Flora, vegetation, and soil in broadleaved ancient and planted woodland, and scrub on RøsnÆs, Denmark

9 small ancient woodlands (>200 years), 9 planted woodland sites (25 — ca. 100 years), and 6 sites in grazed scrub (ca. 100 years) on the RøsnÆs peninsula, Denmark, showed characteristic differences: in ancient woodland, the tree and shrub layer was fairly rich, the field layer rather poor in species. The dominance of spring flowering geophytes, the abundance of Anemone nemorosa , and the occurrence of Corylus avellana and Polygonatum multiflorum were characteristic. pH of the soil was relatively low, organic matter content high, and light intensity at the forest floor in summer low. In planted woodland there was more light, and the field layer was rich in short-lived species, but poor in spring flowering geophytes. Many woodland species were rare in planted woodland, some did not at all occur there, and none were specific for this type of woodland. The scrub was marked by grazing and a strong relief, hence pH was high and organic matter content low. The field layer was rich both in shortlived species and in spring flowering geophytes. — It is suggested that ancient woodland species (i.e. species restricted to or preferably occurring in woodlands existing prior to the enclosure ca. 200 years ago) is a heterogenous group, consisting of a) species favoured by traditional woodland management; b) species restricted to woodlands where specific environmental (e.g. soil) conditions have had sufficient time to develop; c) species with limited ability to spread or establish; or d) species which in ancient woodland are represented by small and scattered populations.     

Green light drives photosynthesis in mosses

Temperate forests are characterised by variable light quality (i.e. spectral composition of light) at or near the forest floor. These understory environments have a high concentration of green light, as red and blue light are preferentially absorbed by upper canopy leaves. Understory species may be well-adapted for using green light to drive photosynthesis. Angiosperms have been shown to use green light for photosynthesis, but this ability has not been demonstrated in shade-dwelling bryophytes. In this study, net photosynthetic rate (P_N) of three temperate understory species of moss (Dichodontium pellucidum (Hedw.) Schimp., Leucobryum albidum (Brid. ex P.Beauv) Lindb. and Amblystegium serpens (Hedw.) Schimp.) was measured under green, red?+?blue, and red?+?blue?+?green light to assess green light use efficiency. All three species were capable of photosynthesising beyond their respiratory demands using solely green light, with higher green light use efficiency measured in plants collected from areas with greater canopy cover, suggesting growth in a green light concentrated environment increases green light use efficiency. Each species was also collected from sites differing in their degree of canopy cover and grown under three light treatments (high light, low light, and green light). Photosynthetic efficiency (chlorophyll fluorescence), tissue nitrogen and carbon isotope concentrations were assessed after a short growth period. Growth conditions had little effect on leaf chemistry and monochromatic green light did not significantly degrade photosynthetic efficiency. This study provides the first evidence to date of positive net ‘green light photosynthesis’ in mosses.     

Air humidity,soil moisture and soil chemistry as determinants of the herb layer composition in European beech forests

Question: What role does air humidity play as an environmental factor for the abundance and distribution of temperate woodland herbs? Location: Beech forests on calcareous soils in southern lower Saxony, central Germany. Methods: The abundance of woodland herb species and total herb cover were investigated in 60 plots with contrasting exposure, slope angle and relief type. On all plots, air humidity, air temperature, soil moisture, photosynthetically active radiation, pH (H₂O) and concentration of salt‐exchangeable Ca, Mg and K were measured. Species‐environment relationships were analysed with multiple regression analysis and CCA. Results: Air humidity (RH), soil moisture and the concentration of exchangeable Ca and K, but not light, C/N ratio and the concentration of exchangeable Mg were identified as the most important abiotic factors influencing the cover of the most abundant plant species and total herb cover. RH varied substantially across the different forest floor site types and influenced species abundance independent of soil moisture. In several species (including Mercurialis perennis and Impatiens noli‐tangere), RH was found to be a key environmental factor. Other species such as Aegopodium podagraria and Lamiastrum galeobdolon depended more on elevated soil moisture, while RH was less important. Conclusions: This study showed that the distribution of widespread temperate woodland herb species depends on high air humidity, and that certain sensitive species do not occur at sites with reduced air humidity even though soil moisture is high. Thus, high air humidity and ample soil moisture are key abiotic factors in beech forests on calcareous soils. Shade level (PAR) was found to be of secondary importance.     

