The potential for forest canopy litterfall interception by a dense fern understorey, and the consequences for litter decomposition

Most studies on litter decomposition have assumed that all falling plant litter reaches the ground where it then decomposes. In many forests a proportion of this litter may in fact be intercepted by understorey vegetation, but the ecological significance of this remains poorly understood. We performed two experiments in a temperate rainforest in southern New Zealand, in which there was a dense understorey of the crown fern Blechnum discolor. The fronds of this fern originate from a crown, and have a funnel‐like arrangement that can trap falling litter and prevent it from reaching the ground. The first experiment measured the effects of ferns on the spatial distribution of litter accumulation over one year. The ferns intercepted a substantial proportion of the total litterfall, and the fern crowns (from which the fronds originate) retained 10% of the total incoming litterfall (despite occupying only 2% of the ground area). The retained litter had a substantially higher ratio of twig to foliar litter than did the incoming litterfall. Further, much of the litter not retained on the crowns of the ferns accumulated at the base of the fern trunks. The second experiment considered litter decomposition in fern crowns versus the ground under the ferns. The litter that had accumulated in the crowns was characterized by higher microbial basal respiration and active microbial biomass than was the litter that had accumulated on the ground. The use of litterbags revealed that litter decomposition rates were significantly higher on the fern crowns than on the ground at 30 cm and 60 cm from the fern trunks. These results show that litter interception ameliorates the decomposer environment and increases the rate of litter decomposition. In total, this study provides evidence for understorey ferns greatly influencing both the spatial distribution of litterfall and the decomposition of plant litter. Although the ecological role of understorey vegetation in forested ecosystems has received little attention to date, our results point to understorey species as an important driver of forest ecosystem processes.     

Context dependency of litter‐mixing effects on decomposition and nutrient release across a long‐term chronosequence

Litter decomposition is an important driver of terrestrial systems, and factors that determine decomposition rate for individual litter species have been widely studied. Fewer studies have explored the factors that regulate how mixing litters of multiple species affects litter decomposition and nutrient dynamics, and only a handful of studies have investigated how litter‐mixing effects may differ among different habitats or ecosystems, or how they respond to environmental gradients. We used a well‐established retrogressive chronosequence involving thirty lake islands in northern Sweden in which time since fire disturbance increases with decreasing island size; smaller islands therefore have reduced rates of aboveground and belowground ecosystem processes. On each of these islands we utilized plots with and without the long‐term experimental removal of shrubs. Litters from the six most common plant species on the islands were prepared in single‐, three‐ and six‐species litterbags, and placed on both the shrub‐removal and non‐removal plots on each island to decompose for one year. We found significant non‐additive effects of litter mixing on litter decomposition rates, on final litter N and P concentrations, and on litter N loss, but these non‐additive effects varied both in direction and magnitude with changed number of species, and even among litter mixtures with the same number of species. Further, the magnitude of non‐additive effects of litter mixing on both litter decomposition and nutrient dynamics was significantly influenced by both island size and the interaction between island size and shrub‐removal treatment. When shrubs were present, there was a U‐shaped relationship between these non‐additive effects and island size, while the relationship was positive when shrubs were removed. Hence, our results support previous findings that litter mixing may produce non‐additive effects on litter decomposition and nutrient dynamics, and that these effects tend to be idiosyncratic due to the importance of effects of individual species in the mixture. Most importantly, our results show that non‐additive litter‐mixing effects change greatly across environmental gradients, meaning that the biotic and abiotic characteristics of an ecosystem can be a powerful driver of the magnitude and even the direction of litter‐mixing effects on ecosystem processes.     

