Introduced ungulate herbivore alters soil processes after fire

Ungulate herbivory can have profound effects on ecosystem processes by altering organic inputs of leaves and roots as well as changing soil physical and chemical properties. These effects may be especially important when the herbivore is an introduced species. Utilizing large mammal exclosures to prevent access by introduced elk at multiple sites along a fire chronosequence, we examined the effects of elk herbivory and fire on soil microbial activity and nutrient availability. Using time since fire as a co-variate and herbivore exclosures, paired with areas outside of the exclosures, we hypothesized that reductions in plant biomass due to herbivory would reduce organic inputs to soils and impact soil microbial activities and nutrient storage. We found three major patterns: (1) when elk were excluded, surface mineral soils had higher soil organic carbon (C), total nitrogen (N), microbial N pools, and increased extra-cellular enzyme activity of a C-acquiring enzyme across a gradient of time since fire. (2) When introduced elk are present, the activity of some extracellular enzymes as well as NO₃ ⁻ availability are enhanced in the soil but the post-fire patterns described above with respect to nutrient accrual over time are delayed. (3) Herbivory by an introduced ungulate upsets the trajectory of ecosystem “recovery” after wildfire and delays soil C and N dynamics by an estimated 14.5–21 years, respectively. These results suggest that introduced, browsing herbivores significantly decelerate ecosystem processes but herbivory by exotics may also result in unpredictability in specific soil responses.     

An invasive frog, Eleutherodactylus coqui, increases new leaf production and leaf litter decomposition rates through nutrient cycling in Hawaii

A frog endemic to Puerto Rico, Eleutherodactylus coqui, invaded Hawaii in the late 1980s, where it can reach densities of 50,000 individuals ha⁻¹. Effects of this introduced insectivore on invertebrate communities and ecosystem processes, such as nutrient cycling, are largely unknown. In two study sites on the Island of Hawaii, we studied the top-down effects of E. coqui on aerial, herbivorous, and leaf litter invertebrates; herbivory, plant growth, and leaf litter decomposition rates; and leaf litter and throughfall chemistry over 6 months. We found that E. coqui reduced all invertebrate communities at one of the two study sites. Across sites, E. coqui lowered herbivory rates, increased NH₄⁺ and P concentrations in throughfall, increased Mg, N, P, and K in decomposing leaf litter, increased new leaf production of Psidium cattleianum, and increased leaf litter decomposition rates of Metrosideros polymorpha. In summary, E. coqui effects on invertebrates differed by site, but E. coqui effects on ecosystem processes were similar across sites. Path analyses suggest that E. coqui increased the number of new P. cattleianum leaves and leaf litter decomposition rates of M. polymorpha by making nutrients more available to plants and microbes rather than through changes in the invertebrate community. Results suggest that E. coqui in Hawaii has the potential to reduce endemic invertebrates and increase nutrient cycling rates, which may confer a competitive advantage to invasive plants in an ecosystem where native species have evolved in nutrient-poor conditions.     

Effects of elevated CO2 and O3 on leaf damage and insect abundance in a soybean agroecosystem

By altering myriad aspects of leaf chemistry, increasing concentrations of CO₂ and O₃ in the atmosphere derived from human activities may fundamentally alter the relationships between insect herbivores and plants. Because exposure to elevated CO₂ can alter the nutritional value of leaves, some herbivores may increase consumption rates to compensate. The effects of O₃ on leaf nutritional quality are less clear; however, increased senescence may also reduce leaf quality for insect herbivores. Additionally, changes in secondary chemistry and the microclimate of leaves may render plants more susceptible to herbivory in elevated CO₂ and O₃. Damage to soybean (Glycine max L.) leaves and the size and composition of the insect community in the plant canopy were examined in large intact plots exposed to elevated CO₂ (~550 μmol mol⁻¹) and elevated O₃ (1.2*ambient) in a fully factorial design with a Soybean Free Air Concentration Enrichment system (SoyFACE). Leaf area removed by folivorous insects was estimated by digital photography and insect surveys were conducted during two consecutive growing seasons, 2003 and 2004. Elevated CO₂ alone and in combination with O₃ increased the number of insects and the amount of leaf area removed by insect herbivores across feeding guilds. Exposure to elevated CO₂ significantly increased the number of western corn rootworm (Diabrotica virgifera) adults (foliage chewer) and soybean aphids (Aphis glycines; phloem feeder). No consistent effect of elevated O₃ on herbivory or insect population size was detected. Increased loss of leaf area to herbivores was associated with increased carbon-to-nitrogen ratio and leaf surface temperature. Soybean aphids are invasive pests in North America and new to this ecosystem. Higher concentrations of CO₂ in the atmosphere may increase herbivory in the soybean agroecosystem, particularly by recently introduced insect herbivores. Handling editor: Gary Felton.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

