Influence of Colophospermum mopane canopy cover on litter decomposition and nutrient dynamics in a semi‐arid African savannah

The objective of this study was to investigate the influence of mopane canopy cover on litter decomposition in a semi‐arid African savannah. We used a randomized block design with five blocks of 100 × 100 m demarcated in a 10‐ha pocket of open mopane woodland. Litterbags were placed beneath large (8.3 m crown diameter) and small mopane trees (2.7 m crown diameter) and in the intercanopy area. Decomposition was fastest in the intercanopy area exposed to solar radiation (k = 0.35 year^?1), intermediate beneath small trees (k = 0.28 year^?1) and slowest beneath large trees (k = 0.23 year^?1). Soil temperatures beneath small and large trees were 3–5 and 6–9°C lower than in the intercanopy area, respectively. Bacterial and fungal counts were significantly higher (P < 0.05) beneath large than small trees and in the intercanopy area. The amount of N and P released did not vary significantly among sampling sites. Soil moisture in the dry season was similar among sampling sites but rainy‐season soil moisture was significantly greater (P < 0.05) beneath large than small trees and in the intecanopy area. Mopane canopy cover retarded litter decomposition suggesting that photodegradation could be an important factor controlling carbon turnover in semi‐arid African savannahs.     

Conversion of secondary broadleaved forest into Chinese fir plantation alters litter production and potential nutrient returns

Litter production, litter standing crop, and potential nutrient return via litterfall to soil were studied during a 4-year period (January 2004–December 2007) in a Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) plantation and a secondary broadleaved forest in Hunan Province in subtropical China. Mean annual litterfall in the sampling sites varied from 358 g m⁻² in the pure plantation to 669 g m⁻² in the secondary broadleaved forest. Total litterfall followed a bimodal distribution pattern for both forests. Amount of litterfall was also related to the air temperature in both forests. During the period under this study, annual variation in the total litterfall in the pure plantation was significantly higher than that in the secondary broadleaved forest. Litterfall was markedly seasonal in the both forests. Leaf proportions of litterfall in the pure plantation and secondary broadleaved forest were 58.1 and 61.7%, respectively. Total potential nutrient returns to the soil through litterfall in the pure plantation were only 46.2% of those in the secondary broadleaved forest. Total litter standing crop was 913 and 807 g m⁻² in the pure plantation and secondary broadleaved forest, respectively. Our results confirm that conversion from a secondary broadleaved forest into a pure coniferous plantation changes the functioning of the litter system.     

Litterfall dynamics in carbonate and deltaic mangrove ecosystems in the Gulf of Mexico

From 1996 to 2002, we measured litterfall, standing litter crop, and litter turnover rates in scrub, basin, fringe and riverine forests in two contrasting mangrove ecosystems: a carbonate-dominated system in the Southeastern Everglades and a terrigenous-dominated system in Laguna de Terminos (LT), Mexico. We hypothesized that litter dynamics is driven by latitude, geomorphology, hydrology, soil fertility and soil salinity stress. There were significant temporal patterns in LT with litterfall rates higher during the rainy season (2.4 g m⁻² day⁻¹) than during the dry season (1.8 g m⁻² day⁻¹). Total annual litterfall was significantly higher in the riverine forest (12.8 Mg ha⁻² year⁻¹) than in the fringe and basin forests (9.7 and 5.2 Mg ha⁻² year⁻¹, respectively). In Southeastern Everglades, total annual litterfall was also significantly higher during the rainy season than during the dry season. Spatially, the scrub forest had the lowest annual litterfall (2.5 Mg ha⁻² year⁻¹), while the fringe and basin had the highest (9.1 and 6.5 Mg ha⁻² year⁻¹, respectively). In LT, annual standing litter crop was 3.3 Mg ha⁻¹ in the fringe and 2.2 Mg ha⁻¹ in the basin. Litter turnover rates were significantly higher in the fringe mangrove forest (4.1 year⁻¹) relative to the basin forests (2.2 year⁻¹). At Southeastern Everglades there were significant differences in annual standing litter crop: 1.9, 3.3 and 4.5 Mg ha⁻¹ at scrub, basin and fringe mangrove sites, respectively. Furthermore, turnover rates were similar at both basin and fringe mangrove types (2.1 and 2.0 year⁻¹, respectively) but significantly higher than scrub mangrove forest (1.3 year⁻¹). These findings suggest that litter export is important in regulating litter turnover rates in frequently flooded riverine and fringe forests, while in infrequently flooded basin forests, in situ litter decomposition controls litter turnover rates.     

