Leaf herbivory and decomposability in a Malaysian tropical rain forest

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.     

Predicting leaf physiology from simple plant and climate attributes: a global GLOPNET analysis.

Knowledge of leaf chemistry, physiology, and life span is essential for global vegetation modeling, but such data are scarce or lacking for some regions, especially in developing countries. Here we use data from 2021 species at 175 sites around the world from the GLOPNET compilation to show that key physiological traits that are difficult to measure (such as photosynthetic capacity) can be predicted from simple qualitative plant characteristics, climate information, easily measured ("soft") leaf traits, or all of these in combination. The qualitative plant functional type (PFT) attributes examined are phylogeny (angiosperm or gymnosperm), growth form (grass, herb, shrub, or tree), and leaf phenology (deciduous vs. evergreen). These three PFT attributes explain between one-third and two-thirds of the variation in each of five quantitative leaf ecophysiological traits: specific leaf area (SLA), leaf life span, mass-based net photosynthetic capacity (Amass), nitrogen content (N(mass)), and phosphorus content (P(mass)). Alternatively, the combination of four simple, widely available climate metrics (mean annual temperature, mean annual precipitation, mean vapor pressure deficit, and solar irradiance) explain only 5-20% of the variation in those same five leaf traits. Adding the climate metrics to the qualitative PFTs as independent factors in the model increases explanatory power by 3-11% for the five traits. If a single easily measured leaf trait (SLA) is also included in the model along with qualitative plant traits and climate metrics, an additional 5-25% of the variation in the other four other leaf traits is explained, with the models accounting for 62%, 65%, 66%, and 73% of global variation in N(mass), P(mass), A(mass), and leaf life span, respectively. Given the wide availability of the summary climate data and qualitative PFT data used in these analyses, they could be used to explain roughly half of global variation in the less accessible leaf traits (A(mass), leaf life span, N(mass), P(mass)); this can be augmented to two-thirds of all variation if climatic and PFT data are used in combination with the readily measured trait SLA. This shows encouraging possibilities of progress in developing general predictive equations for macro-ecology, global scaling, and global modeling.     

Leaf traits are good predictors of plant performance across 53 rain forest species

We compared the leaf traits and plant performance of 53 co-occurring tree species in a semi-evergreen tropical moist forest community. The species differed in all leaf traits analyzed: leaf life span varied 11-fold among species, specific leaf area 5-fold, mass-based nitrogen 3-fold, mass-based assimilation rate 13-fold, mass-based respiration rate 15-fold, stomatal conductance 8-fold, and photosynthetic water use efficiency 4-fold. Photosynthetic traits were strongly coordinated, and specific leaf area predicted mass-based rates of assimilation and respiration; leaf life span predicted many other leaf characteristics. Leaf traits were closely associated with growth, survival, and light requirement of the species. Leaf investment strategies varied on a continuum trading off short-term carbon gain against long-term leaf persistence that, in turn, is linked to variation in whole-plant growth and survival. Leaf traits were good predictors of plant performance, both in gaps and in the forest understory. High growth in gaps is promoted by cheap, short-lived, and physiologically active leaves. High survival in the forest understory is enhanced by the formation of long-lived well protected leaves that reduce biomass loss by herbivory, mechanical disturbance, or leaf turnover. Leaf traits underlay this growth-survival trade-off; species with short-lived, physiologically active leaves have high growth but low survival. This continuum in leaf traits, through its effect on plant performance, in turn gives rise to a continuum in species' light requirements.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

Leaf structural diversity is related to hydraulic capacity in tropical rain forest trees

The hydraulic resistance of the leaf (R1) is a major bottleneck in the whole plant water transport pathway and may thus be linked with the enormous variation in leaf structure and function among tropical rain forest trees. A previous study found that R1 varied by an order of magnitude across 10 tree species of Panamanian tropical lowland rain forest. Here, correlations were tested between R1 and 24 traits relating to leaf venation and mesophyll structure, and to gross leaf form. Across species, R1 was related to both venation architecture and mesophyll structure. R1 was positively related to the theoretical axial resistivity of the midrib, determined from xylem conduit numbers and dimensions, and R1 was negatively related to venation density in nine of 10 species. R1 was also negatively related to both palisade mesophyll thickness and to the ratio of palisade to spongy mesophyll. By contrast, numerous leaf traits were independent of R1, including area, shape, thickness, and density, demonstrating that leaves can be diverse in gross structure without intrinsic trade-offs in hydraulic capacity. Variation in both R1-linked and R1-independent traits related strongly to regeneration irradiance, indicating the potential importance of both types of traits in establishment ecology.     

Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.     

Trait differences between naturalized and invasive plant species independent of residence time and phylogeny

The ability to predict which alien plants will transition from naturalized to invasive prior to their introduction to novel regions is a key goal for conservation and has the potential to increase the efficacy of weed risk assessment (WRA). However, multiple factors contribute to plant invasion success (e.g., functional traits, range characteristics, residence time, phylogeny), and they all must be taken into account simultaneously in order to identify meaningful correlates of invasion success. We compiled 146 pairs of phylogenetically paired (congeneric) naturalized and invasive plant species in Australia with similar minimum residence times (i.e., time since introduction in years). These pairs were used to test for differences in 5 functional traits (flowering duration, leaf size, maximum height, specific leaf area [SLA], seed mass) and 3 characteristics of species’ native ranges (biome occupancy, mean annual temperature, and rainfall breadth) between naturalized and invasive species. Invasive species, on average, had larger SLA, longer flowering periods, and were taller than their congeneric naturalized relatives. Invaders also exhibited greater tolerance for different environmental conditions in the native range, where they occupied more biomes and a wider breadth of rainfall and temperature conditions than naturalized congeners. However, neither seed mass nor leaf size differed between pairs of naturalized and invasive species. A key finding was the role of SLA in distinguishing between naturalized and invasive pairs. Species with high SLA values were typically associated with faster growth rates, more rapid turnover of leaf material, and shorter lifespans than those species with low SLA. This suite of characteristics may contribute to the ability of a species to transition from naturalized to invasive across a wide range of environmental contexts and disturbance regimes. Our findings will help in the refinement of WRA protocols, and we advocate the inclusion of quantitative traits, in particular SLA, into the WRA schemes.     

Ecology of hemiepiphytism in fig species is based on evolutionary correlation of hydraulics and carbon economy

Woody hemiepiphytic species (Hs) are important components of tropical rain forests, and they have been hypothesized to differ from non-hemiepiphytic tree species (NHs) in adaptations relating to water relations and carbon economy; but few studies have been conducted comparing ecophysiological traits between the two growth forms especially in an evolutionary context. Using common-garden plants of the genus Ficus, functional traits related to plant hydraulics and carbon economy were compared for seven NHs and seven Hs in their adult terrestrial "tree-like" growth phase. We used phylogenetically independent contrasts to test the hypothesis that differences in water availability selected for contrasting suites of traits in Hs and NHs, driving evolutionary correlations among functional traits including hydraulic conductivity and photosynthetic traits. Species of the two growth forms differed in functional traits; Hs had substantially lower xylem hydraulic conductivity and stomatal conductance, and higher instantaneous photosynthetic water use efficiency. Leaf morphological and structural traits also differed strikingly between the two growth forms. The Hs had significantly smaller leaves, higher leaf mass per area (LMA), and smaller xylem vessel lumen diameters. Across all the species, hydraulic conductivity was positively correlated with leaf gas exchange indicating high degrees of hydraulic-photosynthetic coordination. More importantly, these correlations were supported by correlations implemented on phylogenetic independent contrasts, suggesting that most trait correlations arose through repeated convergent evolution rather than as a result of chance events in the deep nodes of the lineage. Vatiation in xylem hydraulic conductivity was also centrally associated with a suite of other functional traits related to carbon economy and growth, such as LMA, water use efficiency, leaf nutrient concentration, and photosynthetic nutrient use efficiency, indicating important physiological constraints or trade-offs among functional traits. Shifts in this trait cluster apparently related to the adaptation to drought-prone canopy growth during the early life cycle of Hs and clearly affected ecophysiology of the later terrestrial stage of these species. Evolutionary flexibility in hydraulics and associated traits might be one basis for the hyper-diversification of Ficus species in tropical rain forests.     