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.     

Recovery of ecosystem carbon fluxes and storage from herbivory

The carbon (C) sink strength of arctic tundra is under pressure from increasing populations of arctic breeding geese. In this study we examined how CO₂ and CH₄ fluxes, plant biomass and soil C responded to the removal of vertebrate herbivores in a high arctic wet moss meadow that has been intensively used by barnacle geese (Branta leucopsis) for ca. 20 years. We used 4 and 9 years old grazing exclosures to investigate the potential for recovery of ecosystem function during the growing season (July 2007). The results show greater above- and below-ground vascular plant biomass within the grazing exclosures with graminoid biomass being most responsive to the removal of herbivory whilst moss biomass remained unchanged. The changes in biomass switched the system from net emission to net uptake of CO₂ (0.47 and −0.77 μmol m⁻² s⁻¹ in grazed and exclosure plots, respectively) during the growing season and doubled the C storage in live biomass. In contrast, the treatment had no impact on the CH₄ fluxes, the total litter C pool or the soil C concentration. The rapid recovery of the above ground biomass and CO₂ fluxes demonstrates the plasticity of this high arctic ecosystem in terms of response to changing herbivore pressure.     

Litter production, leaf litter decomposition and nutrient return in Cunninghamia lanceolata plantations in south China: effect of planting conifers with broadleaved species

This study compared litter production, litter decomposition and nutrient return in pure and mixed species plantations. Dry weight and N, P, K, Ca, Mg quantities in the litterfall were measured in one pure Cunninghamia lanceolata plantation (PC) and two mixed-species plantations of C. lanceolata with Alnus cremastogyne (MCA) and Kalopanax septemlobus (MCK) in subtropical China. Covering 6 years of observations, mean annual litter production of MCA (4.97 Mg·ha⁻¹) and MCK (3.97 Mg·ha⁻¹) was significantly higher than that of PC (3.46 Mg·ha⁻¹). Broadleaved trees contributed 42% of the total litter production in MCA and 31% in MCK. Introduction of broadleaved tree species had no significant effect on litterfall pattern. Total litterfall was greatest in the dry season from November to March. Nutrient returns to the forest floor through leaf litter were significantly higher in both MCA and MCK than in PC (P < 0.05). The amounts of N, K, and Mg returned to the forest floor through leaf litter were highest in the MCA, and P and Ca returns were highest in the MCK. Percent contribution of broadleaf litter to total nutrient returns ranged from 41.7% to 86.9% in MCA and from 49.3% to 74.8% in MCK. The decomposition rate of individual leaf litter increased in the order: C. lanceolata < K. septemlobus < A. cremastogyne. Litter mixing had a positive effect on decomposition rate of the more recalcitrant litter and promoted nutrient return. Relative to mass loss of A. cremastogyne decomposing alone, higher mass loss of the mixture of C. lanceolata and A. cremastogyne was observed after 330 days of decomposition. These results indicate that mixed plantations of different tree species have advantages over monospecific plantations with regards to nutrient fluxes and these advantages have relevance to restoration of degraded sites. Responsible Editor: Alfonso Escudero.     