Temporal and structural effects of stands on litter production in Melaleuca quinquenervia dominated wetlands of south Florida

Melaleuca quinquenervia dominates large areas of the Florida Everglades in the southeastern USA where it has transformed sedge-dominated marshes into melaleuca forests. Despite its prevalence, very little is known about the ecology and stand dynamics of this invasive tree. We delineated large-, intermediate-, and small-tree stands in non-flooded, seasonally flooded and permanently flooded areas of Florida in 1997, measured their biological attributes, and then quantified litterfall components for 3–4 year periods. Melaleuca wood components and mature seed-capsules comprised the largest and the smallest portions of aboveground biomass, respectively, while leaves, fine stems, mature fruits, bud scales, floral structures, and residues represented decreasingly smaller fractions of the litter during the succeeding year. Dry weight proportion of leaves in litter was greatest (80.9%) in non-flooded and least (69.1%) in permanently flooded habitats. It was also greatest in small (85.6%) and least in large (64.7%) tree stands. Reproductive structures and mature-fruit fractions in litter were highest in large-tree stands whereas the bud-scale fraction showed no relationship to tree size. Seasonally flooded habitats had the most litterfall, wherein small-, intermediate-, and large-tree stands generated 0.662, 0.882, and 1.128 kg m⁻² yr⁻¹, respectively. Dry weight of stems, leaves, bud–scales, floral structures, and mature fruit fractions in litter increased as the predominant size of the trees in the stand increased. Total annual litter production was highest during 1999–2000. Leaf fall occurred year-round with maximal amount during April, July, and October. Highest amounts of bud scales and floral structures fell during October–January, which corresponded with flushes of vegetative growth and major flowering events. Overall, melaleuca alone accounted for nearly 99% of the total litterfall dry weight in all habitats and months sampled. The amount of non-melaleuca litter was greater in small-tree stands than in intermediate- or large-tree stands. Litterfall data of this nature will be helpful in detecting changes occurring in melaleuca canopies in response to biological control impact and in prescribing site-specific management strategies.     

High litterfall in old-growth and secondary upper montane forest of Costa Rica

Tropical upper montane forests usually comprise trees of small stature with a relatively low aboveground productivity. In contrast to this rule, in the Cordillera de Talamanca (Costa Rica), tall trees (>35 m in height and more than 60 cm in diameter) are characteristic for the upper montane old-growth oak forests which are growing at an altitude of almost 3,000 m close to the alpine timberline. For these exceptional forests, productivity data are not yet available. In this study, we analyzed litterfall and its components (tree leaves, litter of epiphytic vascular and non-vascular plants, mistletoes, twigs and other canopy debris) in three forest stands belonging to different successional stages and related seasonal changes in litterfall to micrometeorological variables. The studied stands were early-successional forest (10–15-year-old), mid-successional forest (40-year-old), and old-growth forest. The stands are dominated by Quercus copeyensis and are located at 2,900-m altitude. Total litterfall was highest in the mid-successional forest (1,720 g m⁻² y⁻¹), and reached 1,288 g m⁻² y⁻¹ in the old-growth forest and 934 g m⁻² y⁻¹ in the early-successional forest. Litter mass was dominated by leaves in all stages (56–84% of total litterfall). In the old-growth forest, however, twigs and small canopy debris particles (33%), epiphytes (6%), and mistletoes (5%) also contributed substantially to litter mass. Leaf litterfall showed a clear seasonal pattern with a negative correlation to monthly precipitation and highest values in the dry season (January–April). However, the strongest correlation existed with minimum air temperature (negative), probably because temperatures already dropped at the end of the rainy season, when precipitation had not yet declined and leaf shedding already increased. In contrast, litterfall of epiphyte mass, and twigs and other debris was mostly dependent on occasional strong winds. We conclude that the upper montane oak forests of the Cordillera de Talamanca are exceptional with respect to the large tree size and the relatively high productivity as indicated by litterfall. Litter mass was especially high in the mid-successional and old-growth forests, where the observed annual totals are among the highest recorded for tropical forests so far.     

