Litterfall and nutrient return in five tree species in a common garden experiment

Canopy litterfall is a significant pathway for return of nutrients and carbon (C) to the soil in forest ecosystems. Litterfall was studied in five even-aged stands of Norway spruce, Sitka spruce, Douglas-fir, European beech and common oak at three different locations in Denmark; two sandy sites, Ulborg and Lindet in Jutland, and one loamy site, Frederiksborg on Zealand. Litterfall was collected during three years from 1994 to 1996 in all five species and during six years from 1994 to 1999 in Norway spruce, Sitka spruce and European beech. The average total litterfall was in the range of 3200–3700 kg ha⁻¹ yr⁻¹ and did not differ significantly among tree species. There were no significant differences in total litterfall among sites during the short period, but during the longer period the richer site Frederiksborg had significantly higher total and foliar litterfall amounts compared to the more nutrient-poor sites Lindet and Ulborg. There were close relationships between foliar and total litterfall suggesting that foliar litterfall can be reliably estimated from total litterfall. Beech and oak bud scale litter was significantly related to foliar litterfall. The amount of branch and twig litter was significantly higher in oak than in other tree species. The average foliar litterfall was well related to the annual volume increment. The relationship differed markedly from previously reported relationships based on global litterfall data suggesting that such relationships are better evaluated at the regional level. Nutrient concentrations and fluxes in foliar litterfall were not significantly different among the five tree species. However, there was a significant effect of site on most nutrient concentrations of the three litterfall fractions, and foliar fluxes of P, Ca and Mn were all significantly highest at Frederiksborg and lowest at Ulborg. The similarity in litterfall inputs to the forest floor under these five tree species suggested that previous reports of large variability in forest floor accumulation should primarily be attributed to differences in litter decomposition.     

The influence of the forest canopy on nutrient cycling

Rates of key soil processes involved in recycling of nutrients in forests are governed by temperature and moisture conditions and by the chemical and physical nature of the litter. The forest canopy influences all of these factors and thus has a large influence on nutrient cycling. The increased availability of nutrients in soil in clearcuts illustrates how the canopy retains nutrients (especially N) on site, both by storing nutrients in foliage and through the steady input of available C in litter. The idea that faster decomposition is responsible for the flush of nitrate in clearcuts has not been supported by experimental evidence. Soil N availability increases in canopy gaps as small as 0.1 ha, so natural disturbances or partial harvesting practices that increase the complexity of the canopy by creating gaps will similarly increase the spatial variability in soil N cycling and availability within the forest. Canopy characteristics affect the amount and composition of leaf litter produced, which largely determines the amount of nutrients to be recycled and the resulting nutrient availability. Although effects of tree species on soil nutrient availability were thought to be brought about largely through differences in the decomposition rate of their foliar litter, recent studies indicate that the effect of tree species can be better predicted from the mass and nutrient content of litter produced, hence total nutrient return, than from litter decay rate. The greater canopy complexity in mixed species forests creates similar heterogeneity in nutritional characteristics of the forest floor. Site differences in slope position, parent material and soil texture lead to variation in species composition and productivity of forests, and thus in the nature and amount of litter produced. Through this positive feedback, the canopy accentuates inherent differences in site fertility.     

Contrasting net primary productivity and carbon distribution between neighboring stands of Quercus robur and Pinus sylvestris

Standing biomass, net primary production (NPP) and soil carbon (C) pools were studied in a 67-year-old pedunculate oak (Quercus robur L.) stand and a neighboring 74-year- old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Despite a 14% lower tree density and a lower tree height in the oak stand, standing biomass was slightly higher than in the pine stand (177 and 169 Mg ha(-1) in oaks and pines, respectively), indicating that individual oak trees contained more biomass than pine trees of similar diameter. Moreover, NPP in the oak stand was more than double that in the pine stand (17.7 and 8.1 Mg ha(-1) year(-1), respectively). Several observations indicated that soil organic matter accumulated at higher rates under pines than under oaks. We therefore hypothesized that the pines were exhibiting an age-related decline in productivity due to nutrient limitation. The poor decomposability of pine litter resulted in the observed accumulation of organic matter. The subsequent immobilization of nutrients in the organic matter, combined with the already nutrient-poor soil conditions, resulted in a decrease in total NPP over time, as well as in a substantial shift in the allocation of NPP toward fine roots. In the oak stand, litter is less recalcitrant to decay and soil acidity is less severe; hence, organic matter does not accumulate and nutrients are recycled. This probably explains why NPP was much higher in the oaks than in the pines and why only a small proportion of NPP was allocated to oak fine roots.     

