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Effects of nitrogen deposition and empirical nitrogen critical loads for ecoregions of the United States

Human activity in the last century has led to a significant increase in nitrogen (N) emissions and atmospheric deposition. This N deposition has reached a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. One approach for quantifying the deposition of pollution that would be harmful to ecosystems is the determination of critical loads. A critical load is defined as the input of a pollutant below which no detrimental ecological effects occur over the long-term according to present knowledge. The objectives of this project were to synthesize current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and to estimate associated empirical N critical loads. The receptors considered included freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Ecosystem impacts included: (1) biogeochemical responses and (2) individual species, population, and community responses. Biogeochemical responses included increased N mineralization and nitrification (and N availability for plant and microbial uptake), increased gaseous N losses (ammonia volatilization, nitric and nitrous oxide from nitrification and denitrification), and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root : shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species. The range of critical loads for nutrient N reported for U.S. ecoregions, inland surface waters, and freshwater wetlands is 1-39 kg N.ha -1.yr -1, spanning the range of N deposition observed over most of the country. The empirical critical loads for N tend to increase in the following sequence for different life forms: diatoms, lichens and bryophytes, mycorrhizal fungi, herbaceous plants and shrubs, and trees. The critical load approach is an ecosystem assessment tool with great potential to simplify complex scientific information and communicate effectively with the policy community and the public. This synthesis represents the first comprehensive assessment of empirical critical loads of N for major ecoregions across the United States. ?? 2011 by the Ecological Society of America.

Ecological Applications↗

Land cover characterization and land surface parameterization research

The understanding of land surface processes and their parameterization in atmospheric, hydrologic, and ecosystem models has been a dominant research theme over the past decade. For example, many studies have demonstrated the key role of land cover characteristics as controlling factors in determining land surface processes, such as the exchange of water, energy, carbon, and trace gases between the land surface and the lower atmosphere. The requirements for multiresolution land cover characteristics data to support coupled-systems modeling have also been well documented, including the need for data on land cover type, land use, and many seasonally variable land cover characteristics, such as albedo, leaf area index, canopy conductance, surface roughness, and net primary productivity. Recently, the developers of land data have worked more closely with the land surface process modelers in these efforts.

Ecological Applications↗

NDVI, C3 and C4 production, and distributions in Great Plains grassland land cover classes

The distributions of C 3 and C 4 grasses were used to interpret the distribution, seasonal performance, and potential production of grasslands in the Great Plains of North America. Thirteen major grassland seasonal land cover classes were studied with data from three distinct sources. Normalized Difference Vegetation Index (NDVI) data derived from the National Oceanic and Atmospheric Administration (NOAA) Advanced Very High Resolution Radiometer (AVHRR) sensor were collected for each pixel over a 5-yr period (1989–1993), analyzed for quantitative attributes and seasonal relationships, and then aggregated by land cover class. Data from the State Soil Geographic (STATSGO) database were used to identify dominant plant species contributing to the potential production in each map unit. These species were identified as C 3 or C 4 , and contributions to production were aggregated to provide estimates of the percentage of C 3 and C 4 production for each intersection of the STATSGO map units and the seasonal land cover classes. Carbon isotope values were obtained at specific sites from the soil organic matter of the upper horizon of soil cores and were related to STATSGO estimates of potential production. The grassland classes were distributed with broad northwest-to-southeast orientations. Some classes had large variations in C 3 and C 4 composition with high proportions of C 4 species in the south and low proportions in the north. This diversity of photosynthetic types within land cover classes that cross regions of different temperature and precipitation results in similar seasonal patterns and magnitudes of NDVI. The easternmost class, 65, containing tallgrass prairie components, bluestem, Indiangrass, and switchgrass, possessed the highest maximum NDVI and time-integrated NDVI values each year. Grassland classes varied over 5 yr from a high integrated NDVI mean of 4.9 in class 65 in the east to a low of 1.2 in class 76 (sand sage, blue grama, wheatgrass, and buffalograss) in the southwest. Although environmental conditions varied widely during the 5 yr, the rankings of class performance were consistent across years for these NDVI metrics. Land cover classes were less consistent in time of onset, which was often earlier in areas in the north dominated by C 3 grasses than in areas to the south dominated by C 4 grasses. At the level of seasonal land cover classes, no significant relationship was found between the proportions of C 3 and C 4 species and estimates of potential production derived from the STATSGO database or inferred from the seasonal patterns of NDVI. The isotopic data from specific sites and the potential production data from STATSGO suggest similar patterns of high proportional production by C 4 species throughout the south and a decline in proportional production north of the central Great Plains. The land cover classes integrate ecosystem units that encompass a wide diversity of species and C 3 and C 4 proportions and provide a classification that consistently captures significant ecosystem parameters for the Great Plains.

