranging from perforating the road barrier for wildlife crossings to closing certain roads, offer promise for reducing this enormous ecological effect.     

A Resistant-Kernel Model of Connectivity for Amphibians that Breed in Vernal Pools

Abstract:  Pool-breeding amphibian populations operate at multiple scales, from the individual pool to surrounding upland habitat to clusters of pools. When metapopulation dynamics play a role in long-term viability, conservation efforts limited to the protection of individual pools or even pools with associated upland habitat may be ineffective over the long term if connectivity among pools is not maintained. Connectivity becomes especially important and difficult to assess in regions where suburban sprawl is rapidly increasing land development, road density, and traffic rates. We developed a model of connectivity among vernal pools for the four ambystomatid salamanders that occur in Massachusetts and applied it to the nearly 30,000 potential ephemeral wetlands across the state. The model was based on a modification of the kernel estimator (a density estimator commonly used in home range studies) that takes landscape resistance into account. The model was parameterized with empirical migration distances for spotted salamanders ( Ambystoma maculatum ), dispersal distances for marbled salamanders ( A. opacum ), and expert-derived estimates of landscape resistance. The model ranked vernal pools in Massachusetts by local, neighborhood, and regional connectivity and by an integrated measure of connectivity, both statewide and within ecoregions. The most functionally connected pool complexes occurred in southeastern and northeastern Massachusetts, areas with rapidly increasing suburban development. In a sensitivity analysis estimates of pool connectivity were relatively insensitive to uncertainty in parameter estimates, especially at the local and neighborhood scales. Our connectivity model could be used to prioritize conservation efforts for vernal-pool amphibian populations at broader scales than traditional pool-based approaches.     

公路绿化带降噪效应及其影响因素研究

以山西省典型公路绿化带为研究对象,在路侧设置样带分梯度布设噪音测点,研究绿化带对交通噪音的衰减效应、绿化带的噪音衰减格局及绿化带特征因子与降噪效应的关系,并通过拟合回归模型,分析绿化带的降噪机理。结果表明,公路绿化带噪音衰减格局呈单调递减趋势,模型拟合不同水平梯度的噪音衰减效应,以三次多项式拟合度最高;绿化带10m内是最佳降噪范围,10～60m为有效降噪范围,60～80m是次级降噪范围,40～60m宽的绿化带可满足绿化带的降噪要求;合理栽植绿化树种,使其成林后郁闭度≥0.6,可使噪音值从路基处至80m处降低31.0～36.4dB,绿化带树木高大并且栽植乔灌草复层结构,会增大林带的降噪效应;绿化带降噪效应与绿化带宽度、郁闭度、100m绿化带胸高断面积等因子正相关,与疏透度负相关,因子的互作对降噪效应起综合作用。     

Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.