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Human activities are increasing wildfires and livestock activity in arid ecosystems with potential implications for the spread of invasive grasses. The objective of this study was to test whether fire history and cattle activity alter soil resource gradients, thereby affecting patterns of Bromus rubens L. (red brome) invasion. Six paired burned and unburned transect lines (1-km long) were established in the northeast Mojave Desert along the boundaries of four independent wildfire scars. At 100-m transect increment points, we measured the distance to the two nearest cowpats, and two random points and measured the density, height, biomass, and seed production of red brome, soil moisture and inorganic nitrogen (N). Cattle activity was 29% greater along burned transects compared to unburned transects (P < 0.05). Red brome height, density, and seed production were 11-34% greater along burned transects than unburned transects (P < 0.05). Red brome height, biomass, density, and seed production were twofold to tenfold greater next to cowpats compared to random points (P < 0.05). Soils along burned transects and beneath cowpats had greater soil inorganic N (P < 0.05), which was positively correlated with red brome density, height, biomass, and seed production (R 2  = 0.60-0.85, P < 0.0001). Transgenerational effects were evident as seeds from red brome next to cowpats had 27% higher germination than seeds collected from random points. Positive responses of red brome to increased inorganic N related to fire and cattle activity may contribute fine fuel infill that drives invasive grass-fire cycles in deserts., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Nutrient resorption is a fundamental physiological process in plants, with important ecological controls over numerous ecosystem functions. However, the role of community assembly in driving responses of nutrient resorption to perturbation remains largely unknown. Following the Price equation framework and the Community Assembly and Ecosystem Function framework, we quantified the contribution of species loss, species gain, and shared species to the reduction of community-level nutrient resorption efficiency in response to multi-level nitrogen (N) addition in a temperate steppe, after continuous N addition for seven years. Reductions of both N and phosphorus (P) resorption efficiency (NRE and PRE, respectively) were positively correlated with N addition levels. The dissimilarities in species composition between N-enriched and control communities increased with N addition levels, and N-enriched plots showed substantial species losses and gains. Interestingly, the reduction of community-scale NRE and PRE mostly resulted from N-induced decreases in resorption efficiency for the shared species in the control and N-enriched communities. There were negative correlations between the contributions of species richness effect and species identity effect and between the number and identity of species gained for the changes in both NRE and PRE following N enrichment. By simultaneously considering N-induced changes in species composition and in species-level resorption, our work presents a more complete picture of how different community assembly processes contribute to N-induced changes in community-level resorption., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Drought and nutrient input are two main global change drivers that threaten ecosystem function and services. Resolving the interactive effects of human-induced stressors on individual species is necessary to improve our understanding of community and ecosystem responses. This study comparatively assessed how different nutrient conditions affect whole-plant drought responses across 13 common temperate grassland species. We conducted a fully factorial drought-fertilization experiment to examine the effect of nutrient addition [nitrogen (N), phosphorus (P), and combined NP] on species' drought survival, and on drought resistance of growth as well as drought legacy effects. Drought had an overall negative effect on survival and growth, and the adverse drought effects extended into the next growing season. Neither drought resistance nor legacy effects exhibited an overall effect of nutrients. Instead, both the size and the direction of the effects differed strongly among species and between nutrient conditions. Consistently, species performance ranking under drought changed with nitrogen availability. The idiosyncratic responses of species to drought under different nutrient conditions may underlie the seemingly contradicting effects of drought in studies on grassland composition and productivity along nutrient and land-use gradients-ranging from amplifying to dampening. Differential species' responses to combinations of nutrients and drought, as observed in our study, complicate predictions of community and ecosystem responses to climate and land-use changes. Moreover, they highlight the urgent need for an improved understanding of the mechanisms that render species more or less vulnerable to drought under different nutrients., (© 2023. The Author(s).)

Human activities cause a multitude of environmental issues, including increased temperatures and altered precipitation patterns associated with climate change, air pollution, and other impacts of urbanization. One area highly affected by these issues is the Sonoran Desert, specifically the Phoenix metropolitan area where urbanization is among the most rapid in the United States. Most studies investigate these multiple environmental change factors independently or sometimes in pairs, but rarely all together as co-occurring forms of change. We examined how the simultaneous manipulation of increasing temperatures, altered precipitation patterns, nitrogen deposition, and urbanization influenced soil respiration and mineral N pools in the Sonoran Desert. Soil was collected from urban and exurban sites, from both nitrogen-fertilized and control plots. To simulate projected climate change, the soils were incubated in microcosm at the annual average Phoenix temperature as well a 2 ℃ increase under a factorial precipitation treatment of decreased frequency and increased pulse size. Our results show that C and N dynamics were altered by all four forms of environmental change. However, the dominance of significant 3- and 4-way interactions among the four environmental factors for both respiration and mineral N pools demonstrates that the impact of any given form of environmental change will depend on the levels of the other environmental factors. In other words, the cumulative effect of altered precipitation, fertilization, temperature, and urbanization on soil biogeochemical processes is not necessarily predictable from their individual impact., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Tropical forests are threatened by anthropogenic activities such as conversion into agricultural land, logging and fires. Land-use change and disturbance affect ecosystems not only aboveground, but also belowground including the ecosystems' carbon and nitrogen cycle. We studied the impact of different types of land-use change (intensive and traditional agroforestry, logging) and disturbance by fire on fine root biomass, dynamics, morphology, and related C and N fluxes to the soil via fine root litter across different ecosystems at different elevational zones at Mt. Kilimanjaro (Tanzania). We found a decrease in fine root biomass (80-90%), production (50%), and C and N fluxes to the soil via fine root litter (60-80%) at all elevation zones. The traditional agroforestry 'Chagga homegardens' (lower montane zone) showed enhanced fine root turnover rates, higher values of acquisitive root morphological traits, but similar stand fine root production, C and N fluxes compared to the natural forest. The decrease of C and N fluxes with forest disturbance was particularly strong at the upper montane zone (60 and 80% decrease, respectively), where several patches of Podocarpus forest had been disturbed by fire in the previous years. We conclude that changes on species composition, stand structure and land management practices resulting from land-use change and disturbance have a strong impact on the fine root system, modifying fine root biomass, production and the C and N supply to the soil from fine root litter, which strongly affects the ecosystems' C and N cycle in those East African tropical forest ecosystems., (© 2023. The Author(s).)

In the boreal forests, feather mosses such as Hylocomium splendens and Pleurozium schreberi are colonized by cyanobacteria, which provide large amounts of nitrogen to forest ecosystems through nitrogen fixation. Although these feather mosses are also ubiquitous in subalpine forests of East Asia, little is known regarding their associated cyanobacteria and their ability to fix nitrogen. In this study, we investigated (1) whether cyanobacteria co-exist and fix nitrogen in the two species of feather mosses that cover the ground surface in a subalpine forest of Mt. Fuji, (2) whether cyanobacteria belonging to a common cluster with boreal forests are found in feather mosses in Mt. Fuji, and (3) whether moss-associated nitrogen fixation rates differed among moss growing substrates, canopy openness, and moss nitrogen concentrations in the same forest area. Our results showed that cyanobacteria colonized feather mosses in the subalpine forests of Mt. Fuji and acetylene reduction rates as an index of nitrogen fixation tended to be higher in H. splendens than in P. schreberi. Based on analysis of the nifH gene, 43 bacterial operational taxonomic units (OTUs) were identified, 28 of which represented cyanobacteria. Among the five clusters of cyanobacteria classified based on their nifH gene and identified in northern Europe, four (Nostoc cluster I, Nostoc cluster II, Stigonema cluster, and nifH2 cluster) were also found at Mt. Fuji. The acetylene reduction rate differed depending on the moss growing substrate and the total nitrogen concentration of moss shoots, and a strong negative correlation was observed with the total nitrogen concentration., (© 2023. The Author(s).)