Colorado, Kansas, Montana, Nebraska, New Mexico, N↗

Conditions for the return and simulation of the recovery of burrowing mayflies in western Lake Erie

In the 1950s, burrowing mayflies, Hexagenia spp. ( H. limbata and H. rigida ), were virtually eliminated from the western basin of Lake Erie (a 3300 km 2 area) because of eutrophication and pollution. We develop and present a deterministic model for the recolonization of the western basin by Hexagenia to pre-1953 densities. The model was based on the logistic equation describing the population growth of Hexagenia and a presumed competitor, Chironomus (dipteran larvae). Other parameters (immigration, low oxygen, toxic sediments, competition with Chironomus, and fish predation) were then individually added to the logistic model to determine their effect at different growth rates. The logistic model alone predicts 10–41 yr for Hexagenia to recolonize western Lake Erie. Immigration reduced the recolonization time by 2–17 yr. One low-oxygen event during the first 20 yr increased recovery time by 5–17 yr. Contaminated sediments added 5–11 yr to the recolonization time. Competition with Chironomus added 8–19 yr to recovery. Fish predators added 4–47 yr to the time required for recolonization. The full model predicted 48–81 yr for Hexagenia to reach a carrying capacity of ≈350 nymphs/m 2 , or not until around the year 2038 if the model is started in 1990. The model was verified by changing model parameters to those present in 1970, beginning the model in 1970 and running it through 1990. Predicted densities overlapped almost completely with actual estimated densities of Hexagenia nymphs present in the western basin in Lake Erie in 1990. The model suggests that recovery of large aquatic ecosystems may lag substantially behind remediation efforts.

Michigan, Ohio, Ontario↗

Flood dependency of cottonwood establishment along the Missouri River, Montana, USA

Flow variability plays a central role in structuring the physical environment of riverine ecosystems. However, natural variability in flows along many rivers has been modified by water management activities. We quantified the relationship between flow and establishment of the dominant tree (plains cottonwood, Populus deltoides subsp. monilifera ) along one of the least hydrologically altered alluvial reaches of the Missouri River: Coal Banks Landing to Landusky, Montana. Our purpose was to refine our understanding of how local fluvial geomorphic processes condition the relationship between flow regime and cottonwood recruitment. We determined date and elevation of tree establishment and related this information to historical peak stage and discharge over a 112-yr hydrologic record. Of the excavated trees, 72% were established in the year of a flow >1400 m 3 /s (recurrence interval of 9.3 yr) or in the following 2 yr. Flows of this magnitude or greater create the necessary bare, moist establishment sites at an elevation high enough to allow cottonwoods to survive subsequent floods and ice jams. Almost all cottonwoods that have survived the most recent flood (1978) were established >1.2 m above the lower limit of perennial vegetation (active channel shelf). Most younger individuals were established between 0 and 1.2 m, and are unlikely to survive over the long term. Protection of riparian cottonwood forest along this National Wild and Scenic section of the Missouri River depends upon maintaining the historical magnitude, frequency, and duration of floods >1400 m 3 /s. Here, a relatively narrow valley constrains lateral channel movement that could otherwise facilitate cottonwood recruitment at lower flows. Effective management of flows to promote or maintain cottonwood recruitment requires an understanding of locally dominant fluvial geomorphic processes.

