A comparison of fungal endophytic community diversity in tree leaves of rural and urban temperate forests of Kanto district,eastern Japan

To clarify the effects of forest fragmentation and a change in tree species composition following urbanization on endophytic fungal communities, we isolated fungal endophytes from the foliage of nine tree species in suburban (Kashiwa City, Chiba) and rural (Mt. Wagakuni, Ibaraki; Mt. Takao, Tokyo) forests and compared the fungal communities between sites and host tree species. Host specificity was evaluated using the index of host specificity (Si), and the number of isolated species, total isolation frequency, and the diversity index were calculated. From just one to several host-specific species were recognized in all host tree species at all sites. The total isolation frequency of all fungal species on Quercus myrsinaefolia, Quercus serrata, and Chamaecyparis obtusa and the total isolation frequency of host-specific species on Q. myrsinaefolia, Q. serrata, and Eurya japonica were significantly lower in Kashiwa than in the rural forests. The similarity indices (nonmetric multidimensional scaling (NMS) and C_MH) of endophytic communities among different tree species were higher in Kashiwa, as many tree species shared the same fungal species in the suburban forest. Endophytic fungi with a broad host range were grouped into four clusters suggesting their preference for conifer/broadleaves and evergreen/deciduous trees. Forest fragmentation and isolation by urbanization have been shown to cause the decline of host-specific fungal species and a decrease in β diversity of endophytic communities, i.e., endophytic communities associated with tree leaves in suburban forests were found to be depauperate.     

Pine stumps act as hotspots for seedling regeneration after pine dieback in a mixed natural forest dominated by <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis>

Over the past few decades, rural forest ecosystems in Japan have experienced dynamic vegetation changes due to forest dieback and changes in land use, leading to the loss of local species populations and biodiversity. The aim of this study was to evaluate the importance of pine (Pinus densiflora) stumps and logs for tree seedling regeneration in a mixed natural forest in Kyoto Prefecture, Japan, that had previously experienced severe pine dieback, and to determine which factors most greatly affect seedling establishment. Seedlings of 17 tree species were recorded on pine stumps and logs in later stages of decay, among which Chamaecyparis obtusa and Rhododendron reticulatum were most dominant. Both of these species had a greater density on pine stumps than on logs or soil, despite stumps covering less than 0.5% of the study area. In addition, the seedling densities of both species were positively associated with moss cover on coarse woody debris, but negatively associated with wood pH. Brown rot in the sapwood and heartwood, which occurred more frequently in stumps than in logs, also positively associated with the seedling densities of both species. Predictive modelling showed that C. obtusa seedlings exhibited a stronger response to pH in stumps than in logs. Therefore, since brown-rotted wood is acidic due to fungal decay activities, brown-rotted pine stumps may present hotspots of C. obtusa seedling regeneration at the study site.     

Temporal variation in different life stages of moths (Lepidoptera) from a southwestern temperate forest of Korea: A preliminary study

Two study sites in a temperate forest in Mt. Seungdal-san, Muan, Jeonnam in southwestern Korea were selected for the collection of caterpillars and adult moths (Lepidoptera). Here, we determined (1) the characteristics of moth fauna based on lepidopteran larvae and adults, and (2) the differences in temporal variation between these two developmental stages. In addition, we investigated the life history of moth species collected at the surveyed sites. Of the 206 caterpillars collected, 17 moth species from 9 families were successfully reared. A total of 342 adult moth species and 2108 individuals from 15 families were collected. Twenty-two plants were identified as host plants of 188 caterpillars. Two tree species, Prunus serrulata var. spontanea and Quercus serrata, were widely distributed in the study sites. Leaf water content (%) of Quercus serrata and Carpinus laxiflorawas the highest in June, whereas the leaf water content of Platycarya strobilacea was the highest in August. We postulated that many of the variables in the temperate forest, such as host plant availability, parasitoids, disease, and weather, affected the lepidopteran caterpillar populations. There were differences in species composition among the sampling methods which suggest the need for diverse methods to uncover total biodiversity. However, it should be noted that a long term and well-planned study with efficient rearing of caterpillars is needed to better understand these correlations.     

