Exploitative competition between invasive herbivores benefits a native host plant

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).     

Modeling range dynamics in heterogeneous landscapes: invasion of the hemlock woolly adelgid in eastern North America

Range expansion by native and exotic species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. Still unanswered, however, is the question of how well this new generation of dynamic models will anticipate range expansion. Here we develop a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in climate and habitat to model range expansion of the hemlock woolly adelgid (HWA; Adelges tsugae). We parameterize this model using multiyear data sets describing population and dispersal dynamics of HWA and apply it to eastern North America over a 57-year period (1951-2008). To evaluate the model, the observed pattern of spread of HWA during this same period was compared to model predictions. Our model predicts considerable heterogeneity in the risk of HWA invasion across space and through time, and it suggests that spatiotemporal variation in winter temperature, rather than hemlock abundance, exerts a primary control on the spread of HWA. Although the simulations generally matched the observed current extent of the invasion of HWA and patterns of anisotropic spread, it did not correctly predict when HWA was observed to arrive in different geographic regions. We attribute differences between the modeled and observed dynamics to an inability to capture the timing and direction of long-distance dispersal events that substantially affected the ensuing pattern of spread.     

Nitrogen dynamics in an Australian semiarid grassland soil

We conducted a four-week laboratory incubation of soil from a Themeda triandra Forsskal grassland to clarify mechanisms of nitrogen (N) cycling processes in relation to carbon (C) and N availability in a hot, semiarid environment. Variation in soil C and N availability was achieved by collecting soil from either under tussocks or the bare soil between tussocks, and by amending soil with Themeda litter. We measured N cycling by monitoring: dissolved organic nitrogen (DON), ammonium (NH4+), and nitrate (NO3-) contents, gross rates of N mineralization and microbial re-mineralization, NH4+ and NO3- immobilization, and autotrophic and heterotrophic nitrification. We monitored C availability by measuring cumulative soil respiration and dissolved organic C (DOC). Litter-amended soil had cumulative respiration that was eightfold greater than non-amended soil (2000 compared with 250 microg C/g soil) and almost twice the DOC content (54 compared with 28 microg C/g soil). However, litter-amended soils had only half as much DON accumulation as non-amended soils (9 compared with 17 microg N/g soil) and lower gross N rates (1-4 compared with 13-26 microg N x [g soil](-1) x d(-1)) and NO3- accumulation (0.5 compared with 22 microg N/g soil). Unamended soil from under tussocks had almost twice the soil respiration as soil from between tussocks (300 compared with 175 microg C/g soil), and greater DOC content (33 compared with 24 microg C/g soil). However, unamended soil from under tussocks had lower gross N rates (3-20 compared with 17-31 microg N x [g soil](-1) d(-1)) and NO3- accumulation (18 compared with 25 microg N/g soil) relative to soil from between tussocks. We conclude that N cycling in this grassland is mediated by both C and N limitations that arise from the patchiness of tussocks and seasonal variability in Themeda litterfall. Heterotrophic nitrification rate explained >50% of total nitrification, but this percentage was not affected by proximity to tussocks or litter amendment. A conceptual model that considers DON as central to N cycling processes provided a useful initial framework to explain results of our study. However, to fully explain N cycling in this semiarid grassland soil, the production of NO3- from organic N sources must be included in this model.     