Montana↗

Modeling changes in growth and diet on polychlorinated biphenyl bioaccumulation in Coregonus hoyi

Restrictions on the release of PCBs into the environment began in the early 1970s, resulting in reduced PCB concentrations in Lake Michigan bloaters ( Coregonus hoyi ). However, since 1980, PCB concentrations in bloater have declined only slightly. The bloater population also increased 40-fold during 1970 through 1984, resulting in a diet shift and a density-dependent decline in growth. Our goal was to determine how these changes in diet and growth may have affected PCB accumulation in Lake Michigan bloater. We evaluated the consequences of these two changes on bloater PCB concentrations by developing a bioenergetics-based PCB bioaccumulation model. Bloater PCB concentrations increased little when we increased the amount of Diporeia hoyi, the most contaminated prey, in the bloater diet. By comparing constant growth (similar to those of the 1970s) to observed decline in growth rates, we found that lower growth rates during the 1980s placed older, more contaminated bloaters in the size range most vulnerable to predators and in the size range sampled by PCB monitoring programs. Bloater PCB concentration trends may be influenced by the sampling methodology of the United States Geological Survey. If fish PCB trends are to be used as an indicator of system-level PCB trends, sampling should include a representative sample of fish of known age from the population.

Ecological Applications↗

Biogeochemical effects of seawater restoration to diked salt marshes

We conducted greenhouse microcosm experiments to examine the biogeochemical effects of restoring seawater to historically diked Cape Cod salt marshes. Peat cores from both seasonally flooded and drained diked marshes were waterlogged with seawater, and porewater chemistry was subsequently monitored for 21 mo. The addition of seawater to highly organic, seasonally flooded peat caused the death of freshwater wetland plants, 6–8 cm of sediment subsidence, and increased N and P mineralization. Also, sulfides and alkalinity increased 10-fold, suggesting accelerated decomposition by sulfate reduction. Addition of seawater to the low-organic-content acidic peat from the drained marsh increased porewater pH, alkalinity, PO 4 -P, and Fe(II), which we attribute to the reestablishment of SO 4 and Fe(III) mineral reduction. Increased cation exchange contributed to 6-fold increases in dissolved Fe(II) and Al and 60-fold increases in NH 4 -N within 6 mo of salination. Seawater reintroductions to seasonally flooded diked marshes will cause porewater sulfides to increase, likely reducing the success of revegetation efforts. Sulfide toxicity is of less concern in resalinated drained peats because of the abundance of Fe(II) to precipitate sulfides, and of NH 4 -N to offset sulfide inhibition of N uptake. Restoration of either seasonally flooded or drained diked marshes could stimulate potentially large nutrient and Fe(II) releases, which could in turn increase primary production and lower oxygen in receiving waters. These findings suggest that tidal restoration be gradual and carefully monitored.

Massachusetts↗

Multiscale sampling of plant diversity: Effects of minimum mapping unit size

Only a small portion of any landscape can be sampled for vascular plant diversity because of constraints of cost (salaries, travel time between sites, etc.). Often, the investigator decides to reduce the cost of creating a vegetation map by increasing the minimum mapping unit (MMU), and/or by reducing the number of vegetation classes to be considered. Questions arise about what information is sacrificed when map resolution is decreased. We compared plant diversity patterns from vegetation maps made with 100-ha, 50-ha, 2-ha, and 0.02-ha MMUs in a 754-ha study area in Rocky Mountain National Park, Colorado, United States, using four 0.025-ha and 21 0.1-ha multiscale vegetation plots. We developed and tested species–log(area) curves, correcting the curves for within-vegetation type heterogeneity with Jaccard’s coefficients. Total species richness in the study area was estimated from vegetation maps at each resolution (MMU), based on the corrected species–area curves, total area of the vegetation type, and species overlap among vegetation types. With the 0.02-ha MMU, six vegetation types were recovered, resulting in an estimated 552 species (95% ci = 520–583 species) in the 754-ha study area (330 plant species were observed in the 25 plots). With the 2-ha MMU, five vegetation types were recognized, resulting in an estimated 473 species for the study area. With the 50-ha MMU, 439 plant species were estimated for the four vegetation types recognized in the study area. With the 100-ha MMU, only three vegetation types were recognized, resulting in an estimated 341 plant species for the study area. Locally rare species and keystone ecosystems (areas of high or unique plant diversity) were missed at the 2-ha, 50-ha, and 100-ha scales. To evaluate the effects of minimum mapping unit size requires: (1) an initial stratification of homogeneous, heterogeneous, and rare habitat types; and (2) an evaluation of within-type and between-type heterogeneity generated by environmental gradients and other factors. We suggest that at least some portions of vegetation maps created at a coarser level of resolution be validated at a higher level of resolution.