Evolutionary dietary shifts have major ecological consequences. One likely consequence is a change in nutrient limitation-some nutrients become more abundant in the diet, others become more scarce. Individuals' behavior should change accordingly to match this new limitation regime: they should seek out nutrients that are deficient in the new diet. We investigated the relationship between diet and responses to nutrients using adult Costa Rican butterflies with contrasting feeding habits, testing the hypothesis that animals will respond more positively to nutrients that are scarcer in their diets. Via literature searches and our own data, we showed that nitrogen and sodium are both at lower concentration in nectar than in fruit. We therefore assessed butterflies' acceptance of sodium and four nitrogenous compounds that ranged in complexity from inorganic nitrogen (ammonium chloride) to protein (albumin). We captured wild butterflies, offered them aqueous solutions of each substance, and recorded whether they accepted (drank) or rejected each substance. Support for our hypothesis was mixed. Across the sexes, frugivores were four times more likely to accept amino acids (hydrolyzed casein) than nectivores, in opposition to expectation. In males, nectivores accepted sodium almost three times more frequently than frugivores, supporting expectations. Together, these results suggest that in butterflies, becoming frugivorous is associated with an increased receptivity to amino acids and decreased receptivity to sodium. Nectivory and frugivory are widespread feeding strategies in organisms as diverse as insects, birds, and bats; our results suggest that these feeding strategies may put different pressures on how animals fulfill their nutritional requirements. [ABSTRACT FROM AUTHOR]

Environmental changes are expected to shift the distribution of functional trait values in plant communities through a combination of species turnover and intraspecific variation. The strength of these shifts may depend on the availability of individuals with trait values adapted to new environmental conditions, represented by the functional diversity (FD) of existing community residents or dispersal from the regional species pool. We conducted a 3-year nutrient- and seed-addition experiment in old-field plant communities to examine the contributions of species turnover and intraspecific variation to community trait shifts, focusing on four key plant functional traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We further examined the influence of initial FD and seed availability on the strength of these shifts. Community mean height, leaf area, and SLA increased in response to fertilization, and these shifts were driven almost entirely by intraspecific variation. The strength of intraspecific shifts in height and leaf area was positively related to initial intraspecific FD in these traits. Intraspecific trait responses to fertilization varied among species, with species of short stature displaying stronger shifts in SLA and LDMC but weaker shifts in leaf area. Trait shifts due to species turnover were generally weak and opposed intraspecific responses. Seed addition altered community taxonomic composition but had little effect on community trait shifts. These results highlight the importance of intraspecific variation for short-term community functional responses and demonstrate that the strength of these responses may be mediated by community FD. [ABSTRACT FROM AUTHOR]

Human drivers are often proposed to be stronger than biophysical drivers in influencing ecosystem structure and function in highly urbanized areas. In residential land cover, private yards are influenced by individual homeowner preferences and actions while also experiencing large-scale human and biophysical drivers. We studied plant nitrogen (%N) and N stable isotopic composition (δN) in residential yards and paired native ecosystems in seven cities across the US that span major ecological biomes and climatic regions: Baltimore, Boston, Los Angeles, Miami, Minneapolis-St. Paul, Phoenix, and Salt Lake City. We found that residential lawns in three cities had enriched plant δN ( P < 0.03) and in six cities higher plant N (%) relative to the associated native ecosystems ( P < 0.05). Plant δN was progressively depleted across a gradient of urban density classes in Baltimore and Boston ( P < 0.05). Lawn fertilization was associated with depleted plant δN in Boston and Los Angeles ( P < 0.05), and organic fertilizer additions were associated with enriched plant δN in Los Angeles and Salt Lake City ( P < 0.04). Plant δN was significantly enriched as a function of housing age in Baltimore ( r = 0.27, P < 0.02), Boston ( r = 0.27, P < 0.01), and Los Angeles ( r = 0.34, P < 0.01). These patterns in plant δN and plant N (%) across these cities suggests that N sources to lawns, as well as greater rates of N cycling combined with subsequent N losses, may be important drivers of plant N dynamics in lawn ecosystems at the national scale. [ABSTRACT FROM AUTHOR]

Even though drought impacts on tree physiology have been identified, whether drought affects leaf litter chemistry that, in turn, influences litter decay rates is still poorly understood. We compared litter quality and decomposition for two cohorts of leaves from five co-occurring seasonally deciduous tree species: Acer saccharum, Tilia americana, Quercus rubra, Quercus alba, and Ostrya virginiana. One cohort experienced a growing-season drought, and the other cohort came from the same trees in the ensuing, post-drought growing season. Leaf litter production was greater for drought litter than post-drought litter for all five species. Specific leaf area and nitrogen concentrations were 20% greater for the drought cohort than the post-drought cohort. Concentrations of non-structural carbohydrates were about 14% greater for the drought cohort, except for greater values for post-drought A. saccharum litter. Pectin in the middle lamella of leaf litter was 31% lower for the drought cohort compared to post-drought cohort. We found few differences in litter decay rates between drought and post-drought cohorts, although Q. rubra litter had more decomposition for the post-drought cohort than the drought cohort, whereas A. saccharum litter had more decomposition for the drought cohort than the post-drought cohort. Leaf litter decay rates for the drought cohort were related to litter nitrogen and lignin concentrations, whereas decay rates for the post-drought cohort were related to litter carbohydrate concentrations. Our findings suggest that the role of drought events on seasonally deciduous forest ecosystems must recognize species-specific, idiosyncratic responses in leaf litter quality and decomposition., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Water balance influences soil development, and consequently plant communities, by driving weathering of soil minerals and leaching of plant nutrients from the soil. Along gradients in water balance, soils exhibit process domains where chemical properties are relatively stable punctuated by pedogenic thresholds where soil chemical properties change rapidly with little additional change in water balance. We ask if plant macronutrient concentrations in leaves also exhibit non-linear trends along water balance gradients, and if so, how these non-linearities relate to those in soils. We analyze foliar nutrient concentrations and foliar N:P ratios from eight species that span a range of growth forms along three water balance gradients (three of the species are found on multiple gradients). The gradients are located on basaltic substrate of different ages and have previously been characterized by studies on soil development. We find that maximum concentrations of foliar macronutrients occur at an intermediate water balance. As with soil nutrients, time mediates the effect of water balance on foliar nutrients, such that plants on older soils attain maximum nutrient concentrations at a lower water balance. On both a young, 20 ky and an old, 4100 ky water balance gradient, foliar nutrients reach peak concentrations at a water balance greater than the threshold for depletion of rock-derived nutrients in surface soils. Our findings suggest that plant acquisition of essential nutrients is imperfectly predicted by overall soil nutrient availability because the regulation of internal nutrient pools by plants makes nutrient pools within leaves partially independent of soil nutrient availability., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Increases in nitrogen (N) inputs to the biosphere can exacerbate the introduction and spread of invasive non-native plant species. Often, with elevated soil N levels, invasive plants establish and further enrich soil N pools, changing overall ecosystem function. This study examined the relationship between soil N cycling and an increasingly prevalent, invasive plant species, tall oatgrass (Arrhenatherum elatius subsp. elatius), in foothills ecosystems between the Colorado Rocky Mountains and the Denver-Boulder Metropolitan area-similar to many Western US grasslands and woodlands. It focused on investigating differences in soil N transformations, inorganic N pools, and vegetation characteristics across invaded and uninvaded plots at three sites in two seasons (summer and autumn). There was a statistically significant effect of invasion on rates of net N mineralization, but it was dependent on site and season (p = 0.046). Site had a statistically significant effect on soil moisture and aboveground biomass C:N (p < 0.04). The interactions of invasion x site were statistically significant for ammonium pools (p < 0.03). These findings suggest that A. elatius invasion can be associated with accelerated N cycling, but that the nature of the relationship differs by location and season in the foothills. More broadly, this study contributes to determining how the N cycle is shifting in grassland ecosystems subject to increasing pressures from anthropogenic change., (© 2022. The Author(s).)

