UV‐B effect on Quercus robur leaf litter decomposition persists over four years

The effects of elevated UV‐B (280–315 nm) radiation on the long‐term decomposition of Quercus robur leaf litter were assessed at an outdoor facility in the UK by exposing saplings to elevated UV‐B radiation (corresponding to a 30% increase above the ambient level of erythemally weighted UV‐B, equivalent to that resulting from a c. 18% reduction in ozone column) under arrays of cellulose diacetate‐filtered fluorescent UV‐B lamps that also produced UV‐A radiation (315–400 nm). Saplings were also exposed to elevated UV‐A radiation alone under arrays of polyester‐filtered fluorescent lamps and to ambient solar radiation under arrays of nonenergized lamps. After 8 months of irradiation, abscised leaves were placed into litter bags and allowed to decompose in the litter layer of a mixed deciduous woodland for 4.08 years. The dry weight loss of leaf litter from saplings irradiated with elevated UV‐B and UV‐A radiation during growth was 17% greater than that of leaf litter irradiated with elevated UV‐A radiation alone. Annual fractional weight loss of litter (k), and the estimated time taken for 95% of material to decay (3/k) were respectively increased and decreased by 27% for leaf litter exposed during growth to elevated UV‐B and UV‐A radiation, relative to that exposed to UV‐A alone. The present data corroborate those from a previous study indicating that UV‐B radiation applied during growth accelerates the subsequent decomposition of Q. robur leaf litter in soil, but indicate that this effect persists for over four years after abscission.     

Aphids decelerate litter nitrogen mineralisation through changes in litter quality

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Effects of the litter layer of Pteridium aquilinum on seed banks under experimental restoration

Questions: Does the litter layer of Pteridium aquilinum (bracken) act as a barrier to certain species in the seed bank? Does bracken control/restoration treatment affect seed transfer through the litter layer? Location: Five experiments at three sites across the UK covering two major vegetation types; acid‐grassland and heath‐land. Methods: At each experiment a range of bracken control and vegetation restoration treatments were applied for about ten years. The seed bank was sampled in both the bracken litter and the soil. The cover (%) of each species in the vegetation and the bracken litter abundance (cover and depth) was also estimated. Results: The bracken litter layer acts as an inert barrier as it contained a large proportion of seeds available in the litter‐soil profile (38%– 67% of the total). Bracken litter depth and cover also influenced significantly the seed bank composition in both the bracken litter and the soil. These effects were site‐specific, and species‐specific. The application of treatments changed significantly the balance between seed inputs and outputs in the bracken litter layer for some species. This was either a positive or negative response relative to the untreated control plots. Conclusion: For heathland and acid‐grassland restoration, the bracken litter layer may be an important seed source, but it must be disturbed particularly before seed addition.     

Vegetative and reproductive phenology of the mangrove Kandelia obovata

Mangroves in the subtropical area of Japan are growing to their northern limits, yet little is known of their phenology. The aim of the present study was to understand both vegetative and reproductive phenology patterns, such as leaf emergence, leaf fall, bud setting, flowering, fruiting and propagule setting, in the mangrove Kandelia obovata. The phenology of this species was assessed using litter‐fall data for 5 years. Leaf and stipule litter‐falls continued with a clear monthly pattern throughout the years. New leaf production and leaf fall peaked in summer, immediately after the propagules fell. Leaf and stipule litter‐falls were linked to monthly sunshine hour, and monthly mean air temperature and monthly mean air relative humidity, respectively. Kandelia obovata had a distinct flowering period, with the flowering phenophase starting in spring and continuing into summer. Fruit initiation started at the end of summer and continued into autumn, whereas propagule production occurred during winter and spring. Flowering of K. obovata was influenced by monthly sunshine hour and monthly mean air temperature, whereas fruit and propagule litter‐falls were not linked to any climatic factors. The present results showed that a small portion (4.4%) of flowers developed into propagules. The average development period from flower buds to mature propagules was approximately 11 months. Kendall's consistency coefficient suggested that the monthly trends in vegetative and reproductive litter‐fall components, except for branches, did not change significantly among years.     