Effects of coppicing on demographic structure, fruit and seed set in Orchis mascula

Opening of the forest canopy may result in higher fitness of understory plants due to increased light intensity on the forest soil and increased nutrient availability. Especially, tuberous orchids may profit from increased light as photosynthetic assimilation products and nutrients are accumulated in their belowground storage organs. We investigated the effects of coppicing on demographic structure and fruit and seed set in 15 populations of the tuberous, perennial orchid Orchis mascula, seven of which were located in undisturbed, shaded woodland and eight in recently coppiced woodland. Coppicing resulted in a massive increase in flowering and in increased fruit set. On average, 42.9% of all individuals consisted of flowering plants in coppiced woodland, whereas in undisturbed woodland only 20.8% of all individuals flowered. The percentage fruit set varied between 20% and 55% in coppiced woodland and was strongly related to population size (measured as the number of flowering individuals). Mean fruit weight and fruit size were also significantly higher in populations located in coppiced woodland than in undisturbed woodland, whereas the proportion of viable seeds was not related to woodland type. Given that O. mascula is dependent on seed set to fill space with new individuals, these results suggest that the long-term survival of this species strongly depends on frequent opening of the canopy.     

Environmental effects on photosynthesis, nitrogen-use efficiency, and metabolite pools in leaves of sun and shade plants

Effects of varying light intensity and nitrogen nutrition on photosynthetic physiology and biochemistry were examined in the sun plant Phaseolus vulgaris (common bean) and in the shade plant Alocasia macrorrhiza (Australian rainforest floor species). In both Phaseolus and Alocasia, the differing growth regimes produced large changes in photosynthetic capacity and composition of the photosynthetic apparatus. CO₂-saturated rates of photosynthesis were linearly related to leaf nitrogen (N) content in both species but photosynthesis per unit leaf N was markedly higher for Phaseolus than for Alocasia. Photosynthetic capacity was also higher in Phaseolus per unit ribulose 1,5-bisphosphate (RuBP) carboxylase (RuBPCase) protein. The leaf content of RuBPCase was linearly dependent on leaf N content in the two species. However, the proportion of leaf N which was RuBPCase was greater in Phaseolus than in Alocasia and was more sensitive to growth conditions, ranging from 6% of leaf N at low light to 20% at high light. In Alocasia, this range was much less, 6 to 11%. However, chlorophyll content was much more sensitive to light intensity in Alocasia. Thus, the RuBPCase/chlorophyll ratio was quite responsive to N availability and light intensity in both species (but for different reasons), ranging from 6 grams per gram for Phaseolus and 2 grams per gram for Alocasia at high leaf N and 1.5 gram per gram for Phaseolus and 0.5 gram per gram for Alocasia at low leaf N. These large changes in the proportions of components of the photosynthetic apparatus had marked effects on the sensitivity of these species to photoinhibition. These environmental effects also caused changes in the absolute levels of metabolites of the photosynthetic carbon reduction cycle. Concentrations of RuBP and P-glycerate were approximately 2-fold higher in high light-grown than low light-grown Phaseolus and Alocasia when expressed on a leaf area basis. However, if metabolite pool sizes are expressed on the basis of the RuBPCase catalytic site concentration, then they were little affected by the marked changes in leaf makeup. There appears to be fundamental differences between these species in the mechanism of sun-shade adaptation and N partitioning in the photosynthetic apparatus that result in significant differences in the N-use efficiency of photosynthesis between Phaseolus and Alocasia but similar RuBPCase:substrate:product ratios despite these differences.     

Pathogen induced disturbance and succession in temperate forests: Evidence from a 100-year data set in southern Sweden

Major tree species are declining in many temperate forests due to changing disturbance regimes, including invasive pests and pathogens. We examined the interaction of secondary succession and Dutch elm disease in the Swedish temperate forest reserve Dalby Söderskog, based on five tree surveys made between 1909 and 2011. The forest is characterized by the coexistence of four major European tree species: wych elm (Ulmus glabra), European ash (Fraxinus excelsior), European beech (Fagus sylvatica) and pedunculate oak (Quercus robur). After protection of the forest in 1918, lack of disturbance mainly favoured elm, while the oak population declined due to mortality of old oaks and lack of regeneration. Dutch elm disease has caused high and continuous elm mortality since 1988. As a result, increased light availability at the forest floor favoured abundant regeneration of ash, beech, and lately also oak. The recent arrival of an invasive fungal pathogen causing ash dieback may once again change the course of succession. Open space emerging from loss of elm and ash in forest reserves may be used by reserve managers to favour oak regeneration and biodiversity of semi-open woodlands once lost during succession to closed forest. We conclude that winners and losers change places as an effect of invasive pathogens, resulting in unexpected successions and both losses and gains in valuable ecological niches and habitat structures in temperate broadleaf forests.     