Changes in western wheatgrass foliage quality following defoliation: consequences for a graminivorous grasshopper

We determined the effects of defoliation by a graminivorous grasshopper on the foliage quality of the C3 plant, western wheatgrass (Pascopyrum smithii [Rydb] A. Love). Additionally, we determined the effects of this defoliation upon the subsequent feeding of the graminivorous grasshopper Phoetaliotes nebrascensis Thomas (Orthoptera: Acrididae). In field and greenhouse studies, graminivorous grasshopper herbivory altered the quality of remaining western wheatgrass foliage. In the greenhouse, severe (50% foliage removal) grasshopper grazing (638 grasshoppers/m² for 72h) resulted in decreased foliar nitrogen (–12%), carbohydrate (–11%) and water (–2.5%) concentrations, and increased phenolic concentrations (+43%). These changes were associated with decreased adult female grasshopper mass gain, consumption rate, approximate digestibility, and food conversion efficiencies. In the field, moderate (14% foliage removal) grasshopper grazing (20 grasshoppers/m² for 20 days) led to a 10% reduction in foliar nitrogen concentrations. Foliage quality changes in the field were not associated with any reductions in grasshopper mass gain, consumption rates, food digestibility, or conversion efficiencies. The results presented here are consistent with the hypothesis that defoliation leads to a reallocation of carbon and nitrogen compounds within the plant such that foliage quality for P. nebrascensis is reduced.     

Changes in fungal diversity and composition along a chronosequence of Eucalyptus grandis plantations in Ethiopia

Eucalyptus tree species are widely used in Ethiopian plantations, but the impact of these plantations on the soil fungal communities is still unknown. We assessed the changes in diversity, species composition and ecological guilds of the soil fungal communities across tree ages of Eucalyptus grandis plantations by DNA metabarcoding of ITS2 amplicons. Changes in soil fungal species composition, diversity and ecological guilds were related to stand age but also to fertility changes. The relative abundance of saprotrophs and pathogens were negatively correlated with stand age, and positively with soil fertility. In contrast, the relative abundance and diversity of ectomycorrhizal species were higher in older, less fertile stands, including well-known cosmopolitan species but also species associated with Eucalyptus, such as Scleroderma albidum and Descomyces albellus. We show that soil fungal community changes are linked to progressive soil colonization by tree roots but are also related to soil fertility changes.     

Short and long term consequences of increases in exotic species richness on water filtration by marine invertebrates

Although recent research has considered the consequences of global declines in the number of species, less attention has focused on the aggregate effects of regional increases in species richness as a result of human-mediated introductions. Here we examine several potential ecosystem consequences of increasing exotic species diversity of suspension feeding marine invertebrates. First, we experimentally manipulated native and non-native suspension feeder richness and measured its effect on short-term phytoplankton clearance rates. Multispecies communities all performed similarly, regardless of whether they were dominated by natives, exotics, or an even mix of the two. Individual species varied considerably in filtration rates, but non-native species often filtered less than the most similar native. Second, we determined potential changes in integrated function over time by comparing seasonal patterns of recruitment as a proxy for the ability to quickly recover filtration capacity after a disturbance. We found that exotic species have complementary seasonal phenologies both to native species and each other. Our results suggest that the consequences of local increases in species richness due to invasions may be manifest over long (annual to interannual) time scales, even when short term changes in ecosystem function are negligible.     

Variation in protein complexation capacity among and within six plant species across a boreal forest chronosequence

We investigated among and within species variation in several litter chemical properties, including protein complexation capacity (PCC), for six plant species across a boreal forest chronosequence in northern Sweden across which stand fertility declines sharply with stand age. We hypothesized (1) that evergreen species which dominate in late-successional stands would exhibit higher PCCs than deciduous species that dominate in young stands, (2) that individual species would increase their PCCs in response to nutrient limitation as succession proceeds, and (3) that differences in PCC among litter types would determine their interactive effects with proteins on soil N and C mineralization. The data demonstrated a high PCC, but a low PCC per unit of soluble phenol, for two deciduous species that dominate in early-successional high fertility stands, providing mixed support for our first hypothesis. No species demonstrated a significant correlation between their PCC and stand age, which did not support our second hypothesis. Finally, a soil incubation assay revealed that litter extracts for three of the six species had negative interactive effects with added proteins on N mineralization rates, and that all six species demonstrated positive interactive effects with protein on C mineralization. This pattern did not provide strong support for our third hypothesis, and suggests that N immobilization was likely a more important factor regulating N mineralization than stabilization of proteins into tannin complexes. These data suggest that multiple interactive mechanisms between litter extracts and proteins likely occur simultaneously to influence the availability of N in soils.     