Ecological Applications↗

Effects of ultraviolet radiation on boreal toads in Colorado

Field exposures of Bufo boreas embryos to fractions of ambient UV-B radiation at two sites in Rocky Mountain National Park, Colorado, USA, were conducted to evaluate UV-B as a possible cause of recent severe declines of this species. There were no differences in hatching success of B. boreas embryos exposed to 0–100% of ambient UV-B radiation at either study site, results that are different from those of recent studies in Oregon that found increased mortality of B. boreas embryos exposed to ambient UV-B. The reasons for these differing results are not apparent, and several possible explanations exist, including differences in experimental design, presence or absence of a pathogenic fungus, and geographic genetic variation. Bufo boreas embryos were probably not receiving higher doses of UV-B radiation during the experiments in Oregon compared to the experiments in this study. Results of this study do not support UV-B radiation alone as the cause of the decline of B. boreas during the past 20 yr in the southern Rocky Mountains, but UV-B cannot be dismissed because of the contradictory results from other studies.

Colorado↗

Nitrogen excess in North American ecosystems: Predisposing factors, ecosystem responses, and management strategies

Most forests in North America remain nitrogen limited, although recent studies have identified forested areas that exhibit symptoms of N excess, analogous to overfertilization of arable land. Nitrogen excess in watersheds is detrimental because of disruptions in plant/soil nutrient relations, increased soil acidification and aluminum mobility, increased emissions of nitrogenous greenhouse gases from soil, reduced methane consumption in soil, decreased water quality, toxic effects on freshwater biota, and eutrophication of coastal marine waters. Elevated nitrate ( NO 3 − ) loss to groundwater or surface waters is the primary symptom of N excess. Additional symptoms include increasing N concentrations and higher N:nutrient ratios in foliage (i.e., N:Mg, N:P), foliar accumulation of amino acids or NO 3 − , and low soil C:N ratios. Recent nitrogen-fertilization studies in New England and Europe provide preliminary evidence that some forests receiving chronic N inputs may decline in productivity and experience greater mortality. Long-term fertilization at Mount Ascutney, Vermont, suggests that declining and slow N-cycling coniferous stands may be replaced by fast-growing and fast N-cycling deciduous forests. Symptoms of N saturation are particularly severe in high-elevation, nonaggrading spruce–fir ecosystems in the Appalachian Mountains and in eastern hardwood watersheds at the Fernow Experimental Forest near Parsons, West Virginia. In the Los Angeles Air Basin, mixed conifer forests and chaparral watersheds with high smog exposure are N saturated and exhibit the highest streamwater NO 3 − concentrations for wildlands in North America. High-elevation alpine watersheds in the Colorado Front Range and a deciduous forest in Ontario, Canada, are N saturated, although N deposition is moderate (∼8 kg·ha −1 ·yr −1 ). In contrast, the Harvard Forest hardwood stand in Massachusetts has absorbed >900 kg N/ha during 8 yr of N amendment studies without significant NO 3 − leaching, illustrating that ecosystems vary widely in the capacity to retain N inputs. Overly mature forests with high N deposition, high soil N stores, and low soil C:N ratios are prone to N saturation and NO 3 − leaching. Additional characteristics favoring low N retention capacity include a short growing season (reduced plant N demand) and reduced contact time between drainage water and soil (i.e., porous coarse-textured soils, exposed bedrock or talus). Temporal patterns of hydrologic fluxes interact with biotic uptake and internal cycling patterns in determining ecosystem N retention. Soils are the largest storage pool for N inputs, although vegetation uptake is also important. Recent studies indicate that nitrification may be widespread in undisturbed ecosystems, and that microbial assimilation of NO 3 − may be a significant N retention mechanism, contrary to previous assumptions. Further studies are needed to elucidate the sites, forms, and mechanisms of N retention and incorporation into soil organic matter, and to test potential management options for mitigating N losses from forests. Implementation of intensive management practices in N-saturated ecosystems may only be feasible in high-priority areas and on a limited scale. Reduction of N emissions would be a preferable solution, although major reductions in the near future are unlikely in many areas due to economic, energy-use, policy, and demographic considerations.