Dynamics of soil respiration in sparse <Emphasis Type="Italic">Ulmus pumila</Emphasis> woodland under semi-arid climate

Sparse Ulmus pumila woodlands play an important role in contributing to ecosystem function in semi-arid grassland of northern China. To understand the key attributes of soil carbon cycling in U. pumila woodland, we studied dynamics of soil respiration in the canopy field (i.e., the projected crown cover area) and the open field at locations differing in distance (i.e., at 1–1.5, 3–4, 10, and >15 m) to tree stems from July through September of 2005, and measured soil biotic factors (e.g., fine root mass, soil microbial biomass, and activity) and abiotic factors [e.g., soil water content (SWC) and organic carbon] in mid-August. Soil respiration was further separated into root component and microbial component at the end of the field measurement in September. Results showed that soil respiration had a significant exponent relationship with soil temperature at 10-cm depth. The temperature sensitivity index of soil respiration, Q ₁₀, was lower than the global average of 2.0, and declined significantly (P < 0.05) with distance. The rate of soil respiration was generally greater in the canopy field than in the open field; monthly mean of soil respiration was 305.5–730.8 mg CO₂ m⁻² h⁻¹ in the canopy field and 299.6–443.1 mg CO₂ m⁻² h⁻¹ in the open field from July through September; basal soil respiration at 10°C declined with distance, and varied from ~250 mg CO₂ m⁻² h⁻¹ near tree stems to <200 mg CO₂ m⁻² h⁻¹ in the open field. Variations in soil respiration with distance were consistent with patterns of SWC, fine root mass, microbial biomass and activities. Regression analysis indicated that soil respiration was tightly coupled with microbial respiration and only weakly related to root respiration. Overall, variations in SWC, soil nutrients, microbial biomass, and microbial activity are largely responsible for the spatial heterogeneity of soil respiration in this semi-arid U. pumila woodland.     

Litterfall deposition and leaf litter nutrient return in different locations at Northeastern Mexico

The aim of this study was to determine the litterfall production and macronutrient (Ca, K, Mg, N, and P) deposition through leaf litter in four sites with different types of vegetation. Site one (Bosque Escuela) was located at 1600 m a.s.l. in a pine forest mixed with deciduous trees, second site (Crucitas at 550 m a.s.l.) in the ecotone of a Quercus spp. forest and the Tamaulipan thornscrub and third and fourth sites (Campus at 350 m a.s.l. and Cascajoso at 300 m a.s.l., respectively) were in the Tamaulipan thornscrub. Litter constituents (leaves, reproductive structures, twigs, and miscellaneous residues) were collected at 15-day intervals from December 21, 2006, throughout December 20, 2007. Collections were carried out in ten litter traps (1.0 × 1.0 m) randomly situated at each site of approximately 2,500 m². Total annual litterfall deposition was 4407, 7397, 6304, and 6527 kg ha⁻¹ y⁻¹ for Bosque Escuela, Crucitas, Campus and Cascajoso, respectively. Of total annual litter production, leaves were higher varying from 74 (Bosque Escuela) to 86% (Cascajoso) followed by twigs from 4 (Cascajoso) to 14% (Crucitas), reproductive structures from 6 (Bosque Escuela) to 10% (Crucitas), and miscellaneous litterfall from <1 (Campus) to 12% (Bosque Escuela). The Ca annual deposition was significantly higher in Cascajoso (232.7 kg ha⁻¹ y⁻¹), followed by Campus (182.3), Crucitas (130.5) and Bosque Escuela (30.3). The K (37.5, 32.5, 24.8, 7.2, respectively), Mg (22.6, 17.7, 13.7, 4.5, respectively) followed the same pattern as Ca. However, N was higher in Campus (85.8) followed by Crucitas (85.1), Cascajoso (68.3), and Bosque Escuela (18.3). The P was higher in Campus and Crucitas (4.0) followed by Cascajoso (3.4) and Bosque Escuela (1.4). On an annual basis for all sites, the order of nutrient deposition through leaf litter was Ca > N> K > Mg > P, whereas on site basis of total nutrient deposition (Ca + N + K + Mg + P), the order was Cascajoso > Campus > Crucitas > Bosque Escuela. Ca, K, Mg, N, and P nutrient use efficiency values in leaf litter were higher in Bosque Escuela, while lower figures were acquired in Cascajoso and Crucitas sites. It seems that the highest litterfall deposition was found in the ecotone of a Quercus spp. forest and the Tamaulipan thornscrub; however, the Tamaulipan thornscrub vegetation alone had better leaf litter nutrient return.     

