Increased Litter Mass Research Articles

Litter accumulation drives grassland plant community composition and functional diversity via leaf traits

Direct physical interference from litter is an important mechanism driving changes in the diversity and composition of plant communities dominated by perennial species. While the consequences of litter for species diversity and composition are well known, the mechanisms shaping how litter acts as a filter on leaf and shoot functional traits to alter community structure and functional diversity remain unclear. These questions were examined using a large field experiment in native mixed-grass prairie grasslands in the brown and dark brown soil zones in central Saskatchewan, Canada. Litter additions ranging from 0 to 2,290 g m−2 were applied to plots in a response surface design, and responses including community structure, above-ground functional diversity, and community-weighted mean values for leaf functional traits were measured. Species richness, functional richness, and functional dispersion declined with increasing litter mass, but functional evenness increased. Community composition became more variable because increased litter acted as a filter on the perennial species previously present in the plots; the remaining species in a given plot were the most litter-tolerant subset of the original community in the plot. Increased litter mass affected graminoid and forb species differently. Increased litter mass selected for graminoid species characterized by greater height, leaf area, specific leaf area (SLA), leaf thickness, and leaf tissue density. In contrast, increased litter selected for forb species characterized by larger, thicker leaves, lower SLA, and lower leaf tissue density. Forbs with cauline leaves were also selected for over acaulescent growth forms. Physical interference by litter likely reduces grassland diversity and alters species composition in diverse grassland communities by forcing convergence in community trait structure, but not convergence in species composition.

Interactive responses of grass litter decomposition to warming, nitrogen addition and detritivore access in a temperate old field.

Plant litter decomposition has been studied extensively in the context of both climate warming and increased atmospheric N deposition. However, much of this research is based on microbial responses, despite the potential for detritivores to contribute substantially to litter breakdown. We measured litter mass-loss responses to the combined effects of warming, N addition and detritivore access in a grass-dominated old field. We concurrently assessed the roles of litter treatment origin vs. microenvironment (direct warming and N-addition effects) to elucidate the mechanisms through which these factors affect decomposition. After 6 weeks, mass loss increased in N-addition plots, and it increased with detritivore access in the absence of warming. After 1 year, warming, N addition, and detritivore access all increased litter mass loss, although the effects of N addition and warming were non-additive in the detritivore-access plots. For the litter-origin experiment, mass loss after 6 weeks increased in litter from N-addition plots and warmed plots, but unlike the overall decomposition response, the N-addition effect was enhanced by detritivore access. Conversely, for the microenvironment experiment, detritivore access only increased mass loss in unfertilized plots. After 1 year, detritivore access increased mass loss in the litter-origin and microenvironment experiments, but there were no warming or N-addition effects. Overall, our results provide support for a substantial role of detritivores in promoting litter mass loss in our system. Moreover, they reveal important interactions between litter origin, microclimate and detritivores in determining decomposition responses to global change.

Top-Down Regulation of Litter Invertebrates by a Terrestrial Salamander

Terrestrial plethodontid salamanders are abundant predators within the forest floor litter of eastern North America, and are hypothesized to regulate soil and litter invertebrate density and species composition. I tested this hypothesis during a 6-yr study of the effects of the Eastern Red-backed Salamander (Plethodon cinereus) on the invertebrate community of a forest site in northeast Ohio. Salamander surface density, invertebrate abundance, and community composition were monitored within 30 open, circular plots. Variation in plot occupancy by P. cinereus was achieved by supplying plots with differing amounts of artificial cover (0, 1, or 4 ceramic tiles) that served as refuges for the salamanders. Salamander plot occupancy, invertebrate density, leaf litter mass, and leaf litter moisture were quantified each spring and fall from 2003 through 2008. Statistically significant effects of salamander plot-occupancy on invertebrate densities were found for several taxa of mesofauna, including several Collembola taxa, oribatid mites, pseudoscorpions, and psocoptera. The strength and direction of salamander effects varied among taxa and included negative, positive, and no effects on invertebrate densities. The magnitude and sign of salamander effects on invertebrate densities were predicted by seasonal and interannual variation in leaf litter mass and, to a lesser extent, litter moisture content. Salamander effects decreased with increasing litter mass and were more often negative when litter mass was high, whereas positive effects on invertebrate densities were more likely when litter mass was low. For several taxa, the positive effect of P. cinereus also increased with litter moisture. I propose two mechanistic hypotheses for these dynamics that integrate behavioral ecology of salamander prey selection and territorial defense with variation in litter mass and litter moisture.