Ecological Applications↗

Assessing simulated ecosystem processes for climate variability research at Glacier National Park, USA

Glacier National Park served as a test site for ecosystem analyses that involved a suite of integrated models embedded within a geographic information system. The goal of the exercise was to provide managers with maps that could illustrate probable shifts in vegetation, net primary production (NPP), and hydrologic responses associated with two selected climatic scenarios. The climatic scenarios were (a) a recent 12-yr record of weather data, and (b) a reconstituted set that sequentially introduced in repeated 3-yr intervals wetter–cooler, drier–warmer, and typical conditions. To extrapolate the implications of changes in ecosystem processes and resulting growth and distribution of vegetation and snowpack, the model incorporated geographic data. With underlying digital elevation maps, soil depth and texture, extrapolated climate, and current information on vegetation types and satellite-derived estimates of leaf area indices, simulations were extended to envision how the park might look after 120 yr. The predictions of change included underlying processes affecting the availability of water and nitrogen. Considerable field data were acquired to compare with model predictions under current climatic conditions. In general, the integrated landscape models of ecosystem processes had good agreement with measured NPP, snowpack, and streamflow, but the exercise revealed the difficulty and necessity of averaging point measurements across landscapes to achieve comparable results with modeled values. Under the extremely variable climate scenario significant changes in vegetation composition and growth as well as hydrologic responses were predicted across the park. In particular, a general rise in both the upper and lower limits of treeline was predicted. These shifts would probably occur along with a variety of disturbances (fire, insect, and disease outbreaks) as predictions of physiological stress (water, nutrients, light) altered competitive relations and hydrologic responses. The use of integrated landscape models applied in this exercise should provide managers with insights into the underlying processes important in maintaining community structure, and at the same time, locate where changes on the landscape are most likely to occur.

Montana↗

Human influences on trophic cascades along rocky shores

A three-trophic-level interaction among American Black Oystercatchers ( Haematopus bachmani ), limpets ( Lottia spp.), and erect fleshy algae in rocky intertidal communities of central and southern California was documented via manipulative and “natural” experiments. Removal of the territorial limpet ( Lottia gigantea ) initially caused large increases in the percent cover of erect fleshy algae, followed by a more gradual increase in density of small limpets ( Lottia spp.) and a decline in algal cover. Algal cover increased following the removal of small limpets at the sites from which L. gigantea had been removed earlier, thus demonstrating that the large and small limpets had similar inhibitory effects on plant populations. A comparison of sites with and without oystercatchers showed that L. gigantea occupied substrate inclinations in proportion to their availability at sites where oystercatchers were rare, whereas the distribution of L. gigantea was skewed toward vertically inclined substrates where oystercatchers were common. Survival rates of limpets translocated to horizontal and vertical substrates were similar in sites lacking oystercatcher predation, but were much lower on horizontal substrates where oystercatchers were common. Our results are consistent with those from several prior studies in demonstrating that shorelines frequented by humans typically lack oystercatchers. Humans also exploit L. gigantea and reduce populations to low densities of small individuals. These findings may explain why the midlittoral zone of rocky intertidal communities in western North America are so often dominated by high population densities of small limpets.