Cloud forests have been found to lose more nitrogen in stream discharge than they gain from atmospheric deposition. They also support a large diversity and biomass of tree epiphytes, predominately composed of cryptogams. Since cryptogam epiphytes harbor nitrogen fixing cyanobacteria, they may help make up for the nitrogen loss from ecosystems. We assessed cryptogam biomass on the ground, boles and branches in Quercus costaricensis dominated stands near the tree line in the Cordillera de Talamanca, Costa Rica. Nitrogen fixation was assayed using 15 N 2 uptake. Total cryptogam biomass was 2 977 kg ha -1 , with 67% being found on the lower branches. Bryophytes and chlorolichens made up 53% and 44%, respectively, of the biomass. Half of the bryophyte mass was composed of the liverwort Plagiochila heterophylla, and 66% of the chlorolichen of Lobariella pallida. There were no significant differences in nitrogen fixation rates between the cryptogam species, with a mean rate of 5.04 µg N g -1 day -1 during the predominantly wet condition in the forest. The overall nitrogen input from fixation was 6.1 kg N ha -1  year -1 , of which 78% came from bryophytes, 18% from chlorolichens, and 4% from cyanolichens. Only 2.0% of the fixation occurred in cryptogams on the ground, whereas 67%, 24%, and 7% occurred on the lower branches, boles, and upper branches, respectively. These results show that tree epiphytes constitute a significant source of nitrogen for these forests, due to the trees' large surface area, and can make up for the nitrogen lost from these ecosystems.

The impacts of invasive species on biodiversity may be mitigated or exacerbated by abiotic environmental changes. Invasive plants can restructure soil fungal communities with important implications for native biodiversity and nutrient cycling, yet fungal responses to invasion may depend on numerous anthropogenic stressors. In this study, we experimentally invaded a long-term soil warming and simulated nitrogen deposition experiment with the widespread invasive plant Alliaria petiolata (garlic mustard) and tested the responses of soil fungal communities to invasion, abiotic factors, and their interaction. We focused on the phytotoxic garlic mustard because it suppresses native mycorrhizae across forests of North America. We found that invasion in combination with warming, but not under ambient conditions or elevated nitrogen, significantly reduced soil fungal biomass and ectomycorrhizal relative abundances and increased relative abundances of general soil saprotrophs and fungal genes encoding for hydrolytic enzymes. These results suggest that warming potentially exacerbates fungal responses to plant invasion. Soils collected from uninvaded and invaded plots across eight forests spanning a 4 °C temperature gradient further demonstrated that the magnitude of fungal responses to invasion was positively correlated with mean annual temperature. Our study is one of the first empirical tests to show that the impacts of invasion on fungal communities depends on additional anthropogenic pressures and were greater in concert with warming than under elevated nitrogen or ambient conditions.

Plant growth in northern forest ecosystems is considered to be primarily nitrogen limited. Nitrogen deposition is predicted to change this towards co-limitation/limitation by other nutrients (e.g., phosphorus), although evidence of such stoichiometric effects is scarce. We utilized two forest fertilization experiments in southern Sweden to analyze single and combined effects of nitrogen and phosphorus on the productivity, composition, and diversity of the ground vegetation. Our results indicate that the productivity of forest ground vegetation in southern Sweden is co-limited by nitrogen and phosphorus. Additionally, the combined effect of nitrogen and phosphorus on the productivity was larger than when applied solely. No effects on species richness of any of these two nutrients were observed when applied separately, while applied in combination, they increased species richness and changed species composition, mainly by promoting more mesotrophic species. All these effects, however, occurred only for the vascular plants and not for bryophytes. The tree layer in a forest has a profound impact on the productivity and diversity of the ground vegetation by competing for both light and nutrients. This was confirmed in our study where a combination of nitrogen and high tree basal area reduced cover of the ground vegetation compared to all the other treatments where basal area was lower after stand thinning. During the past decades, nitrogen deposition may have further increased this competition from the trees for phosphorus and gradually reduced ground vegetation diversity. Phosphorus limitation induced by nitrogen deposition may, thus, contribute to ongoing changes in forest ground vegetation.

Litter 'quality' or decomposability has historically been estimated through measuring chemical attributes, such as concentrations of nitrogen or 'lignin'. More recently, foliar functional traits, which may incorporate indications of the physical structures of tissues, have been found to correlate with litter mass loss rates. However, these traits may not be adequate to predict early rates of mass loss, in which two factors are crucial: the amount of material quickly lost through leaching, and the ease of access of decomposer organisms to the more labile tissues in the interior of the litter. We investigated relationships among physical and chemical traits in foliage and litter of 12 species native to British Columbia and then observed how these traits related to mass loss during the first 3 months (Phase I) and between 3 and 12 months (Phase II). Novel traits measured in this study include cuticle thickness, litter leaching loss, and litter water uptake. Foliar and litter traits both co-varied along spectra, but several chemical traits, such as nitrogen concentration, changed from foliage to litter, i.e., during senescence. Phase I mass loss was best predicted by leaching loss and traits relating to leaching, such as cuticle thickness and specific leaf area. Phase II mass loss was predicted by traits that may relate to decomposer access and activity, such as leaf dry matter content and foliar nitrogen. Physical traits predicted mass loss as well or better than chemical traits, suggesting that physical characteristics of litter are important in determining early rates of decomposition.

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC-potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L -1 ), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal lakes.

Alpine plant communities are predicted to face range shifts and possibly extinctions with climate change. Fine-scale environmental variation such as nutrient availability or snowmelt timing may contribute to the ability of plant species to persist locally; however, variation in nutrient availability in alpine landscapes is largely unmeasured. On three mountains around Davos, Switzerland, we deployed Plant Root Simulator probes around 58 Salix herbacea plants along an elevational and microhabitat gradient to measure nutrient availability during the first 5 weeks of the summer growing season, and used in situ temperature loggers and observational data to determine date of spring snowmelt. We also visited the plants weekly to assess performance, as measured by stem number, fruiting, and herbivory damage. We found a wide snowmelt gradient which determined growing season length, as well as variations of an order of magnitude or more in the accumulation of 12 nutrients between different microhabitats. Higher nutrient availability had negative effects on most shrub performance metrics, for instance decreasing stem number and the proportion of stems producing fruits. High nutrient availability was associated with increased herbivory damage in early-melting microhabitats, but among late-emerging plants this pattern was reversed. We demonstrate that nutrient availability is highly variable in alpine settings, and that it strongly influences performance in an alpine dwarf shrub, sometimes modifying the response of shrubs to snowmelt timing. As the climate warms and human-induced nitrogen deposition continues in the Alps, these factors may contribute to patterns of local plants persistence.

A basic assumption of the trait-based approach in plant ecology is that differences in functional trait values are greater between species than within species. We questioned this assumption by assessing (1) the relative extent of inter- and intraspecific leaf trait variation throughout a complete growing season (phenological variation) in a group of deciduous and evergreen woody species, and (2) whether species rankings based on leaf traits were maintained across the growing season. We analysed leaf mass per area (LMA) and leaf nutrient concentrations (C, N, P), including the C:N and N:P ratios. Intraspecific trait variation (ITV) due to phenology was significantly greater than interspecific variation for leaf N concentration on a mass basis (Nm; 68.90 %) and for the leaf C:N ratio (60.60 %), whereas interspecific variation was significantly higher than ITV for LMA (62.30 %) and for leaf C concentration on a mass (Cm) and area (Ca) basis (Cm 70.40 %; Ca 65.30 %). ITV was particularly low for LMA (<20 %). Species rankings were highly modified by phenology for a number of leaf traits (Pm, N:P ratio) but were relatively well conserved throughout the growing season for others (LMA, Nm). Patterns of ITV across the growing season differed significantly between deciduous and evergreen species for all traits except leaf P but did not vary between native and exotic species. Overall, our results show that intraspecific phenological variation in leaf traits may be similar to or greater than interspecific variation and that temporal patterns of ITV vary considerably among traits and species, especially for leaf nutrient concentrations, factors which can potentially affect quantitative interspecific relationships.