Herbivore responses to nutrient enrichment and landscape heterogeneity in a mangrove ecosystem

Complex gradients in forest structure across the landscape of offshore mangrove islands in Belize are associated with nutrient deficiency and flooding. While nutrient availability can affect many ecological processes, here we investigate how N and P enrichment interact with forest structure in three distinct zones (fringe, transition, dwarf) to alter patterns of herbivory as a function of folivory, loss of yield, and tissue mining. The effects of nutrient addition and zone varied by functional feeding group or specific herbivore. Folivory ranged from 0 to 0.4% leaf area damaged per month, but rates did not vary by either nutrient enrichment or zone. Leaf lifetime damage ranged from 3 to 10% of the total leaf area and was caused primarily by the omnivorous tree crab Aratus pisonii. We detected two distinct spatial scales of response by A. pisonii that were unrelated to nutrient treatment, i.e., most feeding damage occurred in the fringe zone and crabs fed primarily on the oldest leaves in the canopy. Loss of yield caused by the bud moth Ecdytolopha sp. varied by zone but not by nutrient treatment. A periderm-mining Marmara sp. responded positively to nutrient enrichment and closely mirrored the growth response by Rhizophora mangle across the tree height gradient. In contrast, a leaf-mining Marmara sp. was controlled by parasitoids and predators that killed >89% of its larvae. Thus, nutrient availability altered patterns of herbivory of some but not all mangrove herbivores. These findings support the hypothesis that landscape heterogeneity of the biotic and abiotic environment has species-specific effects on community structure and trophic interactions. Predicting how herbivores respond to nutrient over-enrichment in mangrove ecosystems also requires an assessment of habitat heterogeneity coupled with feeding strategies and species-specific behavior measured on multiple scales of response.     

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.     

The interaction of plant genotype and herbivory decelerate leaf litter decomposition and alter nutrient dynamics

We examined how plant genetic variation and a common herbivore (the leaf-galling aphid, Pemphigus betae ) influenced leaf litter quality, decomposition, and nutrient dynamics in a dominant riparian tree ( Populus spp .). Based on both observational studies and a herbivore exclusion experiment using trees of known genotype, we found four major patterns: 1) the quality of galled vs non-galled or gall-excluded litter significantly differed in the concentration of condensed tannins, lignin, nitrogen and phosphorus; 2) the difference in litter quality resulted in galled litter decomposing at rates 34 to 40% slower than non-galled litter; 3) plant genotype and herbivory had similar effects on the magnitude of decomposition rate constants; and 4) plant genotype mediated the herbivore effects on leaf litter quality and decomposition, as there were genotype-specific responses to herbivory independent of herbivore density. In contrast to other studies that have demonstrated accelerated ecosystem properties in response to arthropod herbivory, our findings argue that herbivore-induced secondary compounds decelerated ecosystem properties though their "after-life" effects on litter quality. Furthermore, these data are among the first to suggest that genotype-specific responses to herbivores can have a major impact on decomposition and nutrient flux, which likely has important consequences for the spatial distribution of nutrients at the landscape level. Due to the magnitude of these effects, we contend that it is important to incorporate a genetic perspective into ecosystem studies.     

Invasion of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Enhanced Ecosystem Carbon and Nitrogen Stocks in the Yangtze Estuary,China

Whether plant invasion increases ecosystem carbon (C) stocks is controversial largely due to the lack of knowledge about differences in ecophysiological properties between invasive and native species. We conducted a field experiment in which we measured ecophysiological properties to explore the response of the ecosystem C stocks to the invasion of Spartina alterniflora (Spartina) in wetlands dominated by native Scirpus mariqueter (Scirpus) and Phragmites australis (Phragmites) in the Yangtze Estuary, China. We measured growing season length, leaf area index (LAI), net photosynthetic rate (Pn), root biomass, net primary production (NPP), litter quality and litter decomposition, plant and soil C and nitrogen (N) stocks in ecosystems dominated by the three species. Our results showed that Spartina had a longer growing season, higher LAI, higher Pn, and greater root biomass than Scirpus and Phragmites. Net primary production (NPP) was 2.16 kg C m⁻² y⁻¹ in Spartina ecosystems, which was, on average, 1.44 and 0.47 kg C m⁻² y⁻¹ greater than that in Scirpus and Phragmites ecosystems, respectively. The litter decomposition rate, particularly the belowground decomposition rate, was lower for Spartina than Scirpus and Phragmites due to the lower litter quality of Spartina. The ecosystem C stock (20.94 kg m⁻²) for Spartina was greater than that for Scirpus (17.07 kg m⁻²), Phragmites (19.51 kg m⁻²) and the mudflats (15.12 kg m⁻²). Additionally, Spartina ecosystems had a significantly greater N stock (698.8 g m⁻²) than Scirpus (597.1 g m⁻²), Phragmites ecosystems (578.2 g m⁻²) and the mudflats (375.1 g m⁻²). Our results suggest that Spartina invasion altered ecophysiological processes, resulted in changes in NPP and litter decomposition, and ultimately led to enhanced ecosystem C and N stocks in the invaded ecosystems in comparison to the ecosystems with native species.     

