The estimation of leaf litter turnover is often limited to early-stage decomposition using unrepresentative models and litter types. In tropical secondary forests, particularly exotic-dominated novel forests, the characterisation of litter turnover remains poor. This study estimated the annual turnover of in-situ leaf litter across four forest successional types in Singapore using a Weibull residence time model. Litter turnover and nutrient dynamics diverged between young secondary and old-growth forests. In particular, within novel forests, annual phosphorus return via leaf litterfall was three times that of primary forests, while the mass loss of in-situ leaf litter was highest among all forest successional types, estimated at 92.8% annually with a mean residence time of 176 days, resulting in a litter pool size a third that of primary forests. Our findings suggest that tree species composition and species-specific effects shaped the observed variations in litter turnover and nutrient dynamics across forest successional types and forest stands, whereas tree species richness, canopy structure, soil nutrient levels, and microclimate were found to be non-predictors. Taken together, our study provides an insight into litter turnover in human-modified tropical landscapes increasingly characterised by novel forests, potentially leading to a reduction in surface litter and soil organic carbon pools.

Litter decomposition plays an important role in the carbon cycle and is affected by many factors in forest ecosystems. This study aimed to quantify the rhizosphere priming effect on litter decomposition in subtropical forest southwestern China. A litter decomposition experiment including control and trenching treatments was conducted using the litter bag method, and the litter decomposition rate was calculated by litter dry mass loss. Trenching did not change soil temperature, but increased the soil water content by 14.5%. In this study, the interaction of soil temperature and soil water content controlled the litter decomposition rate, and explained 87.4 and 85.5% of the variation in litter decomposition in the control and trenching treatments, respectively. Considering changes in soil environmental factors due to trenching, the litter decomposition rates were corrected by regression models. After correction, the litter decomposition rates of the control and trenching treatments were 32.47 ± 3.15 and 25.71 ± 2.72% year–1, respectively, in the 2-year period. Rhizosphere activity significantly primed litter decomposition by 26.3%. Our study suggested a priming effect of rhizosphere activity on litter decomposition in the subtropical forest. Combining previous interaction effect results, we estimated the contributions of total soil organic matter (SOM) decomposition, total litter decomposition, and root respiration to soil respiration in the subtropical forest, and our new method of estimating the components of soil respiration provided basic theory for SOM decomposition research.

A critical question in invasion biology involves the relative importance of propagule rain and community invasibility. For plant invasions, invasibility is often related to disturbance, but few studies of forest invaders have simultaneously investigated both canopy and ground-level disturbance. We investigated the relative importance of seed rain, canopy disturbance, and soil disturbance in a mature forest in Maryland on the recruitment of four invasive species: wine raspberry (Rubus phoenicolasius Maxim.), Japanese barberry (Berberis thunbergii DC), multiflora rose (Rosa multiflora Thunb.), and Japanese stiltgrass [Microstegium vimineum (Trin.) A. Camus]. Using complete censuses of a 9-ha plot at two points in time (2011–12 and 2014), we mapped new recruits, and related their locations to canopy and soil disturbance, as well as to a seed rain index based on locations of reproducing plants and seed-dispersal kernels.

We found that propagule rain, as measured by the seed rain index, was a significant predictor of recruitment for B. thunbergii, R. phoenicolasius, and M. vimineum. For R. multiflora, seed sources were not located, precluding assessment of propagule rain, but recruitment was linked to canopy disturbance, as was recruitment of M. vimineum. However, because reproduction of R. phoenicolasius and, in some years, of B. thunbergii is higher in treefall gaps, these gaps experience higher propagule rain, with the result that recruitment is indirectly associated with these gaps. Ground-layer disturbance was an important predictor of recruitment only for B. thunbergii. Our findings reveal that the importance of propagule rain is the most consistent driver of recruitment, but canopy or ground-layer disturbance promotes recruitment of some invasive plant species.