Changes in stable nitrogen and carbon isotope ratios of plants and soil across a boreal forest fire chronosequence

Background and Aim

Nitrogen (N) and carbon (C) isotopic signatures (δ¹⁵N and δ¹³C) serve as powerful tools for understanding temporal changes in ecosystem processes, but how these signatures change across boreal forest chronosequences is poorly understood.

Methods

The δ¹⁵N, δ¹³C, and C/N ratio of foliage of eight dominant plant species, including trees, understory shrubs, and a moss, as well as humus, were examined across a 361 years fire-driven chronosequence in boreal forest in northern Sweden.

Results

The δ¹³C and C/N ratio of plants and humus increased along the chronosequence, suggesting increasing plant stress through N limitation. Despite increasing biological N fixation by cyanobacteria associated with feather mosses, δ¹⁵N showed an overall decline, and δ¹⁵N of the feather moss and associated vascular plants diverged over time from that of atmospheric N₂.

Conclusions

Across this chronosequence the N fixed by cyanobacteria is unlikely to be used by mosses and vascular plants without first undergoing mineralization and mycorrhizal transport, which would cause a change in δ¹⁵N signature due to isotopic fractionation. The decreasing trend of δ¹⁵N suggests that as the chronosequence proceeds, the plants may become more dependent on N transferred from mycorrhizal fungi or from N deposition.     

The effects of the moss layer on the decomposition of intercepted vascular plant litter across a post-fire boreal forest chronosequence

Aims

Feather mosses form a thick ground layer in boreal forests that can intercept incoming litter fall. This interception may influence the decomposition of incoming litter but this has been little explored. We investigated how the moss layer influences decomposition of intercepted litter along a 362-year fire driven forest chronosequence in northern Sweden across which soil fertility declines.

Methods

We placed leaf litter from three plant species into plots in which mosses and dwarf shrubs were either experimentally removed or left intact, at each of ten stands across the chronosequence. After one year we measured litter mass loss, and litter nitrogen and phosphorous.

Results

Litter decomposed consistently faster, and had higher nitrogen and phosphorus, in the presence of mosses and at greater depth in the moss layer. Despite an increase in moss depth across the chronosequence we did not find consistent increases in effects of moss removal on litter decomposition or on litter N or P.

Conclusions

Our findings identify a clear role of the moss layer in boreal forests in promoting the decomposition of intercepted leaf litter, and highlight that this role is relatively consistent across chronosequence stages that vary greatly in productivity and moss depth.     

Changes in the chemical composition of alder,poplar and willow leaves during decomposition in a river

1. As there was a paucity of information on the aquatic decomposition of leaves of willow (Salix alba), poplar (Populus gr. nigra) and alder (Alnus glutinosa), the net-bag technique was used to study chemical changes in the leaves over a period of six months in the Garonne (France).
2. The disappearance rates of foliar organic matter were about similar for the three species (k = 0.0065 day^?1 for alder, k = 0.0054 day^?1 for poplar and k = 0.0050 day^?1 for willow). Changes in carbon amount were comparable to those of organic matter. Nitrogen and organic phosphorus accumulated during the first months of decomposition. The C/N ratios of foliar matter changed in the same way in the three species, falling steeply for the first month, then levelling off. Amounts of sugar disappeared very rapidly, particularly for alder and willow. Cellulose concentrations were constant throughout the whole decomposition process whilst an increase in the amounts of lignin was observed during the first months; however the latter could be due to an interference by microbial compounds.
3. The changes of some constituents, such as carbon, sugars and cellulose, were described with exponential or multiexponential models. Despite the very different initial ratios of C/N and lignin/N, these changes associated with leaf decomposition were similar for the three species.