Insect herbivory, organic matter deposition and effects on belowground organic matter fluxes in a central European oak forest

Apart from the forest floor, the canopy of forested ecosystems functions as the second most important source for dissolved and particulate fractions of organic and inorganic C and N compounds. However, under mass outbreak situations of insect herbivores this flux path of organic matter is considerably intensified clearly exceeding C and N fluxes from the forest floor. In this paper we report on herbivore-altered C and N fluxes from the canopy to the forest floor and effects on forest floor nutrient fluxes during severe defoliating herbivory of the winter moth (Operophtera brumata) and the mottled umber moth (Eranis defoliaria) in an oak forest in Germany. Over the course of 6.5 months we followed the C and N fluxes with bulk deposition, throughfall solution, insect frass deposits (green-fall together with insect faeces) and with forest floor solution in an 117-yr-old oak (Quercus petraea) forest. Compared to the control, herbivore defoliation significantly enhanced throughfall inputs of total and dissolved organic carbon and nitrogen by a factor of 3 and 2.5 (for TOC and DOC), and by 1.4 and 1.3 times (for TNb and DNb), respectively. Frass plus green-fall C and N fluxes peaked in May with 592 kg C?ha^?1 and 33.5 kg N?ha^?1 representing 79.6% (for C) and 78.3% (for N) of the total C and N input over 2.5 months. The quantitative and qualitative C and N input via faeces and litter deposition significantly differ between the insect affected and non-affected site. However, the C and N fluxes with throughfall did not significantly correlate with forest floor leachates. In this context, forest floor fluxes of TOC, DOC and NO₃-N were significantly lower at the infested site compared to the control, whereas fluxes of NH₄-N together with DON were significantly higher. The study demonstrates the importance of linking the population and associated frass dynamics of herbivorous insects with the cycling of nutrients and organic matter in forest ecosystems, highlighting the remarkable alterations in the timing, amounts and nature of organic matter dynamics on the ecosystem level. Consequently, the ecology of phytophagous insects allows partly to explain temporal-spatial alterations in nutrient cycling and thus ecosystem functioning.     

Productivity and mineral cycling in an oak coppice forest 2. Annual net production of the forest

Annual net production was estimated in the secondary coppice forest near Tokyo, which was dominated by a deciduous oak,Quercus serrata Thunb. Lateral growth of stems and old branches was directly estimated by examining the annual rings for 35 shoots in a clear-cut quadrat of 10m×10m. Phytomasses of current organs were also weighed in the quadrat. Preharvest losses of current organs were determined by twelve 0.5 m² litter traps for fine litter and twelve 6 m² quadrats for woody litter. Branch production was also assessed indirectly by use of the stem-branch allometry and death of branches. The results of the indirect method were in sufficient agreement with the result of the direct one. Grazing loss of leaves from the canopy was estimated directly from the loss in leaf area and indirectly from the animal faeces caught by the litter traps. Net production of the canopy trees was 149 kg a⁻¹ year⁻¹, in which leaf production was 36.9 kg. Animals grazed about 14% of the leaf area by the end of the growing season. True consumption of leaves by animals was 7.6% of leaf production or 10% of leaf mass. Production of undergrowth, mainly a dwarf bamboo,Pleioblastus chino Makino, was 28 kg a⁻¹ year⁻¹, being 15% of the total stand production. Productivity of this forest was significantly higher than that of cool-temperate deciduous broadleaf forests.     