Concentrations of hexazinone and two metabolites in vegetation were determined for 2 yr after broadcast application of a 10% granular formulation of hexazinone at 2 and 4 kg ai ha−1 rates in August 1986 in a boreal forest. Prewinter concentrations of hexazinone in stems of trembling aspen, Saskatoon berry, and willow ranged from 0.02 to 0.05 μg−1 dry wt at 64 d after treatment (DAT). Absorption of hexazinone accelerated during spring thaw (1987), and residues in foliage of several woody and herbaceous species peaked during May to July. Patterns of accumulation of hexazinone and its metabolites varied with the species. Foliar concentrations diminished significantly by the end of the first growing season in 1987 (372 DAT) and were undetectable or extremely low at the end of the second growing season in 1988 (707 DAT). Based on the highest residue concentrations detected in several plant species, it is estimated that wildlife would ingest a maximum of 16, 28, and 24 mg of hexazinone, metabolite A, and metabolite B, respectively, for every kg of dry matter consumed. Reported LD50 values suggest that application of hexazinone at the 4 kg ai ha−1 rate or less poses no toxicological threat to wildlife.

Plant communities and ecosystem processes in seres at multiple altitudes in the tropical montane forest zone of northern Borneo were studied to understand the patterns and mechanisms of the secondary succession after shifting cultivation. A total of 25 stands (and additional three stands) were sampled with stand ages ranging from 2 to 55 y after slash and burn at altitudes between 900 and 1400 m asl. Plant species composition, above-ground biomass (AGB), chemical properties of soils, litter and foliar samples were investigated in each stand. A TWINSPAN analysis classified five plant communities primarily as a sere but with two altitudinal communities in the later successional phase. AGB accumulated steadily at the rate of 2.42 Mg ha−1 y−1 during the succession for the first 55 y due to the ontogenetic development of plants as well as plant community shifts. At the onset of secondary succession, pool of soil NO3-N and soil total P was high probably because burning caused flushes of minerals originating from the burnt plant materials. Pool of soil NO3-N and soil total P decreased with increasing stand age during the succession. Leaf-litter N:P ratios of dominant species significantly increased with increasing stand age suggesting disproportionately greater P deficiency than N deficiency in the later successional phase. It is suggested that tree species shifted to those of greater P-use efficiency during succession in response to decreasing soil P availability. We conclude that the interaction of altitude with the reduction of soil N and P availability was related to the altitudinal split of plant communities in the later phase, while pioneer communities were wide-ranging across altitudes reflecting richer soil nutrients.

Species-specific traits of trees affect ecosystem dynamics, defining forest structure and understorey development. Ochroma pyramidale is a fast-growing tree species, with life-history traits that include low wood density, short-lived large leaves and a narrow open thin crown. We evaluated forest succession in O. pyramidale-dominated secondary forests, diverse secondary forests, both 10–15 y since abandonment, and rain forests by comparing height, density and basal area of all trees (> 5 cm dbh). Furthermore, we compared species richness of understorey trees and shrubs, and basal area and density of trees of early- and late-successional species (< 5 cm dbh) between forest types. We found that tree basal area (mean ± SD: 32 ± 0.9 m2 ha−1) and height (12.4 ± 1.8 m) of canopy trees were higher, and density (1450 ± 339 ha−1) lower in O. pyramidale forests than in diverse forests, and more similar to rain forest. Understorey shrub diversity and tree seedling density and diversity were lower in O. pyramidale forests than in diverse forests, but these forest types had a similar density of early- and late-successional trees. Canopy openness (> 15%) and leaf litter (> 10 cm) were both highest in O. pyramidale forests, which positively affected density of understorey trees and shrubs and negatively affected density of late-successional trees. In conclusion, O. pyramidale forests presented structural features similar to those of rain forest, but this constrained the establishment of understorey tree species, especially late-successional species, decreasing successional development.

The decomposition of leaf litter is governed, in part, by litter invertebrates. In tropical rain forests, ants are dominant predators in the leaf litter and may alter litter decomposition through the action of a top-down control of food web structure. The role of ants in litter decomposition was investigated in a Costa Rican lowland rain forest with two experiments. In a mesocosm experiment, we manipulated ant presence in 50 ambient leaf-litter mesocosms. In a litterbag gradient experiment, Cecropia obtusifolia litter was used to measure decomposition rate constants across gradients in nutrients, ant density and richness, with 27 separate litterbag treatments for total arthropod exclusion or partial arthropod exclusion. After 2 mo, mass loss in mesocosms containing ants was 30.9%, significantly greater than the 23.5% mass loss in mesocosms without ants. In the litter bags with all arthropods excluded, decomposition was best accounted by the carbon: phosphorus content of soil (r2 = 0.41). In litter bags permitting smaller arthropods but excluding ants, decomposition was best explained by the local biomass of ants in the vicinity of the litter bags (r2 = 0.50). Once the microarthropod prey of ants are permitted to enter litterbags, the biomass of ants near the litterbags overtakes soil chemistry as the regulator of decomposition. In concert, these results support a working hypothesis that litter-dwelling ants are responsible for accelerating litter decomposition in lowland tropical rain forests.