Enhanced ozone exposure of European beech (Fagus sylvatica) stimulates nitrogen mobilization from leaf litter and nitrogen accumulation in the soil

Abstract

In a lysimeter study with young beech trees, the effects of elevated ozone concentration on the decomposition and fate of nitrogen in ¹⁵N‐labeled leaf litter were analyzed after one growing season. Nitrogen in the litter was dominated by a relatively inert, residual fraction, but easily decomposable nitrogen was present in substantial amounts. Nitrogen loss was significantly higher at twice‐ambient ozone which was largely attributed to an enhanced mobilization of residual nitrogen. Enhanced mobilization of nitrogen from litter at twice‐ambient ozone exposure resulted in additional ¹⁵N incorporation into the soil down to 30 cm depth. Only 0.41–0.62% of the nitrogen in the litter was incorporated into plant material at both ozone concentrations. Twice‐ambient ozone exposure changed the distribution of the nitrogen taken up from litter inside the beech trees in favor of the shoot, where it may have been used in biosynthetic processes required for defense reactions.     

Evaluating litter decomposition in earth system models with long‐term litterbag experiments: an example using the Community Land Model version 4 (CLM4)

Decomposition is a large term in the global carbon budget, but models of the earth system that simulate carbon cycle‐climate feedbacks are largely untested with respect to litter decomposition. We tested the litter decomposition parameterization of the community land model version 4 (CLM4), the terrestrial component of the community earth system model, with data from the long‐term intersite decomposition experiment team (LIDET). The LIDET dataset is a 10‐year study of litter decomposition at multiple sites across North America and Central America. We performed 10‐year litter decomposition simulations comparable with LIDET for 9 litter types and 20 sites in tundra, grassland, and boreal, conifer, deciduous, and tropical forest biomes using the LIDET‐provided climatic decomposition index to constrain temperature and moisture effects on decomposition. We performed additional simulations with DAYCENT, a version of the CENTURY model, to ask how well an established ecosystem model matches the observations. The results show large discrepancy between the laboratory microcosm studies used to parameterize the CLM4 litter decomposition and the LIDET field study. Simulated carbon loss is more rapid than the observations across all sites, and nitrogen immobilization is biased high. Closer agreement with the observations requires much lower decomposition rates, obtained with the assumption that soil mineral nitrogen severely limits decomposition. DAYCENT better replicates the observations, for both carbon mass remaining and nitrogen, independent of nitrogen limitation. CLM4 has low soil carbon in global earth system simulations. These results suggest that this bias arises, in part, from too rapid litter decomposition. More broadly, the terrestrial biogeochemistry of earth system models must be critically tested with observations, and the consequences of particular model choices must be documented. Long‐term litter decomposition experiments such as LIDET provide a real‐world process‐oriented benchmark to evaluate models.     

Testing a ‘genes‐to‐ecosystems’ approach to understanding aquatic–terrestrial linkages

A ‘genes‐to‐ecosystems’ approach has been proposed as a novel avenue for integrating the consequences of intraspecific genetic variation with the underlying genetic architecture of a species to shed light on the relationships among hierarchies of ecological organization (genes → individuals → communities → ecosystems). However, attempts to identify genes with major effect on the structure of communities and/or ecosystem processes have been limited and a comprehensive test of this approach has yet to emerge. Here, we present an interdisciplinary field study that integrated a common garden containing different genotypes of a dominant, riparian tree, Populus trichocarpa, and aquatic mesocosms to determine how intraspecific variation in leaf litter alters both terrestrial and aquatic communities and ecosystem functioning. Moreover, we incorporate data from extensive trait screening and genome‐wide association studies estimating the heritability and genes associated with litter characteristics. We found that tree genotypes varied considerably in the quality and production of leaf litter, which contributed to variation in phytoplankton abundances, as well as nutrient dynamics and light availability in aquatic mesocosms. These ‘after‐life’ effects of litter from different genotypes were comparable to the responses of terrestrial communities associated with the living foliage. We found that multiple litter traits corresponding with aquatic community and ecosystem responses differed in their heritability. Moreover, the underlying genetic architecture of these traits was complex, and many genes contributed only a small proportion to phenotypic variation. Our results provide further evidence that genetic variation is a key component of aquatic–terrestrial linkages, but challenge the ability to predict community or ecosystem responses based on the actions of one or a few genes.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Does decomposition of standard materials differ among grassland patches maintained by livestock?