Ecophysiological studies of bryophytes have generally been conducted at the shoot or canopy scale. However, their growth forms are diverse, and knowledge of whether bryophytes with different shoot structures have different functional trait levels and scaling relationships is limited. We collected 27 bryophyte species and categorised them into two groups based on their growth forms: erect and prostrate species. Twenty-one morphological, nutrient and photosynthetic traits were quantified. Trait levels and bivariate trait scaling relationships across species were compared between the two groups. The two groups had similar mean values for shoot mass per area (SMA), light saturation point and mass-based nitrogen (N(mass)) and phosphorus concentrations. Erect bryophytes possessed higher values for mass-based chlorophyll concentration (Chl(mass)), light-saturated assimilation rate (A(mass)) and photosynthetic nitrogen/phosphorus use efficiency. N(mass), Chl(mass) and A(mass) were positively related, and these traits were negatively associated with SMA. Furthermore, the slope of the regression of N(mass) versus Chl(mass) was steeper for erect bryophytes than that for prostrate bryophytes, whereas this pattern was reversed for the relationship between Chl(mass) and A(mass). In conclusion, erect bryophytes possess higher photosynthetic capacities than prostrate species. Furthermore, erect bryophytes invest more nitrogen in chloroplast pigments to improve their light-harvesting ability, while the structure of prostrate species permits more efficient light capture. This study confirms the effect of growth form on the functional trait levels and scaling relationships of bryophytes. It also suggests that bryophytes could be good models for investigating the carbon economy and nutrient allocation of plants at the shoot rather than the leaf scale.

High daily temperature range of soil (DTRsoil) negatively affects soil microbial biomass and activity, but its interaction with seasonal soil moisture in regulating ecosystem function remains unclear. For our 5-year field study in the Chihuahuan Desert, we suspended shade cloth 15 cm above the soil surface to reduce daytime temperature and increase nighttime soil temperature compared to unshaded plots, thereby reducing DTRsoil (by 5 ºC at 0.2 cm depth) without altering mean temperatures. Microbial biomass production was primarily regulated by seasonal precipitation with the magnitude of the response dependent on DTRsoil. Reduced DTRsoil more consistently increased microbial biomass nitrogen (MBN; +38%) than microbial biomass carbon (MBC) with treatment responses being similar in spring and summer. Soil respiration depended primarily on soil moisture with responses to reduced DTRsoil evident only in wetter summer soils (+53%) and not in dry spring soils. Reduced DTRsoil had no effect on concentrations of dissolved organic C, soil organic matter (SOM), nor soil inorganic N (extractable NO3 (-)-N + NH4 (+)-N). Higher MBN without changes in soil inorganic N suggests faster N cycling rates or alternate sources of N. If N cycling rates increased without a change to external N inputs (atmospheric N deposition or N fixation), then productivity in this desert system, which is N-poor and low in SOM, could be negatively impacted with continued decreases in daily temperature range. Thus, the future N balance in arid ecosystems, under conditions of lower DTR, seems linked to future precipitation regimes through N deposition and regulation of soil heat load dynamics.

Soil N fertility has an effect on belowground C allocation, but the physiological and morphological responses of individual fine root segments to variations in N availability under field conditions are still unclear. In this study, the direction and magnitude of the physiological and morphological function of fine roots in response to variable in situ soil N fertility in a forest site were determined. We measured the specific root respiration (Rr) rate, N concentration and morphology of fine root segments with 1-3 branching orders in a 100-year-old coniferous forest of Chamaecyparis obtusa. Higher soil N fertility induced higher Rr rates, root N concentration, and specific root length (SRL), and lower root tissue density (RTD). In all fertility levels, the Rr rates were significantly correlated positively with root N and SRL and negatively with RTD. The regression slopes of respiration with root N and RTD were significantly higher along the soil N fertility gradient. Although no differences in the slopes of Rr and SRL relationship were found across the levels, there were significant shifts in the intercept along the common slope. These results suggest that a contrasting pattern in intraspecific relationships between specific Rr and N, RTD, and SRL exists among soils with different N fertility. Consequently, substantial increases in soil N fertility would exert positive effects on organ-scale root performance by covarying the Rr, root N, and morphology for their potential nutrient and water uptake.

While parasites are increasingly recognized as important components of ecosystems, we currently know little about how they alter ecosystem nutrient availability via host-mediated nutrient cycling. We examined whether infection alters the flow of nutrients through hosts and whether such effects depend upon host diet quality. To do so, we compared the mass specific nutrient (i.e., nitrogen and phosphorus) release rates, ingestion rates, and elemental composition of uninfected Daphnia to those infected with a bacterial parasite, P. ramosa. N and P release rates were increased by infection when Daphnia were fed P-poor diets, but we found no effect of infection on the nutrient release of individuals fed P-rich diets. Calculations based on the first law of thermodynamics indicated that infection should increase the nutrient release rates of Daphnia by decreasing nutrient accumulation rates in host tissues. Although we found reduced nutrient accumulation rates in infected Daphnia fed all diets, this reduction did not increase the nutrient release rates of Daphnia fed the P-rich diet because infected Daphnia fed this diet ingested nutrients more slowly than uninfected hosts. Our results thus indicate that parasites can significantly alter the nutrient use of animal consumers, which could affect the availability of nutrients in heavily parasitized environments.

Our study investigated the carbon:nitrogen:phosphorus (C:N:P) stoichiometry of mangrove island of the Mesoamerican Barrier Reef (Twin Cays, Belize). The C:N:P of abiotic and biotic components of this oligotrophic ecosystem was measured and served to build networks of nutrient flows for three distinct mangrove forest zones (tall seaward fringing forest, inland dwarf forests and a transitional zone). Between forest zones, the stoichiometry of primary producers, heterotrophs and abiotic components did not change significantly, but there was a significant difference in C:N:P, and C, N, and P biomass, between the functional groups mangrove trees, other primary producers, heterotrophs, and abiotic components. C:N:P decreased with increasing trophic level. Nutrient recycling in the food webs was highest for P, and high transfer efficiencies between trophic levels of P and N also indicated an overall shortage of these nutrients when compared to C. Heterotrophs were sometimes, but not always, limited by the same nutrient as the primary producers. Mangrove trees and the primary tree consumers were P limited, whereas the invertebrates consuming leaf litter and detritus were N limited. Most compartments were limited by P or N (not by C), and the relative depletion rate of food sources was fastest for P. P transfers thus constituted a bottleneck of nutrient transfer on Twin Cays. This is the first comprehensive ecosystem study of nutrient transfers in a mangrove ecosystem, illustrating some mechanisms (e.g. recycling rates, transfer efficiencies) which oligotrophic systems use in order to build up biomass and food webs spanning various trophic levels.

Leaf traits are closely associated with nutrient use by plants and can be utilized as a proxy for nutrient cycling processes. However, open questions remain, in particular regarding the variability of leaf traits within and across seasonally dry tropical forests. To address this, we considered six leaf traits (specific area, thickness, dry matter content, N content, P content and natural abundance (15)N) of four co-occurring tree species (two that are not associated with N2-fixing bacteria and two that are associated with N2-fixing bacteria) and net N mineralization rates and inorganic N concentrations along a precipitation gradient (537-1036 mm per year) in the Yucatan Peninsula, Mexico. Specifically we sought to test the hypothesis that leaf traits of dominant plant species shift along a precipitation gradient, but are affected by soil N cycling. Although variation among different species within each site explains some leaf trait variation, there is also a high level of variability across sites, suggesting that factors other than precipitation regime more strongly influence leaf traits. Principal component analyses indicated that across sites and tree species, covariation in leaf traits is an indicator of soil N availability. Patterns of natural abundance (15)N in foliage and foliage minus soil suggest that variation in precipitation regime drives a shift in plant N acquisition and the openness of the N cycle. Overall, our study shows that both plant species and site are important determinants of leaf traits, and that the leaf trait spectrum is correlated with soil N cycling.