Nutrients return from leaves and litterfall in a mediterranean cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis> L.) forest in southwestern Spain

The knowledge of the cycle of nutrients is fundamental for the correct comprehension of the tree–soil relationship and for an adequate forest management. In order to analyse the nutrients return from leaves and litterfall in a Mediterranean cork oak forest in southwestern Spain, 12 trees were randomly selected and litterfall collected for 2 years. Samples were taken monthly and separated in different fractions (leaves, twigs, catkins, acorns and miscellaneous), then leaves nutrients were analyzed. Simultaneously, we analyzed the nutrient content of living leaves from the same trees in each season during 1 year. The analyzed nutrients were C, N, P, K, Ca, Mg, Fe, Mn, S, Cu, Zn and Mo. Annual patterns of each nutrient in fallen leaves were characterized and compared with seasonal values of these nutrients in living leaves. Leaves fall has two annual maximum, first and most important in spring around April coinciding with renewal of foliar cover and second around October. Main concentration patterns of N, P and K are related with phenological patterns, in consequence minimum concentration in leaves fall were obtained in periods of growing and maximum litterfall. Concentrations of Ca, Fe and Mn increase with the age of the leaves and maximum concentrations were obtained before periods of maximum litterfall while concentrations of Cu, Mo and Mg stay stable. Seasonal analysis of nutrients in living leaves collected from the same trees in four different periods of the year allowed to corroborate the patterns of leaves fall and the probable osmotic function of K.     

章古台沙地不同树种年龄和树种的生态系统叶片和土壤养分研究

对辽宁省章古台地区的几种主要针叶树种的针叶养分及其林分表层(0-15cm)土壤养分进行研究。结果表明,不同年龄樟子松(45、29和20年)(Pinussylvestrisvar.mongolica)林地表层土壤养分并无显著差别,而29年樟子松林分表层土壤全N、全P和N:P比显著高于相近年龄的赤松(P.densiflora)(29年)和油松林(P.tabulaeformis)(31年)。随着樟子松年龄的增加,叶片P含量逐渐减少,而叶片N和K含量则未表现出随年龄而变化的趋势;对于年龄相近的三个树种,叶片N和P含量表现为:樟子松>赤松>油松,而叶片K含量在三个树种中并无显著差别。表层土壤的全N和全P,有机质和全P以及叶片N和P含量之间表现出了显著的相关性,但是,林分表层土壤养分和树木针叶养分之间并无显著相关关系,这可能与当地对凋落物的过度收集,过度放牧以及土壤水分含量低有关。最后,根据国外研究结果引入N:P比阈值来分析研究区域的N,P养分状况,基于章古台地区这几种针叶树种针叶N:P比,我们认为研究区域的樟子松随着年龄的增加受N限制的可能性降低,29年的赤松林整体表现出不受N和P养分限制,而31年的油松林则表现为系统N和P都供应不足。图3表3参45。     

Effects of prescribed fire on canopy foliar chemistry and suitability for an insect herbivore

We assessed the effects of a prescribed fire on the phytochemical characteristics and vigor of overstory chestnut oak, scarlet oak, and red maple, and measured the impact of potential changes on herbivore fitness. We compared foliar carbohydrates, tannins, nutrients, and fiber concentrations in foliage from burned and non-burned forest canopies. There were significant differences in most foliar characteristics between tree species. Total non-structural carbohydrate concentrations in scarlet oak and red maple foliage, and calcium levels in red maple foliage, were significantly lower in burned plots, but other phytochemical characteristics were largely unaffected by burning. Tree growth also varied with species. Burning increased chestnut oak relative growth, decreased scarlet oak growth, and had no affect on red maple growth. Scarlet oak and red maple foliage from burned and non-burned forest canopies were assayed for gypsy moth performance. Caterpillars fed foliage from scarlet oak, the preferred host, grew larger and developed more rapidly than did those fed red maple foliage. There were no significant burn treatment differences in caterpillar development within either tree species, suggesting that managers using prescribed fire to promote oak regeneration need not worry about enhancing forest stand susceptibility to gypsy moth. However, the fire in this study was of low to moderate intensity; more intense fires may alter foliar palatability.     