In this study, we investigated whether landscape-scale variation of soil P accounts for 13C and 15N composition of detrital invertebrates in a lowland tropical rain forest in Costa Rica. The top 10-cm soil, leaf-litter samples and plant foliage were collected among 18 plots representing a three-fold soil P gradient during 2007–2009. Body tissue of litter invertebrates (extracted from leaf-litter samples) along with soil, leaf litter and green foliage were analysed for total C, total N, δ13C and δ15N values. Differences in δ13C and δ15N signatures across plots and relative trophic distances of detrital food webs (Δ δ15N), and their variation with soil P gradient were evaluated. We found soil P gradient had a significantly positive correlation with δ15N of Asterogyne martiana foliage, leaf litter, collembolans and oribatid mites. The δ15N of the collembolans and pseudoscorpions positively correlated to leaf-litter δ15N. Δ δ15N between the trophic levels remained consistent across the soil P gradient. Higher δ15N in the collembolans and oribatid mites might be derived from their consumption on 15N-enriched decayed debris or fungal hyphae growing on it. It suggests that fine-scale soil P variation can affect trophic dynamics of detrital arthropods via regulation of microbial community and nutrient dynamics.

Secondary forests occupy a growing portion of the tropical landscape mosaic due to regeneration on abandoned pastures and other disturbed sites (Asner et al. 2009). Tropical secondary forests and degraded old-growth forests now account for more than half of the world's tropical forests (Chazdon 2003), and provide critical ecosystem services (Brown & Lugo 1990, Guariguata & Ostertag 2001).

Decomposition of leaf litter is an important process that releases energy and nutrients in both terrestrial and aquatic environments (Moore et al. 2004, Wallace et al. 1997); therefore, the physical, chemical and biological processes controlling leaf-litter decomposition rates can affect nutrient cycling and productivity in these systems (Cross et al. 2007, Wood et al. 2009). Several studies have shown that leaf decomposition is faster in aquatic than in terrestrial habitats due to relatively constant temperatures, continuous leaching and the physical breakdown of leaves by flowing water (Hutchens & Wallace 2002, Langhans & Tockner 2006, Langhans et al. 2008). Yet, comparatively few studies have examined these relationships in tropical systems with flooded forests. Flooding is a predominant feature of the upper Amazon Basin, but its occurrence and magnitude is complex and not strictly seasonal (Junk et al. 1989). To identify the dominant energy pathways and understand the nutrient dynamics of upper Amazon rain forests, it is imperative to investigate organic matter processing in the aquatic/terrestrial transition zones of these ecosystems.

Gastropod abundance was quantified in forest litter around 33 trees harbouring Lobaria pulmonaria in southern Norway. In total, 1709 snails representing 28 species were found, and the number of snail species strongly increased with the total number of specimens found. Number of snail species, as well as snail abundance, was highest around trees on high pH soils. There was a positive relationship between number of snail specimens and cover of grazing traces on L. pulmonaria, presumably because calcareous soils facilitate both litter dwelling and climbing gastropods. The results suggest that gastropods may limit the distribution of L. pulmonaria in calcareous broad-leaved forests.

Litter standing crop (LSC) is the quantity of plant detritus on the floor in forested environments. Knowledge of LSC is important in understanding many ecological phenomena. These include studies of litterfall, decomposition/litter turnover rates and nutrient cycling (Anderson et al. 1983, Dent et al. 2006), general plant performance (Benítez-Malvido & Kossmann-Ferraz 1999), other ecosystem processes such as the effects of fire (Odiwe & Muoghalu 2003) and fauna (Frith & Frith 1990, Giaretta et al. 1999, Levings & Windsor 1985). The determination of accurate annual average LSC data, may require monitoring over long periods due to seasonality and sometimes sporadic nature of litterfall and decomposition rates (Clark et al. 2001). Furthermore, the effects of topography and water movement create the need for both representative site selection and sufficient spatial coverage.