Estimating trophic structures is a common approach used to retrieve information regarding energy pathways, predation, and competition in complex ecosystems. The application of amino acid (AA) compound-specific nitrogen (N) isotope analysis (CSIA) is a relatively new method used to estimate trophic position (TP) and feeding relationships in diverse organisms. Here, we conducted the first meta-analysis of δ(15)N AA values from measurements of 359 marine species covering four trophic levels, and compared TP estimates from AA-CSIA to literature values derived from food items, gut or stomach content analysis. We tested whether the AA trophic enrichment factor (TEF), or the (15)N enrichment among different individual AAs is constant across trophic levels and whether inclusion of δ(15)N values from multiple AAs improves TP estimation. For the TEF of glutamic acid relative to phenylalanine (Phe) we found an average value of 6.6‰ across all taxa, which is significantly lower than the commonly applied 7.6‰. We found that organism feeding ecology influences TEF values of several trophic AAs relative to Phe, with significantly higher TEF values for herbivores compared to omnivores and carnivores, while TEF values were also significantly lower for animals excreting urea compared to ammonium. Based on the comparison of multiple model structures using the metadata of δ(15)N AA values we show that increasing the number of AAs in principle improves precision in TP estimation. This meta-analysis clarifies the advantages and limitations of using individual δ(15)N AA values as tools in trophic ecology and provides a guideline for the future application of AA-CSIA to food web studies.

While ecologists primarily focus on the immediate impact of ecological subsidies, understanding the importance of ecological subsidies requires quantifying the long-term temporal dynamics of subsidies on recipient ecosystems. Deciduous leaf litter transferred from terrestrial to aquatic ecosystems exerts both immediate and lasting effects on stream food webs. Recently, deciduous leaf additions have also been shown to be important subsidies for planktonic food webs in ponds during autumn; however, the inter-seasonal effects of autumn leaf subsidies on planktonic food webs have not been studied. We hypothesized that autumn leaf drop will affect the spring dynamics of freshwater pond food webs by altering the availability of resources, water transparency, and the metabolic state of ponds. We created leaf-added and no-leaf-added field mesocosms in autumn 2012, allowed mesocosms to ice-over for the winter, and began sampling the physical, chemical, and biological properties of mesocosms immediately following ice-off in spring 2013. At ice-off, leaf additions reduced dissolved oxygen, elevated total phosphorus concentrations and dissolved materials, and did not alter temperature or total nitrogen. These initial abiotic effects contributed to higher bacterial densities and lower chlorophyll concentrations, but by the end of spring, the abiotic environment, chlorophyll and bacterial densities converged. By contrast, zooplankton densities diverged between treatments during the spring, with leaf additions stimulating copepods but inhibiting cladocerans. We hypothesized that these differences between zooplankton orders resulted from resource shifts following leaf additions. These results suggest that leaf subsidies can alter both the short- and long-term dynamics of planktonic food webs, and highlight the importance of fully understanding how ecological subsidies are integrated into recipient food webs.

Previous studies comparing invaded and non-invaded sites suggest that cheatgrass (Bromus tectorum L.) causes soil N cycling to increase. Unfortunately, these correlative studies fail to distinguish whether cheatgrass caused the differences in N cycling, or if cheatgrass simply invaded sites where N availability was greater. We measured soil C and N concentrations and net and gross N-cycling rates on 24-year-old replicated field plots in a sagebrush-steppe ecosystem that had been plowed, fumigated, and seeded to different plant communities in 1984. Laboratory assays of soil collected throughout the soil profiles (0-60 cm) showed that soil NO3 (-), organic C and N, and net N mineralization, net nitrification, and soil respiration rates were all greater beneath cheatgrass than in sagebrush-perennial grass plots. In surface soils (0-10 cm), field and lab assays on five sampling dates during 2 years showed gross N mineralization, net N mineralization, and net nitrification rates were all faster beneath cheatgrass than in sagebrush-perennial grass plots. Modeling analyses based on soil respiration and gross N-cycling rates suggest that cheatgrass provides soil microbes with lower C:N substrates and that this could explain the faster N-cycling rates beneath cheatgrass. This is the first long-term replicated field study to conclusively show that cheatgrass created greater soil organic N pool sizes and stimulated N-cycling rates compared to similar-aged stands of sagebrush and native perennial grasses. Increased N-cycling rates may represent a positive plant-soil feedback that promotes continued dominance by cheatgrass, even in the absence of soil disturbance or fire.

Tree species diversity has been reported to increase forest ecosystem above-ground biomass and productivity, but little is known about below-ground biomass and production in diverse mixed forests compared to single-species forests. For testing whether species richness increases below-ground biomass and production and thus complementarity between forest tree species in young stands, we determined fine root biomass and production of trees and ground vegetation in two experimental plantations representing gradients in tree species richness. Additionally, we measured tree fine root length and determined species composition from fine root biomass samples with the near-infrared reflectance spectroscopy method. We did not observe higher biomass or production in mixed stands compared to monocultures. Neither did we observe any differences in tree root length or fine root turnover. One reason for this could be that these stands were still young, and canopy closure had not always taken place, i.e. a situation where above- or below-ground competition did not yet exist. Another reason could be that the rooting traits of the tree species did not differ sufficiently to support niche differentiation. Our results suggested that functional group identity (i.e. conifers vs. broadleaved species) can be more important for below-ground biomass and production than the species richness itself, as conifers seemed to be more competitive in colonising the soil volume, compared to broadleaved species.

A functional traits-based theory of organismal communities is critical for understanding the principles underlying community assembly, and predicting responses to environmental change. This is particularly true for terrestrial arthropods, of which only 20% are described. Using epigaeic ant assemblages, we asked: (1) can we use morphological variation among species to predict trophic position or preferred microhabitat; (2) does the strength of morphological associations suggest recent trait divergence; (3) do environmental variables at site scale predict trait sets for whole assemblages? We pitfall-trapped ants from a revegetation chronosequence and measured their morphology, trophic position [using C:N stoichiometry and stable isotope ratios (δ)] and characteristics of microhabitat and macrohabitat. We found strong associations between high trophic position (low C:N and high δ(15)N) in body tissue and morphological traits: predators were larger, had more laterally positioned eyes, more physical protection and tended to be monomorphic. In addition, morphological traits were associated with certain microhabitat features, e.g. smaller heads were associated with the bare ground microhabitat. Trait-microhabitat relationships were more pronounced when phylogenetic adjustments were used, indicating a strong influence of recent trait divergences. At the assemblage level, our fourth corner analysis revealed associations between the prevalence of traits and macrohabitat, although these associations were not the same as those based on microhabitat associations. This study shows direct links between species-level traits and both diet and habitat preference. Trait-based prediction of ecological roles and community structure is thus achievable when integrating stoichiometry, morphology and phylogeny, but scale is an important consideration in such predictions.

Anthropogenic eutrophication impacts ecosystems worldwide. Here, we use a vegetation dataset from semi-natural grasslands on acidic soils sampled along a gradient in north-western Europe to examine the response of species frequency to nitrogen (N) deposition, controlling for the effects of other environmental variables. A second dataset of acidic grasslands from Germany and the Netherlands containing plots from different time periods was analysed to examine whether the results of the spatial gradient approach coincided with temporal changes in the abundance of species. Out of 44 studied species, 16 were affected by N deposition, 12 of them negatively. Soil pH and phosphorus (P) influenced 24 and 14 species, respectively, predominantly positively. Fewer species were related to the soil contents of NO3(-) or NH4(+), with no significant differences between the number of positive and negative effects. Whereas the temporal change of species was unrelated to their responses to pH, species responding negatively to N deposition, soil P and NO3(-) showed a significant decline over time in both countries. Species that were negatively affected by high N deposition and/or high soil P also showed a negative temporal trend and could be characterised by short stature and slow growth. The results confirm the negative role of N deposition for many plant species in semi-natural acidic grasslands. The negative temporal trends of species sensitive to high N deposition and soil P values clearly show a need for maintaining low soil nutrient status and for restoring the formerly infertile conditions in nutrient-enriched grasslands.