Grazing can modify vegetation structure and species composition through selective consumption, modifying plant litter quality and hence decomposability. In most grasslands, moderate stocking rates maintain a mosaic of high‐quality patches, preferentially used by herbivores (‘grazing lawns’), and low‐quality tall patches, which are avoided. In grazing lawns decomposition rates can be accelerated because of the higher litter quality of its component species and, besides, through the indirect effect of increased nutrient availability in soil. We aimed at testing this indirect effect using standard materials, comparing their decomposition in grazing lawns, open and closed tall tussock grasslands. We selected 10 patches of each type and sampled floristic composition, soil variables and cattle dung deposition. Standard materials were filter paper and Poa stuckertii litter. We prepared litterbags of 0.3 mm (thin mesh) and 1 mm mesh size (coarse mesh). Samples were incubated for 65 days in two ways: above‐ground (thin and coarse mesh) and below‐ground (only thin mesh), aiming at analysing the conditions for decomposition for surface litter and buried litter or dead roots, respectively. Physical and chemical soil variables did not differ among patch types, despite the differences in species composition. Closed tussock grasslands showed the lowest dung deposition, confirming the less intense use of these patches. Soil nitrogen availability (N‐NO₃^‐ and N‐NH₄⁺) was not significantly different among patch types. Each standard material followed a different decomposition pattern across patch types. For above‐ground incubated samples, Poa litter decomposed significantly faster in lawns, and slower in open tussock grasslands. Filter paper decomposed significantly faster in closed tussock grasslands than in the other two patch types. Decomposition of below‐ground incubated samples did not significantly differ among patch types, in line with results for soil variables. Above‐ground differences in decomposition may be associated with differences in microclimatic conditions resulting from differences in vegetation structure.     

Pine Litter from Afforestations Hinders the Establishment of Endemic Plants in Semiarid Scrubby Habitats of Natura 2000 Network

A sowing experiment was carried out using three endemic secondary successional species that commonly co‐occur with Pinus halepensis in afforestations in southeast Spain. The experiment simulated the natural arrangement of needle litter over soil to test the litter's effect on seedling emergence and early growth of dwarf‐shrubs and annual species, the main components of some European Habitats of Community Interest in Spain. Three sowing treatments were compared in a growth chamber: (1) soil without litter taken from clearings in a pine plantation (BARE); (2) intact soil and litter forming two distinct layers as in the field (PINE); and (3) soil and pine litter mixed in the laboratory (PINEMX). Seedling emergence of Diplotaxis harra and Thymus zygis were seven and two times higher, respectively, on BARE than on the pine litter treatments. Pine litter significantly decreased seedling biomass, length, and number of leaves of D. harra and T. zygis. Thymus capitatum was weakly affected by the presence of pine litter. We suggest that plant recruitment in the understory of P. halepensis is partially controlled by interference of pine litter, which might affect dynamics of the studied species. We propose that management practices counteract this negative effect by manipulating the pine litter layer to achieve successful ecological restoration and conservation of endemic‐rich secondary communities in Natura 2000 sites in semiarid western Mediterranean areas.     

Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes

Climate and litter quality have been identified as major drivers of litter decomposition at large spatial scales. However, the role played by soil fauna remains largely unknown, despite its importance for litter fragmentation and microbial activity. We synthesised litterbag studies to quantify the effect sizes of soil fauna on litter decomposition rates at the global and biome scales, and to assess how climate, litter quality and soil fauna interact to determine such rates. Soil fauna consistently enhanced litter decomposition at both global and biome scales (average increment ~ 27%). However, climate and litter quality differently modulated the effects of soil fauna on decomposition rates between biomes, from climate‐driven biomes to those where climate effects were mediated by changes in litter quality. Our results advocate for the inclusion of biome‐specific soil fauna effects on litter decomposition as a mean to reduce the unexplained variation in large‐scale decomposition models.     

On a likelihood-based goodness-of-fit test of the beta-binomial model

When faced with proportion data that exhibit extra-binomial variation, data analysts often consider the beta-binomial distribution as an alternative model to the more common binomial distribution. A typical example occurs in toxicological experiments with laboratory animals, where binary observations on fetuses within a litter are often correlated with each other. In such instances, it may be of interest to test for the goodness of fit of the beta-binomial model; this effort is complicated, however, when there is large variability among the litter sizes. We investigate a recent goodness-of-fit test proposed by Brooks et al. (1997, Biometrics 53, 1097-1115) but find that it lacks the ability to distinguish between the beta-binomial model and some severely non-beta-binomial models. Other tests and models developed in their article are quite useful and interesting but are not examined herein.     

A Maximum Likelihood Estimator for Shape Parameters of S‐Distributions

The S‐distribution is a four‐parameter distribution that is defined in terms of a differential equation, in which the cumulative is represented as the dependent variable: The article proposes a maximum likelihood estimator for the shape parameters of this distribution.     