     

Nutrient fluxes via litterfall and leaf litter decomposition vary across a gradient of soil nutrient supply in a lowland tropical rain forest

The extent to which plant communities are determined by resource availability is a central theme in ecosystem science, but patterns of small-scale variation in resource availability are poorly known. Studies of carbon (C) and nutrient cycling provide insights into factors limiting tree growth and forest productivity. To investigate rates of tropical forest litter production and decomposition in relation to nutrient availability and topography in the absence of confounding large-scale variation in climate and altitude we quantified nutrient fluxes via litterfall and leaf litter decomposition within three distinct floristic associations of tropical rain forest growing along a soil fertility gradient at the Sepilok Forest Reserve (SFR), Sabah, Malaysia. The quantity and nutrient content of small litter decreased along a gradient of soil nutrient availability from alluvial forest (most fertile) through sandstone forest to heath forest (least fertile). Temporal variation in litterfall was greatest in the sandstone forest, where the amount of litter was correlated negatively with rainfall in the previous month. Mass loss and N and P release were fastest from alluvial forest litter, and slowest from heath forest litter. All litter types decomposed most rapidly in the alluvial forest. Stand-level N and P use efficiencies (ratios of litter dry mass to nutrient content) were greatest for the heath forest followed by the sandstone ridge, sandstone valley and alluvial forests, respectively. We conclude that nutrient supply limits productivity most in the heath forest and least in the alluvial forest. Nutrient supply limited productivity in sandstone forest, especially on ridge and hill top sites where nutrient limitation may be exacerbated by reduced rates of litter decomposition during dry periods. The fluxes of N and P varied significantly between the different floristic communities at SFR and these differences may contribute to small-scale variation in species composition.     

Vegetation dynamics across a chronosequence of created wetland sites in Virginia, USA

Plant species composition, dominance, richness, and diversity were measured across a 15-year chronosequence of created wetland sites in Virginia, USA. Using an age-class categorization (1?C2?years, 3?C5?years, 6?C10?years, or 11?C15?years), all classes had a predominance of herbaceous species with perennial life history strategy, and perennials contributed 68.6?% to the overall dominance measure (importance value; IV) averaged across all sites. There was no significant difference in species richness or diversity among age classes. Analysis of Similarity (ANOSIM) indicated that herbaceous species composition was similar between the youngest and oldest age classes, but not the intermediate classes. For woody shrubs and saplings, planted species were more prevalent in the youngest age classes, and volunteer species predominated in the oldest age classes. These results suggest that perennial herbaceous species are important in early plant development on created wetland sites, and may be influential in observed patterns of species composition over time. In the context of plant development in newly created wetlands, dominance shifts from planted to volunteer woody species suggest that planting early successional species, or species with reproductive strategies attuned to created wetland site management, may favor survivorship and recruitment of other species over time.     

Variation in the herb species response and the humus quality across a 200-year chronosequence of beech and oak plantations in Belgium

The present study aimed at exploring the long-term impact of pure and mixed beech Fagus sylvatica and oak Quercus robur stands on the forest floor by documenting changes in the herb species' behaviour and in humus index across a 200-yr chronosequence of forest stands. The research was conducted in central Belgium, in a 4383 ha beech-dominated forest. Analyses were carried out in stands which are replicated, of the same age, managed in the same way, and growing on the same soil type with the same land-use history. The results of this study indicate that stand aging is an important determinant of herb species occurrence in the studied area. Most of the species studied show a different response to stand age in pure compared to mixed stands. Our results clearly show a decrease of the humus quality with age in pure stands (beech as well as oak). On the other hand, we found that mixing beech and oak maintained or improved the humus status along the chronosequence according to the proportion of each tree. So the addition of some oak to the beech made it possible to keep a constant quality of the humus. We found that, even if the understory tree species is very scarce, it may be sufficient to maintain the humus status on the long term. In the present study, a cover of 1% oak in a beech stand was sufficient to show an effect of the minor species on these soils. This pattern contrasts with the widespread idea that substantial effects of the minor tree species on soils might not develop if the ratio of major/minor species is low.     