Response of leaf, sheath and stem nutrient resorption to 7 years of N addition in freshwater wetland of Northeast China

Background and Aims Increased N availability induced by agricultural fertilization applications and atmospheric N deposition may affect plant nutrient resorption in temperate wetlands. However, the relationship between nutrient resorption and N availability is still unclear, and most studies have focused on leaf nutrient resorption only. The aim of our study was to examine the response of leaf and non-leaf organ nutrient resorption to N enrichment in a temperate freshwater wetland. Methods We conducted a 7-year N addition experiment to investigate the effects of increased N loading on leaf, sheath and stem nutrient (N and P) resorption of two dominant species (Deyeuxia angustifolia and Glyceria spiculosa )i n af reshwater marsh in the Sanjiang Plain, Northeast China. Results Our results showed that, for both leaf and nonleaf organs (sheath and stem), N addition decreased N resorption proficiency and hence increased litter N concentration. Moreover, the magnitude of N addition effect on N resorption proficiency varied with fertilization rates for D. angustifolia sheaths and stems, and G. spiculosa leaves. However, increased N loading produced inconsistent impacts on N and P resorption efficiencies and P resorption proficiency, and the effects only varied with species and plant organs. In addition, N enrichment increased litter mass and altered litter allocation among leaf, sheath and stem. Conclusions Our results highlight that leaf and nonleaf organs respond differentially to N addition regarding N and P resorption efficiencies and P resorption proficiency, and also suggest that N enrichment in temperate freshwater wetlands would alter plant internal nutrient cycles and increase litter quality and quantity, and thus substantially influence ecosystem carbon and nutrient cycles.

Effect of nitrogen addition on decomposition of Calamagrostis angustifolia litters from freshwater marshes of Northeast China

Wetland ecosystems store a large amount of organic carbon (C) in soils, due to the slow decomposition rates of plant litter and soil organic matter. Increased nitrogen (N) availability induced by human activities and global warming may accelerate litter decomposition and affect soil organic C dynamics in wetlands. In the present study, we investigated the effect of N addition on decomposition of Calamagrostis angustifolia litters from freshwater marshes in the Sanjiang Plain of Northeast China under field and laboratory conditions. First, we assessed the changes in initial litter chemical composition and subsequent decomposition following three years of N addition at the rate of 24 g N m −2 year −1 under field conditions. Our results showed that N addition increased litter N concentration and decreased C/N ratio, and thus stimulated litter decomposition. Secondly, we examined the effect of increased N availability (0, 25, 50 and 100 mg N g −1 litter) on litter decomposition under laboratory conditions. Increased exterior N availability also enhanced microbial respiration and increased litter mass loss under both waterlogging and non-waterlogging conditions. In addition, waterlogging conditions inhibited microbial respiration and suppressed litter mass loss. These findings demonstrated that N addition increased litter decomposition rates through improved litter quality and enhanced microbial activity in freshwater marshes of Northeast China. This implies that increased N availability accelerates litter decomposition rates, and thus may cause substantial losses of soil C and diminish and even reverse the C sink function of wetlands in the Sanjiang Plain of Northeast China.

Carbon quality rather than stoichiometry controls litter decomposition in a tropical rain forest

Summary 1. Ecological stoichiometry predicts important control of the relative abundance of the key elements carbon (C), nitrogen (N) and phosphorus (P) on trophic interactions. In a nutrient‐poor Amazonian lowland rain forest of French Guiana, we tested the hypothesis that decomposers exploit stoichiometrically diverse plant litter more efficiently, resulting in faster litter decomposition compared to litter with a uniform stoichiometry. 2. In a field experiment in the presence or absence of soil macrofauna, we measured litter mass loss, and N and P dynamics from all possible combinations of leaf litter from four common tree species which were distinctly separated along a C:N and along a N:P gradient. 3. Mean litter mass remaining after 204 days of field exposure varied between 25.2% and 71.3% among litter treatments. Fauna increased litter mass loss by 18%, N loss by 21% and P loss by 14%. Litter species richness had no effect on litter mass loss or nutrient dynamics. In contrast, litter mass and nutrient losses increased with increasing stoichiometric dissimilarity of litter mixtures in presence of fauna, suggesting faster decomposition of a stoichiometrically more heterogeneous litter. 4. However, the effect of stoichiometric dissimilarity was smaller than the strong C quality related litter composition effect and disappeared in the absence of fauna. Increasing proportions of litter that is relatively rich in accessible C compounds (non‐structural carbohydrates, phenolics) and relatively poor in recalcitrant C (condensed tannins, lignin), correlated best with litter mass loss irrespective of fauna presence. No correlation was found for any of the nutrient related litter quality parameters and decomposition. 5. Synthesis. Our results suggest that Amazonian decomposer communities studied here are primarily limited by energy, and only secondarily by litter stoichiometry. Tropical tree species might thus influence decomposers and detritivores by the production of litter of specific C quality with potentially important feedback effects on ecosystem nutrient dynamics and availability.