In a I-ha plot all ground herb species were recorded. Within the plot selected properties were investigated in three 20 m × 20 m subplots. The number of ground herb species and individuals in the subplots were lowest near to a local hilltop and they increased towards the lowest part of the plot. There was no significant difference in canopy openness, or in thickness of leaf litter layer, between the upper and lower subplots. In the upper subplot the soil layer was thin and the weathered bedrock was close to the surface. In the lower subplot the soil was much deeper. The pH and concentrations of Mg were significantly lower and %loss-on-ignition, total P and total N concentrations were significantly higher in the upper subplot. Elevation and soil moisture were significantly correlated in the upper 10 cm of soil. It is not possible to select any one abiotic property as being the critical determinant of the gradient of ground herbs. However, it is suspected that the availability of water in the topsoil, particularly after droughts, may be more important than the soil nutrients in limiting herb distribution.

A study of the benthic leaf litter community of the Tarumazinho, a central Amazonian stream (03° S, 60° W) is presented. This small tributary of the Rio Negro is of the blackwater type with a very low nutrient content. The sampling stations were towards the mouth of the tributary where it flowed through annually inundated forest and the work was carried out during the low water season. Previous studies concluded that these acid waters were of very low productivity and held impoverished faunas. However, this study showed that leaf litter banks held a diverse aquatic community of unexpectedly high biomass. This was because the banks supplied food and shelter and were available during the low water period when other habitats had dried out. The macrofauna was dominated by fish and shrimps. From an examination of gut contents a food web was produced. Primary consumers were found to take mainly detritus and fungi and it was concluded that allochthonous inputs were the main source of energy and nutrients. No feeding specialists were found and all the predators took prey from more than one trophic level.

Soil processes are essential in enabling forest regeneration in disturbed landscapes. Little is known about whether litterfall from dominating pioneer species in secondary rain forest is functionally equivalent to that of mixed rain-forest litter in terms of contribution to soil processes. This study used the litterbag technique to quantify the decomposition and nutrient dynamics of leaf litter characteristic of three wet tropical forest communities in the Paluma Range National Park, Queensland, Australia over 511 d. These were: undisturbed primary rain forest (mixed rain-forest species), selectively logged secondary rain forest (pioneer Alphitonia petriei) and tall open eucalypt forest (Eucalyptus grandis). Mass loss, total N, total P, K, Ca and Mg dynamics of the decaying leaves were determined, and different mathematical models were used to explain the mass loss data. Rainfall and temperature data were also collected from each site. The leaves of A. petriei and E. grandis both decomposed significantly slower in situ than the mixed rain-forest species (39%, 38% and 29% ash-free dry mass remaining respectively). Nitrogen and phosphorus were immobilized, with 182% N and 134% P remaining in E. grandis, 127% N and 132% P remaining in A. petriei and 168% N and 121% P remaining in the mixed rain-forest species. The initial lignin:P ratio and initial lignin:N ratio exerted significant controls on decomposition rates. The exceptionally slow decomposition of the pioneer species is likely to limit soil processes at disturbed tropical rain-forest sites in Australia.

The widespread use of forest litter as animal bedding in central Europe for many centuries gave rise to the first litter manipulation studies, and their results demonstrated that litter and its decomposition are a vital part of ecosystem function. Litter plays two major roles in forest ecosystems: firstly, litterfall is an inherent part of nutrient and carbon cycling, and secondly, litter forms a protective layer on the soil surface that also regulates microclimatic conditions. By reviewing 152 years of litter manipulation experiments, I show that the effects of manipulating litter stem from changes in one, or both, of these two functions, and interactions between the variables influenced by the accumulation of litter can result in feedback mechanisms that may intensify treatment effects or mask responses, making the interpretation of results difficult.

Long-term litter removal increased soil bulk density, overland flow, erosion, and temperature fluctuations and upset the soil water balance, causing lower soil water content during dry periods. Soil pH increased or decreased in response to manipulation treatments depending on forest type and initial soil pH, but it is unclear why there was no uniform response. Long-term litter harvesting severely depleted the forests of nutrients. Decreases in the concentrations of available P, Ca, Mg, and K in the soil occurred after only three to five years. The decline in soil N occurred over longer periods of time, and the relative loss was greater in soils with high initial nitrogen concentration. Tree growth declined with long-term litter removal, probably due to lower nutrient availability. Litter manipulation also added or removed large amounts of carbon thereby affecting microbial communities and altering soil respiration rates.

