Pulsed seaweed subsidies drive sequential shifts in the effects of lizard predators on island food webs

Most prominent theories of food web dynamics imply the simultaneous action of bottom–up and top–down forces. However, transient bottom‐up effects resulting from resource pulses can lead to sequential shifts in the strength of top–down predator effects. We used a large‐scale field experiment (32 small islands sampled over 5 years) to probe how the frequency and magnitude of pulsed seaweed inputs drives temporal variation in the top–down effects of lizard predators. Short‐term weakening of lizard effects on spiders and plants (the latter via a trophic cascade) were associated with lizard diet shifts, and were more pronounced with larger seaweed inputs. Long‐term strengthening of lizard effects was associated with lizard numerical responses and plant fertilisation. Increased pulse frequency reinforced the strengthening of lizard effects on spiders and plants. These results underscore the temporally variable nature of top–down effects and highlight the role of resource pulses in driving this variation.     

Pulses of marine subsidies amplify reproductive potential of lizards by increasing individual growth rate

Pulsed resource subsidies can have profound effects on recipient communities. The effects of resource pulses are often mediated by increases in the density of consumer populations. Here we investigate several mechanisms linking experimental pulses of seaweed deposition to population‐level responses in the brown anole Anolis sagrei. Subsidized lizards grew approximately 30% faster than lizards in seaweed‐removal plots, but there was no effect of seaweed subsidies on survival or body condition. Breeding is strongly seasonal in A. sagrei, resulting in a limited reproductive window of opportunity. Accelerated growth allows subsidized females to reach sexual maturity earlier and thereby exploit more of this window, which is projected to double fecundity in their first year of life. These results show how changes in an individual trait can translate pulses of resource input into reproductive output. Further, they highlight the importance of seasonal timing in mechanistically linking individual‐, population‐ and community‐level responses to pulsed resource subsidies.     

Nutrient presses and pulses differentially impact plants, herbivores, detritivores and their natural enemies

Anthropogenic nutrient inputs into native ecosystems cause fluctuations in resources that normally limit plant growth, which has important consequences for associated food webs. Such inputs from agricultural and urban habitats into nearby natural systems are increasing globally and can be highly variable, spanning the range from sporadic to continuous. Despite the global increase in anthropogenically-derived nutrient inputs into native ecosystems, the consequences of variation in subsidy duration on native plants and their associated food webs are poorly known. Specifically, while some studies have examined the effects of nutrient subsidies on native ecosystems for a single year (a nutrient pulse), repeated introductions of nutrients across multiple years (a nutrient press) better reflect the persistent nature of anthropogenic nutrient enrichment. We therefore contrasted the effects of a one-year nutrient pulse with a four-year nutrient press on arthropod consumers in two salt marshes. Salt marshes represent an ideal system to address the differential impacts of nutrient pulses and presses on ecosystem and community dynamics because human development and other anthropogenic activities lead to recurrent introductions of nutrients into these natural systems. We found that plant biomass and %N as well as arthropod density fell after the nutrient pulse ended but remained elevated throughout the nutrient press. Notably, higher trophic levels responded more strongly than lower trophic levels to fertilization, and the predator/prey ratio increased each year of the nutrient press, demonstrating that food web responses to anthropogenic nutrient enrichment can take years to fully manifest themselves. Vegetation at the two marshes also exhibited an apparent tradeoff between increasing %N and biomass in response to fertilization. Our research emphasizes the need for long-term, spatially diverse studies of nutrient enrichment in order to understand how variation in the duration of anthropogenic nutrient subsidies affects native ecosystems.     

Marine subsidies change short‐term foraging activity and habitat utilization of terrestrial lizards