Ecological Applications↗

Effects of landcover, water redistribution, and temperature on ecosystem processes in the South Plate Basin

Over one-third of the land area in the South Platte Basin of Colorado, Nebraska, and Wyoming, has been converted to croplands. Irrigated cropland now comprises 8% of the basin, while dry croplands make up 31%. We used the RHESSys model to compare the changes in plant productivity and vegetation-related hydrological processes that occurred as a result of either land cover alteration or directional temperature changes (&minus;2&deg;C, +4&deg;C). Land cover change exerted more control over annual plant productivity and water fluxes for converted grasslands, while the effect of temperature changes on productivity and water fluxes was stronger in the mountain vegetation. Throughout the basin, land cover change increased the annual loss of water to the atmosphere by 114 mm via evaporation and transpiration, an increase of 37%. Both irrigated and nonirrigated grains became active earlier in the year than shortgrass steppe, leading to a seasonal shift in water losses to the atmosphere. Basin-wide photosynthesis increased by 80% due to grain production. In contrast, a 4&deg;C warming scenario caused annual transpiration to increase by only 3% and annual evaporation to increase by 28%, for a total increase of 71 mm. Warming decreased basin-wide photosynthesis by 16%. There is a large elevational range from east to west in the South Platte Basin, which encompasses the western edge of the Great Plains and the eastern front of the Rocky Mountains. This elevational gain is accompanied by great changes in topographic complexity, vegetation type, and climate. Shortgrass steppe and crops found at elevations between 850 and 1800 m give way to coniferous forests and tundra between 1800 and 4000 m. Climate is increasingly dominated by winter snow precipitation with increasing elevation, and the timing of snowmelt influences tundra and forest ecosystem productivity, soil moisture, and downstream discharge. Mean annual precipitation of <500 mm on the plains below 1800 m is far less than potential evapotranspiration of 1000&ndash;1500 mm and is insufficient for optimum plant productivity. The changes in water flux and photosynthesis from conversion of steppe to cropland are the result of redistribution of snowmelt water from the mountains and groundwater pumping through irrigation projects.

Ecological Applications↗

Estimating rates of local species extinction, colonization and turnover in animal communities

Species richness has been identified as a useful state variable for conservation and management purposes. Changes in richness over time provide a basis for predicting and evaluating community responses to management, to natural disturbance, and to changes in factors such as community composition (e.g., the removal of a keystone species). Probabilistic capture-recapture models have been used recently to estimate species richness from species count and presence-absence data. These models do not require the common assumption that all species are detected in sampling efforts. We extend this approach to the development of estimators useful for studying the vital rates responsible for changes in animal communities over time; rates of local species extinction, turnover, and colonization. Our approach to estimation is based on capture-recapture models for closed animal populations that permit heterogeneity in detection probabilities among the different species in the sampled community. We have developed a computer program, COMDYN, to compute many of these estimators and associated bootstrap variances. Analyses using data from the North American Breeding Bird Survey (BBS) suggested that the estimators performed reasonably well. We recommend estimators based on probabilistic modeling for future work on community responses to management efforts as well as on basic questions about community dynamics.

