Local-scale determinants of elemental stoichiometry of soil in an old-growth temperate forest

Plant and Soil - Determining which strategies confer a competitive advantage in variable environments with strong short-term resource fluctuations (i.e., seasonal drought and flooding) is critical...     

Apparent climatically induced increase of tree mortality rates in a temperate forest

We provide a first detailed analysis of long-term, annual-resolution demographic trends in a temperate forest. After tracking the fates of 21 338 trees in a network of old-growth forest plots in the Sierra Nevada of California, we found that mortality rate, but not the recruitment rate, increased significantly over the 22 years of measurement (1983–2004). Mortality rates increased in both of two dominant taxonomic groups ( Abies and Pinus ) and in different forest types (different elevational zones). The increase in overall mortality rate resulted from an increase in tree deaths attributed to stress and biotic causes, and coincided with a temperature-driven increase in an index of drought. Our findings suggest that these forests (and by implication, other water-limited forests) may be sensitive to temperature-driven drought stress, and may be poised for die-back if future climates continue to feature rising temperatures without compensating increases in precipitation.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

Influence of micro-landforms on forest structure,tree death and recruitment in a Japanese temperate mixed forest

The micro-landform unit system offers an effective way of analyzing vegetation–geomorphology relationships at a 10-m scale in areas such as the hilly regions of Japan. We analyzed relationships between micro-landforms and tree population parameters over a 9-year interval to elucidate the influence of geomorphic processes on vegetation dynamics. A 2.16-ha permanent plot was established in a temperate mixed forest. Each 5m×5m quadrat within this plot was classified according to six types of micro-landform units: (i) crest slope (CS); (ii) upper sideslope (US); (iii) head hollow (HH); (iv) lower sideslope (LS); (v) foot slope (FS); and (vi) river bed (RB). All living trees larger than 10cm in diameter at breast height (d.b.h.) were identified, mapped and marked in 1989 and resurveyed in 1998. Almost all of the 23 common tree species persisted in their own core habitats (i.e. the micro-landforms) between the two surveys. The species distribution in both surveys showed that the six micro-landforms could be combined into two larger groups: upper and lower hillslope areas. The upper hillslope area had higher tree densities and larger basal areas than the lower hillslope area. It is possible that these differences result from the longer lifespans of trees on the upper hillslope area rather than from differences in mortality and recruitment rates. In addition, the different ways in which trees die in the different micro-landform units may affect the regeneration process in hilly regions through different gap formation. The effects of different geomorphic processes are reflected in the lifespans of the trees and may result in different forest structure and dynamics among micro-landform units.     

Determinants of tree survival at local scale in a sub-tropical forest

Tree survival is a critical driver of stand dynamics, influencing forest structure and composition. Many local-scale drivers (tree size, abiotic and biotic factors) have been proposed as being important in explaining patterns of tree survival, but their contributions are still unknown. We examined the relative importance of these local drivers on tree survival using generalized linear mixed models in an old-growth sub-tropical forest in south China at three levels (community, guild, and species). Among the variables tested, tree size was typically the most important driver of tree survival, followed by abiotic and then biotic variables. Tree size has a strongly positive effect on tree survival for small trees (10–30 cm dbh) and shade-tolerant tree species. Of the abiotic factors tested, elevation tended to be more important in affecting tree survival than other topographic variables. Abiotic factors generally influenced survival of species with relatively high abundances, for individuals in smaller size classes and for mid-tolerant species. Among biotic factors, we found that the mortality of tree species was not driven by density- and frequency-dependent effects in this sub-tropical forest, as indicated by the results of both total basal area of neighbors and the proportion of conspecific neighbors in our study. We conclude that the relative importance of variables driving patterns of tree survival varied greatly among tree size classes, species guilds, shade tolerance, density, and abundance classes in this sub-tropical forest. These results also provide critical information for future studies of forest dynamics and offer insight into forest management in this region.     

The effect of tree species diversity on fine-root production in a young temperate forest

The phenomenon of overyielding in species-diverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5–6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0–30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fine-root production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0–15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15–30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fine-root production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.     

Photographic assessment of temperate forest understory phenology in relation to springtime meteorological drivers

Phenology shows sensitive responses to seasonal changes in atmospheric conditions. Forest understory phenology, in particular, is a crucial component of the forest ecosystem that interacts with meteorological factors, and ecosystem functions such as carbon exchange and nutrient cycling. Quantifying understory phenology is challenging due to the multiplicity of species and heterogeneous spatial distribution. The use of digital photography for assessing forest understory phenology was systematically tested in this study within a temperate forest during spring 2007. Five phenology metrics (phenometrics) were extracted from digital photos using three band algebra and two greenness percentage (image binarization) methods. Phenometrics were compared with a comprehensive suite of concurrent meteorological variables. Results show that greenness percentage cover approaches were relatively robust in capturing forest understory green-up. Derived spring phenology of understory plants responded to accumulated air temperature as anticipated, and with day-to-day changes strongly affected by estimated moisture availability. This study suggests that visible-light photographic assessment is useful for efficient forest understory phenology monitoring and allows more comprehensive data collection in support of ecosystem/land surface models.     