Resource pulses are brief periods of unusually high resource abundance. While population and community responses to resource pulses have been relatively well studied, how individual consumers respond to resource pulses has received less attention. Local consumers are often the first to respond to a resource pulse, and the form and timing of individual responses may influence how the effects of the pulse are transmitted throughout the community. Previous studies in Bahamian food webs have shown that detritivores associated with pulses of seaweed wrack provide an alternative prey source for lizards. When seaweed is abundant, lizards (Anolis sagrei) shift to consuming more marine‐derived prey and increase in density, which has important consequences for other components of the food web. We hypothesized that the diet shift requires individuals to alter their habitat use and foraging activity and that such responses may happen very rapidly. In this study, we used recorded video observations to investigate the immediate responses of lizards to an experimental seaweed pulse. We added seaweed to five treatment plots for comparison with five control plots. Immediately after seaweed addition, lizards decreased average perch height and increased movement rate, but these effects persisted for only 2 days. To explore the short‐term nature of the response, we used our field data to parametrize heuristic Markov chain models of perch height as a function of foraging state. These models suggest a “Synchronized‐satiation Hypothesis,” whereby lizards respond synchronously and feed quickly to satiation in the presence of a subsidy (causing an initial decrease in average perch height) and then return to the relative safety of higher perches. We suggest that the immediate responses of individual consumers to resource pulse events can provide insight into the mechanisms by which these consumers ultimately influence community‐level processes.     

Root fungal symbionts interact with mammalian herbivory, soil nutrient availability and specific habitat conditions

Herbivory, competition and soil fertility interactively shape plant communities and exhibit an important role in modifying conditions for host-dependent fungal symbionts. However, field studies on the combined impacts of natural herbivory, competition and soil fertility on root fungal symbionts are rare. We asked how mammalian herbivory, fertilization, liming and plant–plant competition affect the root colonization of arbuscular mycorrhizal fungi (AMF) and dark septate endophytic (DSE) fungi of the dicot herb, Solidago virgaurea. The 2-year full-factorial experiment was conducted in two contrasting habitats: non-acidic and acidic mountain tundra. We found that herbivory increased arbuscular colonization (i.e. the site of resource exchange) at fertile non-acidic sites, where vegetation was rich in species having AMF symbionts, whereas at infertile acidic sites, where plants having AMF symbiont are scarce, the response was the opposite. Herbivory of the host plant negatively affected DSE hyphal and sclerotial colonization in unfertilized plots, possibly due to reduced carbon flow from the host plant while there was no effect of herbivory in fertilized plots. DSE colonization was highest in unfertilized exclosures where soil nutrient concentrations were also lowest. Liming had a negative effect on DSE hyphal colonization, and its effect also interacted with herbivory and the habitat. Biomass removal of the neighboring plants did not affect the root colonization percent of either arbuscules or DSE. Our results show that the impacts of aboveground mammalian herbivory, soil nutrient availability and specific habitat conditions on belowground root fungal symbionts are highly dependent on each other. Arbuscule response to herbivory appeared to be regulated by specific habitat conditions possibly caused by differences in the AMF availability in the soil while DSE response was associated with availability of host-derived carbon. Our result of the relationship between herbivory and soil nutrients suggests an important role of DSE in ecosystem processes.     

Location Is Everything: Evaluating the Effects of Terrestrial and Marine Resource Subsidies on an Estuarine Bivalve

Estuaries are amongst the world’s most productive ecosystems, lying at the intersection between terrestrial and marine environments. They receive substantial inputs from adjacent landscapes but the importance of resource subsidies is not well understood. Here, we test hypotheses for the effects of both terrestrial- and salmon-derived resource subsidies on the diet (inferred from stable isotopes of muscle tissue), size and percent nitrogen of the soft-shell clam (Mya arenaria), a sedentary estuarine consumer. We examine how these relationships shift across natural gradients among 14 estuaries that vary in upstream watershed size and salmon density on the central coast of British Columbia, Canada. We also test how assimilation and response to subsidies vary at smaller spatial scales within estuaries. The depletion and enrichment of stable isotope ratios in soft-shell clam muscle tissue correlated with increasing upstream watershed size and salmon density, respectively. The effects of terrestrial- and salmon-derived subsidies were also strongest at locations near stream outlets. When we controlled for age of individual clams, there were larger individuals with higher percent nitrogen content in estuaries below larger watersheds, though this effect was limited to the depositional zones below river mouths. Pink salmon exhibited a stronger effect on isotope ratios of clams than chum salmon, which could reflect increased habitat overlap as spawning pink salmon concentrate in lower stream reaches, closer to intertidal clam beds. However, there were smaller clams in estuaries that had higher upstream pink salmon densities, possibly due to differences in habitat requirements. Our study highlights the importance of upstream resource subsidies to this bivalve species, but that individual responses to subsidies can vary at smaller scales within estuaries.     