Total Belowground Carbon Allocation in a Fast-growing Eucalyptus Plantation Estimated Using a Carbon Balance Approach

Trees allocate a large portion of gross primary production belowground for the production and maintenance of roots and mycorrhizae. The difficulty of directly measuring total belowground carbon allocation (TBCA) has limited our understanding of belowground carbon (C) cycling and the factors that control this important flux. We measured TBCA over 4 years using a conservation of mass, C balance approach in replicate stands of fast growing Eucalyptus saligna Smith with different nutrition management and tree density treatments. We measured TBCA as surface carbon dioxide (CO₂) efflux (“soil” respiration) minus C inputs from aboveground litter plus the change in C stored in roots, litter, and soil. We evaluated this C balance approach to measuring TBCA by examining (a) the variance in TBCA across replicate plots; (b) cumulative error associated with summing components to arrive at our estimates of TBCA; (c) potential sources of error in the techniques and assumptions; (d) the magnitude of changes in C stored in soil, litter, and roots compared to TBCA; and (e) the sensitivity of our measures of TBCA to differences in nutrient availability, tree density, and forest age. The C balance method gave precise estimates of TBCA and reflected differences in belowground allocation expected with manipulations of fertility and tree density. Across treatments, TBCA averaged 1.88 kg C m⁻² y⁻¹ and was 18% higher in plots planted with 10⁴ trees/ha compared to plots planted with 1111 trees/ha. TBCA was 12% lower (but not significantly so) in fertilized plots. For all treatments, TBCA declined linearly with stand age. The coefficient of variation (CV) for TBCA for replicate plots averaged 17%. Averaged across treatments and years, annual changes in C stored in soil, the litter layer, and coarse roots (−0.01, 0.06, and 0.21 kg C m⁻² y⁻¹, respectively) were small compared with surface CO₂ efflux (2.03 kg C m⁻² y⁻¹), aboveground litterfall (0.42 kg C m⁻² y⁻¹), and our estimated TBCA (1.88 kg C m⁻² y⁻¹). Based on studies from similar sites, estimates of losses of C through leaching, erosion, or storage of C in deep soil were less than 1% of annual TBCA. Received 6 March 2001; accepted 7 January 2002.     

The influence of mistletoes on nutrient cycling in a semi-arid savanna, southwest Zimbabwe

Parasitic plants are increasingly becoming the focus of research in many ecosystems. They have been shown to alter litterfall properties and decomposition rates in environments where they occur. Despite this recognition, the role of mistletoes in nutrient cycling in semi-arid savanna remains poorly understood. We investigated the litter input, element returns, and associated below-canopy soil nutrient concentrations of three mistletoe species (Erianthemum ngamicum, Plicosepalus kalachariensis, and Viscum verrucosum) that parasitize Acacia karroo trees in a semi-arid savanna, southwest Zimbabwe. Element concentrations in mistletoe leaf litter were enriched relative to the host. Litterfall from mistletoes significantly increased overall litterfall by up to 173 %, with E. ngamicum and P. kalachariensis having greater litterfall than their host trees. Associated with these changes in litterfall was an increase in element returns and the below-canopy soil nutrient concentrations. The increase in nutrient returns was due to both the effect of enriched mistletoe litter and increased volumes of litterfall beneath host trees. Litterfall, element returns, and the below-canopy soil nutrient concentrations were significantly influenced by mistletoe density, with higher values at high mistletoe density. Overall, E. ngamicum and P. kalachariensis had greater influence on litterfall, element returns, and soil nutrient concentrations than V. verrucosum. These findings are consistent with current understanding of enhanced nutrient cycling in the presence of parasitic plants particularly in nutrient-poor ecosystems. We conclude that the introduction of nutrients and associated increase in resource heterogeneity play an important role in determining ecosystem structure and function in semi-arid savannas.     

Soil Nutrients Limit Fine Litter Production and Tree Growth in Mature Lowland Forest of Southwestern Borneo