长白山各植被带主要树种凋落物分解速率及模型模拟的试验研究

2001～2003年在长白山自然保护区内3个垂直植被带的典型群落红松阔叶林、云冷杉林和岳桦林内,利用网袋埋藏法对群落内的6个主要优势乔木树种凋落物进行埋藏分解试验,研究凋落物分解速率及其变化动态;同时利用分解模型,模拟预测凋落物的分解进展,为深入研究这6个树种的营养策略、群落养分循环等奠定基础,也为森林生态系统管理提供理论依据.研究结果表明,所研究的6个树种凋落物都表现出随时间进程失重率增大的现象,但失重率并不与时间呈线性相关.在分解的638d（1.75a）后,6种叶凋落物的分解速率明显升高.到分解实验结止时（699d）,叶凋落物干重剩余率从小至大依次为白桦（24.56%）、紫椴（24.81%）、红松（38.48%）、鱼鳞云杉（41.15%）、岳桦（41.53%）和臭冷杉（42.62%）.枝凋落物分解速率明显低于叶,枝干重剩余率从小至大依次为紫椴（44.98%）、臭冷杉（64.62%）、红松（72.07%）、鱼鳞云杉（73.51%）、白桦（77.37%）和岳桦（80.35%）.在同一海拔高度,阔叶树种叶凋落物分解速率大于针叶树种.并且随着海拔的升高,叶凋落物分解速率逐渐减慢.模型分析预测结果表明,长白山北坡各垂直植被带的优势树种叶凋落物分解95%需4.5～8.0a;年分解系数为紫椴（0.686）〉白桦（0.624）〉 红松（0.441）〉 鱼鳞云杉（0.406）〉 臭冷杉（0.397）〉岳桦（0.385）;枝凋落物分解95%需7.8～29.3a,不同树种间的差异明显.枝年分解系数为紫椴（0.391）〉臭冷杉（0.204）〉红松（0.176）＞鱼鳞云杉（0.157）〉白桦（0.148）〉 岳桦（0.102）.     

Shifts in symbiotic associations in plants capable of forming multiple root symbioses across a long‐term soil chronosequence

Changes in soil nutrient availability during long‐term ecosystem development influence the relative abundances of plant species with different nutrient‐acquisition strategies. These changes in strategies are observed at the community level, but whether they also occur within individual species remains unknown. Plant species forming multiple root symbioses with arbuscular mycorrhizal (AM) fungi, ectomycorrhizal (ECM) fungi, and nitrogen‐(N) fixing microorganisms provide valuable model systems to examine edaphic controls on symbioses related to nutrient acquisition, while simultaneously controlling for plant host identity. We grew two co‐occurring species, Acacia rostellifera (N₂‐fixing and dual AM and ECM symbioses) and Melaleuca systena (AM and ECM dual symbioses), in three soils of contrasting ages (c. 0.1, 1, and 120 ka) collected along a long‐term dune chronosequence in southwestern Australia. The soils differ in the type and strength of nutrient limitation, with primary productivity being limited by N (0.1 ka), co‐limited by N and phosphorus (P) (1 ka), and by P (120 ka). We hypothesized that (i) within‐species root colonization shifts from AM to ECM with increasing soil age, and that (ii) nodulation declines with increasing soil age, reflecting the shift from N to P limitation along the chronosequence. In both species, we observed a shift from AM to ECM root colonization with increasing soil age. In addition, nodulation in A. rostellifera declined with increasing soil age, consistent with a shift from N to P limitation. Shifts from AM to ECM root colonization reflect strengthening P limitation and an increasing proportion of total soil P in organic forms in older soils. This might occur because ECM fungi can access organic P via extracellular phosphatases, while AM fungi do not use organic P. Our results show that plants can shift their resource allocation to different root symbionts depending on nutrient availability during ecosystem development.     

Responses of litter decomposition to temperature along a chronosequence of tropical montane rainforest in a microcosm experiment

We conducted a microcosm experiment for studying the decomposition of Altingia obovata leaf litter by the decomposer community at 20 and 30°C from three forest stands (namely a 35-year-old secondary forest, a 47-year-old secondary forest, and a primary forest) of a tropical montane rainforest. Our results showed that rank-order of the litter decomposition among the three forest stands was not parallel to the stand age. At 20°C, the mass loss of A. obovata leaf litter from the primary forest was higher than those from the two secondary forests, of which the younger stand showed higher mass loss than did the older one. However, there were no differences in mass loss among these three stands at 30°C. The mass loss for the two secondary forest stands, but not for the primary forest stand, increased significantly from 20 to 30°C. The level of lignin decomposition among the three stands at 20°C corresponded to their forest stand age, i.e., the primary forest > the 47-year-old secondary forest > the 35-year-old secondary forest. A rise of 10°C in temperature significantly increased lignin decomposition for the two secondary forests, while the reverse was true for the primary forest. Carbohydrate decomposition was positively related to the temperature but not to the stand age. The different responses of litter decomposition to the forest stand age and temperature might be due to the differences in the microbial activities among the three forest stands.     