Rapid shifts in the chemical composition of aspen forests: an introduced herbivore as an agent of natural selection

The global ecological impacts of introduced and exotic species can be dramatic, leading to losses in biodiversity and ecosystem “meltdown”, however, the evolutionary impacts of introduced species are much less understood. Further, very few studies have examined whether mammalian herbivores can act as agents of natural selection for plant traits. We examined the hypothesis that variation in aspen phytochemistry resulted in selective herbivory by Cervus elaphus (elk), an introduced mammalian herbivore. With the experimental removal of a large elk exclosure, elk selectively eliminated 60% of an aspen population previously protected from herbivory resulting in a dramatic shift in the phytochemical composition of the aspen forest. Selection gradients (β) varied from 0.52 to 0.66, well above average relative to other studies of selection. These results indicate that introduced herbivores can have rapid evolutionary consequences even on long lived native species. Because there are fundamental links between phytochemistry, biodiversity and ecosystem processes, the effects of an introduced herbivore are likely to have cascading impacts on the services ecosystems provide.     

Variations of soil enzyme activities in a temperate forest soil

Soil enzyme activities (dehydrogenase, urease, phosphatase, and arylsulfatase) in a temperate forest soil were determined in relation to landscape position and seasons. Overstory of the area is dominated by Quercus mongolica, Kalopanax pictus, Carpicus cordata, and Acer pseudo-sieboldianum. The activities were measured in three patches, namely a north-facing backslope, a ridge, and a south-facing backslope in autumn and spring over 2 years. In addition, spatially more detailed analysis for phosphatase was conducted before and after litterfall production in six patches. Dehydrogenase, urease, phosphatase, and arylsulfatase activities varied 1.8–18.5 μg INT-formazan g⁻¹ h⁻¹, 45.4–347.0 μg NH₄ ⁺ g⁻¹ h⁻¹, 0.9–4.5 mmol pNP g⁻¹ h⁻¹, and 0.7–2.6 mmol pNP g⁻¹ h⁻¹, respectively. In general, higher enzyme activities were found in the northern aspect than in the southern aspect. This variation appears to be related to differences in chemical properties (e.g., Fe, Al, and Mg) of soil as well as distribution of leaf litter. Two patterns were discernible in relation to seasonal variations. Dehydrogenase and urease exhibited a positive correlation with mean air temperature, suggesting that temperature would be a major controlling variable for those enzymes. In contrast, higher activities were detected in autumn for phosphatase and arylsulfatase activities, which appeared to be closely related to litter production and distribution. Overall results of this study indicate that soil enzyme activities in a forest floor are influenced by several variables such as temperature, nutrient availability, and input of leaf litter, which are closely related to landscape position.     

Estimates of biomass and primary productivity in a high-altitude maple forest of the west central Himalayas

The paper describes the biomass and productivity of maple (Acer cappadocicum) forest occurring at an altitude of 2,750 m in the west central Himalayas. Total vegetation biomass was 308.3 t ha⁻¹, of which the tree layer contributed the most, followed by herbs and shrubs. The seasonal forest-floor litter mass varied between 5.4 t ha⁻¹ (in rainy season) and 6.6 t ha⁻¹ (in winter season). The annual litter fall was 6.2 t ha⁻¹, of which leaf litter contributed the largest part (59% of the total litter fall). Net primary productivity of total vegetation was 19.5 t ha⁻¹ year⁻¹. The production efficiency of leaves (net primary productivity/leaf mass) was markedly higher (2.9 g g⁻¹ foliage mass year⁻¹) than those of the low-altitude forests of the region.     

Influence of stand age on nutrient and energy release through decomposition in alder-cardamom agroforestry systems of the eastern Himalayas