Effect of warming and grazing on litter mass loss and temperature sensitivity of litter and dung mass loss on the Tibetan plateau

AbstractKnowledge about the role of litter and dung decomposition in nutrient cycling and response to climate change and grazing in alpine ecosystems is still rudimentary. We conducted two separate studies to assess the relative role of warming and grazing on litter mass loss and on the temperature sensitivity of litter and dung mass loss. Experiments were conducted for 1–2 years under a controlled warming–grazing system and along an elevation gradient from 3200 to 3800 m. A free‐air temperature enhancement system (FATE) using infrared heaters and grazing significantly increased soil temperatures (average 0.5–1.6 °C) from 0 to 40 cm depth, but neither warming nor grazing affected soil moisture except early in the growing seasons at 30 cm soil depth. Heaters caused greater soil warming at night‐time compared with daytime, but grazing resulted in greater soil warming during daytime compared with night‐time. Annual average values of the soil temperature at 5 cm were 3.2, 2.4 and 0.3 °C at 3200, 3600 and 3800 m, respectively. Neither warming nor grazing caused changes of litter quality for the first year of the controlled warming–grazing experiment. The effects of warming and grazing on litter mass losses were additive, increasing litter mass losses by about 19.3% and 8.3%, respectively, for the 2‐year decomposition periods. The temperature sensitivity of litter mass losses was approximately 11% °C−1 based on the controlled warming–grazing experiment. The annual cumulative litter mass loss was approximately 2.5 times that of dung along the elevation gradient. However, the temperature sensitivity (about 18% °C−1) of the dung mass loss was about three times that of the litter mass loss. These results suggest greater warming at night‐time compared with daytime may accelerate litter mass loss, and grazing will enhance carbon loss to atmosphere in the region through a decrease of litter biomass and an increase of dung production with an increase of stocking rate in future warmer conditions.

Species shift drives decomposition rates following invasion by hemlock woolly adelgid

Insect and disease outbreak is an important cause of selective species removal and accompanying functional change in North American forests. Outbreak of hemlock woolly adelgid, Adelgies tsugae– HWA, is causing selective removal of eastern hemlock Tsuga canadensis at a regional scale. Impacts of outbreak-caused canopy mortality and shifts in dominant species on litter decay were compared across sites that range in HWA-caused canopy damage and subsequent canopy dominance by black birch Betula lenta. Senescent litter from eastern hemlock, black birch, and equal litter mixes were decomposed in the field for 36 months within nine sites in Connecticut and Massachusetts USA. Mass loss and % N accumulation of black birch was 65% and 52% greater compared to eastern hemlock. In contrast, outbreak related canopy damage increased litter mass loss by 11.5% in high mortality stands relative to uninfested stands but canopy damage had no impact on % N dynamics. Non-additive effects of litter mixing influenced chemical dynamics of decaying litter; black birch accumulated less N and eastern hemlock accumulated more N compared to each species decaying alone. However, these changes offset and mixed litter bags overall showed no differences in N dynamics compared to values from each species decaying alone. In eastern hemlock stands invaded by hemlock woolly adelgid, canopy damage influences the rates and dynamics of decay but species differences between hemlock and black birch leaf litter are the dominant mechanisms of decomposition changes and a long-lasting driver of increased N cycling rates. Species shifts may be the dominant driver of altered ecosystem processes for other insect outbreaks, particularly when replacement species have very different characteristics regulating decomposition and N cycling.