Tree functional diversity affects litter decomposition and arthropod community composition in a tropical forest

Disturbance can alter tree species and functional diversity in tropical forests, which in turn could affect carbon and nutrient cycling via the decomposition of plant litter. However, the influence of tropical tree diversity on forest floor organisms and the processes they mediate are far from clear. We investigated the influence of different litter mixtures on arthropod communities and decomposition processes in a 60‐year‐old lowland tropical forest in Panama, Central America. We used litter mixtures representing pioneer and old growth tree species in experimental mesocosms to assess the links between litter types, decomposition rates, and litter arthropod communities. Overall, pioneer species litter decomposed most rapidly and old growth species litter decomposed the slowest but there were clear non‐additive effects of litter mixtures containing both functional groups. We observed distinct arthropod communities in different litter mixtures at 6 mo, with greater arthropod diversity and abundance in litter from old growth forest species. By comparing the decay of different litter mixtures in mesocosms and conventional litterbags, we demonstrated that our mesocosms represent an effective approach to link studies of litter decomposition and arthropod communities. Our results indicate that changes in the functional diversity of litter could have wider implications for arthropod communities and ecosystem functioning in tropical forests.     

Seed–litter–position drives seedling establishment in grassland species under recurrent drought

Changes in land use and climate interfere with grassland ecosystem processes. Here I experimentally investigated the combined effects of land‐use change related litter cover and contrasting water supply on seedling emergence. In this context, the role of the initial relative position of seeds, i.e. seeds on top of the litter versus seeds beneath the litter in interaction with water supply has not been investigated so far. I hypothesised that facilitative effects of litter on seedling emergence occur when seeds are covered by litter and deteriorate when litter covers the ground and seeds fall on it (seeds on top of the litter). Further, I hypothesised that the importance of seed position for seedling emergence will increase under conditions of recurrent drought. I performed a controlled pot experiment on seedling emergence of three common European grassland species (Pimpinella saxifraga, Leontodon autumnalis, Sanguisorba officinalis) by experimental manipulations of litter and water availability. Seedling emergence under moist conditions showed no significant differences between each litter position compared to the control across species. In contrast, under recurrent drought, seedling emergence was significantly higher below the litter compared to seeds on top of the litter and the control (i.e. no litter). In abandoned land, seedling emergence may be limited when seeds fall on ground‐covering litter. In contrast, in grasslands with regular low‐intensity land use, seedling emergence may be enhanced when a moderate level of litter covers seeds at the end of the growing season. Protective mechanisms that occur with seeds positioned beneath litter are particularly important under recurrent drought.     

Invasive plants differentially affect soil biota through litter and rhizosphere pathways: a meta‐analysis

Invasive plants affect soil biota through litter and rhizosphere inputs, but the direction and magnitude of these effects are variable. We conducted a meta‐analysis to examine the different effects of litter and rhizosphere of invasive plants on soil communities and nutrient cycling. Our results showed that invasive plants increased bacterial biomass by 16%, detritivore abundance by 119% and microbivore abundance by 89% through litter pathway. In the rhizosphere, invasive plants reduced bacterial biomass by 12%, herbivore abundance by 55% and predator abundance by 52%, but increased AM fungal biomass by 36%. Moreover, CO₂ efflux, N mineralisation rate and enzyme activities were all higher in invasive than native rhizosphere soils. These findings indicate that invasive plants may support more decomposers that in turn stimulate nutrient release via litter effect, and enhance nutrient uptake by reducing root grazing but forming more symbioses in the rhizosphere. Thus, we hypothesise that litter‐ and root‐based loops are probably linked to generate positive feedback of invaders on soil systems through stimulating nutrient cycling, consequently facilitating plant invasion. Our findings from limited cases with diverse contexts suggest that more studies are needed to differentiate litter and rhizosphere effects within single systems to better understand invasive plant‐soil interactions.     

Proteome analysis of fungal and bacterial involvement in leaf litter decomposition

Fungi and bacteria are key players in the decomposition of leaf litter, but their individual contributions to the process and their interactions are still poorly known. We combined semi‐quantitative proteome analyses (1‐D PAGE‐LC‐MS/MS) with qualitative and quantitative analyses of extracellular degradative enzyme activities to unravel the respective roles of a fungus and a bacterium during litter decomposition. Two model organisms, a mesophilic Gram‐negative bacterium (Pectobacterium carotovorum) and an ascomycete (Aspergillus nidulans), were grown in both, pure culture and co‐culture on minimal medium containing either glucose or beech leaf litter as sole carbon source. P. carotovorum grew best in co‐culture with the fungus, whereas growth of A. nidulans was significantly reduced when the bacterium was present. This observation suggests that P. carotovorum has only limited capabilities to degrade leaf litter and profits from the degradation products of A. nidulans at the expense of fungal growth. In accordance with this interpretation, our proteome analysis revealed that most of the extracellular biodegradative enzymes (i.e. proteases, pectinases, and cellulases) in the cultures with beech litter were expressed by the fungus, the bacterium producing only low levels of pectinases.     