Light, nutrients and the growth of herbaceous forest species

The herb layer of forests planted on former agricultural land often differs from that of old-growth forest. This study investigates if the expected increased nutrient availability in the shaded conditions of newly planted forests and the plasticity of the species to adjust their biomass allocation to different levels of light and nutrients help to explain these differences in the herb layers of the two forest types. In a greenhouse experiment biomass distribution and production of two species characteristic for the highly shaded forest floor, Circaea lutetiana and Mercurialis perennis, and two species more common in the forest-edge, Aegopodium podagraria and Impatiens parviflora were studied at different levels of light (2%, 8% and 66% of the full light level) and nutrients (30 and 300 kg N ha^–1 per year). The main factor affecting allocation and biomass production was light availability. Nutrient supply only had a significant effect at the higher light levels. Species were mainly plastic to changes in light and the two species from the forest floor showed to be more rigid in allocation pattern than the species from the forest-edge. So, although the species from the forest-edge were more plastic, they did not profit from the increased nutrient supply because the main factor affecting biomass distribution and production was light availability.     

Ecological studies on the timberline of Mt. Fuji

An ecological study of dry matter production was made in a dwarf forest dominated byAlnus maximowiczii at the timberline of Mt. Fuji. Annual gross production was estimated by two methods, namely the summation method using stem analysis and total photosynthesis calculated from leaf area and photosynthetic rate per leaf area. Seasonal changes in relative light intensity and in leaf area were measured in a quadrat. Photosynthesis and respiration rates of samples were measured in temperature-regulated assimilation chambers. The phytomass was 2,989 g d.w.m^?2, and those of stems and branches, leaves, and roots were 1,672 g, 293 g, and 1,024 g respectively. The growing period of this plant was about four months and this plant expanded leaves quickly. The maximum gross photosynthetic rate was 21 mg CO₂dm^?2 h^?1 on September 1. Annual net production estimated by examining the annual rings was 922 g d.w.m^?2 year^?1 and annual respiration was 735 g. Annual gross production estimated from photosynthetic rates was 1,747 g d.w.m^?2 year^?1. The sum of annual net production by stem analysis and respiration agree closely with gross production estimated from photosynthetic rate. Gross production of this dwarf forest is comparable to the beech forest of the upper cool temperate zone owing to the high photosynthetic rate ofAlnus maximowiczii.     

Does Acacia dealbata express shade tolerance in Mediterranean forest ecosystems of South America?

The distribution of Acacia dealbata Link (Fabaceae) in its non-native range is associated with disturbed areas. However, the possibility that it can penetrate the native forest during the invasion process cannot be ruled out. This statement is supported by the fact that this species has been experimentally established successfully under the canopy of native forest. Nonetheless, it is unknown whether A. dealbata can express shade tolerance traits to help increase its invasive potential. We investigated the shade tolerance of A. dealbata under the canopy of two native forests and one non-native for three consecutive years, as well as its early growth and photosynthetic performance at low light intensities (9, 30, and 70 μmol m⁻²sec⁻¹) under controlled conditions. We found many A. dealbata plants surviving and growing under the canopy of native and non-native forests. The number of plants of this invasive species remained almost constant under the canopy of native forests during the years of study. However, the largest number of A. dealbata plants was found under the canopy of non-native forest. In every case, the distribution pattern varied with a highest density of plants in forest edges decreasing progressively toward the inside. Germination and early growth of A. dealbata were slow but successful at three low light intensities tested under controlled conditions. For all tested light regimes, we observed that in this species, most of the energy was dissipated by photochemical processes, in accordance with the high photosynthetic rates that this plant showed, despite the really low light intensities under which it was grown. Our study reveals that A. dealbata expressed shade tolerance traits under the canopy of native and non-native forests. This behavior is supported by the efficient photosynthetic performance that A. dealbata showed at low light intensities. Therefore, these results suggest that Mediterranean forest ecosystems of South America can become progressively invaded by A. dealbata and provide a basis for estimating the possible impacts that this invasive species can cause in these ecosystems in a timescale.     