The influence of stand age (5, 10, 15, 20, 30 and 40 years) on the decomposition of litter fractions, nutrient and energy release of mixtures of N₂-fixing alder (Alnus nepalensis) and non-N₂-fixing large cardamom (Amomum subulatum) systems was compared. Seasonal decomposition rates were distinct with the highest rate in the first 6 months followed by subsequent seasons. The decomposition rate was substantially high in younger stands (10- to 15-years) and declined in the older stands. Heat sink from the stand floor litter increased from 171 × 10⁶ kJ year⁻¹ in 5 years to 299 × 10⁶ kJ year⁻¹ at 15 years and then considerably decreased with advancing age. However, energy and nutrient releases were slow at a high initial lignin-to-initial N ratio and C-to-N ratio, and there was an inverse relationship between the k-value of ash-free-mass and N expressed as a function of the C-to-N ratio. Quantities of nutrient release and energy loss per unit area in 24 months of decomposition were highest in 15 years and subsequently they lowered with advancing age. Nutrient loss indicated approximately uniform absolute and relative rates. Absolute energy consistently decreased by 81–88% in 24 months. Ash-free mass of decomposing litter remaining at different retrieval dates was associated with a narrowing of the C-to-N ratio. The relative loss rate of ash-free mass, nutrients and energy content was strongly related to the C-to-N ratio, litter temperature and litter moisture. The influence of Alnus in the younger stands on nutrient and energy releases were rapid, indicating accelerated nutrient cycling and energy dynamics. The intensity of the processes was highly phenomenal and considerably high in younger stands up to 20 years. Thus, an appropriate management cycle of the Alnus-cardamom system for sustainability is 15–20 years.     

Tree Species Composition and Harvest Intensity Affect Herbivore Density and Leaf Damage on Beech,Fagus sylvatica,in Different Landscape Contexts

Most forests are exposed to anthropogenic management activities that affect tree species composition and natural ecosystem processes. Changes in ecosystem processes such as herbivory depend on management intensity, and on regional environmental conditions and species pools. Whereas influences of specific forest management measures have already been addressed for different herbivore taxa on a local scale, studies considering effects of different aspects of forest management across different regions are rare. We assessed the influence of tree species composition and intensity of harvesting activities on arthropod herbivores and herbivore-related damage to beech trees, Fagus sylvatica, in 48 forest plots in three regions of Germany. We found that herbivore abundance and damage to beech trees differed between regions and that – despite the regional differences - density of tree-associated arthropod taxa and herbivore damage were consistently affected by tree species composition and harvest intensity. Specifically, overall herbivore damage to beech trees increased with increasing dominance of beech trees – suggesting the action of associational resistance processes – and decreased with harvest intensity. The density of leaf chewers and mines was positively related to leaf damage, and several arthropod groups responded to beech dominance and harvest intensity. The distribution of damage patterns was consistent with a vertical shift of herbivores to higher crown layers during the season and with higher beech dominance. By linking quantitative data on arthropod herbivore abundance and herbivory with tree species composition and harvesting activity in a wide variety of beech forests, our study helps to better understand the influence of forest management on interactions between a naturally dominant deciduous forest tree and arthropod herbivores.     

The impact of elevated CO2 on plant-herbivore interactions: experimental evidence of moderating effects at the community level

Surprisingly little research has been published on the responses to elevated [CO₂] at the community level, where herbivores can select their preferred food. We investigated the combined effects of atmospheric [CO₂] and herbivory on synthesised plant communities growing on soils of different fertility. Factorial combinations of two [CO₂] (350 or 700 l l⁻¹), two fertility (fertilised or non-fertilised), and two herbivory (herbivores present or absent) treatments were applied to a standard mixture of seven fast- and eight slow-growing plants in outdoor microcosms. The herbivores used were the grain aphid (Sitobion avenae) and the garden snail (Helix aspersa). We measured plant biomass, foliar nitrogen and soluble tannin concentration, aphid fecundity, and snail growth, fecundity, and feeding preferences over one growing season. Elevated [CO₂] did not have a significant impact on (1) the combined biomass of fast-growing or slow-growing plants, (2) herbivore feeding preferences, or (3) herbivore fitness. There was, however, a significant biomass increase of Carex flacca (which represented in all cases less than 5% of total live biomass), and some chemical changes in unpalatable plants under elevated [CO₂]. The herbivory treatment significantly increased the biomass of slow-growing plants over fast-growing plants, whereas fertilisation significantly increased the abundance of fast-growing plants over slow-growing plants. Predictions on the effects of elevated [CO₂] based on published single-species experiments were not supported by the results of this microcosm study. Received: 30 November 1997 / Accepted: 24 July 1998