Variation in stand structure,light and seedling abundance across a tropical moist forest chronosequence,Panama

Abstract. We asked whether forest structure and understory light environments across a tropical moist forest chronosequence followed predictions of a 4‐phase model of secondary succession (establishment, thinning, transition and steady‐state) and whether seedling density and diversity were functions of light availability as predicted by this model. Using aerial photographs, we identified eight second‐growth stands (two each aged ca. 20, 40, 70, and 100 yr) and two old‐growth stands within Barro Colorado Nature Monument, Panama. Trees and seedlings were sampled in nested, contiguous quadrats in 2 160‐m transects in each stand. Light was measured as percent transmittance of diffuse photosynthetically active radiation (TPAR) at each seedling quadrat and by estimation of percent total incident radiation during the growing season from hemispherical canopy photographs. Basal area, tree density, and canopy height followed predictions of the 4‐phase model. Percent total radiation, but not TPAR, declined with stand age as did seedling density. While seedlings were more likely to occur in quadrats at higher light levels, much variation in seedling density was not related to light availability. Seedling patch sizes were small irrespective of light patches, estimated as semivariance ranges. Seedling species richness was a function of seedling density; estimates of species diversity unbiased by density did not vary systematically as a function of stand age. Proximate seed sources, efficient dispersal mechanisms, and appropriate establishment conditions can promote establishment of species‐rich communities early in successions of heterogeneous tropical moist forest.     

Links between plant and fungal communities across a deforestation chronosequence in the Amazon rainforest

Understanding the interactions among microbial communities, plant communities and soil properties following deforestation could provide insights into the long-term effects of land-use change on ecosystem functions, and may help identify approaches that promote the recovery of degraded sites. We combined high-throughput sequencing of fungal rDNA and molecular barcoding of plant roots to estimate fungal and plant community composition in soil sampled across a chronosequence of deforestation. We found significant effects of land-use change on fungal community composition, which was more closely correlated to plant community composition than to changes in soil properties or geographic distance, providing evidence for strong links between above- and below-ground communities in tropical forests.     

Biomass, litterfall and decomposition rates for the fringed rhizophora mangle forest lining the Bon Accord Lagoon, Tobago

The mangrove forest that fringes the Bon Accord Lagoon measures 0.8 km(2) and is dominated by red mangrove (Rhizophora mangle). This forest forms the landward boundary of the Buccoo Reef Marine Park in Southwest Tobago, and is part of a mangrove-seagrass-coral reef continuum. Biomass and productivity, as indicated by litterfall rates, were measured in seven 0.01 ha monospecific plots from February 1998 to February 1999, and decomposition rates were determined. Red mangrove above-ground biomass ranged between 2.0 and 25.9 kg (dry wt.) m(-2). Mean biomass was 14.1+/-8.1 kg (dry wt.) m(-2) yielding a standing crop of 11 318+/-6 488 t. Litterfall rate varied spatially and seasonally. It peaked from May to August (4.2-4.3 g dry wt. m(-2) d(-1)) and was lowest from October to December (2.3-2.8 g dry wt. m(-2) d(-1)). Mean annual litterfall rate was 3.4+/-0.9 g dry wt. m(-2) d(-1). Leaf degradation rates ranged from 0.3% loss d(-1) in the upper intertidal zone to 1% loss d(-1) at a lower intertidal site flooded by sewage effluent. Mean degradation rate was 0.4+/-1% loss d(-1) . The swamp produces 2.8 t dry wt. of litterfall and 12 kg dry wt. of decomposed leaf material daily. Biomass and litterfall rates in Bon Accord Lagoon were compared to five similar sites that also participate in the Caribbean Coastal Marine Productivity Programme (CARICOMP). The Bon Accord Lagoon mangrove swamp is a highly productive fringed-forest that contributes to the overall productivity of the mangrove-seagrass-reef complex.