鼎湖山森林演替序列水文过程中总有机碳(TOC)变化规律及其对土壤碳平衡的贡献

森林水文过程中的总有机碳转运对土壤有机碳平衡起着重要的作用,但我们对于水文过程对碳平衡的贡献机理所知甚少.本研究针对鼎湖山季风常绿阔叶林演替序列不同森林生态系统(马尾松林、针阔混交林和季风常绿阔叶林(简称季风林))的大气降水、穿透水、树干流、凋落物淋洗水以及地表径流中的总有机碳(TOC)进行了三年(2002年4月-2005年5月)观测,以此来分析水文学过程中TOC的变化规律和水文学过程对不同成熟度森林生态系统土壤有机碳积累的贡献.每场雨后进行水样的采集,采集的水样装入棕色玻璃瓶中,加硫酸至pH值小于2,放置于实验室冰箱冷藏待测.TOC用日本岛津公司生产的5000A型TOC-V分析仪测定.研究结果及推论如下:鼎湖山森林水文学过程中TOC浓度和总量变化呈现规律性的变化.大气降水中的TOC浓度和总量分别为(3.65±0.59)mg·L~(-1)和51.8104 kg·hm~(-2)·a~(-1),大气降水是鼎湖山森林生态系统水文循环过程中TOC的主要来源.穿透水(DTF)中TOC浓度和总量均为:松林>混交林>季风林,其中季风林TOC浓度显著低于其他两种林型.松林树干流的TOC浓度显著高于混交林和季风林.凋落物淋洗水TOC浓度和总量大小依次均为:松林>混交林>季风林,且三林型间存在显著差异(p<0.05).径流中TOC浓度和总量均较小,且无明显差异.在湿季5月份,穿透水、树干流、凋落物淋洗水的TOC浓度呈现下降趋势.干季(10月)开始以后,穿透水、树干流、凋落物淋洗水中的TOC浓度又逐步回升.地表径流中TOC浓度干湿季变化趋势不明显.干季中各水文学分量TOC浓度大于湿季,但TOC总量呈现相反趋势.在森林水文学过程中,凋落物淋洗水所携带的有机碳量是土壤有机碳输入的最大项,季风林、混交林、松林中TOC总量分别为246.983 kg·hm~(-2)·a~(-1),255.187kg·hm~(-2)·a~(-1)和261.876kg·hm~(-2)·a~(-1);其次是直接到达土壤表面的穿透水,季风林、混交林、松林中TOC总量分别为28.152kg·hm~(-2)·a~(-1),37.410kg·hm~(-2)·a~(-1)和43.176kg·hm~(-2)·a~(-1);树干流中有机碳浓度虽高,但总量很微小,季风林、混交林、松林中TOC总量分别为4.663kg·hm~(-2)·a~(-1),5.910kg·hm~(-2)·a~(-1)和4.566kg·hm~(-2)·a~(-1),所以对土壤有机碳收入贡献不大.径流所携带的TOC总量很小,季风林、混交林、松林中分别为8.707kg·hm~(-2)·a~(-1),9.318kg·hm~(-2)·a~(-1),7.220kg·hm~(-2)·a~(-1).由此可知,水文过程输入土壤的TOC总量远大于径流所带走的TOC总量,导致了水文过程中的TOC存留在土壤中,对土壤有机碳(SOC)的积累起着重要作用.季风林、混交林和马尾松林土壤每年通过水文学过程净输入的有机碳量分别为(27.1+1.65)g·m~(-2),(28.9±2.79)g·m~(-2)和(30.2±2.65)g·m~(-2).水文学过程中的这部分有机碳由于占总有机碳比例较小往往被忽视,但是正是由于水分在土壤中的下渗使得有机碳的分布趋于均匀,这将更加利于SOC的积累和保存.     

Examining the coupling of carbon and nitrogen cycles in Appalachian streams: the role of dissolved organic nitrogen