Drought enhances symbiotic dinitrogen fixation and competitive ability of a temperate forest tree

General circulation models project more intense and frequent droughts over the next century, but many questions remain about how terrestrial ecosystems will respond. Of particular importance, is to understand how drought will alter the species composition of regenerating temperate forests wherein symbiotic dinitrogen (N₂)-fixing plants play a critical role. In experimental mesocosms we manipulated soil moisture to study the effect of drought on the physiology, growth and competitive interactions of four co-occurring North American tree species, one of which (Robinia pseudoacacia) is a symbiotic N₂-fixer. We hypothesized that drought would reduce growth by decreasing stomatal conductance, hydraulic conductance and increasing the water use efficiency of species with larger diameter xylem vessel elements (Quercus rubra, R. pseudoacacia) relative to those with smaller elements (Acer rubrum and Liriodendron tulipifera). We further hypothesized that N₂ fixation by R. pseudoacacia would decline with drought, reducing its competitive ability. Under drought, growth declined across all species; but, growth and physiological responses did not correspond to species’ hydraulic architecture. Drought triggered an 80 % increase in nodule biomass and N accrual for R. pseudoacacia, improving its growth relative to other species. These results suggest that drought intensified soil N deficiency and that R. pseudoacacia’s ability to fix N₂ facilitated competition with non-fixing species when both water and N were limiting. Under scenarios of moderate drought, N₂ fixation may alleviate the N constraints resulting from low soil moisture and improve competitive ability of N₂-fixing species, and as a result, supply more new N to the ecosystem.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

Comparison of lepidopteran larval communities among tree species in a temperate deciduous forest, Japan

Abstract.  1. The enormous diversity of phytophagous insects in forest canopies is hypothesised to be supported by the number of herbivorous species per host tree species or host specificity. It is therefore necessary to examine the effect of host plant species on compositional changes in the herbivore communities.
2. The lepidopteran larval communities were examined in the canopies of 10 tree species in a temperate deciduous forest of Japan. The phylogeny and leaf flush phenology of host plant species were taken into account as factors affecting the herbivore community assembly.
3. Examination of seasonal changes in the larval community structures on each tree species showed that larval species richness, abundance, and evenness decreased significantly from spring to summer. Larval species richness and abundance were characterised by family-level phylogenetic differences among tree species, whereas evenness was determined at a higher taxonomic level.
4. Compositional changes in the larval communities among tree species showed a remarkable pattern, with a phylogenetic effect at a high taxonomic level in spring, similar to evenness, but a phenological effect in summer. This suggests that host specificity could support the lepidopteran larval diversity in spring.
5. These results suggest that the differences in host utilisation of the herbivore, which reflects the phylogenetic effect of the host plants, can be important as a factor affecting the diversity of lepidopteran larval communities in temperate forests.     

Heterogeneous dispersal of tree litterfall corresponding with patchy canopy structure in a temperate mixed forest

Spatial dispersal patterns of fine litter were analyzed in relation to forest structure within a temperate mixed stand, in northeastern Japan. The canopy layer was a mosaic of two types of patches, those of shade-tolerant Abies firma (estimated age being 81–146 yrs.) and early successional deciduous broad-leaved trees (about 45 yrs. of age). Litter input to the forest floor was observed for a period of three years using 38 traps set at a height of 1.2 m. The average annual total of fine litter was 6.233 (dw.) metric tons hectare^-1, 85.7% of the total litterfall. Fine litter was mostly leaves from the overstory (about 64%). Litter distribution maps were drawn, using the Davis's grid method, of the total fine litter and its two main elements, namely A. firma (a 36.1% share) and deciduous broad-leaved species (36.7%). The mosaic distribution of heterogeneous patches of litter was seem to corresponded with distribution of the canopy structure.     

Environmental correlates of tree species distributions vary among age classes in a northern temperate forest

Processes involved in the structuring of forest communities include: (1) ecological sorting, where species poorly suited to local conditions are subject to environmental filtering and competitive displacement; (2) disturbance, resulting in stochastic removal of individuals and reinitiating successional regimes and (3) dispersal limitation, inhibiting the infiltration of species into preferred sites. Temporal dynamics in these processes lead to difficulty inferring causal landscape?Cbiota correlations. Complicating factors include potential for ontogenetic variation in habitat preferences among age classes, and inherent ambiguity regarding severity and coverage of historical disturbance events. Sorting species into age groups can provide relevant temporal information. Fundy National Park is a northern, mixed-temperate forest in Atlantic Canada (Acadian forest type), which was pervasively altered upon European settlement. Species frequency data for three tree age classes (saplings, juveniles and adults) in permanent sample plots (400?m², n?=?33) were compared to environmental data, including soil chemistry, understory light conditions, physiography and disturbance history using ordination and randomization techniques. Abiotic and disturbance-related predictors of species distributions differed among life stages. Specifically, in the adult stage, stand age was a critical predictor of distribution, whereas in younger age classes environmental variables such as nutrient availability and soil moisture and drainage were key drivers of distribution. It is concluded that older populations were increasingly less constrained by environmental conditions, suggesting that adult populations bear the legacy of stochastic landscape alteration, thus appearing randomly distributed along environmental gradients. As younger populations gradually expand in distribution, they are filtered into preferred conditions by ecological sorting. These findings indicate the importance of considering age-class effects and site history in further assessments of interactions between landscapes and flora.     