Andreaea Hartmanii Thed.—A histological study

Abstract

Photosynthetic responses to light intensity were studied under laboratory conditions in seven bryophyte species from evergreen laurel forest, a threatened habitat, on Terceira island in the Azores. Four mosses (Andoa berthelotiana, Echinodium prolixum, Fissidens serrulatus, Myurium hochstetteri) and three liverworts (Bazzania azorica, Frullania tamarisci, Lepidozia cupressina) were selected to encompass a range of potential responses to variations in the forest light environment. Carbon dioxide exchange measurements were made, using an infra-red gas-analyser, at photosynthetic photon flux densities (PPFD) of 0-900 µmol m^-2 s^-1 and a mean temperature of 21°C in fully hydrated shoots. Most species achieved light saturation of photosynthesis below 30 µmol m^-2 s^-1, the lowest value being for A. berthelotiana (20 µmol m^-2 s^-1) and the highest for M. hochstetteri (68 µmol m^-2 s^-1). The liverwort F. tamarisci had the highest maximum photosynthetic rate (P_max, 23 µmol CO₂ g^-1 h^-1) whereas P_max was lowest in the mosses E. prolixum and M. hochstetteri (10 µmol CO₂ g^-1 h^-1). Dark respiration rate, a critical factor in toleration of shade by forest floor plants, was highest in the species with the highest values for P_max. Compensation point was extremely low (7 µmol photons m^-2 s^-1) in Fissidens serrulatus, a species found in the deep shade of forest ravines and caves, and highest in M. hochstetteri a moss restricted to better illuminated habitats within and outside the forest. No photoinhibition was detected during the relatively short exposures to high irradiances. Comparison of these responses with data on the forest light environment indicates that, despite the possession of considerable shade adaptations, during winter in the evergreen laurel forest, low light levels may often limit photosynthetic rates of the bryophytes.     

Seasonal changes in photosynthesis of four understory herbs in deciduous forests

Seasonal patterns of photosynthesis and respiration of single leaves of four understory perennial herbs in deciduous forests were investigated in relation to their leaf growth and light conditions on the forest floor.Anemone flaccida shows rapid growth of leaf area and high rates of gross photosynthesis at light saturation (Psat) in its early stage of development. Its photosynthetic activity is restricted to a brief period of high light intensity before the closure of overstory canopies.Disporum smilacinum possesses light-photosynthesis curves of the shade-leaf type throughout its whole growing period. A shading experiment has shown that this plant is low-light adapted and can utilize weak light efficiently. The light-photosynthesis curve ofSyneilesis palmata shifts from the sun-leaf type to the shade-leaf type in response to the seasonal change of light regime on the forest floor. Evergreen leaves ofPyrola japonica have three year longevity, and light-photosynthesis curves of the shade-leaf type. They maintain some photosynthetic activity even in late autumn and winter.     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Seasonal occurrence and distribution of myxomycetes on different types of leaf litter in a warm temperate forest of western Japan

The seasonal occurrence and distribution of myxomycetes on different types of newly defoliated leaf litter were examined in a secondary forest in a warm temperate region of western Japan. The two types of leaf litter (deciduous trees, Prunus verecunda and Quercus variabilis, and evergreen trees, Q. glauca and Cinnamomum camphora) were incubated in trays on the forest floor. A total of 45 myxomycete species were recorded from 3021 collected samples that occurred at the July peak during the warmest and humidest season from April to November. The occurrence of species was significantly related to the changes in mean temperature and minimum temperature on both leaf types under humid conditions. Myxomycete assemblages were divided into three seasonal phases. Most of the species occurred in June–September, while a few species demonstrated characteristic distributions; i.e., Didymium melanospermum appeared in April– May and Diderma umbilicatum appeared in October– November. The respective leaf types supported the reproduction of myxomycetes with high species richness and diversity, with 34 species and H’ = 2.59 on deciduous trees and similarly 30 species and H’ = 2.49 on evergreen trees. Several species, however, exhibited a preference for either the deciduous tree or evergreen tree leaves. Thus, a mixed forest that defoliates during different two seasons yields a greater species diversity of myxomycete assemblage.     