The importance of biotic factors in predicting global change effects on decomposition of temperate forest leaf litter

Increasing atmospheric CO(2) and temperature are predicted to alter litter decomposition via changes in litter chemistry and environmental conditions. The extent to which these predictions are influenced by biotic factors such as litter species composition or decomposer activity, and in particular how these different factors interact, is not well understood. In a 5-week laboratory experiment we compared the decomposition of leaf litter from four temperate tree species (Fagus sylvatica, Quercus petraea, Carpinus betulus and Tilia platyphyllos) in response to four interacting factors: elevated CO(2)-induced changes in litter quality, a 3 degrees C warmer environment during decomposition, changes in litter species composition, and presence/absence of a litter-feeding millipede (Glomeris marginata). Elevated CO(2) and temperature had much weaker effects on decomposition than litter species composition and the presence of Glomeris. Mass loss of elevated CO(2)-grown leaf litter was reduced in Fagus and increased in Fagus/Tilia mixtures, but was not affected in any other leaf litter treatment. Warming increased litter mass loss in Carpinus and Tilia, but not in the other two litter species and in none of the mixtures. The CO(2)- and temperature-related differences in decomposition disappeared completely when Glomeris was present. Overall, fauna activity stimulated litter mass loss, but to different degrees depending on litter species composition, with a particularly strong effect on Fagus/Tilia mixtures (+58%). Higher fauna-driven mass loss was not followed by higher C mineralization over the relatively short experimental period. Apart from a strong interaction between litter species composition and fauna, the tested factors had little or no interactive effects on decomposition. We conclude that if global change were to result in substantial shifts in plant community composition and macrofauna abundance in forest ecosystems, these interacting biotic factors could have greater impacts on decomposition and biogeochemical cycles than rising atmospheric CO(2) concentration and temperature.

Badger (Taxidea taxus) disturbances increase soil heterogeneity in a degraded shrub-steppe ecosystem

In the western United States, overgrazing, weed invasion and wildfire have resulted in the conversion of shrub-steppe to annual grasslands, with substantial effects on ecosystem function. In these landscapes, badgers disturb large areas of soil while foraging for fossorial animals. Mounds created by badgers contained the lowest concentrations of total carbon, nitrogen and sulphur, mineral nitrogen and mineralizable nitrogen, inter-mound soils had the highest concentrations, and excavation pits had intermediate levels. Soil C:N ratio and pH were greater, and electrical conductivity and soluble Ca2+, Mg2+ and K+ were lower on mound soils compared with either pit or inter-mound soils. Larger pits generally trapped more litter, and increased litter mass equated with greater concentrations of active carbon, but only at the burned sites. Older mounds supported more vascular plants and cryptogamic crusts. Our results demonstrate reduced levels of nutrients and a higher C:N ratio on the mounds compared with either the pits or inter-mounds. Alteration to the homogeneous post-fire landscape by badgers contributes to patchiness in soils and vegetation, which is critical to the functioning of arid systems. Given their effect on soil C:N ratios, mounds may be important sites for recovery of indigenous shrub-steppe plant species.

The burden of motherhood: gliding locomotion in mammals influences maternal reproductive investment

Body mass is usually positively associated with reproductive output in females, with larger females having more offspring. However, in gliding or flying species, large body mass is constrained as a result of wing loading, defined as the ratio of body mass to the surface area available to generate lift. Gliding animals may be particularly susceptible to increased wing loading resulting from added mass during pregnancy. We compiled reproductive and morphological data for gliding and related nongliding mammals in all extant taxa where this type of locomotion occurs to test the predictions that gliders will have proportionally lighter litters than related nongliders, and relative litter mass will be negatively associated with wing loading. Contrary to our 1st prediction, gliders had heavier offspring than did nongliders in all taxa examined. Consistent with our 2nd prediction, however, greater relative litter mass was associated with lower wing loading. Maintaining the ability to glide when pregnant may influence investment in reproduction. However, gliding locomotion appears to be associated with increased litter mass, perhaps because of smaller litters of heavy young. Thus, we suggest that gliders may be characterized by K-selected life-history traits such as low mortality, slow metabolism, and fewer offspring per litter, but increased investment in individual young. The findings of this comparative study highlight how major shifts in locomotor mode can have a profound influence on patterns of life history.