Although regional and global models of nitrogen (N) cycling typically focus on nitrate, dissolved organic nitrogen (DON) is the dominant form of nitrogen export from many watersheds and thus the dominant form of dissolved N in many streams. Our understanding of the processes controlling DON export from temperate forests is poor. In pristine systems, where biological N limitation is common, N contained in recalcitrant organic matter (OM) can dominate watershed N losses. This recalcitrant OM often has moderately constrained carbon:nitrogen (C:N) molar ratios (approximately 25-55) and therefore, greater DON losses should be observed in sites where there is greater total dissolved organic carbon (DOC) loss. In regions where anthropogenic N pollution is high, it has been suggested that increased inorganic N availability can reduce biological demand for organic N and therefore increase watershed DON losses. This would result in a positive correlation between inorganic and organic N concentrations across sites with varying N availability. In four repeated synoptic surveys of stream water chemistry from forested watersheds along an N loading gradient in the southern Appalachians, we found surprisingly little correlation between DON and DOC concentrations. Further, we found that DON concentrations were always significantly correlated with watershed N loading and stream water [NO3-] but that the direction of this relationship was negative in three of the four surveys. The C:N molar ratio of dissolved organic matter (DOM) in streams draining watersheds with high N deposition was very high relative to other freshwaters. This finding, together with results from bioavailability assays in which we directly manipulated C and N availabilities, suggests that heterotrophic demand for labile C can increase as a result of dissolved inorganic N (DIN) loading, and that heterotrophs can preferentially remove N-rich molecules from DOM. These results are inconsistent with the two prevailing hypotheses that dominate interpretations of watershed DON loss. Therefore, we propose a new hypothesis, the indirect carbon control hypothesis, which recognizes that heterotrophic demand for N-rich DOM can keep stream water DON concentrations low when N is not limiting and heterotrophic demand for labile C is high.     

Constitutive and herbivore‐induced volatiles in pear,alder and hawthorn trees

Summary. Qualitative and quantitative differences among pear cultivars were found in constitutive and Cacopsylla-induced volatiles, depending on experimental treatment of the trees (i.e., uninfested and partly or completely infested by psyllids). Blend differences were also found between pear cultivars and wild-type pear, alder and hawthorn–the latter trees are frequently present in pear orchard hedgerows. ?Interesting differences were found in the presence of methyl salicylate and (E,E)-α-farnesene, two compounds previously found to mediate attraction of predatory bugs towards psyllid-infested pear trees. Methyl salicylate is expressed constitutively and is induced systemically by infestation in the whole plant of all four cultivars. (E,E)-α-farnesene on the other hand showed also systemic induction in Bartlett, NY10355 and Beurré Hardy, but in partially infested Conference trees it was induced locally, only in herbivore-damaged leaves. No methyl salicylate or (E,E)-α-farnesene were identified in honeydew. In field collected headspace samples of alder leaves infested by aphids and leaf beetles we found methyl salicylate but no (E,E)-α-farnesene, whereas in uninfested hawthorn neither were identified. Insight in the variability of damage-related pear volatiles will have important implications for integrated pest management in the field. Received 27 August 2002; accepted 28 November 2002 R1D=" Correspondence to: Petru Scutareanu, e-mail:scuterea@science.uva.nl     

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.     

Leaf volatiles and polyphenols in pear trees infested byPsylla pyricola. Evidence of simultaneously induced responses

Summary Feeding by the homopteranPsylla pyricola on leaves of pear trees induces the production of volatile compounds, such as (E,E)--farnesene and methyl-salicylate, as well as the production of polyphenols. The inference on induction is based on GC-MS and HPLC chromatograms from the same samples ofPsylla infested leaves, leaves from the same pear tree beforePsylla infestation and uninfested leaves from other pear trees.Psylla infestation greatly enhanced the production of volatiles ((E,E)--farnesene, methyl-salicylate and others) and triggered the production of new polyphenols, characterized by much longer retention times.However, the responses to infestation depend critically on leaf age (defined by leaf distance to apex). With respect to the leaf volatiles it appears that infested, old leaves produce fewer compounds and lower amounts of the volatiles than infested, young leaves. Moreover, there seem to be differences in pattern. Relative to (E,E)--farnesene, methyl-salicylate was found in much lower amounts in heavily infested, old leaves. With respect to polyphenols it was found that infested old leaves collected in August have polyphenols with the same retention times, but more or less equal amounts as uninfested young leaves collected in May. This shows thatPsylla infestation causes the induced response mostly in young leaves.The induced leaf volatiles may act as synomones to heteropteran bugs. As shown elsewhere,Anthocoris nemoralis responds significantly to (E,E)--farnesene and methyl-salicylate when offered in pure form against clean air in a Y-tube olfactometer. The effect of polyphenols on the performance ofP. pyricola is not yet known. Hence, a role in direct defence is still to be investigated.     