Theory of grazing optimization in which herbivory improves photosynthetic ability

Herbivory had been generally considered to have a negative effect on plants, but a lot of studies have recently indicated that continuous herbivory pressure has a positive effect on plant performance, known as "grazing optimization." Based on field observations, we analytically examined a hypothesis of grazing optimization in which herbivory improves the photosynthetic ability of individual plants. We examined plant performance under various herbivory pressures and considered the evolution of plant phenology in response to a given herbivory pressure. First, we compared plant performances measured under their native conditions with specific herbivory levels. This was called the long-term response. Second, we examined the performances of plants adapting to a certain level of herbivory pressure under a non-native herbivory intensity. This was called the short-term response. According to numerical analysis, in realistic situations, grazing optimization is unlikely to be observed as a long-term response. However, grazing optimization can occur as short-term response if a plant is adapted to a certain level of herbivory pressure and the photosynthetic ability decreases significantly with the increasing size of vegetative parts. Our results suggest that improved photosynthetic ability by herbivory can result in grazing optimization, although it is constrained by the functional form of photosynthetic ability, native conditions, and experimental design.     

Grazing tolerance and mycorrhizal colonization: Effects of resource manipulation and plant size in biennial Gentianella campestris

As herbivory usually leads to loss of photosynthesizing biomass, its consequences for plants are often negative. However, in favorable conditions, effects of herbivory on plants may be neutral or even beneficial. According to the compensatory continuum hypothesis plants can tolerate herbivory best in resource-rich conditions. Besides herbivory, also primarily positive biotic interactions like mycorrhizal symbiosis, bear carbon costs. Tritrophic plant–fungus–herbivore interaction further complicates plant's cost-benefit balance, because herbivory of the host plant is expected to cause decline in mycorrhizal colonization under high availability of soil nutrients when benefits of symbiosis decline in relation to costs. To gain insight into above interactions we tested the effects of plant size and resource manipulation (simulated herbivory and fertilization) on both above-ground performance and on root fungal colonization of the biennial Gentianella campestris.Clipping caused allocation shift from height growth to branches in all groups except in large and fertilized plants. For large plants nutrient addition may have come too late, as the number of meristems was most likely determined already before the fertilization. Clipping decreased the amount of DSE (dark septate endophytic) fungi which generally are not considered to be mycorrhizal. The effect of clipping on total fungal colonization and colonization by arbuscular mycorrhizal (AM) fungal coils were found to depend on host size and resource level. Dissimilar mycorrhizal response to simulated herbivory in small vs. large plants could be due to more intensive light competition in case of small plants. Carbon limited small plants may not be able to maintain high mycorrhizal colonization, whereas large clipped plants allocate extra resources to roots and mycorrhizal fungi at the expense of above-ground parts. Our results suggest that herbivory may increase carbon limitation that leads re-growing shoots and fungal symbionts to function as competing sinks for the limited carbon reserves.     

Phytoplankton Community Composition in the Tri‐Lakes Area of Central Wisconsin,USA

Tri‐Lakes (Upper and Lower Camelot, Sherwood, Arrowhead) in Adams County, WI, USA are man‐made impoundments draining substantial agricultural lands and surrounded by considerable shoreline residential development. The planktonic algal community, as sampled from June to November 2000, was sparse‐to‐moderately dense, fairly diverse (69 genera from six divisions basin‐wide), and unremarkable in taxonomic composition. All sites sampled displayed the general algal successional trends expected from northern‐temperate, mildly eutrophic waters. These included sparse but taxonomically diverse communities in the spring; a late spring pulse of diatoms; a late summer pulse of green algae; and a steadily increasing component of Cyanobacteria leading to their community dominance by the end of the growing season. Upper Lake Camelot (55 genera) best represented this pattern. Lower Lake Camelot (53 genera) had a large green algal pulse but only a small diatom pulse. Lake Sherwood was the most taxonomically diverse body (63 genera) and had the most extreme pulses of diatoms and greens. Lake Arrowhead had the lowest taxonomic diversity (39 genera), was the most dominated by Cyanobacteria, and had only minor pulses of diatoms and greens. The algal communities indicate a mesotrophic to slightly eutrophic lake status. Continued agricultural and residential inputs of fertilizers and pesticides will likely exacerbate the cyanobacterial dominance leading to further reductions in aquatic health and aesthetic values. Previous chemical treatment and macrophyte removal have achieved limited success, and might have altered algal community dynamics. Remediation approaches that might improve water quality include: reducing upstream inputs via sediment traps or lagoons; reducing in‐lake nutrients via sediment removal; reducing residential inputs via improved septic/sanitation systems; and shoreline vegetation filter strips.     