In northern forests, large amounts of missing N that dominate N balances at scales ranging from small watersheds to large regional drainage basins may be related to N-gas production by soil microbes. We measured denitrification rates in forest soils in northeastern North America along a N deposition gradient to determine whether N-gas fluxes were a significant fate for atmospheric N inputs and whether denitrification rates were correlated with N availability, soil O2 status, or forest type. We quantified N2 and N2O fluxes in the laboratory with an intact-core method and monitored soil O2, temperature and moisture in three forests differing in natural and anthropogenic N enrichment: Turkey Lakes Watershed, Ontario; Hubbard Brook Experimental Forest, New Hampshire; and Bear Brook Watershed, Maine (fertilized and reference plots in hardwood and softwood stands). Total N-gas flux estimates ranged from <1 in fertilized hardwood uplands at Bear Brook to >100 kg N ha(-1) year(-1) in hardwood wetlands at Turkey Lakes. N-gas flux increased systematically with natural N enrichment from soils with high nitrification rates (Bear Brook < Hubbard Brook < Turkey Lakes) but did not increase in the site where N fertilizer has been added since 1989 (Bear Brook). Our results show that denitrification is an important and underestimated term (1-24% of atmospheric N inputs) in N budgets of upland forests in northeastern North America, but it does not appear to be an important sink for elevated anthropogenic atmospheric N deposition in this region.

Atmospheric inputs of N and S in bulk deposition (open collectors) and throughfall (beneath canopy collectors) were measured in and adjacent to two Class 1 wilderness areas of the northeastern US. In general, atmospheric S inputs followed our expectations with throughfall S fluxes increasing with elevation in the White Mountains, New Hampshire and throughfall S fluxes being greater in coniferous than deciduous stands in both sites. In contrast, throughfall N fluxes decreased significantly with elevation. Throughfall NO3 (-) fluxes were greater in coniferous than deciduous stands of Lye Brook, Vermont, but were greater in deciduous than coniferous stands of the White Mountains. We found overlap in the range of values for atmospheric N inputs between our measurements and monitoring data [National Atmospheric Deposition Program (NADP) and Clean Air Status and Trends Network (CASTNET)] for wet and total (wet + dry) deposition at Lye Brook. However, our measurements of total S deposition in the White Mountains and bulk (wet) deposition at both Lye Brook and the White Mountains were significantly lower than NADP plus CASTNET, and NADP data, respectively. Natural abundance (18)O in throughfall and bulk deposition were not significantly different, suggesting that there was no significant biological production of [Formula: see text] via nitrification in the canopy. NO3 (-) concentrations in streams were low and had natural abundance (18)O values consistent with microbial production, demonstrating that atmospheric N is being biologically transformed while moving through these watersheds and that these forested watersheds are unlikely to be N saturated.

Research in warm-climate biomes has shown that invasion by symbiotic dinitrogen (N2)-fixing plants can transform ecosystems in ways analogous to the transformations observed as a consequence of anthropogenic, atmospheric nitrogen (N) deposition: declines in biodiversity, soil acidification, and alterations to carbon and nutrient cycling, including increased N losses through nitrate leaching and emissions of the powerful greenhouse gas nitrous oxide (N2O). Here, we used literature review and case study approaches to assess the evidence for similar transformations in cold-climate ecosystems of the boreal, subarctic and upper montane-temperate life zones. Our assessment focuses on the plant genera Lupinus and Alnus, which have become invasive largely as a consequence of deliberate introductions and/or reduced land management. These cold biomes are commonly located in remote areas with low anthropogenic N inputs, and the environmental impacts of N2-fixer invasion appear to be as severe as those from anthropogenic N deposition in highly N polluted areas. Hence, inputs of N from N2 fixation can affect ecosystems as dramatically or even more strongly than N inputs from atmospheric deposition, and biomes in cold climates represent no exception with regard to the risk of being invaded by N2-fixing species. In particular, the cold biomes studied here show both a strong potential to be transformed by N2-fixing plants and a rapid subsequent saturation in the ecosystem's capacity to retain N. Therefore, analogous to increases in N deposition, N2-fixing plant invasions must be deemed significant threats to biodiversity and to environmental quality.

Global change phenomena, such as forest disturbance and land-use change, significantly affect elemental balances as well as the structure and function of terrestrial ecosystems. However, the importance of shifts in soil nutrient stoichiometry for the regulation of belowground biota and soil food webs have not been intensively studied for tropical ecosystems. In the present account, we examine the effects of land-use change and soil depth on soil and microbial stoichiometry along a land-use sequence (natural forest, pastures of different ages, secondary succession) in the tropical mountain rainforest region of southern Ecuador. Furthermore, we analyzed (PLFA-method) whether shifts in the microbial community structure were related to alterations in soil and microbial stoichiometry. Soil and microbial stoichiometry were affected by both land-use change and soil depth. After forest disturbance, significant decreases of soil C:N:P ratios at the pastures were followed by increases during secondary succession. Microbial C:N ratios varied slightly in response to land-use change, whereas no fixed microbial C:P and N:P ratios were observed. Shifts in microbial community composition were associated with soil and microbial stoichiometry. Strong positive relationships between PLFA-markers 18:2n6,9c (saprotrophic fungi) and 20:4 (animals) and negative associations between 20:4 and microbial N:P point to land-use change affecting the structure of soil food webs. Significant deviations from global soil and microbial C:N:P ratios indicated a major force of land-use change to alter stoichiometric relationships and to structure biological systems. Our results support the idea that soil biotic communities are stoichiometrically flexible in order to adapt to alterations in resource stoichiometry.

In terrestrial angiosperms, defense and resistance mechanisms against herbivores have been studied extensively; yet this topic is poorly understood in aquatic angiosperms. We investigated induced response mechanisms in Myriophyllum spicatum to the generalist insect herbivore Acentria ephemerella in three independent experiments. Various morphological and chemical response variables were examined in grazed apical shoots and compared to undamaged controls. We further estimated plant palatability of induced and non-induced apices in choice assays, and assessed the growth response of Acentria larvae in no-choice feeding assays. Leaves of induced apices were splayed out horizontally and changed in color from green to red. The dry matter content and thus plant toughness increased by up to 19%, but silica levels stayed constant. Induced apices exhibited a decline in chlorophyll content of up to 34%, reflected also by a 10% decrease in nitrogen levels, while nitrogen increased by 14% in lower parts of grazed shoots. Also, herbivore-deterring total phenolic compounds increased by up to 20% in apices. In choice trials, Acentria larvae strongly avoided grazed tips, and growth was reduced by 25% on induced apices. In total, we observed five different induced resistance and defensive traits in grazed apices: changes in appearance, increased plant toughness, delocalization of N-containing metabolites, increased polyphenols, and reduced nutritional value. The observed changes prevent herbivore damage and loss of apical tissue, which are most valuable for plant fitness. Our study presents the first evidence of multiple, parallel defense strategies including constitutive and induced defense mechanisms in a freshwater angiosperm.

Young larval stages of many organisms represent bottlenecks in the life-history of many species. The high mortality commonly observed in, for example, decapod larvae has often been linked to poor nutrition, with most studies focussing on food quantity. Here, we focus instead on the effects of quality and have investigated its effects on the nutritional condition of lobster larvae. We established a tri-trophic food chain consisting of the cryptophyte Rhodomonas salina, the calanoid copepod Acartia tonsa and larvae of the European lobster Homarus gammarus. In a set of experiments, we manipulated the C:N:P stoichiometry of the primary producers, and accordingly those of the primary consumer. In a first experiment, R. salina was grown under N- and P-limitation and the nutrient content of the algae was manipulated by addition of the limiting nutrient to create a food quality gradient. In a second experiment, the effect on lobster larvae of long- and short-term exposure to food of varying quality during ontogenetic development was investigated. The condition of the lobster larvae was negatively affected even by subtle N- and P-nutrient limitations of the algae. Furthermore, younger lobster larvae were more vulnerable to nutrient limitation than older ones, suggesting an ontogenetic shift in the capacity of lobster larvae to cope with low quality food. The results presented here might have substantial consequences for the survival of lobster larvae in the field, as, in the light of future climate change and re-oligotrophication of the North Sea, lobster larvae might face marked changes in temperature and nutrient conditions, thus significantly altering their condition and growth.