Effect of fertilization on the litter fall of pinus sylvestris and betula pubescens on drained peatlands

NPK fertilization on a dwarf shrub pine bog initially increased the amount and nutrient content of the tree litter. Eight years after fertilization, however, the amount of micronutrients decreased compared to the amount of N and P in the litter. Fertilization on a fertile mire also increased the nutrient content of the litter fall, especially in the mature pine and birch stands. The amount of nutrients in the litter fall of the birch stands was considerably greater than that in the pine stand of the same volume.     

两种混交度的比较分析

混交度是描述混交林中树种混交程度的重要空间结构指数。由于传统混交度仅以对象木与最近邻木之间的树种异同为基础,并不考虑最近邻木之间的树种差异,不能完全反映树种的空间隔离程度。本文在传统混交度的基础上,提出一种新的混交度即树种多样性混交度。通过比较,树种多样性混交度更能反映树种相互隔离程度,是一个较好的林分空间结构指数。     

Species differences in timing of leaf fall and foliage chemistry modify nutrient resorption efficiency in deciduous temperate forest stands

Extensive variation in fractional resorption of mineral elements from plant leaves is still not fully understood. In multi-species forest stands, species leaf fall phenology and leaf constitution may significantly modify the timing of nutrient return to the soil and overall plant nutrient loss. We studied leaf fall and nutrient loss kinetics, and leaf composition in three natural, temperate, deciduous broadleaf forest stands to determine the role of timing of leaf abscission and nutrient immobilization in cell walls on nutrient resorption efficiency of senescing leaves. Nitrogen (N), phosphorus and potassium contents decreased continuously in attached leaves after peak physiological activity during mid-season. Changes in nutrient contents of attached leaves were paralleled by decreases in nutrient contents in freshly fallen leaf litter. In different species and for different nutrients, resorption of nutrients from senescing leaves proceeded with different kinetics. The maximum nutrient resorption efficiency (the fraction of specific nutrient resorbed from the leaves at the end of leaf fall) did not depend on the mid-seasonal nutrient concentration. Species with earlier leaf fall resorbed leaf nutrients at a faster rate, partly compensating for the earlier leaf fall. Nevertheless, the litter-mass weighted mean nutrient contents in leaf litter were still larger in species with earlier leaf fall, demonstrating an inherent trade-off between early leaf fall and efficient nutrient resorption. This trade-off was most important for N. Losses of the non-mobile nutrients calcium and magnesium were unaffected by the timing of leaf fall. There was large variation in the maximum N resorption efficiency among species. Correlations among leaf chemical variables suggested that the maximum N resorption efficiency decreased with the increasing fraction of cell walls in the leaves, possibly due to a greater fraction of N occluded in cell wall matrix. We conclude that species leaf fall phenology and leaf chemistry modify the timing and quantities of plant nutrient losses, and that more diverse forest stands supporting a spectrum of species with different phenologies and leaf types produce litter with more variable chemical characteristics than monotypic stands.     

不同林龄樟子松叶片养分含量及其再吸收效率

树木叶片的养分再吸收效率能够反映树木对养分保存、利用以及对养分贫瘠环境的适应能力。以科尔沁沙地东南缘章古台地区樟子松人工林为研究对象,分析了11、20、29、45年生树木叶片的基本特征、养分含量及其再吸收效率。结果表明:叶片衰老后其质量和面积明显减少;叶片凋落前的平均养分含量没有表现出随樟子松年龄增加而出现有规律的变化;凋落叶片中的N、P、K、Mg含量表现出随年龄增加而增加的趋势,而Ca的趋势与之相反;11年生和20年生的樟子松叶片N、P、K的再吸收效率相似,都显著高于29年生和45年生樟子松(P<0.05),而樟子松叶片对Mg的再吸收效率表现出随年龄增大而显著降低,Ca随叶片的衰老而不断累积,再吸收效率表现为负值,20年生的樟子松叶片Ca再吸收效率最大,11年生和45年生最低。樟子松叶片的N、P、K、Mg养分再吸收效率随年龄增加而降低的趋势表明,随年龄增加樟子松对贫瘠养分生境的适应能力逐渐降低,反映了樟子松养分保存方面的衰退特征。     