Multivariate identification of plant functional response and effect traits in an agricultural landscape

Plant functional traits have been proposed as a linkage between the environmental control of vegetation and ecosystem function. Identification of traits that mediate the response of plant species to the environment is well established, but the identification of effect traits and the linkage between the two sets is less developed. This was attempted for a study of eight contrasting land uses in a marginal agricultural landscape where data on vegetation, management controls of the disturbance regime, and soil characteristics, including nitrogen release, were measured simultaneously with measures of ecosystem function such as litter decomposition rates and primary productivity on 30 sites. Trait data were assembled from databases, and an iterative multivariate approach using the three table (species, trait, environment) method RLQ was employed to identify a parsimonious set of traits that predict plant species responses to the environment and a parsimonious set of traits that link vegetation to ecosystem function. The lists of response and effect traits were similar, and where differences were observed, traits were usually highly correlated with at least one trait in the other list. This approach identified a small number of traits (canopy height, leaf dry matter content, leaf size, and specific leaf area) that provide a means of linking vegetation responses to environmental change with changes in ecosystem function. Other response traits included vegetative spread strategy, start of flowering, and seed terminal velocity, but within the system studied these traits were all significantly correlated to the traits shared between the response and effect lists.     

Leaf habit and woodiness regulate different leaf economy traits at a given nutrient supply

The large variation in the relationships between environmental factors and plant traits observed in natural communities exemplifies the alternative solutions that plants have developed in response to the same environmental limitations. Qualitative attributes, such as growth form, woodiness, and leaf habit can be used to approximate these alternative solutions. Here, we quantified the extent to which these attributes affect leaf trait values at a given resource supply level, using measured plant traits from 105 different species (254 observations) distributed across 50 sites in mesic to wet plant communities in The Netherlands. For each site, soil total N, soil total P, and water supply estimates were obtained by field measurements and modeling. Effects of growth forms, woodiness, and leaf habit on relations between leaf traits (SLA, specific leaf area; LNC, leaf nitrogen concentration; and LPC, leaf phosphorus concentration) vs. nutrient and water supply were quantified using maximum-likelihood methods and Bonferroni post hoc tests. The qualitative attributes explained 8-23% of the variance within sites in leaf traits vs. soil fertility relationships, and therefore they can potentially be used to make better predictions of global patterns of leaf traits in relation to nutrient supply. However, at a given soil fertility, the strength of the effect of each qualitative attribute was not the same for all leaf traits. These differences may imply a differential regulation of the leaf economy traits at a given nutrient supply, in which SLA and LPC seem to be regulated in accordance to changes in plant size and architecture while LNC seems to be primarily regulated at the leaf level by factors related to leaf longevity.     

Decomposition of diverse litter mixtures in streams

In view of growing interest in understanding how biodiversity affects ecosystem functioning, we investigated effects of riparian plant diversity on litter decomposition in forest streams. Leaf litter from 10 deciduous tree species was collected during natural leaf fall at two locations (Massif Central in France and Carpathians in Romania) and exposed in the field in litter bags. There were 35 species combinations, with species richness ranging 1-10. Nonadditive effects on the decomposition of mixed-species litter were minor, although a small synergistic effect was observed in the Massif Central stream where observed litter mass remaining was significantly lower overall than expected from data on single-species litter. In addition, variability in litter mass remaining decreased with litter diversity at both locations. Mean nitrogen concentration of single- and mixed-species litters (0.68-4.47% of litter ash-free dry mass) accounted for a large part of the variation in litter mass loss across species combinations. For a given species or mixture, litter mass loss was also consistently faster in the Massif Central than in the Carpathians, and the similarity in general stream characteristics, other than temperature, suggests that this effect was largely due to differences in thermal regimes. These results support the notion that decomposition of litter mixtures is primarily driven by litter quality and environmental factors, rather than by species richness per se. However, the observed consistent decrease in variability of decomposition rate with increasing plant species richness indicates that conservation of riparian tree diversity is important even when decomposition rates are not greatly influenced by litter mixing.     

杉木林凋落物产量、分解率与储量的关系

在福建三明采用2种方法测定27a生杉木林的杉木叶和小枝凋落物的分解率．采用网袋法(孔径0．5mm尼龙网袋)测定的杉木叶和小枝凋落物残留率Olson指数模型的分解系数(k值)分别为0．7692a^-1和0．2501a^-1，相应的第1年的分解率分别为53．66％和22．13％；另一种测定矗值的方法较为准确，是通过计算凋落物年产量与地面凋落物储量的比值得到，采用该方法计算的杉木叶和小枝凋落物的矗值分别为1．788a^-1和0．8622a^-1，分别是网袋法测定值的2．32和3．54倍，相应的第1年的分解率分别为83．27％和57．78％，分别是网袋法测定值的1．55倍和2．61倍．采用后一种方法测定的凋落物分解率可以解决尼龙网袋法测定分解速率结果偏低的问题．     