The effect of macro-invertebrates and plant litter of different quality on the release of N from litter to plant on alpine pastureland

The effect of 15N-labelled litter of different quality (Luzula sylvatica, a grass species, Vaccinium gaultheroides, a deciduous dwarf shrub, and Calluna vulgaris, a hardy dwarf shrub) and the presence of macro-decomposers (Lumbricus rubellus, Lumbricidae, and Enantiulus nanus, Diplopoda) on the growth of Dactylis glomerata (Poaceae), a grass species abundant on alpine pastureland, was investigated. After 4 months, the presence of soil animals significantly increased litter mass loss of L. sylvatica, V. gaultheroides and C. vulgaris by 27%, 11% and 40%, respectively. Soil animals generally reduced microbial biomass but significantly increased it in treatments where either L. sylvatica or C. vulgaris was present. The presence of soil animals significantly increased shoot and root biomass of D. glomerata by 48% and 64%, respectively. L. rubellus increased the transfer of 15N from the litter into plants. We conclude that macro-decomposers increased nutrient mobilization and plant uptake of nutrients mineralized from recalcitrant litter materials. Litter of L. sylvatica contributed most to the 15N uptake by D. glomerata, suggesting that litter quality is crucial for the cycling of nutrients on abandoned alpine pastureland.

Differences in litter mass change mite assemblage structure on a deciduous forest floor

Few mechanisms that determine the assemblage structure of mites have been identified. Whilst the relative abundance of soil fauna is known to change with humus form, the degree to which the quantity of litter inputs play a part in these changes has not been investigated. We tested the response of oribatid and mesostigmatid mites in litter and soil layers to increasing levels of birch Betula pubescens litter to test whether litter mass could affect the mite assemblage. Six litter treatments (1, 2, 4, 8 and 12×natural litter mass and complete litter removal) were established in November 2004 and the soil and litter communities sampled in October 2005. Species composition of oribatids was distinct for the soil and litter. There was no apparent effect of increasing litter mass on the soil mite assemblage. In the litter layer, the number of oribatids g−1 of litter showed a strong negative relationship with increasing litter mass whilst the number of mesostigmatids g−1 of litter was unresponsive to litter mass. Hence, the relative abundance of these two groups altered with increasing litter mass. The response of the oribatid groups Oppiidae and Poronota followed this negative relationship with litter mass but Phthiracaridae appeared less affected. Consequently, there was a subtle shift in the relative abundance of these groups with increasing litter mass. Our results demonstrate that oribatids as a whole and within groups respond in a predictable manner to increases in litter mass whilst mesostigmatids are unresponsive. Whilst there are undoubtedly biological and physical aspects that vary with litter mass, litter mass itself, is able to explain some patterns in the assemblage of oribatid mites.

Amounts and degradability of dissolved organic carbon from foliar litter at different decomposition stages

Litter is one of the main sources of dissolved organic carbon (DOC) in forest soils and litter decomposition is an important control of carbon storage and DOC dynamics. The aim of our study was to evaluate (i) effects of tree species on DOC production and (ii) relationships between litter decomposition and the amount and quality of DOC. Five different types of leaves and needles were exposed in litterbags at two neighboring forest sites. Within 12 months we sampled the litterbags five times and leached aliquots of field moist litter in the laboratory. In the collected litter percolates we measured DOC concentrations and recorded UV and fluorescence spectra in order to estimate the aromaticity and complexity of the organic molecules. Furthermore, we investigated the biodegradability of DOC from fresh and decomposed litter during 6 weeks incubations. Fresh sycamore maple litter released the largest amounts of DOC reaching about 6.2% of litter C after applying precipitation of 94 mm. We leached 3.9, 1.6, 1.0 and 3.3% carbon from fresh mountain ash, beech, spruce and pine litter, respectively. In the initial phase of litter decomposition significantly decreasing DOC amounts were released with increasing litter mass loss. However, after mass loss exceeds 20% DOC production from needle litter tended to increase. UV and fluorescence spectra of percolates from pine and spruce litter indicated an increasing degree of aromaticity and complexity with increasing mass loss as often described for decomposing litter. However, for deciduous litter the relationship was less obvious. We assume that during litter decomposition the source of produced DOC in coniferous litter tended toward a larger contribution from lignin-derived compounds. Biodegradability of DOC from fresh litter was very high, ranging from 30 to 95% mineralized C. DOC from degraded litter was on average 34% less mineralizable than DOC from fresh litter. Taking into account the large DOC production from decomposed needles we can assume there is an important role for DOC in the accumulation of organic matter in soils during litter decomposition particularly in coniferous forests.