Efforts to improve models of terrestrial productivity and to understand the function of tropical forests in global carbon cycles require a mechanistic understanding of spatial variation in aboveground net primary productivity (ANPP) across tropical landscapes. To help derive such an understanding for Borneo, we monitored aboveground fine litterfall, woody biomass increment and ANPP (their sum) in mature forest over 29 months across a soil nutrient gradient in southwestern Kalimantan. In 30 (0.07 ha) plots stratified throughout the watershed (∼340 ha, 8–190 m a.s.l.), we measured productivity and tested its relationship with 27 soil parameters. ANPP across the study area was among the highest reported for mature lowland tropical forests. Aboveground fine litterfall ranged from 5.1 to 11.0 Mg ha⁻¹ year⁻¹ and averaged 7.7 ± 0.4 (mean ± 95 C.I.). Woody biomass increment ranged from 5.8 to 23.6 Mg ha⁻¹ year⁻¹ and averaged 12.0 ± 2.0. Growth of large trees (≥60 cm dbh) contributed 38–82% of plot-wide biomass increment and explained 92% of variation among plots. ANPP, the sum of these parameters, ranged from 11.1 to 32.3 Mg ha⁻¹ year⁻¹ and averaged 19.7 ± 2.2. ANPP was weakly related to fine litterfall (r ² = 0.176), but strongly related to growth of large trees at least 60 cm dbh (r ² = 0.848). Adjusted ANPP after accounting for apparent “mature forest bias” in our sampling method was 17.5 ± 1.2 Mg ha⁻¹ year⁻¹.Relating productivity measures to soil parameters showed that spatial patterning in productivity was significantly related to soil nutrients, especially phosphorus (P). Fine litterfall increased strongly with extractable P (r ² = 0.646), but reached an asymptote at moderate P levels, whereas biomass increment (r ² = 0.473) and ANPP (r ² = 0.603) increased linearly across the gradient. Biomass increment of large trees was more frequently and strongly related to nutrients than small trees, suggesting size dependency of tree growth on nutrients. Multiple linear regression confirmed the leading importance of soil P, and identified Ca as a potential co-limiting factor. Our findings strongly suggest that (1) soil nutrients, especially P, limit aboveground productivity in lowland Bornean forests, and (2) these forests play an important, but changing role in carbon cycles, as canopy tree logging alters these terrestrial carbon sinks. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differences in litter characteristics and macroinvertebrate assemblages between litter patches in pools and riffles in a headwater stream

We compared the abundance of litter categories (coarse particulate organic matter 1–16 mm, leaves >16 mm, and small woody detritus 16–100 mm) and macroinvertebrate assemblages between natural litter patches in pools and riffles in a headwater stream. Litter patches in pools were formed under conditions of almost no current (<6 cm s⁻¹), but in riffles they were formed under variable current velocities (13–89 cm s⁻¹). Although the abundance of each litter category exhibited seasonal change, leaves were more abundant in riffles, and coarse particulate organic matter and small woody detritus were more abundant in pools throughout the study period. Macroinvertebrate assemblages in pools and riffles also changed seasonally but distinctly differed from each other. Shredders, collector-gatherers, and predators were the dominant functional groups in abundance in both pools and riffles, but the dominant shredders were caddisflies in pools and stoneflies in riffles. It is considered that the hydraulic conditions affected macroinvertebrate assemblages directly and indirectly through influences on the characteristics of litter retained in the patches. Our results suggest that the relative abundance of litter patches in pools and riffles largely affects the macroinvertebrate community structure of the headwater stream. Received: July 19, 2001 / Accepted: December 19, 2001     

Culm recruitment,dry matter dynamics and carbon flux in recently harvested and mature bamboo savannas in the Indian dry tropics

Culm recruitment, standing crop biomass, net production and carbon flux were estimated in mature (5 years after last harvest) and recently harvested bamboo (Dendrocalamus strictus (Roxb.) Nees) savanna sites in the dry tropics. During the 2 study years bamboo shoot recruitment was 1711–3182 and 1432–1510 shoots ha⁻¹ in harvested and mature sites, respectively. Corresponding shoot mortality was 66–93% and 62–69%, respectively. Total biomass was 34.9 t ha⁻¹ at the harvested site and 47.4 t ha⁻¹ at the mature site. Harvesting increased the relative contribution of belowground bamboo biomass. Annual litter input to soil was 2.7 and 5.9 t ha⁻¹ year⁻¹ at the harvested and mature sites, respectively. The bulk of the annual litterfall (78–88%) occurred in the cool dry season (November to February). The mean litter mass on the savanna floor ranged from 3.1 to 3.3 t ha⁻¹; at the harvested site wood litter contributed 70% of the litter mass and at the mature site leaves formed 77% of the litter mass. The mean total net production (TNP) for the two annual cycles was 15.8 t ha⁻¹ year⁻¹ at the harvested site and 19.3 t ha⁻¹ year⁻¹ at the mature site. Nearly half (46–57%) of the TNP was allocated to the belowground parts. Short lived components (leaves and fine roots) contributed about four-fifths of the net production of bamboo. Total carbon storage in the system was 64.4 t ha⁻¹ at the harvested site and 75.4 t ha⁻¹ at the mature site, of which 23–28% was distributed in vegetation, 2% in litter and 70–75% in soil. Annual net carbon deposition was 6.3 and 8.7 t ha⁻¹ year⁻¹ at harvested and mature sites, respectively.     