Recent land‐use changes affect stream ecosystem processes in a subtropical island in Brazil

Land‐use changes such as conversion of natural forest to rural and urban areas have been considered as main drivers of ecosystem functions decline, and a large variety of indicators has been used to investigate these effects. Here, we used a replicated litter‐bag experiment to investigate the effects of land‐use changes on the leaf‐litter breakdown process and leaf‐associated invertebrates along the forest–pasture–urban gradient located in a subtropical island (Florianópolis, SC, Brazil). We identified the invertebrates and measured the litter breakdown rates using the litter bags approach. Litter bags containing 3 g of dry leaf of Alchornea triplinervia were deployed on forest rural and urban streams. Principal component analysis, based on physico‐chemical variables which, confirmed a gradient of degradation from forest to urban streams with intermediate values in rural areas. In accordance, shredder richness and abundance were lower in rural and urban than in forest streams. The land‐use changes led also to the dominance of tolerant generalist taxa (Chironomidae and Oligochaeta) reducing the taxonomic and functional diversity in these sites. Leaf‐litter breakdown rates decreased from forest to rural and finally to urban areas and were associated with changes in pH, water velocity, dissolved oxygen and abundance of leaf‐shredding invertebrates, although global decomposition rates did not differ between rural and urban streams. Overall, this study showed that land‐use changes, namely to rural and urban areas, have a strong impact on tropical streams ecosystems, in both processes and communities composition and structure. Despite of being apparently a smaller transformation of landscape, rural land use is comparable to urbanisation in terms of impact in stream functioning. It is thus critical to carefully plan urban development and maintain forest areas in the island of Florianópolis in order to preserve its natural biodiversity and aquatic ecosystems functioning.     

Plant litter chemistry alters the content and composition of organic carbon associated with soil mineral and aggregate fractions in invaded ecosystems

Through the input of disproportionate quantities of chemically distinct litter, invasive plants may potentially influence the fate of organic matter associated with soil mineral and aggregate fractions in some of the ecosystems they invade. Although context dependent, these native ecosystems subjected to prolonged invasion by exotic plants may be instrumental in distinguishing the role of plant–microbe–mineral interactions from the broader edaphic and climatic influences on the formation of soil organic matter (SOM). We hypothesized that the soils subjected to prolonged invasion by an exotic plant that input recalcitrant litter (Japanese knotweed, Polygonum cuspidatum) would have a greater proportion of plant‐derived carbon (C) in the aggregate fractions, as compared with that in adjacent soil inhabited by native vegetation that input labile litter, whereas the soils under an invader that input labile litter (kudzu, Pueraria lobata) would have a greater proportion of microbial‐derived C in the silt‐clay fraction, as compared with that in adjacent soils that receive recalcitrant litter. At the knotweed site, the higher C content in soils under P. cuspidatum, compared with noninvaded soils inhabited by grasses and forbs, was limited to the macroaggregate fraction, which was abundant in plant biomarkers. The noninvaded soils at this site had a higher abundance of lignins in mineral and microaggregate fractions and suberin in the macroaggregate fraction, partly because of the greater root density of the native species, which might have had an overriding influence on the chemistry of the above‐ground litter input. At the kudzu site, soils under P. lobata had lower C content across all size fractions at a 0–5 cm soil depth despite receiving similar amounts of Pinus litter. Contrary to our prediction, the noninvaded soils receiving recalcitrant Pinus litter had a similar abundance of plant biomarkers across both mineral and aggregate fractions, potentially because of the higher surface area of soil minerals at this site. The plant biomarkers were lower in the aggregate fractions of the P. lobata‐invaded soils, compared with noninvaded pine stands, potentially suggesting a microbial co‐metabolism of pine‐derived compounds. These results highlight the complex interactions among litter chemistry, soil biota, and minerals in mediating soil C storage in unmanaged ecosystems; these interactions are particularly important under global changes that may alter plant species composition and hence the quantity and chemistry of litter inputs in terrestrial ecosystems.