Ecological Applications↗

Estimating the effectiveness of further sampling inspecies inventories

Estimators of the number of additional species expected in the next Δ n samples offer a potentially important tool for improving cost-effectiveness of species inventories but are largely untested. We used Monte Carlo methods to compare 11 such estimators, across a range of community structures and sampling regimes, and validated our results, where possible, using empirical data from vascular plant and beetle inventories from Glacier National Park, Montana, USA. We found that B. Efron and R. Thisted’s 1976 negative binomial estimator was most robust to differences in community structure and that it was among the most accurate estimators when sampling was from model communities with structures resembling the large, heterogeneous communities that are the likely targets of major inventory efforts. Other estimators may be preferred under specific conditions, however. For example, when sampling was from model communities with highly even species-abundance distributions, estimates based on the Michaelis-Menten model were most accurate; when sampling was from moderately even model communities with S = 10 species or communities with highly uneven species-abundance distributions, estimates based on Gleason’s (1922) species–area model were most accurate. We suggest that use of such methods in species inventories can help improve cost-effectiveness by providing an objective basis for redirecting sampling to more-productive sites, methods, or time periods as the expectation of detecting additional species becomes unacceptably low.

Ecological Applications↗

Of elephants and blind men: Deer management in the U.S. National Parks

Overabundant populations of white-tailed deer ( Odocoileus virginianus ) are becoming common in the eastern United States. Faced with burgeoning deer populations in eastern parks, the National Park Service (NPS) formulated policy based on its long experience with ungulate management in western parks. That the NPS failed to find a management solution acceptable to its many constituencies was inevitable. Like blind men touching different parts of an elephant and disagreeing about its form, those engaged in the debate about deer management in parks are viewing different parts of the ecological system. None has seen the entire system, and consequently, there is neither common agreement on the nature of the problem nor on the solutions. We explore the quandary of deer management in eastern parks by addressing three questions: (1) Can the National Park Service reconcile its management goals with those of its neighbors? (2) Can thresholds be identified for determining when to intervene in natural processes? (3) Is there a scientific foundation for proceeding with effective management of deer? We argue that reconciling the NPS management with that of state conservation agencies is not possible because management policy guides these agencies in opposite directions: the NPS is charged with limiting human impact on ecological processes, and state agencies are charged with exerting human control over population abundance. Questions about thresholds and a scientific basis for management arise from concern that irrupting deer populations are a manifestation of disrupted natural processes. Several population growth paradigms are at the heart of this ecological question. The science provides no consensus about which of these paradigms are appropriate to deer in eastern ecosystems. Thus, it is premature to expect science to identify if or when natural processes have been disrupted. While the NPS cannot effectively achieve its goals without better science, neither can it wait for science to fully understand the dynamics of plant–herbivore interactions. The best hope for resolving both the biological and political dilemmas surrounding deer management is through an adaptive management approach.

Ecological Applications↗

How grazing and soil quality affect native and exotic plant diversity in Rocky Mountain grasslands