Contribution of agroforestry trees to nutrient requirements of intercropped plants

A major tenet of agroforestry, that trees maintain soil fertility, is based primarily on observations of higher crop yields near trees or where trees were previously grown. Recently objective analyses and controlled experiments have addressed this topic. This paper examines the issues of tree prunings containing sufficient nutrients to meet crop demands, the timing of nutrient transfer from decomposition to intercrops, the percent of nutrients released that are taken up by the crop, and the fate of nutrients not taken up by the crop.The amount of nutrients provided by prunings are determined by the production rate and nutrient concentrations, both depending on climate, soil type, tree species, plant part, tree density and tree pruning regime. A large number of screening and alley cropping trials in different climate-soil environments indicate that prunings of several tree species contain sufficient nutrients to meet crop demand, with the notable exception of phosphorus. Specific recommendations for the appropriate trees in a given environment await synthesis of existing data, currently only general guidelines can be provided.Tree biomass containing sufficient nutrients to meet crop demand is not enough, the nutrients must be supplied in synchrony to crop needs. Nutrient release patterns from organic materials are, in part, determined by their chemical composition, or quality. Leguminous materials release nitrogen immediately, unless they contain high levels of lignin or polyphenols. Nonlegumes and litter of both legumes and nonlegumes generally immobilize N initially. There is little data on release patterns of other nutrients. Indices that predict nutrient release patterns will assist in the selection of species for synchronizing with crop demand and improve nutrient use-efficiency.Field trials with agroforestry species ranging in quality show that as much as 80% of the nutrients are released during the course of annual crop growth but less than 20% is captured by the crop, a low nutrient-use efficiency. There are insufficient data to determine how much of the N not captured by the crop is captured by the trees or is in the soil organic matter, the availability of that N to subsequent crops, or how much of that N is lost through leaching, volatilization or denitrification. Longer term trials are needed.     

The Growth of Sitka Spruce on Poorly Drained Soils in Northern Ireland

A soil survey of Lisnaskea Forest, Co. Fermanagh, showed thatgrowth of Sitka spruce was far from uniform on areas of apparentlyuniform soil. The relation between soil factors, tree growth,and foliar nutrient status on two blocks of trees growing ongleyed soils was therefore studied. Simple regression showed that tree growth was poor when thelevels of foliar nutrients were low and when there was a matof undecomposed needles on the forest floor. It is thereforeconsidered that lack of organic matter breakdown in the soilis causing poor tree nutrition and growth. Multivariate statistical analysis also showed highly significantregressions between tree growth, level of foliar nutrients,and depth of litter layer. The correlations between soil factorsalone and tree growth were barely significant and the multivariatetechnique did not give an acceptable method of predicting treegrowth from soil measurements. Methods of improving soil conditions are discussed.     

Nutrient cycling under mixed-species tree systems in southeast Asia

Eucalyptus and Acacia are two tree genera that are commonly used in industrial plantations and as components of agroforestry systems in southeast Asia. These fast-growing trees are mostly grown in monocultures. However, questions are now being raised about the long-term sustainability of their growth as well as their effects on site quality. Losses of N and P from the site through biomass harvest and during site preparation constitute a major nutrient drain. As an alternative to monocultures, mixed-species cultures which include at least one N2-fixing tree species can improve plant productivity and soil N dynamics. Among the various aspects of N dynamics in such stands, export of N during biomass harvest and inputs by the N₂-fixing component are the most important. Reported estimates of the amount of N fixed by acacia and other N₂-fixing trees are highly variable, depending on inherited plant characteristics, tree age, site factors (e.g., drought), soil fertility (e.g., available P, metal toxicities) and unreliable methods of measuring N₂-fixation. Of the available techniques for assessing N₂-fixation by trees, the total N difference method (TND) is the simplest. The contribution of roots to assessments of N₂-fixation is recognized but rarely measured. For short-rotation mixed-species plantations, the amount and time of N transfer from N₂-fixing trees to non-N₂-fixing trees are important issues to consider when attempting to develop productive nutrient management strategies. Based on limited information from trials in southeast Asia, it appears that acacia fixes substantial amounts of N during the first few years of establishment and a significant amount of that N is transferred to adjacent eucalypts, thereby improving the growth and nutrition of the eucalypts. The presumed transfer of N from acacias to eucalypts during the early stages of plantation development probably results from belowground turnover of roots and nodules because aboveground litter decomposition is slight at this stage, and contributes little to the overall N dynamics. The available information on P cycling in mixed-species stands, during the early stages of stand growth, provides inconclusive evidence as to whether the availability of soil P increases, despite indications of higher levels of phosphatase activity in the fine roots of nitrogen-fixing trees. This would imply that additional inputs of P as fertilizer are required to remove any P deficiency in mixed-species stands. Long-term observations are required for better understanding of the nutritional and growth benefits of including N₂-fixing trees in mixed-species stands. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparison of new foliar nutrient thresholds derived from van den Burg’s literature compilation with established central European references