Ecological stoichiometric characteristics of carbon,nitrogen, and phosphorus in three forests in the Lüliang Mountainous Area of Shanxi Province北大核心CSCD

Using a standard plot method, the stoichiometry of carbon (C), nitrogen (N), and phosphorus (P) in leaves, litter, and soil (0-20 cm depth) was investigated for three forest types: Populus davidiana, Larix principis-rupprechtii, and Pinus tabuliformis. The results showed that the stoichiometry of C, N, and P of the same component in the three forests were significantly different. The C and N contents in leaves, litter, and soil in P. davidiana forest were higher than those in L. principis-rupprechtii and P. tabuliformis forests were. However, P in the L. principis-rupprechtii forest was higher than that in the P. davidiana forest and P. tabuliformis forests were. The C, N, and P contents of the components in the three forests were, in order, leaves > litter > soil, and the three nutrient contents were significantly higher in leaves and litter than they were in soil. C:N and C:P in the three forests exhibited a trend of litter > leaves > soil, whereas that for N:P was leaves > soil > litter. There were highly significant positive relationships in N:P between the litter and the soil in the P. davidiana forest. Leaf C:N and litter C:P in the L. principis-rupprechtii forest were significantly negatively correlated, whereas N:P in the leaves and soil was positively correlated. There was a significant positive correlation in N:P between the leaves and the soil in the P. tabuliformis forest. In conclusion, the N contents in leaves and soil exhibited a significant positive correlation, whereas there was no significant correlation between C, N, and P in litter or soil. Environmental factors had a large influence on the stoichiometry of C, N, and P in soil. In particular, latitude and altitude had the most significant effects on C, N, P, C:N, and C:P and were significantly p ositively correlated. T hese results provide a scientific basis for f urther studies on nutrient utilization a nd t he cyclic characteristics of different forests in this area. © 2018 Science Press. All rights reserved.     

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.     

Effects of initial decomposing leaf litter of Cinnamomum camphora on the growth and physiology of Impatiens balsamina北大核心CSCD

A pot experiment was conducted to study the allelopathic effects of initial decomposing leaf litter of Cinnamomum camphora on growth and physiology of Impatiens balsamina. Three leaf litter treatments included 20, 40 and 80 g of C. camphora leaf litter mixed with 8 kg of soil, namely T1, T2, and T3, respectively. In order to test the effect of leaf litter addition on the permeability and ventilation of soil simultaneously, a parallel trial with steamed leaf litter was conducted with the three treatments of the leaf litter. The leaf litter was steamed for 2 d to remove the secondary metabolites as much as possible, dried, and then mixed with 8 kg of soil, namely Z1, Z2, and Z3, respectively. No leaf litter was added in control (CK). The growth parameters of I. balsamina were determined at the 20 d, 60 d, 100 d and 120 d after sowing and the main physiology indicators were determined at the 60 d. The results indicated that: (1) The ground surface diameter and height of I. balsamina were inhibited significantly at 60 d (P < 0.05). Photosynthetic pigments and gas exchange parameters of I. balsamina were inhibited significantly at 60 d, and the inhibition effect was stronger with increased amount of leaf litter addition. The chlorophyll content, Pn and Ls decreased significantly with increased amount of leaf litter (P < 0.05). The activity of superoxide dismutase (SOD) and peroxidase (POD) in leaves of I. balsamina decreased with the increase of leaf litter addition. The content of MDA in treatments T1, T2 and T3 were all higher than that in CK, which indicated that I. balsamina suffered oxidative damage in a certain degree. The content of free proline (Pro) and soluble sugar (SS) in leaves of I. balsamina decreased significantly with the increase of the leaf litter (P < 0.05), while the content of soluble protein (SP) increased. (2) In the parallel trial, 60 d after sowing, no obvious difference was observed between CK and any steamed leaf litter treatment in terms of the morphological and physiological features stated above (P > 0.05). It indicated that the soil physical properties were not greatly influenced by leaf litter addition in the dose interval designed, or that the release of secondary metabolites from decomposing leaf litter was probably a better reason to explain the inhibition of leaf litter treatment to I. balsamina growth. (3)The compound effect (CE) of leaf litter decomposition on I. balsamina was enhanced with increase of the leaf litter, to 0.169, 0.354, and 0.497, respectively, in treatments of T1, T2 and T3. The study indicated that initial decomposition of C. camphora leaf litter in soil reduces the content of photosynthetic pigments, inhibits photosynthetic capacity and resistance physiology of I. balsamina, weakens its adaptability to the environment, and restrains growth of the plant. © 2015, Science Press. All rights reserved.     