Drought-induced events of massive tree mortality appear to be increasing worldwide. Species-specific vulnerability to drought mortality may alter patterns of species diversity and affect future forest composition. We have explored the consequences of the extreme drought of 2005, which caused high sapling mortality (approx. 50 %) among 10-year-old saplings of two coexisting pine species in the Mediterranean mountains of Sierra Nevada (Spain): boreo-alpine Pinus sylvestris and Mediterranean P. nigra. Sapling height growth, leaf δ(13)C and δ(18)O, and foliar nitrogen concentration in the four most recent leaf cohorts were measured in dead and surviving saplings. The foliar isotopic composition of dead saplings (which reflects time-integrated leaf gas-exchange until mortality) displayed sharp increases in both δ(13)C and δ(18)O during the extreme drought of 2005, suggesting an important role of stomatal conductance (g(s)) reduction and diffusional limitations to photosynthesis in mortality. While P. nigra showed decreased growth in 2005 compared to the previous wetter year, P. sylvestris maintained similar growth levels in both years. Decreased growth, coupled with a sharper increase in foliar δ(18)O during extreme drought in dead saplings, indicate a more conservative water use strategy for P. nigra. The different physiological behavior of the two pine species in response to drought (further supported by data from surviving saplings) may have influenced 2005 mortality rates, which contributed to 2.4-fold greater survival for P. nigra over the lifespan of the saplings. This species-specific vulnerability to extreme drought could lead to changes in dominance and distribution of pine species in Mediterranean mountain forests.

Global warming and excess nitrogen deposition can exert strong impacts on aquatic populations, communities, and ecosystems. However, experimental data to establish clear cause-and-effect relationships in naturally complex field conditions are scarce in aquatic environments. Here, we describe the design and performance of a unique outdoor enclosure facility used to simulate warming, increased nitrogen supply, and both factors combined in a littoral freshwater wetland dominated by common reed, Phragmites australis. The experimental system effectively simulated a 2.8 °C climate warming scenario over an extended period, capturing the natural temperature variations in the wetland at diel and seasonal scales with only small deviations. Excess nitrogen supply enhanced nitrate concentrations especially in winter when it was associated with increased concentration of ammonium and dissolved organic carbon. Nitrogen also reduced dissolved oxygen concentrations, particularly in the summer. Importantly, by stimulating biological activity, warming enhanced the nitrogen uptake capacity of the wetland during the winter, emphasizing the need for multifactorial global change experiments that examine both warming and nitrogen loading in concert. Establishing similar experiments across broad environmental gradients holds great potential to provide robust assessments of the impacts of climate change on shallow aquatic ecosystems.

The importance of rock-derived mineral nutrients (P, K, Mn, Mg, and Ca) in plant physiological function is well established. However, one important and relatively unexplored question is whether or not the same rules of plant nutrient use efficiency apply to these essential elements even if they are not limiting to primary production. We examined conifer growth and nutrient use dynamics across sites with contrasting geologies (sedimentary and volcanic) that vary in both rock-derived mineral nutrient and N availability. Differences in bedrock geochemistry generally corresponded to differences in available soil nutrients, such that the volcanic site tended to have greater available nutrients. Foliar nutrient concentrations reflected both differences in bedrock chemistry and indices of available soil nutrients for P, K, and Mn. Aboveground biomass production did not follow expected patterns and was greater for trees growing on low nutrient sites, but only with respect to the annual woody increment. Fine litter production did not differ between sites. Finally, we found evidence for trade-offs between two commonly examined components of nutrient use efficiency (NUE): nutrient productivity (A n) and mean residence time of nutrients. However, we did not find evidence for higher plant NUE in soils with lower nutrient availability for N or rock-derived nutrients.

Estimating the nutritional value of a herbivore's diet is difficult because it requires knowing what the animal eats, the relative quality of each component and how these components interact in relation to animal physiology. Current methods are cumbersome and rely on many assumptions that are hard to evaluate. We describe a new method for estimating relative diet quality directly from faeces that avoids the problems inherent in other methods. We combine this method with near infrared reflectance spectroscopy (NIRS) to analyse many samples and thus provide a technique with immense value in ecological studies. The method stems from the correlation between the concentrations of dietary and faecal nitrogen in herbivores eating a tannin-free diet, but a weaker relationship in browsers that ingest substantial amounts of tannins, which form complexes with proteins. These complexes reduce the availability of nitrogen and may increase faecal nitrogen concentrations. Using the tannin-binding compound, polyethylene glycol, we showed that tannin-bound nitrogen is a significant and variable part of faecal nitrogen in wild common brushtail possums (Trichosurus vulpecula). We developed a technique to measure faecal available nitrogen and found that it predicted the reproductive success of female brushtail possums in northern Australia. Faecal available nitrogen combined with NIRS provides a powerful tool for estimating the relative nutritional value of the diets of browsing herbivores in many ecological systems. It is a better indicator of diet quality than other commonly used single-nutrient measures such as faecal nitrogen and foliage analysis paired with observed feeding behaviour.

Both atmospheric [CO2] and average surface temperatures are predicted to increase with potentially different, additive or opposing, effects on leaf quality and insect herbivore activity. Few studies have directly measured the interactive effects of concurrent changes in [CO2] and temperature on insect herbivores. None have done so over the entire developmental period of a tree-feeding insect, and none have compared responses to low pre-industrial [CO2] and present day [CO2] to estimate responses to future increases. Eucalypt herbivores may be particularly sensitive to climate-driven shifts in plant chemistry, as eucalypt foliage is naturally low in [N]. In this study, we assessed the development of the eucalypt herbivore Doratifera quadriguttata exposed concurrently to variable [CO2] (290, 400, 650 μmol mol(-1)) and temperature (ambient, ambient +4 °C) on glasshouse-grown Eucalyptus tereticornis. Overall, insects performed best on foliage grown at pre-industrial [CO2], indicating that modern insect herbivores have already experienced nutritional shifts since industrialisation. Rising [CO2] increased specific leaf mass and leaf carbohydrate concentration, subsequently reducing leaf [N]. Lower leaf [N] induced compensatory feeding and impeded insect performance, particularly by prolonging larval development. Importantly, elevated temperature dampened the negative effects of rising [CO2] on larval performance. Therefore, rising [CO2] over the past 200 years may have reduced forage quality for eucalypt insects, but concurrent temperature increases may have partially compensated for this, and may continue to do so in the future. These results highlight the importance of assessing plant-insect interactions within the context of multiple climate-change factors because of the interactive and potentially opposing effects of different factors within and between trophic levels.

We examined geographic variation in the structure and function of salt marsh communities along the Atlantic and Gulf coasts of the United States. Focusing on the arthropod community in the dominant salt marsh plant Spartina alterniflora, we tested two hypotheses: first, that marsh community structure varies geographically, and second, that two aspects of marsh function (response to eutrophication and addition of dead plant material) also vary geographically. We worked at eleven sites on the Gulf Coast and eleven sites on the Atlantic Coast, dividing each coast up into two geographic areas. Abiotic conditions (tidal range, soil organic content, and water content, but not soil salinity), plant variables (Spartina nitrogen content, height, cover of dead plant material, but not live Spartina percent cover or light interception), and arthropod variables (proportional abundances of predators, sucking herbivores, stem-boring herbivores, parasitoids, and detritivores, but not total arthropod numbers) varied among the four geographic regions. Latitude and mean tidal range explained much of this geographic variation. Nutrient enrichment increased all arthropod functional groups in the community, consistent with previous experimental results, and had similar effects in all geographic regions, contrary to our hypothesis, suggesting widespread consistency in this aspect of ecosystem function. The addition of dead plant material had surprisingly little effect on the arthropod community. Our results caution against the uncritical extrapolation of work done in one geographic region to another, but indicate that some aspects of marsh function may operate in similar ways in different geographic regions, despite spatial variation in community structure.

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China's West Development.