We used multiscale plots to sample vascular plant diversity and soil characteristics in and adjacent to 26 long-term grazing exclosure sites in Colorado, Wyoming, Montana, and South Dakota, USA. The exclosures were 7–60 yr old (31.2 ± 2.5 yr, mean ± 1 se ). Plots were also randomly placed in the broader landscape in open rangeland in the same vegetation type at each site to assess spatial variation in grazed landscapes. Consistent sampling in the nine National Parks, Wildlife Refuges, and other management units yielded data from 78 1000-m 2 plots and 780 1-m 2 subplots. We hypothesized that native species richness would be lower in the exclosures than in grazed sites, due to competitive exclusion in the absence of grazing. We also hypothesized that grazed sites would have higher native and exotic species richness compared to ungrazed areas, due to disturbance (i.e., the intermediate-disturbance hypothesis) and the conventional wisdom that grazing may accelerate weed invasion. Both hypotheses were soundly rejected. Although native species richness in 1-m 2 subplots was significantly higher ( P < 0.05) in grazed sites, we found nearly identical native or exotic species richness in 1000-m 2 plots in exclosures (31.5 ± 2.5 native and 3.1 ± 0.5 exotic species), adjacent grazed plots (32.6 ± 2.8 native and 3.2 ± 0.6 exotic species), and randomly selected grazed plots (31.6 ± 2.9 native and 3.2 ± 0.6 exotic species). We found no significant differences in species diversity (Hill’s diversity indices, N 1 and N 2), evenness (Hill’s ratio of evenness, E 5), cover of various life-forms (grasses, forbs, and shrubs), soil texture, or soil percentage of N and C between grazed and ungrazed sites at the 1000-m 2 plot scale. The species lists of the long-ungrazed and adjacent grazed plots overlapped just 57.9 ± 2.8%. This difference in species composition is commonly attributed solely to the difference in grazing regimes. However, the species lists between pairs of grazed plots (adjacent and distant 1000-m 2 plots) in the same vegetation type overlapped just 48.6 ± 3.6%, and the ungrazed plots and distant grazed plots overlapped 49.4 ± 3.6%. Differences in vegetation and soils between grazed and ungrazed sites were minimal in most cases, but soil characteristics and elevation were strongly correlated with native and exotic plant diversity in the study region. For the 78 1000-m 2 plots, 59.4% of the variance in total species richness was explained by percentage of silt (coefficient = 0.647, t = 5.107, P < 0.001), elevation (coefficient = 0.012, t = 5.084, P < 0.001), and total foliar cover (coefficient = 0.110, t = 2.104, P < 0.039). Only 12.8% of the variance in exotic species cover (log 10 cover) was explained by percentage of clay (coefficient = −0.011, t = −2.878, P < 0.005), native species richness (coefficient = −0.011, t = −2.156, P < 0.034), and log 10 N (coefficient = 2.827, t = 1.860, P < 0.067). Native species cover and exotic species richness and frequency were also significantly positively correlated with percentage of soil N at the 1000-m 2 plot scale. Our research led to five broad generalizations about current levels of grazing in these Rocky Mountain grasslands: (1) grazing probably has little effect on native species richness at landscape scales; (2) grazing probably has little effect on the accelerated spread of most exotic plant species at landscape scales; (3) grazing affects local plant species and life-form composition and cover, but spatial variation is considerable; (4) soil characteristics, climate, anddisturbances may have a greater effect on plant species diversity than do current levels of grazing; and (5) few plant species show consistent, directional responses to grazing or cessation of grazing.

Ecological Applications↗

A GIS modeling method applied to predicting forest songbird habitat

We have developed an approach for using “presence” data to construct habitat models. Presence data are those that indicate locations where the target organism is observed to occur, but that cannot be used to define locations where the organism does not occur. Surveys of highly mobile vertebrates often yield these kinds of data. Models developed through our approach yield predictions of the amount and the spatial distribution of good-quality habitat for the target species. This approach was developed primarily for use in a GIS context; thus, the models are spatially explicit and have the potential to be applied over large areas. Our method consists of two primary steps. In the first step, we identify an optimal range of values for each habitat variable to be used as a predictor in the model. To find these ranges, we employ the concept of maximizing the difference between cumulative distribution functions of (1) the values of a habitat variable at the observed presence locations of the target organism, and (2) the values of that habitat variable for all locations across a study area. In the second step, multivariate models of good habitat are constructed by combining these ranges of values, using the Boolean operators “and” and “or.” We use an approach similar to forward stepwise regression to select the best overall model. We demonstrate the use of this method by developing species-specific habitat models for nine forest-breeding songbirds (e.g., Cerulean Warbler, Scarlet Tanager, Wood Thrush) studied in southern Ohio. These models are based on species’ microhabitat preferences for moisture and vegetation characteristics that can be predicted primarily through the use of abiotic variables. We use slope, land surface morphology, land surface curvature, water flow accumulation downhill, and an integrated moisture index, in conjunction with a land-cover classification that identifies forest/nonforest, to develop these models. The performance of these models was evaluated with an independent data set. Our tests showed that the models performed better than random at identifying where the birds occurred and provided useful information for predicting the amount and spatial distribution of good habitat for the birds we studied. In addition, we generally found positive correlations between the amount of habitat, as predicted by the models, and the number of territories within a given area. This added component provides the possibility, ultimately, of being able to estimate population sizes. Our models represent useful tools for resource managers who are interested in assessing the impacts of alternative management plans that could alter or remove habitat for these birds.

Ecological Applications↗