Mercury cycling in litter and soil in different forest types in the Adirondack region, New York, USA.

The fate of mercury in decomposing leaf litter and soil is key to understanding the biogeochemistry of mercury in forested ecosystems. We quantified mercury dynamics in decomposing leaf litter and measured fluxes and pools of mercury in litterfall, throughfall, and soil in two forest types of the Adirondack region, New York, USA. The mean content of total mercury in leaf litter increased to 134% of its original mass during two years of decomposition. The accumulation pattern was seasonal, with significant increases in mercury mass during the growing season (+4.9% per month). Litterfall dominated mercury fluxes into the soil in the deciduous forest, whereas throughfall dominated fluxes into the coniferous forest. The increase in mercury mass in decomposing deciduous litter during the growing season was greater than could be accounted for by throughfall inputs during the growing season (P < 0.05), suggesting translocation of mercury from the soil to the decomposing deciduous litter. This internal recycling mechanism concentrates mercury in the organic horizons and retards transport through the soil, thereby increasing the residence time of mercury in the forest floor. A mass balance assessment suggests that the ultimate fate of mercury in the landscape depends upon forest type and associated differences in the delivery and incorporation of mercury into the soil. Our results show that incorporation of mercury into decaying leaf litter increases its residence time in the landscape and may further delay the recovery of surface waters, fish, and associated biota following control of mercury emissions to the atmosphere.     

The Chemistry of Throughfall, Stemflow and Soil Water Beneath Oak Woodland and Moorland Vegetation in Upland Wales

The chemistry of bulk precipitation, throughfall, stemflow and soil waters beneath an oak wood (Quercus petraea) canopy and soil waters under moorland vegetation were measured at two sites on acid brown podzolic soils near Llyn Brianne in rural mid-Wales, UK. Between March 1986 and November 1988, precipitation was 4354 mm and annual interception losses from the oak canopy averaged 13% of incident precipitation. Throughfall and stemflow were more acid and concentrations of most solutes were increased 2- to 4-fold compared with bulk precipitation. Nitrate was the only solute retained within the tree canopy. Throughfall collected beneath patches of bracken on the forest floor was less acidic but contained substantially higher concentrations of major ions than bulk precipitation and oak throughfall. the moorland soil was more acidic, contained more exchangeable calcium but less exchangeable aluminium and potassium than the woodland soil. Soil waters beneath both vegetation types were acidic (mean pH range 4.5-4.9) and dominated by sodium and chloride. with the exception of calcium, soil water solute concentrations were greater beneath oak. These differences are ascribed to larger atmospheric inputs beneath the oak canopy compared with the shorter grasses, combined with the effect of differences in nutrient dynamics and water fluxes. Variations in soil water aluminium chemistry are explained in terms of ion exchange and podzolisa-tion processes. the water quality implications of increased upland afforestation of moorland by broadleaved trees are discussed.     

The Chemistry of Throughfall,Stemflow and Soil Water Beneath Oak Woodland and Moorland Vegetation in Upland Wales

The chemistry of bulk precipitation, throughfall, stemflow and soil waters beneath an oak wood (Quercus petraea) canopy and soil waters under moorland vegetation were measured at two sites on acid brown podzolic soils near Llyn Brianne in rural mid-Wales, UK. Between March 1986 and November 1988, precipitation was 4354 mm and annual interception losses from the oak canopy averaged 13% of incident precipitation. Throughfall and stemflow were more acid and concentrations of most solutes were increased 2- to 4-fold compared with bulk precipitation. Nitrate was the only solute retained within the tree canopy. Throughfall collected beneath patches of bracken on the forest floor was less acidic but contained substantially higher concentrations of major ions than bulk precipitation and oak throughfall. the moorland soil was more acidic, contained more exchangeable calcium but less exchangeable aluminium and potassium than the woodland soil. Soil waters beneath both vegetation types were acidic (mean pH range 4.5-4.9) and dominated by sodium and chloride. with the exception of calcium, soil water solute concentrations were greater beneath oak. These differences are ascribed to larger atmospheric inputs beneath the oak canopy compared with the shorter grasses, combined with the effect of differences in nutrient dynamics and water fluxes. Variations in soil water aluminium chemistry are explained in terms of ion exchange and podzolisa-tion processes. the water quality implications of increased upland afforestation of moorland by broadleaved trees are discussed.     