The globalization of N deposition: ecosystem consequences in tropical environments

Human activities have more than doubled the inputs of nitrogen (N) into terrestrial systems globally. The sources and distribution of anthropogenic N, including N fertilization and N fixed during fossil fuel combustion, are rapidly shifting from the temperate zone to a more global distribution. The consequences of anthropogenic N deposition for ecosystem processes and N losses have been studied primarily in N-limited ecosystems in the temperate zone; there is reason to expect that tropical ecosystems, where plant growth is most often limited by some other resource, will respond differently to increasing deposition. In this paper, we assess the likely direct and indirect effects of increasing anthropogenic N inputs on tropical ecosytem processes. We conclude that anthropogenic inputs of N into tropical forests are unlikely to increase productivity and may even decrease it due to indirect effects on acidity and the availability of phosphorus and cations. We also suggest that the direct effects of anthropogenic N deposition on N cycling processes will lead to increased fluxes at the soilwater and soil-air interfaces, with little or no lag in response time. Finally, we discuss the uncertainties inherent in this analysis, and outline future research that is needed to address those uncertainties.     

Terpene production and growth of three Pacific Northwest conifers in response to simulated browse and nutrient availability

Natural variation in ungulate browsing behavior interferes with the understanding of plant morphological and biochemical responses to herbivory. To investigate mechanisms for recovery from herbivory, we examined growth patterns and biosynthesis of terpenoids under simulated browse (three clipping intensities) and supplemental mineral nutrition (four levels of controlled-release fertilization) for Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco], western hemlock (Tsuga heterophylla Raf. Sarg.), and western red-cedar (Thuja plicata Donn ex D. Don) seedlings on a reforestation site in Northwestern Oregon, USA, that was fenced to exclude ungulates. Higher clipping intensities increased relative height growth (at cost of diameter growth) for all the species. Only western red-cedar showed a decline in monoterpene concentrations with increasing clipping severity, suggesting prioritization in biosynthesis of terpenoids for this species. Douglas-fir and western hemlock responded to fertilization mostly through increased growth. Western red-cedar growth responses to fertilization were less pronounced, but monoterpene concentrations were 2–3 times higher compared to non-fertilized trees. Douglas-fir and western hemlock browse recovery and responses to fertilization consisted primarily of increased growth, while western red-cedar balanced growth promotion with production of chemical defense compounds. Our data suggests the evolution of species-dependent resource allocation strategies in response to both browse and soil nutrient availability.     

Aquatic insect subsidies influence microbial composition and processing of detritus in near-shore subarctic heathland

Insects are major conduits of resources moving from aquatic to terrestrial systems. While the ecological impacts of insect subsidies are well documented, the underlying mechanisms by which these resources change recipient ecosystems remain poorly understood. Most subsidy inputs enter terrestrial systems as detritus; thus, soil microbes will likely influence the processing of insect subsidies, with implications for plant community composition and net primary productivity (NPP). In a subarctic ecosystem near Lake Mývatn, Iceland where midge (Diptera: Chironomidae) deposition to land is high, we investigated how insect subsidies affected litter processing and microbial communities. We also evaluated how those belowground effects related to changes in inorganic nitrogen, plant composition and NPP. We simulated subsidies by adding midge carcasses to 1-m² heathland plots, where we measured effects on decomposition rates and the plant community. We then studied how fertilization treatments (control, KNO₃ and midge-carcass addition) affected graminoid biomass and inorganic nitrogen in greenhouse experiments. Lastly, we conducted a soil-incubation study with a phospholipid fatty acid analysis (PLFA) to examine how midge addition to heathland soils affected microbial respiration, biomass and composition. We found that midge addition to heathland soils increased litter decomposition and graminoid plant cover by 2.6× and 2×, respectively. Greenhouse experiments revealed similar patterns, with midge carcasses increasing graminoid biomass by at least 2× and NH₄⁺ concentrations by 7×. Our soil-incubation study found that midge carcasses elevated microbial respiration by 64%, microbial biomass by 43% and shifted microbial functional composition. Our findings indicate that insect subsidies can stimulate soil microbial communities and litter decomposition in subarctic heathlands, leading to increased NPP and changes in plant community composition.     