Invasion by a Perennial Herb Increases Decomposition Rate and Alters Nutrient Availability in Warm Temperate Lowland Forest Remnants

We determined the impact of the invasive herb, Tradescantia fluminensis Vell., on litter decomposition and nutrient availability in a remnant of New Zealand lowland podocarp–broadleaf forest. Using litter bags, we found that litter beneath mats of Tradescantia decomposed at almost twice the rate of litter placed outside the mat. Values of k (decomposition quotient) were 9.44±0.42 yrs for litter placed beneath Tradescantia and 5.42±0.42 yrs for litter placed in native, non-Tradescantia plots. The impact of Tradescantia on decomposition was evident through the smaller forest floor mass in Tradescantia plots (2.65±1.05 t ha⁻¹) compared with non-Tradescantia plots (5.05±1.05 t ha⁻¹), despite similar quantities of annual leaf litterfall into Tradescantia plots (6.85±0.85 t ha⁻¹ yr⁻¹) and non-Tradescantia plots (7.45±1.05 t ha⁻¹ yr⁻¹). Moreover, there was increased plant nitrate available, as captured on resin bags, in Tradescantia plots (25.77 ± 8.32 cmol(−)/kg resin) compared with non-Tradescantia plots (9.55±3.72 cmol(−)/kg resin). Finally, the annual nutrient uptake by Tradescantia represented a large proportion of nutrients in litterfall (41% N, 61% P, 23% Ca, 46% Mg and 83% K), exceeded the nutrient content of the forest floor (except Ca), but was a small proportion of the topsoil nutrient pools. Taken together, our results show that Tradescantia increases litter decomposition and alters nutrient availability, effects that could influence the long-term viability of the majority of podocarp–broadleaf forest remnants affected with Tradescantia in New Zealand. These impacts are likely mostly due to Tradescantia's vegetation structure (i.e., tall, dense mats) and associated microclimate, compared with native ground covers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Allochthonous litter inputs, organic matter standing stocks, and organic seston dynamics in upland Panamanian streams: potential effects of larval amphibians on organic matter dynamics

Allochthonous inputs of detritus represent an important energy source for streams in forested regions, but dynamics of these materials are not well studied in neotropical headwater streams. As part of the tropical amphibian declines in streams (TADS) project, we quantified benthic organic matter standing stocks and organic seston dynamics in four Panamanian headwater streams, two with (pre-amphibian decline) and two without (post-decline) healthy amphibian assemblages. We also measured direct litterfall and lateral litter inputs in two of these streams. Continuous litterfall and monthly benthic samples were collected for 1 year, and seston was collected 1–3 times/month for 1 year at or near baseflow. Direct litterfall was similar between the two streams examined, ranging from 934–1,137 g DM m⁻² y⁻¹. Lateral inputs were lower, ranging from 140–187 g DM m⁻¹ y⁻¹. Dead leaves (57–60%), wood (24–29%), and green leaves (8–9%) contributed most to inputs, and total inputs were generally higher during the rainy season. Annual habitat-weighted benthic organic matter standing stocks ranged from 101–171 g AFDM m⁻² across the four study reaches, with ∼4 × higher values in pools compared to erosional habitats. Total benthic organic matter (BOM) values did not change appreciably with season, but coarse particulate organic matter (CPOM, >1 mm) generally decreased and very fine particulate organic matter (VFPOM, 1.6–250 μm) generally increased during the dry season. Average annual seston concentrations ranged from 0.2–0.6 mg AFDM l⁻¹ (fine seston, <754 μm >250 μm) and 2.0–4.7 mg AFDM l⁻¹ (very fine, <250 μm >1.6 μm), with very fine particles composing 85–92% of total seston. Quality of fine seston particles in the two reaches where tadpoles were present was significantly higher (lower C/N) than the two where tadpoles had been severely reduced (P = 0.0028), suggesting that ongoing amphibian declines in this region are negatively influencing the quality of particles exported from headwaters. Compared to forested streams in other regions, these systems receive relatively high amounts of allochthonous litter inputs but have low in-stream storage. Handling editor: J. Padisak     

Litter pool sizes,decomposition, and nitrogen dynamics in <Emphasis Type="Italic">Spartina alterniflora</Emphasis>-invaded and native coastal marshlands of the Yangtze Estuary