Effects of prairie restoration on weed invasions

Restoration of late-successional plant communities may reduce weed invasion in non-cropland. In studies of natural succession and invasion, however, the presence of late-successional vegetation is often confounded with lower weed propagule pressure and greater time since disturbance. This paper reports a study in which propagule pressure and time since disturbance were controlled by adding weed species to experimental plots dominated by early and late-successional perennial grass species. Seeds of 12 weed species were added to randomized, 6-year-old plots of restored prairie and non-restored, old-field vegetation. Restoration reduced added weed biomass by 92% in year one and 72% in year two, and reduced biomass of four individual weed species, each by more than 82%. Likely mechanisms through which restoration may have reduced invasion include competition and establishment limitation. To examine these mechanisms, subplots were treated with N addition and burning, to reduce competition and remove litter, respectively. Several patterns suggest that competition may have mediated reductions in weed invasion: restoration increased biomass of resident vegetation, decreased light levels, and decreased weed size. Furthermore, adding N reduced effects of restoration on resident and added weeds. The importance of establishment limitation in reducing invasion is suggested by increased litter mass and decreased weed density with restoration. Burning removed litter, and reduced effects of restoration on added weeds. These results suggest that tallgrass prairie restoration can reduce weed invasion, and that this reduction may involve multiple mechanisms, including competition and establishment limitation.

Response of the Mixed Prairie to Protection from Grazing

The Mixed Prairie plant communities developed with the influences of fire and grazing. Available evidence suggests that removal of these disturbances could cause succession toward a more mesic type with the accumulation or litter or loss in productivity as nutrient turnover is delayed. Exclosures constructed in 1927 in a semiarid Mixed Prairie community provided an opportunity to examine the effects that protection had on vegetation and soils. Fifteen exclosures were selected for detailed examination; of these, 11 were located on Chernozemic soil and 4 on Solonetzic soil. We measured plant and soil variables both inside and outside the exclosures in a test of the hypothesis that protection from grazing will lead to a loss of production potential of the semi-arid. Mixed Prairie communities in the Northern Great Plains of southeastern Alberta. We found little evidence that 70 years of protection from large animal disturbance reduced the production potential of the plant communities. Conversely, most evidence suggested a neutral effect or an improvement as reflected in an increased cover of Pascopyrum smithii Rydb. (Love) (P = 0.049) and increased annual net primary production (P = 0.047). The effect of protection appeared largely driven by the accumulation of litter mass that primarily benefits soil and plant indices of quality on the Chernozemic soil type. Although protection tended to reduce species diversity (P = 0.097) among native plants on the Chernozemic soil type, evenness and richness were not affected (P > 0.10). The potential effect that reduced diversity might have on reducing production stability appears more than compensated for by increased litter mass. DOI:10.2458/azu_jrm_v55i3_willms

Response of the mixed prairie to protection from grazing

The Mixed Prairie plant communities developed with the influences of fire and grazing. Available evidence suggests that removal of these disturbances could cause succession toward a more mesic type with the accumulation or litter or loss in productivity as nutrient turnover is delayed. Exclosures constructed in 1927 in a semiarid Mixed Prairie community provided an opportunity to examine the effects that protection had on vegetation and soils. Fifteen exclosures were selected for detailed examination; of these, 11 were located on Chernozemic soil and 4 on Solonetzic soil. We measured plant and soil variables both inside and outside the exclosures in a test of the hypothesis that protection from grazing will lead to a loss of production potential of the semi-arid. Mixed Prairie communities in the Northern Great Plains of southeastern Alberta. We found little evidence that 70 years of protection from large animal disturbance reduced the production potential of the plant communities. Conversely, most evidence suggested a neutral effect or an improvement as reflected in an increased cover of Pascopyrum smithii Rydb. (Löve) (P = 0.049) and increased annual net primary production (P = 0.047). The effect of protection appeared largely driven by the accumulation of litter mass that primarily benefits soil and plant indices of quality on the Chernozemic soil type. Although protection tended to reduce species diversity (P = 0.097) among native plants on the Chernozemic soil type, evenness and richness were not affected (P > 0.10). The potential effect that reduced diversity might have on reducing production stability appears more than compensated for by increased litter mass.