Soil warming accelerates decomposition of fine woody debris

Aims

There is evidence that increased N inputs to boreal forests, via atmospheric deposition or intentional fertilization, may impact negatively on ectomycorrhizal (ECM) fungi leading to a reduced flux of plant-derived carbon (C) back to the atmosphere via ECM. Our aim was to investigate the impact of N fertilization of a Pinus sylvestris (L.) forest stand on the return of recently photoassimilated C via the ECM component of soil respiration.

Methods

We used an in situ, large-scale, ¹³C-CO₂ isotopic pulse labelling approach and monitored the ¹³C label return using soil gas efflux chambers placed over three different types of soil collar to distinguish between heterotrophic (R_H), autotrophic (R_A; partitioned further into contributions from ECM hyphae and total R_A) and total (R_S) soil respiration.

Results

The impact of N fertilization was to significantly reduce R_A, particularly respiration via extramatrical ECM hyphae. ECM hyphal flux in control plots showed substantial spatial variability, resulting in mean flux estimates exceeding estimates of total R_A, while ECM contributions to R_A in N treated plots were estimated at around 30%.

Conclusion

Significant impacts on soil C cycling may be caused by reduced plant C allocation to ECM fungi in response to increased N inputs to boreal forests; ecosystem models so far lack this detail.     

Competition induces allelopathy but suppresses growth and anti-herbivore defence in a chemically rich seaweed

Many seaweeds and terrestrial plants induce chemical defences in response to herbivory, but whether they induce chemical defences against competitors (allelopathy) remains poorly understood. We evaluated whether two tropical seaweeds induce allelopathy in response to competition with a reef-building coral. We also assessed the effects of competition on seaweed growth and seaweed chemical defence against herbivores. Following 8 days of competition with the coral Porites cylindrica, the chemically rich seaweed Galaxaura filamentosa induced increased allelochemicals and became nearly twice as damaging to the coral. However, it also experienced significantly reduced growth and increased palatability to herbivores (because of reduced chemical defences). Under the same conditions, the seaweed Sargassum polycystum did not induce allelopathy and did not experience a change in growth or palatability. This is the first demonstration of induced allelopathy in a seaweed, or of competitors reducing seaweed chemical defences against herbivores. Our results suggest that the chemical ecology of coral–seaweed–herbivore interactions can be complex and nuanced, highlighting the need to incorporate greater ecological complexity into the study of chemical defence.     

Marine subsidies alter the diet and abundance of insular and coastal lizard populations

We evaluated the impact of marine materials on the diet and abundance of arthropodivorous lizards inhabiting islands and the coast of the Gulf of California. Here, marine materials are brought onto land by seabirds and by tidal action, and both subsidy pathways cause arthropod abundance to increase. We evaluated Uta stansburiana (side-blotched lizard) diets in three habitats defined by having: (1) no marine subsidies available, (2) only seabird-derived subsidies, and (3) only tidal-derived subsidies. Stable isotope data indicated that lizard diets are subsidized indirectly by seabird and tidal activity. For example, in coastal areas we determined that 40% of a lizard's diet contains arthropods that have consumed algae. Such subsidies may explain why we found that lizards in coastal areas occupy a significantly higher trophic position than lizards in unsubsidized areas. We analyzed eight years of survey data on all arthropodivorous lizards to determine if diet subsidies result in increased lizard abundance. We found that lizards were more abundant in coastal areas than they were in inland habitats, and that they were more abundant on islands with seabirds than on islands without seabird populations. This study provides insight into the importance and effect of marine-derived nutrients from multiple sources on vertebrate consumers inhabiting islands and coastal areas.     