Past studies have focused primarily on the effects of invasive plants on litter decomposition at soil surfaces. In natural ecosystems, however, considerable amounts of litter may be at aerial and belowground positions. This study was designed to examine the effects of Spartina alterniflora invasion on the pool sizes and decomposition of aerial, surficial, and belowground litter in coastal marshlands, the Yangtze Estuary, which were originally occupied by two native species, Scirpus mariqueter and Phragmites australis. We collected aerial and surficial litter of the three species once a month and belowground litter once every 2 months. We used the litterbag method to quantify litter decomposition at the aerial, surficial and belowground positions for the three species. Yearly averaged litter mass in the Spartina stands was 1.99 kg m⁻²; this was 250 and 22.8% higher than that in the Scirpus (0.57 kg m⁻²) and Phragmites (1.62 kg m⁻²) stands, respectively. The litter in the Spartina stands was primarily distributed in the air (45%) and belowground (48%), while Scirpus and Phragmites litter was mainly allocated to belowground positions (85 and 59%, respectively). The averaged decomposition rates of aerial, surficial, and belowground litter were 0.82, 1.83, and 1.27 year⁻¹ for Spartina, respectively; these were 52, 62 and 69% of those for Scirpus litter at corresponding positions and 158, 144 and 78% of those for Phragmites litter, respectively. The differences in decomposition rates between Spartina and the two native species were largely due to differences in litter quality among the three species, particularly for the belowground litter. The absolute amount of nitrogen increased during the decomposition of Spartina stem, sheath and root litter, while the amount of nitrogen in Scirpus and Phragmites litter declined during decomposition for all tissue types. Our results suggest that Spartina invasion altered the carbon and nitrogen cycling in the coastal marshlands of China.     

Impacts of the exotic,nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen-cycling in a pine–oak ecosystem

We investigated the influence of the exotic nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen cycling in a pitch pine (Pinus rigida) −scrub oak (Quercus ilicifolia, Q. prinoides) ecosystem. Within paired pine-oak and adjacent black locust stands that were the result of a 20-35 year-old invasion, we evaluated soil nutrient contents, soil nitrogen transformation rates, and annual litterfall biomass and nitrogen concentrations. In the A horizon, black locust soils had 1.3-3.2 times greater nitrogen concentration relative to soils within pine-oak stands. Black locust soils also had elevated levels of P and Ca, net nitrification rates and total net N-mineralization rates. Net nitrification rates were 25-120 times greater in black locust than in pine-oak stands. Elevated net N-mineralization rates in black locust stands were associated with an abundance of high nitrogen, low lignin leaf litter, with 86 kg N ha^–1 yr^–1 in leaf litter returned compared with 19 kg N ha^–1 yr^–1 in pine-oak stands. This difference resulted from a two-fold greater litterfall mass combined with increased litter nitrogen concentration in black locust stands (1.1% and 2.6% N for scrub oak and black locust litter, respectively). Thus, black locust supplements soil nitrogen pools, increases nitrogen return in litterfall, and enhances soil nitrogen mineralization rates when it invades nutrient poor, pine-oak ecosystems. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nutrient transfer by leaf litterfall during a sugar maple decline episode at Lake Clair watershed,Québec,Canada

In a declining sugar maple (SM) stand, we tested the hypothesis that an increasing relative abundance of American beech (AB) and yellow birch (YB) would improve litter quality by providing a higher proportion of litterfall richer in base cations and lower in acidity. From 1989 to 2006, SM leaf fall diminished from 59% (1,718 kg ha⁻¹ year⁻¹) to 36% (915 kg ha⁻¹ year⁻¹) of the total leaf fall biomass. Overall, the increase in AB and YB litterfall compensated for the SM decrease, resulting in constant annual leaf litterfall fluxes (2,803 kg ha⁻¹ year⁻¹) over the period studied. However, because the leaf litter for AB and YB had Ca and Mg concentrations 2–3 times higher than did SM, Ca and Mg concentrations and fluxes in leaf litterfall significantly increased between 1989 and 2006. Leaf litterfall of AB and YB also has a higher base/acid ratio than SM. Consequently, changes in forest composition following SM decline led to a clear improvement in litterfall quality in terms of base cations content and fluxes and acid–base properties.     