Seasonal variations in plant species effects on soil N and P dynamics

It is well established that plant species influence ecosystem processes, but we have little ability to predict which vegetation changes will alter ecosystems, or how the effects of a given species might vary seasonally. We established monocultures of eight plant species in a California grassland in order to determine the plant traits that account for species impacts on nitrogen and phosphorus cycling. Plant species differed in their effects on net N mineralization and nitrification rates, and the patterns of species differences varied seasonally. Soil PO4- and microbial P were more strongly affected by slope position than by species. Although most studies focus on litter chemistry as the main determinant of plant species effects on nutrient cycling, this study showed that plant species affected biogeochemical cycling through many traits, including direct traits (litter chemistry and biomass, live-tissue chemistry and biomass) and indirect traits (plant modification of soil bioavailable C and soil microclimate). In fact, species significantly altered N and P cycling even without litter inputs. It became particularly critical to consider the effects of these multiple traits in order to account for seasonal changes in plant species effects on ecosystems. For example, species effects on potential rates of net N mineralization were most strongly influenced by soil bioavailable C in the fall and by litter chemistry in the winter and spring. Under field conditions, species effects on soil microclimate influenced rates of mineralization and nitrification, with species effects on soil temperature being critical in the fall and species effects on soil moisture being important in the dry spring. Overall, this study clearly demonstrated that in order to gain a mechanistic, predictive understanding of plant species effects on ecosystems, it is critical to look beyond plant litter chemistry and to incorporate the effects of multiple plant traits on ecosystems.     

Modeling leaf dispersal in mixed hardwood forests using a ballistic approach

In mixed-species stands, modeling leaf litter dispersal is important to predict the physical and chemical characteristics of the forest floor, which plays a major role in nutrient cycling and in plant population dynamics. In this study, a spatially explicit model of leaf litterfall was developed and compared with two other models. These three models were calibrated for a mixed forest of oak and beech using litterfall data from mapped forest plots. All models assumed that an allometric equation described individual leaf litter production, but they strongly differed in the modeling of the probability density of leaf shedding with distance from source trees. Two models used a negative exponential function to account for leaf dispersal with distance, and this function was allowed to vary according to wind direction in one of them. In contrast, our approach was based on a simple ballistic equation considering release height, wind speed, wind direction, and leaf fall velocity; the distributions of wind speeds and wind directions were modeled according to a Weibull and a Von Mises distribution, respectively. Using an independent validation data set, all three models provided predictions well correlated to measurements (r > 0.83); however, the two models with a direction-dependent component were slightly more accurate. In addition, parameter estimates of the ballistic model were in close agreement with a foliar litter production equation derived from the literature for beech and with wind characteristics measured during leaf litterfall for both species. Because of its mechanistic background, such a spatially explicit model might be incorporated as a litterfall module in larger models (nutrient cycling, plant population dynamics) or used to determine the manner in which patch size in mixed-species stands influences litter mixture.     

Nematode assemblages from <Emphasis Type="Italic">Avicennia</Emphasis><Emphasis Type="Italic">marina</Emphasis> leaf litter in a temperate mangrove forest in south-eastern Australia

Meiofauna from Avicennia marina leaf litter in a temperate mangrove forest was enumerated, and the nematode assemblages compared on the bases of leaf colour (used as a guide to leaf age) and shore horizon where samples were collected. Twenty-one putative nematode species were collected from 48 leaf litter samples. Univariate analyses indicated that neither the colour of the leaf nor the shore horizon significantly affected abundance of nematodes. However, of the four (2Ƕ) treatment groups, rarefaction curves revealed highest diversity on brown leaves from under the shade of the tree canopy (H'=0.751ǂ.126 SE, n=17). Species diversity of leaf litter nematodes was lower in this temperate mangrove system than reported from tropical mangrove studies. ANOSIM tests confirmed a significant effect of shore horizon on nematode assemblages. The dominant feeding group among nematodes was non-selective deposit feeders (7/21 species, but 77% of all nematodes). Epigrowth grazers were represented by 8/21 species of nematodes, but only 19% of the total number. Excised leaves became skeletonised by about 15 weeks. Shorter temporal scales of life cycles of nematodes compared with leaf degradation, and the dynamic nature of epibiontic assemblages, probably explain the similar assemblage structure on yellow and brown leaves.     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.