Tsuga canadensis (L.) Carr. mortality will impact hydrologic processes in southern Appalachian forest ecosystems.

Eastern hemlock (Tsuga canadensis (L.) Carr.) is one of the principal riparian and cove canopy species in the southern Appalachian Mountains. Throughout its range, eastern hemlock is facing potential widespread mortality from the hemlock woolly adelgid (HWA). If HWA-induced eastern hemlock mortality alters hydrologic function, land managers will be challenged to develop management strategies that restore function or mitigate impacts. To estimate the impact that the loss of this forest species will have on the hydrologic budget, we quantified and modeled transpiration over a range of tree sizes and environmental conditions. We used heat dissipation probes, leaf-level gas-exchange measurements, allometric scaling, and time series modeling techniques to quantify whole-tree and leaf-level transpiration (E(L)) of eastern hemlock. We monitored trees ranging from 9.5 to 67.5 cm in diameter along a riparian corridor in western North Carolina, USA during 2004 and 2005. Maximum rates of daily tree water use varied by diameter and height, with large trees transpiring a maximum of 178-186 kg H2O x tree(-1) x d(-1). Values of E(L) could be predicted from current and lagged environmental variables. We forecasted eastern hemlock E(L) for inventoried stands and estimated a mean annual transpiration rate of 63.3 mm/yr for the hemlock component, with 50% being transpired in the winter and spring. In typical southern Appalachian stands, eastern hemlock mortality would thus reduce annual stand-level transpiration by approximately 10% and reduce winter and spring stand-level transpiration by approximately 30%. Eastern hemlock in the southern Appalachians has two distinct ecohydrological roles: an evergreen tree that maintains year-round transpiration rates and a riparian tree that has high transpiration rates in the spring. No other native evergreen in the southern Appalachians will likely fill the ecohydrological role of eastern hemlock if widespread mortality occurs. With the loss of this species, we predict persistent increases in discharge, decreases in the diurnal amplitude of streamflow, and increases in the width of the variable source area.     

Population dynamics of beech scale (Cryptococcus fagisuga) (Coccina,Pseudococcidae) related to physiological defence reactions of attacked beech trees (Fagus sylvatica)

Summary Changes in the density of beech scale infestation (Cryptococcus fagisuga) of a mature beech stand (Fagus sylvatica) were investigated after gap-cutting in limed and unlimed areas bordering on the gaps and in untreated areas over a 5-year period. Parallelly the concentrations of sucrose, protein amino acids, and procyanidins were examined in the inner and outer bark of non-infested beech trees and beeches infested by beech scale.Irrespective of liming in the areas bordering on the gaps the proportion of beech trees with increasing beech scale infestation was significantly higher, the proportion of beeches with diminishing infestation was significantly lower than in the untreated control areas. Trees with the same infestation tendency occurred in clusters.Physiological defence reactions in the inner and outer bark of the infested beech trees were registrated as an increase in the procyanidin content and a decrease in the protein amino acid content. A change in the outer bark—inner bark—ratio of the infested beech trees showed a transfer of compounds between the inner and outer bark. During the period of observation long-lasting shifts occurred in the pattern of compounds of the bark irrespective of actual infestation intensity.     