In this paper, we suggest new nutrient threshold values and ratios for the major nutrients (N, P, K, Ca, Mg) for four main trees species Pinus sylvestris, Picea abies, Fagus sylvatica and Quercus (Q. robur and Q. petraea) statistically derived from the data collection of van den Burg’s literature compilation (1985, 1990), and compare them with widely used central European literature. The comparison is focused on the normal range of nutrition of major elements. We could show that the new critical foliar nutrient concentrations and ratios do not have the imponderabilities of the established reference systems, like restricted calibration range or data gaps for species in general or some major nutrients in particular. Ranges from the new critical foliar nutrient concentrations are generally smaller, except for broadleaves foliar Ca and Mg values, where another reference provides the narrowest ranges. The practicability of the different systems has been exemplary tested on a foliage data set of Norway spruce from the Bavarian soil survey. This application illustrates that an evaluation based on the new nutrient thresholds is in the majority of cases more cautious than the other reference systems. Moreover, evaluations based on the new concentrations and ratios are quite consistent. Comparisons with nutrient ratios related to the susceptibility to parasite attacks show that the thresholds are close to a general optimum range, indicating health and intact resistance mechanisms of these tree species. Uncertainties of the new threshold values as well as the need to further evaluate and develop reference systems for nutrient status of forest trees are discussed.     

Seasonal changes in litter fall and its quality from three sub-tropical fruit tree species at Nelspruit, South Africa

Litter fall from upper storey trees in agroforestry systems contributes to nutrient cycling for the benefit of all components of the system besides serving as mulch. This study examined the seasonal changes in the quantity and quality of leaf litter fall from three sub-tropical fruit trees viz: avocado (Persea americana L.), mango (Mangifera indica L.) and litchi (Litchi chinensis L.) which have potential for use in agroforestry. Leaf litter production was estimated using nylon mesh litter traps erected over five randomly selected trees of each species in a completely randomised design. Litter quality was determined by analysing ash content and polyphenol, carbon, cellulose, lignin and nutrient concentrations over a 2?year period (2007?C2008). Total annual leaf litter production during the study period (dry matter basis) was 8.3, 6.3 and 5.6?t?ha^?1?year^?1 for litchi, mango and avocado, respectively. In both years, leaf litter fall was greatest during autumn and lowest during winter in all species. There were no significant differences in S, Ca, Mg and Mn concentrations in the leaf litter, but polyphenol, N, P and K concentrations differed significantly (P?<?0.05) between species. It was concluded that litter quality from all three tree species was low and would require appropriate management to improve its quality.     

Corresponding patterns of site quality,decline and tree growth in a sessile oak stand

This study focuses on two rarely studied aspects of oak decline: relations with site characteristics and effects on tree growth. The study was carried out in a 5.5 ha stand in Hungary which is strongly affected by oak decline. The nearly pure sessile oak (Quercus petraea) stand of mostly coppice origin was 90 years old at the beginning of the study. Within-stand site heterogeneity was described by the herbaceous vegetation. Four ecological site types were distinguished by the species composition of herbs, and characterized by the ecological indicator values of the species. Tree growth between 1987 and 1993 was measured, and tree vigour was estimated from visual characteristics five times between 1987 and 1993. Potential volume increment of declining trees was estimated with the growth rates of healthy trees of the same size. Volume increment loss caused by oak decline was also assessed. Significant positive relationships were found between tree vigour and tree size and between tree vigour and tree growth. The growth of seriously declining trees dropped to almost one-half of that of healthy ones. Growth reduction of living trees at the stand level amounted to 5.4%, whereas growth reduction of all trees, including those that died during the observation period, amounted to 19.9% of the potential growth. Tree size and growth were greater on better sites. A strong relationship was also found between tree vigour and site type, but sessile oak was more susceptible to decline at better sites.     