Light and soil moisture effects on biomass and its allocation in Osmorhiza depauperata Philippi (Apiaceae)

Changes in forest openings affect light quality and quantity, and the magnitude of rainfall that reaches the soil surface. Osmorhiza depauperata, a geophyte, acclimates to changes imposed because of forest openings. We studied which changes in biomass allocation allow acclimation of O. depauperata to the various environments that this species inhabits, and where it develops better. Three light intensities (I4 = 4 %, I26 = 26 %, I64 = 64 % of ambient sunlight) and two moisture levels (M40 = 40–60 %, M80 = 80–100 % field capacity) were evaluated on O. depauperata under greenhouse conditions. Plant biomasses per pot were 0.81, 0.56 and 0.48 g at I26, I4 and I64 light intensities, respectively, after one growing season. The biomass allocation to aboveground tissues and leaf area decreased as light intensity increased. Soil moisture modified only belowground biomass and weight of fine roots. The interaction between soil moisture content and light intensity was consistent. This was because of a significant reduction in total plant biomass under high both soil moisture content and high light intensity. Osmorhiza depauperata growth was favored most at medium light intensities. Changes in biomass allocation among various organs allow this species to inhabit forest habitats with different light intensities.     

荆条叶性状对野外不同光环境的表型可塑性

光照是影响植物生长和分布的重要环境因子。对生长在野外5种不同光环境下(林外、阔叶林林缘、阔叶林林下、针叶林林窗和针叶林林下)的荆条的叶片进行取样研究,通过对光合作用光响应曲线、叶绿素荧光、叶绿素含量、叶片氮磷含量以及叶片形态的测量,来反映荆条对不同光环境的表型可塑性。研究结果表明,荆条叶片对于野外不同的光环境具有很好的适应机制,叶片功能性状受到结构性状的调节。低光下通过高的比叶面积(SLA)、单位质量叶绿素含量、光系统II最大量子产量,低的暗呼吸速率、光饱和点、光补偿点、叶绿素a,b的比值来提高对光能的利用效率,维持生长;高光下则通过与SLA有关的叶片结构的变化对光合作用进行调节。大多数的叶性状只受到日光照总量的影响,SLA的大小与日最高光强有关,可以对不同日变化模式的光照做出迅速的响应,是适应不同光照的敏感指标。尽管光照是不同光环境下影响荆条叶性状的主要环境因子,土壤养分含量同样会对叶性状产生影响,高土壤养分下的高叶长与叶柄长的比值体现了植物对资源获取和支撑结构之间分配的权衡。     

Plant species diversity in abandoned coppice forests in a temperate deciduous forest area of central Japan

We investigated plant species diversity as it related to stand structure and landscape parameters in abandoned coppice forests in a temperate, deciduous forest area of central Japan, where Fagus crenata was originally dominant. The species occurring in the study plots were classified into habitat types based on a statistical analysis of their occurrence bias in particular habitats (e.g., primary forest, coniferous plantation) in the landscape studied. The relationships between stand structure, which reflected the gradient of management, and forest floor plant species diversity (H and J) and richness (number of species per unit area) were not significant. However, these factors did influence the forest floor plant composition of the different types of habitat. According to the multiple regression analysis, species diversity and the richness of forest floor plants was affected by landscape parameters rather than by stand structure. For trees, species richness was mainly affected by the relative dominance of F. crenata, which is one of the stand structure parameters that decreases with intensive management. This is probably because many of the tree species that are characteristic of coppice forests increase after F. crenata have been eliminated by management; these species are not dominant in the original forest, where they are suppressed by F. crenata, the shade-tolerant dominant species. The species diversity (H and J) of trees was positively correlated with some landscape parameters, including the road density around the study plot, which may be associated with the intensity of management activity. The number of disturbance-tolerant species increased with increasing road density. Stand structure mainly affected disturbance-intolerant forest floor plant species and disturbance-tolerant tree species. Thus, the species diversity responses differed between forest floor plants and trees. The impact of forest management on species diversity was more prominent for forest floor plants.     

森林光环境对4种乔木幼树光合和光谱反射特性的影响

在林窗、林缘、林下3种自然光环境下,对吉林蛟河阔叶红松林常见树种拧筋槭、白牛槭、色木槭和红松的光合和光谱特性进行对比研究,以期从生理生态角度探讨它们对自然光环境的适应能力和响应机制。结果表明：4种乔木幼树叶片的光饱和点(LSP)和光补偿点(LCP)随着生长环境光强的下降而减小,林下各树种幼树LSP和LCP显著低于林窗内幼树(P<0.05);除白牛槭外林下其他树种最大净光合效率(P_nmax)均显著小于林窗内幼树(P<0.05);不同树种幼树表观量子效率(AQY)和暗呼吸速率(R_d)随光环境的变化并没有出现明显的变化规律,各环境间差异较小。除白牛槭外,其他树种幼树叶绿素归一化指数(Chl NDI)随环境光强的减少而增大,林下树种的光化学反射指数(PRI)普遍高于光条件更好环境下树种,说明林下幼树叶片叶绿素含量和光合速率大于其他光环境;各树种植物叶片水分指数(WI)在不同光环境之间存在显著差异,环境光强越大植物叶片WI越小,植物叶片水分浓度越小。林缘下3种槭属幼树结构不敏感植被光谱指数(SIPI)显著小于其他光环境(P<0.05)...     