Long-term burning interacts with herbivory to slow decomposition

Fires can generate spatial variation in trophic interactions such as insect herbivory. If trophic interactions mediated by fire influence nutrient cycling, they could feed back on the more immediate consequences of fire on nutrient dynamics. Here we consider herbivore-induced effects on oak litter quality and decomposition within a long-term manipulation of fire frequency in central Minnesota, USA. We focused on bur oak (Quercus macrocarpa) trees, which are common across the fire frequency gradient and are often heavily infested with either lace bugs (Corythuca arcuata) or aphids (Hoplochaithropsus quercicola). We used targeted exclusion to test for herbivore-specific effects on litter chemistry and subsequent decomposition rates. Lace bug exclusion led to lower lignin concentrations in litterfall and subsequently accelerated decomposition. In contrast, aphid exclusion had no effect on litterfall chemistry or on decomposition rate, despite heavy infestation levels. Effects of lace bug herbivory on litterfall chemistry and decomposition were similar in burned and unburned areas. However, lace bug herbivory was much more common in burned than in unburned areas, whereas aphid herbivory was more common in unburned areas. These results suggest that frequent fires promote oak-herbivore interactions that decelerate decomposition. This effect should amplify other influences of fire that slow nitrogen cycling.     

Deer herbivory alters forest response to canopy decline caused by an exotic insect pest.

Hemlock woolly adelgid (HWA; Adelges tsugae) infestations have resulted in the continuing decline of eastern hemlock (Tsuga canadensis) throughout much of the eastern United States. While the initial impacts of HWA infestations have been documented, our understanding of forest response to this disturbance remains incomplete. HWA infestation is not occurring in isolation but within a complex ecological context. The role of potentially important interacting factors, such as elevated levels of white-tailed deer herbivory, is poorly understood. Despite the potential for herbivory to alter forest successional trajectories following a canopy disturbance, little is known about herbivory-disturbance interactions, and herbivory is rarely considered in assessing forest response to a co-occurring disturbance. We used repeated censuses of deer exclosures and paired controls (400 paired plots) to quantify the impact of deer herbivory on tree seedling species abundance in 10 eastern hemlock ravines that span a gradient in HWA-induced canopy decline severity. Use of a maximum likelihood estimation framework and information theoretics allowed us to quantify the strength of evidence for alternative models developed to estimate the impacts of herbivory on tree seedling abundance as a function of varying herbivore density and canopy decline severity. The exclusion of deer herbivory had marked impacts on the abundance of the studied seedling species: Acer rubrum, Acer saccharum, Betula lenta, Nyssa sylvatica, Quercus montana, and Tsuga canadensis. For all six species, the relationship between seedling abundance and deer density was either exponential or saturating. Although the functional form of the response varied among seedling species, the inclusion of both deer density and canopy decline severity measures consistently resulted in models with substantially greater support. Canopy decline resulted in higher proportional herbivory impacts and altered the ranking of herbivory impacts by seedling species. Our results suggest that, by changing species' competitive abilities, white-tailed deer herbivory alters the trajectory of forest response to this exotic insect pest and has the potential to shift future overstory composition.     

Dynamics of nitrogen and dissolved organic carbon at the Hubbard brook experimental forest

The factors controlling spatial and temporal patterns in soil solution and streamwater chemistry are highly uncertain in northern hardwood forest ecosystems in the northeastern United States, where concentrations of reactive nitrogen (Nr) in streams have surprisingly declined over recent decades in the face of persistent high rates of atmospheric Nr deposition and aging forests. Reactive nitrogen includes inorganic species (e.g., ammonium [NH4+], nitrate [NO3-]) and some organic forms (e.g., amino acids) available to support the growth of plants and microbes. The objective of this study was to examine controls on the spatial and temporal patterns in the concentrations and fluxes of nitrogen (N) species and dissolved organic carbon (DOC) in a 12-year record of soil solutions and streamwater along an elevational gradient (540-800 m) of a forested watershed at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains of New Hampshire, USA. Dissolved organic N and DOC concentrations were elevated in the high-elevation spruce-fir-white birch (SFB) zone of the watershed, while NO3- was the dominant N species in the lower elevation hardwood portion of the watershed. Within the soil profile, N retention was centered in the mineral horizon, and significant amounts of N were retained between the lower mineral soil and the stream, supporting the idea that near- and in-stream processes are significant sinks for N at the HBEF. Temporal analysis suggested that hydrologic flow paths can override both abiotic and biotic retention mechanisms (i.e., during the non-growing season when most hydrologic export occurs, or during years with high rainfall), there appears to be direct flushing of N from the organic horizons into the stream via horizontal flow. Significant correlations between soil NO3- concentrations, nitrification rates and streamwater NO3- exports show the importance of biological production as a regulator of inorganic N export. The lack of internal production response (e.g., mineralization, nitrification) to a severe ice storm in 1998 reinforces the idea that plant uptake is the dominant regulator of export response to disturbance.     

Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna

Animals that bore into calcareous material can cause considerable damage to molluscan shells. In contrast, smaller microbial phototrophic endoliths have until recently been thought of as relatively benign. Phototrophic endoliths (primarily cyanobacteria) infest the shells of 50 to 80% of midshore populations of the mussel Perna perna (L.) in South Africa. This infestation causes clearly visible shell degradation, and we record here ecologically important lethal and sub-lethal effects (e.g. changes in growth and reproductive output) of the endoliths on their mussel hosts. Endolith infestation reduced the strength of shells significantly and also affected shell growth. In situ marking of shells, using the fluorochrome calcein, showed that infested and non-infested mussels increased in shell length at the same rate. However, the rate of increase in shell thickness (associated with shell repair) was significantly faster in infested than in uninfested individuals. This increase in the rate of shell thickening was not sufficient to compensate for rapid endolith-induced shell degradation and, around the site of adductor muscle attachment, infested shells were thinner than their uninfested counterparts. The shells of 18% of recently dead mussels had holes induced by endolith erosion. This effect was highly size dependent, and the proportion of mortality due to endoliths rose to almost 50% for the largest mussels. The re-routing of energy due to shell repair had important sub-lethal effects on the reproductive rates of mussels. During the reproductive period, mean dried flesh mass for large (>70 mm), non-infested P. perna was substantially higher than for infested individuals. This difference was almost entirely due to differences in gonad mass, which was approximately 100% higher for non-infested mussels. We conclude that, by attacking the shell, phototrophic endoliths reduce both the longevity and reproductive output of large mussels on the midshore. Received: 26 January 1999 / Accepted: 17 August 1999     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

Litter decomposition and nitrogen mineralization from an annual to a monthly model

The forest litter decomposition model (FLDM) described in this paper provides an important basis for assessing the impacts of forest management on seasonal stream water quality and export of dissolved organic carbon (DOC). By definition, models with annual time steps are unable to capture seasonal, within-year variation. In order to simulate seasonal variation in litter decomposition and DOC production and export, we have modified an existing annual FLDM to account for monthly dynamics of decomposition and residual mass in experimental litterbags placed in 21 different forests across Canada.The original annual FLDM was formulated with three main litter pools (fast, slow, and very slow decomposing litter) to address the fact that forest litter is naturally composed of a mixture of organic compounds that decompose at different rates. The annual FLDM was shown to provide better simulations than more complex models like CENTURY and SOMM.The revised monthly model retains the original structure of the annual FLDM, but separates litter decomposition from nitrogen (N) mineralization. In the model, monthly soil temperature, soil moisture, and mean January soil temperature are shown to be the most important controlling variables of within-year variation in decomposition. Use of the three variables in a process-based definition of litter decomposition is a significant departure from the empirical definition in the annual model. The revised model is shown to give similar calculations of residual mass and N concentration as the annual model (r² = 0.91, 0.78), despite producing very different timeseries of decomposition over six years. It is shown from a modelling perspective that (i) forest litter decomposition is independent of N mineralization, whereas N mineralization is dependent on litter decomposition, and (ii) mean January soil temperature defines litter decomposition in the summer because of winter-temperatures’ role in modifying forest-floor microorganism community composition and functioning in the following summer.