Context dependency of animal resource subsidies

The transport of resource subsidies by animals has been documented across a range of species and ecosystems. Although many of these studies have shown that animal resource subsidies can have significant effects on nutrient cycling, ecosystem productivity, and food‐web structure, there is a great deal of variability in the occurrence and strength of these effects. Here we propose a conceptual framework for understanding the context dependency of animal resource subsidies, and for developing and testing predictions about the effects of animal subsidies over space and time. We propose a general framework, in which abiotic characteristics and animal vector characteristics from the donor ecosystem interact to determine the quantity, quality, timing, and duration (QQTD) of an animal input. The animal input is translated through the lens of recipient ecosystem characteristics, which include both abiotic and consumer characteristics, to yield the QQTD of the subsidy. The translated subsidy influences recipient ecosystem dynamics through effects on both trophic structure and ecosystem function, which may both influence the recipient ecosystem's response to further inputs and feed back to influence the donor ecosystem. We present a review of research on animal resource subsidies across ecosystem boundaries, placed within the context of this framework, and we discuss how the QQTD of resource subsidies can influence trophic structure and ecosystem function in recipient ecosystems. We explore the importance of understanding context dependency of animal resource subsidies in increasingly altered ecosystems, in which the characteristics of both animal vectors and donor and recipient ecosystems may be changing rapidly. Finally, we make recommendations for future research on animal resource subsidies, and resource subsidies in general, that will increase our understanding and predictive capacity about their ecosystem effects.     

Collembola respond to aphid herbivory but not to honeydew addition

Abstract. 1. Euedaphic collembola alter their soil distribution in response to above‐ground aphid herbivory of Poa annua L. Graminae, a host grass. 2. Two mechanisms potentially underpin this effect. Carbon‐rich aphid honeydew falling onto the soil surface may affect mycophagous collembola; alternatively aphid‐induced changes in root biomass may be necessary to produce changes in collembola abundance. 3. When compared to a plant‐only control, aphid herbivory increased the number of collembola in the top 5 cm of soil, reduced both foliar and root biomass, and increased shoot/root ratio. Honeydew addition had no effect on collembola numbers or any recorded host‐plant parameter. 4. Honeydew deposition is not responsible for the increased numbers of collembola found in the upper soil after aphid herbivory; aphid‐induced reductions in root biomass may be the most important factor explaining knock‐on effects of aphid herbivory on soil fauna.     

Growth and development rates in a riparian spider are altered by asynchrony between the timing and amount of a resource subsidy

Rapid growth in response to increased prey abundance may be induced by environmental variability associated with resource subsidies. Spiders living in riparian areas are subject to frequent, episodic bursts of aquatic prey (subsidies). These periods of high resource abundance may occur at different points in recipient consumers’ development through variation in emergence patterns of prey between years or across a landscape. We examine how variable timing of subsidy abundance intersects with life history scheduling to produce different growth and development outcomes for individuals within a population. Through a series of controlled feeding experiments, we tested the hypotheses that the spider Tetragnatha versicolor: (1) exhibits compensatory growth in response to subsidy variability, (2) that rapid increases in mass may result in a greater risk of mortality, and (3) that the timing of subsidy resources relative to the development schedule of this spider may produce different outcomes for individual growth patterns and adult condition. Spiders fed at very high rates grew fastest but also showed evidence of increased mortality risk during moulting. T. versicolor is capable of exhibiting strong growth compensation—individuals suffering initial growth restriction were able to catch up completely with animals on a constant diet utilising the same amount of food. Spiders that received an early pulse of resources (simulating an early arrival of an aquatic insect subsidy to riparian forests) did worse on all measures of development and fitness than spiders that received either a constant supply of food or a late pulse of resources. Importantly, receiving large amounts of food early in life appears to actually confer relative disadvantages in terms of later performance compared with receiving subsidies later in development. Subsidies may provide greater benefits to individuals or age cohorts encountering this resource abundance closer to the onset of reproductive efforts than subsidies arriving early in development.     

Rapid plant species loss at high rates and at low frequency of N addition in temperate steppe

Humans are both intentionally (fertilization) and unintentionally (atmospheric nutrient deposition) adding nutrients worldwide. Increasing availability of biologically reactive nitrogen (N) is one of the major drivers of plant species loss. It remains unclear, however, whether plant diversity will be equally reduced by inputs of reactive N coming from either small and frequent N deposition events or large and infrequent N fertilization events. By independently manipulating the rate and frequency of reactive N inputs, our study teases apart these potentially contrasting effects. Plant species richness decreased more quickly at high rates and at low frequency of N addition, which suggests that previous fertilization studies have likely over‐estimated the effects of N deposition on plant species loss. N‐induced species loss resulted from both acidification and ammonium toxicity. Further study of small and frequent N additions will be necessary to project future rates of plant species loss under increasing aerial N deposition.