Nitrogen Turnover in Forest Floors of Coastal Douglas-Fir at Sites Differing in Soil Nitrogen Capital

Nitrogen cycling is generally considered to be more rapid on sites with high availability of N; this is usually associated with differences in tree species composition. We tested whether N cycling in stands of a single tree species increased with increasing mineral soil nitrogen capital. Rates of N cycling in nine stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were estimated by measuring annual N input in litter, N content of the forest floor, and net N mineralization rate in the forest floor. Rates of C and N turnover were estimated from the litterfall:forest floor ratio. The amount of N returned in litter increased as soil N capital increased. The increase in litter N content resulted from both increased litter mass (C content) and increased N concentration in litter. Despite the greater litter input, forest floor mass was smaller at N-rich sites, indicating more rapid turnover of the forest floor on N-rich sites. There was a positive relationship between fractional annual loss of N from the forest floor and soil N capital. Therefore, even without changes in tree species composition, sites with greater soil N capital returned more N in annual litterfall and had faster turnover of N in the forest floor. Fractional annual loss of C also increased with increasing soil N capital, indicating faster decomposition on N-rich sites. The rate of net N mineralization during laboratory incubations of the forest floor was not correlated with soil N capital or N concentrations of litter, but was related to the C:N ratio of the forest floor. Net N mineralization was appreciable only at two sites where forest floor C:N ratios were <35. The rate of net N mineralization or C:N ratio of the forest floor were not good indicators of N availability at these sites. The results of this study are consistent with the hypothesis that rates of N cycling are faster on N-rich sites, and that this effect can occur in the absence of changes in tree species composition.

NITROGEN TURNOVER IN FOREST FLOORS OF COASTAL DOUGLAS-FIR AT SITES DIFFERING IN SOIL NITROGEN CAPITAL

Nitrogen cycling is generally considered to be more rapid on sites with high availability of N; this is usually associated with differences in tree species composition. We tested whether N cycling in stands of a single tree species increased with increasing mineral soil nitrogen capital. Rates of N cycling in nine stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) were estimated by measuring annual N input in litter, N content of the forest floor, and net N mineralization rate in the forest floor. Rates of C and N turnover were estimated from the litterfall:forest floor ratio. The amount of N returned in litter increased as soil N capital increased. The increase in litter N content resulted from both increased litter mass (C content) and increased N concentration in litter. Despite the greater litter input, forest floor mass was smaller at N-rich sites, indicating more rapid turnover of the forest floor on N-rich sites. There was a positive relationship between fractional annual loss of N from the forest floor and soil N capital. Therefore, even without changes in tree species composition, sites with greater soil N capital returned more N in annual litterfall and had faster turnover of N in the forest floor. Fractional annual loss of C also increased with increasing soil N capital, indicating faster decomposition on N-rich sites. The rate of net N mineralization during laboratory incubations of the forest floor was not correlated with soil N capital or N concentrations of litter, but was related to the C:N ratio of the forest floor. Net N mineralization was appreciable only at two sites where forest floor C:N ratios were <35. The rate of net N mineralization or C:N ratio of the forest floor were not good indicators of N availability at these sites. The results of this study are consistent with the hypothesis that rates of N cycling are faster on N-rich sites, and that this effect can occur in the absence of changes in tree species composition.

EFFECTS OF LEAF-LITTER ACCUMULATION ON RIPARIAN PLANT DIVERSITY ACROSS EUROPE

Floods in rivers can deposit large quantities of organic debris in the riparian corridor and cause considerable change to its vegetation. We studied the importance of leaf-litter accumulation on riparian vegetation in different climates by constructing gradients ranging from complete litter removal to addition of up to 6000 g/m2 (35–40 cm deep) of leaf litter and recorded the consequent changes in vegetation. Experiments were located along rivers in northern and southern Sweden, Italy, and Romania. Plant biomass decreased with increasing litter mass but did not vary with site-dependent variables such as climate and soil. Species density for various plant life traits varied with litter mass, summer temperature, and summer precipitation. Litter mass was the most important variable for all life traits except rhizomatous and stoloniferous species, which were most dependent upon summer precipitation. These findings are useful for the prediction of responses in riparian vegetation following future climatic changes.