The influence of a neotropical herbivore (<Emphasis Type="Italic">Lamponius portoricensis</Emphasis>) on nutrient cycling and soil processes

The role of phytophagous insects in ecosystem nutrient cycling remains poorly understood. By altering the flow of litterfall nutrients from the canopy to the forest floor, herbivores may influence key ecosystem processes. We manipulated levels of herbivory in a lower montane tropical rainforest of Puerto Rico using the common herbivore, Lamponius portoricensis (Phasmatidea), on a prevalent understory plant, Piper glabrescens (Piperaceae), and measured the effects on nutrient input to the forest floor and on rates of litter decomposition. Four treatment levels of herbivory generated a full range of leaf area removal, from plants experiencing no herbivory to plants that were completely defoliated (>4,000 cm² leaf area removed during the 76-day study duration). A significant (P<0.05) positive regression was found between all measures of herbivory (total leaf area removed, greenfall production, and frass-related inputs) and the concentration of NO ₃ ⁻ in ion exchange resin bags located in the litter layer. No significant relationship was found between any of the herbivory components and resin bag concentrations of NH ₄ ⁺ or PO ₄ ⁻ . Rates of litter decay were significantly affected by frass-related herbivore inputs. A marginally significant negative relationship was also found between the litter mass remaining at 47 days and total leaf area removed. This study demonstrated a modest, but direct relationship between herbivory and both litter decomposition and NO ₃ ⁻ transfer to the forest floor. These results suggest that insect herbivores can influence forest floor nutrient dynamics and thus merit further consideration in discussions on ecosystem nutrient dynamics.     

Nitrogen and phosphorus economy of the riparian shrub <Emphasis Type="Italic">Salix gracilistyla</Emphasis> in western Japan

Salix gracilistyla is one of the dominant plants in the riparian vegetation of the upper-middle reaches of rivers in western Japan. This species colonizes mainly sandy habitats, where soil nutrient levels are low, but shows high potential for production. We hypothesized that S.␣gracilistyla uses nutrients conservatively within stands, showing a high resorption efficiency during leaf senescence. To test this hypothesis, we examined seasonal changes in nitrogen (N) and phosphorus (P) concentrations in aboveground organs of S. gracilistyla stands on a fluvial bar in the Ohtagawa River, western Japan. The concentrations in leaves decreased from April to May as leaves expanded. Thereafter, the concentrations showed little fluctuation until September. They declined considerably in autumn, possibly owing to nutrient resorption. We converted the nutrient concentrations in each organ to nutrient amounts per stand area on the basis of the biomass of each organ. The resorption efficiency of N and P in leaves during senescence were estimated to be 44 and 46%, respectively. Annual net increments of N and P in aboveground organs, calculated by adding the amounts in inflorescences and leaf litter to the annual increments in perennial organs, were estimated to be 9.9 g and 0.83 g m⁻² year⁻¹, respectively. The amounts released in leaf litter were 6.7 g N and 0.44 g P m⁻². These values are comparable to or larger than those of other deciduous trees. We conclude that S. gracilistyla stands acquire large amounts of nutrients and release a large proportion in leaf litter.     

Pools and fluxes of carbon in three Norway spruce ecosystems along a climatic gradient in Sweden

This paper presents an integrated analysis of organic carbon (C) pools in soils and vegetation, within-ecosystem fluxes and net ecosystem exchange (NEE) in three 40-year old Norway spruce stands along a north-south climatic gradient in Sweden, measured 2001–2004. A process-orientated ecosystem model (CoupModel), previously parameterised on a regional dataset, was used for the analysis. Pools of soil organic carbon (SOC) and tree growth rates were highest at the southernmost site (1.6 and 2.0-fold, respectively). Tree litter production (litterfall and root litter) was also highest in the south, with about half coming from fine roots (<1 mm) at all sites. However, when the litter input from the forest floor vegetation was included, the difference in total litter input rate between the sites almost disappeared (190–233 g C m⁻² year⁻¹). We propose that a higher N deposition and N availability in the south result in a slower turnover of soil organic matter than in the north. This effect seems to overshadow the effect of temperature. At the southern site, 19% of the total litter input to the O horizon was leached to the mineral soil as dissolved organic carbon, while at the two northern sites the corresponding figure was approx. 9%. The CoupModel accurately described general C cycling behaviour in these ecosystems, reproducing the differences between north and south. The simulated changes in SOC pools during the measurement period were small, ranging from −8 g C m⁻² year⁻¹ in the north to +9 g C m⁻² year⁻¹ in the south. In contrast, NEE and tree growth measurements at the northernmost site suggest that the soil lost about 90 g C m⁻² year⁻¹. An erratum to this article can be found at