Indirect facilitation by a liana might explain the dominance of a small tree in a temperate forest

Aims Lianas are expected to influence composition, structure and functioning of forest systems due to unequal distribution across the potential set of host plants. However, our understanding of mechanisms associated with preferences for specific hosts is still limited, and so is our ability to discern between endogenous and exogenous forces driving forest dynamics in the long run. In this paper, we evaluated whether the dominant liana Hedera helix can indirectly contribute to the eventual dominance of the small multi-stemmed tree Corylus avellana in a remnant temperate forest in central Iberian Peninsula from comparatively reduced liana infestation on C. avellana relative to co-occurring woody species.Methods Through principal component analysis and co-occurrence analysis, we studied the distribution and spatial association between woody species and the liana H. helix. We analyzed the relationship between the number of species in a plot and the number of species infested by the liana to test the hypothesis that H. helix is a generalist liana. Through generalized linear mixed models, we tested the dynamic-multi-stemmed growth form of C. avellana as a plausible life strategy to withstand, in the long run, the liana infestation. In particular, we tested (i) the relationship between stem size and the probability of H. helix infestation including all the tree species within plots and (ii) the relationship between stem size and mortality as evidence of the stem turn over in the tree C. avellana .Important findings Our results indicate that H. helix and C. avellana significantly co-occur in mature stands of this remnant temperate forest where pioneer woody species are absent. Hedera helix severely infests all the woody species whenever stem size exceed ≈25 cm perimeter and there is physical contact at the base of the stem. This implies that all the trees in the community are potential hosts for H. helix. Mixed models indicate that both, infestation by H. helix and stem mortality, are positively related to C. avellana stem perimeter. Reduced long-term infestation of the liana by means of a multi-stemmed growth form with high stem turnover in C. avellana might be an advantage with respect to unipodial tree species. Thus, the liana-tree coexistence pattern may be interpreted as an indirect positive interaction that, contrary to previous findings, results here in species dominance instead of species coexistence.     

六种温带森林类型凋落物量长期动态及其环境驱动

阐明凋落物动态及其环境控制机制,可以为森林生态系统生产力及碳汇功能的维持提供重要的数据支持和理论依据。以长白山系余脉张广才岭西坡林龄相近但立地条件不同的4种天然次生林（即硬阔叶林、杨桦林、杂木林和蒙古栎林）和2种人工林（落叶松人工林和红松人工林）为研究对象,对其地上凋落物产量及其组分以及相关环境因子进行了14年（2008-2021年）的连续测定,旨在揭示森林凋落物量及其组分的时空变化（林型间和年际变异）及其环境驱动机制。结果表明：6种森林类型的凋落总量（TL）无显著差异,波动范围为500.5-556.1 g m^-2 a^-1;但其叶凋落量（LL）、繁殖组织凋落量（RT）和其他组织凋落量（OT）均存在显著差异,波动范围依次分别为333.9-391.8 g m^-2 a^-1、8.43-69.93 g m^-2 a^-1和93.4-185.9 g m^-2 a^-1。6种森林类型的TL均存在显著的年际变化;其中LL和OT年际变化的显著性因森林类型而不同,而RT的年际变化不显著。除落叶松人工林外,其余5种森林类型的LL与生长季平均气温、日最低气温均值、土壤10 cm深度处的平均温度、最低温度（Ts_min）和土壤5 cm含水量（Ms）均呈显著正相关。杂木林、硬阔叶林和红松人工林的RT与Ms呈显著负相关;杂木林、杨桦林和硬阔叶林的OT与Ts_min呈显著负相关。样地水平的LL与土壤10 cm处含水量存在显著的正相关关系,而RT和OT则与其呈现显著负相关关系。这些结果表明林龄相似的温带森林地上凋落物总量有趋同趋势,但其通过改变组分分配格局来适应立地条件的变化;土壤湿度和温度变化会引起凋落物量的年际变化,但不同森林类型的凋落物量对环境波动的敏感性不同。     

Local-scale spatial diversity patterns of ectomycorrhizal fungal communities in a subtropical pine-oak forest

This study aimed to analyze spatial patterns of soil ectomycorrhizal fungal (EMF) communities at the local scale in a subtropical pine-oak forest located in the Nearctic-Neotropical transition in central Mexico, to underpin biodiversity conservation strategies in forest fragments of this region. We used a spatially-explicit nested square sampling design with the same sampling representativeness at all scales and replicated three times. We detected 674 EMF OTUs within 19,200 m² and 65 OTUs on average per sample. Seventy percent of OTUs were detected in only 1–4 samples. Average community similarity was below 5%, showed minor change within 14 and 339 m distance and increased with the spatial grain used to compare the data. We found a high species-area relationship and beta diversity coefficients for soil fungi indicating that, at the local scale, increasing area by a constant factor of four represented an increase in OTU richness by a factor of two.