Lead dynamics in the forest floor and mineral soil in south-central Ontario

Recent studies have suggested that the residence time of Pb in the forest floor may not be as long as previously thought, and there is concern that the large pulse of atmospheric Pb deposited in the 1960s and early 1970s may move rapidly through mineral soils and eventually contaminate groundwater. In order to assess Pb mobility at a woodland (JMOEC) in south-central Ontario, a stable Pb isotope tracer ²⁰⁷Pb (8?mg?m^?2) was added to the forest floor in white pine (Pinus strobus) and sugar maple (Acer saccharum) stands, respectively, and monitored over a 2-year period. Excess ²⁰⁷Pb was rapidly lost from the forest floor. Applying first-order rate coefficients (k) of 0.57 (maple) and 0.32 (pine) obtained from the tracer study, and estimates of Pb deposition in the region, current predicted Pb concentrations in the forest floor are 1.5–3.1 and 2.1–5.8?mg?kg^?1 in the maple and pine plots, respectively. These values compare favorably with measured concentrations (corrected for mineral soil contamination) of 3.1–4.3?mg?kg^?1 in the maple stand and 2.6–3.6?mg?kg^?1 in the pine stand. The response time (1/k) of Pb in the forest floor at the sugar maple and white pine plots was estimated to be 1.8 and 3.1 years, respectively. The rapid loss of Pb from the forest floor at the JMOEC is much greater than previously reported, and is probably due to the rapid rate of litter turnover that is characteristic of forests with mull-type forest floors. In a survey of 23 forested sites that border the Precambrian Shield in south-central Ontario, Pb concentrations in the forest floor increased exponentially with decreasing soil pH. Lead concentrations in the forest floor at the most acidic survey sites, which exhibited mor-type forest floors, were approximately 10 times higher (～80?mg?kg^?1) than at the JMOEC, and pollution Pb burdens were up to 25 times greater. Despite the rapid loss of Pb from the forest floor at the JMOEC, the highest pollution Pb concentrations were found in the upper (0–1?cm) mineral soil horizon. Lead concentrations in the upper 30?cm of mineral soil were strongly correlated with organic matter content, indicating that pollution Pb does not move as a pulse down the soil profile, but instead is linked with organic matter distribution, indicating groundwater contamination is unlikely.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

Do Earthworms Have a Greater Influence on Nitrogen Dynamics Than Atmospheric Nitrogen Deposition?

Elevated levels of inorganic nitrogen (N) deposition and earthworm invasion have the potential to alter N dynamics in eastern North American temperate forests. A regional comparison was conducted across 21 sugar maple (Acer saccharum Marsh) stands in southern Ontario, where forest floor C:N ratios ranged from 17 to 38 showed that, similar to many other studies, rates of potential net mineralization and nitrification increased below a forest floor C:N ratio threshold of approximately 25 and that nitrification rates are positively correlated with foliar N concentration. However, detailed measurements at four representative stands, receiving between 9.8 and 19 kg N ha^?1 y^?1 in throughfall, showed that foliar N levels were highest at the site with the lowest N deposition. The primary difference amongst these sites was the presence of invasive earthworms. Specifically, sites without earthworms had significantly higher forest floor N with a lower C:N ratio than the sites with earthworms. There was no significant difference in the rate of sugar maple litter decomposition or chemistry amongst the sites assessed after 540 days using fine (2-mm mesh) litter bags, suggesting that differences in forest floor N levels were most likely due to consumption of litter by large earthworm species and that the lower C:N ratio of the forest floor in sites without earthworms is brought about primarily by a much longer residence time. This work supports the conclusions that forest floor N concentration (or C:N ratio) has a very strong control on N dynamics in forests, but shows that the presence of earthworms can have an impact on forest floor C:N ratio and hence N dynamics that is greater than current levels of atmospheric inorganic N deposition in temperate forests of Ontario.