Dynamics of organic matter and nutrient return from litterfall in stands of ten tropical tree plantation species

We studied the rates and patterns of carbon and nutrient fluxes in litterfall in ten tropical tree plantation species grown at the USDA Forest Service Arboretum in the Luquillo Experimental Forest, Puerto Rico. The stands were 26-years old and grew under similar climatic and edaphic conditions. Individual plantation species ranked differently in terms of their capacity to return mass and specific nutrients to the forest floor, and with respect to their efficiency of nutrient use. The species that returned the most mass did not return the most P, N, or cations. Moreover, species differed according to the amount of N and P resorption before leaf fall. These differences reflect the variation in the ecophysiological response of each species to edaphic and climatic conditions. The difference between average and minimum resorption values of the species studied indicate that other environmental factors, such as heavy winds or the physical effects of heavy rains, can force the shedding of non-senesced leaves. This higher quality material, although not very much in quantity, can provide a small pulse of available nutrients to the forest floor community. The same holds true for other high nutrient/low mass fractions of litterfall such as reproductive parts and miscellaneous materials.In areas with no prevalent or strongly seasonal water limitations, temporal variations of leaf litter on the forest floor are the combined result of the rate of fall and decomposition of the falling material, and the diverse responses of species to different environmental cues. Leaf fall was inversely correlated to reduced water availability in three of the species studied. Leaf fall of the other species was correlated either to daylight duration or minimum temperatures. The results highlight the importance of understanding species performance relative to nutrient and mass metabolism before selection for plantation use, or for rehabilitation of degraded lands.     

Silvopastoralism in New Zealand: review of effects of evergreen and deciduous trees on pasture dynamics

Complex interactions between livestock, trees and pasture occur in silvopastoral systems. Between trees and pasture, competition for soil resources (nutrients and water) occurs, becoming especially relevant when one of them is in scarce supply. Trees reduce light and water reaching the understorey layers according to tree density and canopy size. However, they may ameliorate extreme climatological features (reducing wind speed and evapotranspiration, and alleviating extreme temperatures), and improve soil properties, for example, deciduous tree litter may contribute to increased pH and soil nutrient concentrations. During tree establishment, there are generally negligible effects on pasture, irrespective of tree type. However, there is a decline in pasture production and nutritive value under shade with increasing tree age and higher stand density. Under the same conditions, deciduous trees affect pasture later (extinction point of pasture occurs at 85% of canopy closure) than evergreen trees (about 67% for Pinus radiata D. Don). This is mainly because deciduous trees have a leafless period that enables pasture recovery, and their litter smothers pasture less intensely because of its relatively fast decomposition. Silvopastoral studies conducted in New Zealand are reviewed to discuss these effects, and differences in the effects of evergreen and deciduous trees are shown using the examples of P. radiata, and Populus and Salix spp. respectively, which exist in many temperate countries. Future research needs are outlined.     

Dynamics of some mineral nutrients and heavy metals in decomposing forest litter

The dynamics of three major mineral nutrients (K, Ca, Mg) and six heavy metals (Fe, Mn, Zn, Cu, Pb and Cd) were studied in decomposing natural, unpolluted litter in two forest types: Scots pine needle litter in a pure Scots pine forest and oak‐hornbeam leaf litter in a mixed oak‐hornbeam stand. Of the mineral nutrients, only K in the oak‐hornbeam system showed a significant decrease in concentration during the course of litter decomposition. For Ca and Mg in both systems and for K in the Scots pine system no clear relation between concentration and accumulated litter‐mass loss was observed. On the other hand, for Fe, Zn, Pb and Cd at both sites, as well as for Mn in the oak‐hornbeam stand and Cu in the Scots pine stand highly significant positive relations were found between heavy metal concentrations and accumulated mass loss. A decrease in concentration was noted only in the case of Mn in the Scots pine system. No clear pattern was found for Cu in the oak‐hornbeam stand. Concentrations of heavy metals in the decomposing litter reached levels at or above those reported to retard decomposition in other studies. At the oak‐hornbeam stand increased also the absolute amounts of Fe, Zn, Pb and Cd.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.