Partial mycoheterotrophy, a newly discovered form of mixotrophy in plants, has been described in at least two major lineages of angiosperms, the orchids and ericaceous plants in the tribe Pyroleae. Partial mycoheterotrophy entails carbon gains both directly from photosynthesis and via symbiotic mycorrhizal fungi, but determining the degree of plant dependence on fungal carbon is challenging. The purpose of this study was to determine if two chlorophyllous species of Pyroleae, Chimaphila umbellata and Pyrola picta, were receiving carbon via mycorrhizal networks and, if so, if their proportional dependency on fungal carbon gains increased under reduced light conditions. This was accomplished by a field experiment that manipulated light and plants' access to mycorrhizal networks, and by using the stable carbon isotope composition (δ(13)C) of leaf soluble sugars as a marker for the level of mycoheterotrophy. Based on leaf soluble sugars δ(13)C values, we calculated a site-independent isotope enrichment factor as a measure of fungal contributions to plant C. We found that, under each treatment and over time, the two test species demonstrated different isotopic responses caused by their different intrinsic physiologies. Our data, along with previously published studies, suggest that Chimaphila umbellata is primarily an autotrophic understory plant, while Pyrola picta may be capable of partial mycoheterotrophy. However, in this study, a 50% decrease in light availability did not significantly change the relative dependency of P. picta on carbon gains via mycoheterotrophy.

Ecologists want to explain why populations of animals are not evenly distributed across landscapes and often turn to nutritional explanations. In seeking to link population attributes with food quality, they often contrast nutritionally positive traits, such as the concentration of nitrogen, against negative ones, such as fibre concentration, by using a ratio of these traits. This specific ratio has attracted attention because it sometimes correlates with the biomass of colobine primates across sites in Asia and Africa. Although empirically successful, we have identified problems with the ratio that may explain why it fails under some conditions to predict colobine biomass. First, available nitrogen, rather than total nitrogen, is nutritionally important, while the presence of tannins is the major factor reducing the availability of nitrogen in browse plant species. Second, tannin complexes inflate measures of fibre. Finally, simple ratios may be unsound statistically because they implicitly assume isometric relationships between variables. We used data on the chemical composition of plants from three continents to examine the relationships between the concentrations of nitrogen, available nitrogen, fibre and tannins in foliage and to evaluate the nitrogen to fibre ratio. Our results suggest that the ratio of the concentration of nitrogen to fibre in leaves does sometimes closely correlate with the concentration of available nitrogen. However, the ratio may give misleading results when leaves contain high concentrations of tannins. The concentration of available nitrogen, which incorporates measures of total nitrogen, dry matter digestibility and tannins, should give a better indication of the nutritional value of leaves for herbivorous mammals that can readily be extrapolated to habitats.

This study evaluates acclimation of photosynthesis and stomatal conductance in three evolutionary lineages of C(3), C(3)-C(4) intermediate, and C(4) species grown in the low CO(2) and hot conditions proposed to favo r the evolution of C(4) photosynthesis. Closely related C(3), C(3)-C(4), and C(4) species in the genera Flaveria, Heliotropium, and Alternanthera were grown near 380 and 180 μmol CO(2) mol(-1) air and day/night temperatures of 37/29°C. Growth CO(2) had no effect on photosynthetic capacity or nitrogen allocation to Rubisco and electron transport in any of the species. There was also no effect of growth CO(2) on photosynthetic and stomatal responses to intercellular CO(2) concentration. These results demonstrate little ability to acclimate to low CO(2) growth conditions in closely related C(3) and C(3)-C(4) species, indicating that, during past episodes of low CO(2), individual C(3) plants had little ability to adjust their photosynthetic physiology to compensate for carbon starvation. This deficiency could have favored selection for more efficient modes of carbon assimilation, such as C(3)-C(4) intermediacy. The C(3)-C(4) species had approximately 50% greater rates of net CO(2) assimilation than the C(3) species when measured at the growth conditions of 180 μmol mol(-1) and 37°C, demonstrating the superiority of the C(3)-C(4) pathway in low atmospheric CO(2) and hot climates of recent geological time.

Seasonal oscillations in the carbon (δ(13)C) and nitrogen (δ(15)N) isotope signatures of aquatic algae can cause seasonal enrichment-depletion cycles in the isotopic composition of planktonic invertebrates (e.g., copepods). Yet, there is growing evidence that seasonal enrichment-depletion cycles also occur in the isotope signatures of larger invertebrate consumers, taxa used to define reference points in isotope-based trophic models (e.g., trophic baselines). To evaluate the general assumption of temporal stability in non-zooplankton aquatic invertebrates, δ(13)C and δ(15)N time series data from the literature were analyzed for seasonality and the influence of biotic (feeding group) and abiotic (trophic state, climate regime) factors on isotope temporal patterns. The amplitude of δ(13)C and δ(15)N enrichment-depletion cycles was negatively related to body size, although all size-classes of invertebrates displayed a winter-to-summer enrichment in δ(13)C and depletion in δ(15)N. Among feeding groups, periphytic grazers were more variable and displayed larger temporal changes in δ(13)C than detritivores. For nitrogen, temporal variability and magnitude of directional change of δ(15)N was most strongly related to ecosystem trophic state (eutrophic > mesotrophic, oligotrophic). This study provides evidence of seasonality in the isotopic composition of aquatic invertebrates across very broad geographical and ecological gradients as well as identifying factors that are likely to modulate the strength and variability of seasonality. These results emphasize the need for researchers to recognize the likelihood of temporal changes in non-zooplankton aquatic invertebrate consumers at time scales relevant to seasonal studies and, if present, to account for temporal dynamics in isotope trophic models.

Although the productivity and nitrogen (N)-use traits of mire plants differ dramatically between fens and bogs, soil N richness does not necessarily differ, whereas the soil-water pH is distinctly lower in bogs than in fens. The ecophysiological mechanisms underlying these relations are unclear. To assess the relative availability of N forms in relation to soil-water pH, we focused on the net N uptake rate per unit root weight (NNUR), glutamine synthetase activity and nitrate reductase activity, and performed reciprocal transplant experiments with the seedlings of fen (Carex lyngbyei) and bog (C. middendorffii) sedge species in intact habitat sites. The soil-water pH was clearly lower at the bog site, but the NH(4) (+), NO(3) (-) or dissolved organic-N concentrations did not differ between the fen and bog sites. The activity of both enzymes for inorganic-N assimilation did not differ among the sites and species. However, the fen species grown at bog sites showed a drastic decrease in the NNUR, suggesting a suppression of organic-N uptake. The bog species showed no NNUR difference between the sites. These results indicate that inorganic-N availability does not differ between the two habitats, but organic-N availability is lowered in a low-pH bog, particularly in the case of fen species. Therefore, the relative availability of N forms shows species-specific variations that depend on the differences in the soil-water pH of root zone, even at similar N richness, which would play a key role in plant distribution strategies in relation to the fen-bog gradient.

Global climate change is expected to affect terrestrial ecosystems in a variety of ways. Some of the more well-studied effects include the biogeochemical feedbacks to the climate system that can either increase or decrease the atmospheric load of greenhouse gases such as carbon dioxide and nitrous oxide. Less well-studied are the effects of climate change on the linkages between soil and plant processes. Here, we report the effects of soil warming on these linkages observed in a large field manipulation of a deciduous forest in southern New England, USA, where soil was continuously warmed 5°C above ambient for 7 years. Over this period, we have observed significant changes to the nitrogen cycle that have the potential to affect tree species composition in the long term. Since the start of the experiment, we have documented a 45% average annual increase in net nitrogen mineralization and a three-fold increase in nitrification such that in years 5 through 7, 25% of the nitrogen mineralized is then nitrified. The warming-induced increase of available nitrogen resulted in increases in the foliar nitrogen content and the relative growth rate of trees in the warmed area. Acer rubrum (red maple) trees have responded the most after 7 years of warming, with the greatest increases in both foliar nitrogen content and relative growth rates. Our study suggests that considering species-specific responses to increases in nitrogen availability and changes in nitrogen form is important in predicting future forest composition and feedbacks to the climate system.