Litter manipulation experiments have shown that litter cover acts as a physical barrier to the shoot emergence of small-seeded species; further, the microclimate maintained by the litter layer may be favourable to herbivores and pathogens and is important in determining later seedling survival and performance. Litter manipulation altered the competitive outcomes between tree seedlings and forbs, thereby influencing species composition and diversity; changes in the species composition of understorey vegetation following treatments occurred fairly rapidly. By decreasing substrate availability and altering the microclimate, litter removal changed fungal species composition and diversity and led to a decline in populations of soil fauna. However, litter addition did not provoke a corresponding increase in the abundance or diversity of fungi or soil fauna.

Large-scale long-term studies are still needed in order to investigate the interactions between the many variables affected by litter, especially in tropical and boreal forests, which have received little attention. Litter manipulation treatments present an opportunity to assess the effects of increasing primary production in forest ecosystems; specific research aims include assessing the effects of changes in litter inputs on the carbon and nutrient cycles, decomposition processes, and the turnover of organic matter.

In forested ecosystems leaf litter is generally the primary pathway through which nutrients are cycled from the canopy to the forest floor (other pathways include throughfall, stemflow and animal faeces; Jordan 1985). Consequently, any disturbance that alters the quantity or quality of litter can have dramatic impacts on nutrient cycling and the availability of essential nutrients to plants (Vitousek 1984). Fragmentation of tropical forests has been demonstrated to cause several changes in both the biotic (Cosson et al. 1999, Laurance et al. 1998, Saunders et al. 1991) and abiotic environments (Camargo & Kapos 1995, Debinski & Holt 2000, Laurance 2002, Laurance et al. 2002) and thus may influence litter accumulation in the remnant patches (Carvalho & Vasconcelos 1999, Didham 1998, Laurance et al. 2002).

Plants often modify microsite conditions important for seedling establishment. In tropical moist forest, advance regeneration in the form of shade-suppressed seedlings is a major component of regrowth in new gaps. Tree seedlings may be filtered by the composition and structure of the forest understorey. In a lowland forest in Central Panama, we examined light availability, litter accumulation and the seedling community (abundance, diversity and composition) under and away from the canopies of a common subcanopy, clonal palm, Oenocarpus mapora, and tested whether seedling abundance varies as a function of changes in understorey light conditions and leaf litter depth. We found evidence that leaf litter was five times deeper and light availability 27% lower under O. mapora canopy than where it was absent. The probability of finding a seedling in plots under O. mapora canopies was 33% lower than in plots without the palm. Plots under O. mapora canopies also had 49% fewer species of seedlings and those seedlings came from significantly larger seeds than seedlings in plots not under the palm. Overall, seedling density was positively correlated with light availability and negatively correlated with leaf litter depth. A transplant experiment confirmed that for at least one large-seeded species of dicot, seedlings are negatively affected by the reduction in light availability and increase in leaf litter. Under O. mapora, seedlings of Gustavia superba (Lecythidaceae) had lower survivorship and biomass accumulation than away from the palm. Thus, site conditions produced by large, clonal, multi-layered palms such as O. mapora may reduce establishment of small-seeded and shade-intolerant species, potentially affecting species composition in the understorey seedling pool.

How do animals respond to an unpredictably heterogeneous environment? Ants foraging in the leaf litter of tropical wet forests experience unpredictably fluctuating food resources. To study how an ant species responds to these changes, foragers were tracked to determine home ranges of 51 colonies of Aphaenogaster araneoides, in three sites in a Costa Rican tropical wet forest. Of these colonies 16 were excavated to measure colony size, colony growth, and reproductive investment. These demographic variables were compared with two measures of home range quality: leaf litter dry weight and mass of arthropods. Home range areas of colonies were highly correlated with colony size, and moderately correlated with resource abundance. Colony growth was independent of colony size, as is found in other ants in unpredictable environments. The growth of colonies was closely associated with resource abundance. Production of the male reproductive caste was closely tied to the size of a colony rather than growth, but male production in slow-growing colonies was limited. Colonies foraging within high-quality environments grew at a faster rate, but reproduction was mainly correlated with colony size. Furthermore, it was found that the frequency of foragers in long-term treatment plots with supplemental food and reduced leaf-litter quality was not significantly different from the frequency of foragers in control plots. This rain-forest ant does not modify its home range areas in response to poor environments, and as a result, small-scale environmental heterogeneity strongly determines growth and reproduction.