Community structure of a species-rich temperate forest,Ogawa Forest Reserve,central Japan

Species assembly and niche differentiation were studied, and future species composition was predicted by simple Markov models, in an old-growth deciduous forest at the Ogawa Forest Reserve in central Japan. The dominant species in our 6ha study site are Quercus serrata, Fagus japonica, and F. crenata.An ordination by population parameters revealed four different combination 3 of life forms and regeneration niches. Cluster analysis based on interspecific spatial correlation revealed three groups of species. The species in cluster A, such as F. japonica, occurred at the bottom of the valley, while those in cluster B, such as Q. serrata, occurred along ridges. Species in cluster C, such as F. crenata, did not show any particular habitat preference. Clusters B and C were further divided into three smaller clusters (a-c). Both clusters Ba and Bb included shade intolerant species. Species in cluster Ba had large clump sizes (>1500 m²), reflecting regeneration following large-scale disturbances. Species in cluster Bb had smaller clump sizes (<400 m²) reflecting regeneration following local disturbances. Clusters Ca and Cb mainly included shade tolerant species and shade intolerant species, respectively. Markov models predicted that shade intolerant species, particularly those in cluster Ba, would be eliminated. Thus, species coexist by differentiation of both habitats and regeneration niche in this forest community. Some species such as Quercus serrata, however, regenerate following large-scale disturbances or human activity.     

Distribution of ectomycorrhizal fungi in a Chamaecyparis obtusa stand at different distances from a mature Quercus serrata tree

Ectomycorrhizal fungal distributions at different distances from one mature Quercus serrata tree were demonstrated in a Japanese cypress stand. After 4 months for field-planted and 3 months for laboratory-incubated Q. serrata seedlings, they were evaluated ectomycorrhiza formations. For the field-planted seedlings, molecular discrimination was conducted to clarify the species diversity of ectomycorrhizal fungi. Ectomycorrhiza formation was confirmed at all distances in both methods. However, the formation and associating fungal species were decreased with increasing distance. These results indicate that the density and diversity of ectomycorrhizal fungi decrease rapidly within several tens of meters from arbuscular mycorrhizal forest edges.     

Early seedling growth of pine (Pinus densiflora) and oaks (Quercus serrata,Q. mongolica,Q. variabilis) in response to light intensity and soil moisture

In climatic chambers seed germination and seedling growth of Pinus densiflora Sieb. et Zucc., Quercus serrata Thunb., Quercus mongolica Fisch. ex Turcz. and Quercus variablilis Bl. were investigated as functions of light intensity and soil moisture. In Korea these tree species occur widely and form mixed forests with different distributions. Species clearly differed in the pattern of germination and early seedling growth between light and soil treatments. The germination of pine did not differ between the experimental treatments until the breaking of the primary buds. After that, light intensity was the deciding factor for further development. In the most moist treatment (approx. field capacity) growth of the pine seedlings was strongly inhibited. For the three oak species, differences between experimental treatments first occurred after complete formation of primary leaves. Seed development strongly correlated with the weight of the acorn. The large seeded Q. variabilis (acorns with mean weight of 4.7 g) developed faster and reached larger dimensions towards the end of the experiment than Q. mongolica (2.8 g per acorn) and Q. serrata (0.9 g per acorn). Regarding height and biomass growth, the oak species showed a higher shade tolerance than pine. The proleptic shoot growth was clearly influenced by the light intensity. Root formation was favoured by a high exposure to light. In case of the oak species reduction of soil moisture increased the length of primary roots and the number of secondary roots.     

Comparison of climatic effects on radial growth of evergreen broad-leaved trees at their northern distribution limit and co-dominating deciduous broad-leaved trees and evergreen conifers

Using dendrochronological techniques, this study examined whether tree-ring width of two evergreen broad-leaved species (Cleyera japonica, Eurya japonica) at their inland northern distribution limit in central Japan is more limited by low temperature compared with two co-dominating deciduous broad-leaved species (Fagus japonica, Magnolia hypoleuca) and two evergreen conifer species (Chamaecyparis obtusa, Abies firma), whose distribution limits are further north. The two deciduous broad-leaved species and the two evergreen conifers are tall tree species. Evergreen broad-leaved Cleyera japonica is a sub-canopy species and Eurya japonica is a small tree species. The tree-ring widths of four of the six species (except for Eurya japonica and Magnolia hypoleuca) correlated positively with the March temperature just before the start of the growth period. For deciduous broad-leaved Magnolia hypoleuca, the tree-ring width was correlated positively and negatively with July temperature and precipitation, respectively. However, the other deciduous broad-leaved Fagus japonica showed no such relationships. For the evergreen broad-leaved Cleyera japonica and evergreen conifers Chamaecyparis obtusa and Abies firma, tree-ring widths correlated positively with winter temperatures, probably because evergreen species can assimilate during warm winters. The tree-ring width of Cleyera japonica also correlated positively with temperatures of many months of the growth period. By contrast, the tree-ring width of the other evergreen broad-leaved Eurya japonica showed no positive correlation with the temperature in any month. Most Eurya japonica trees were suppressed by tall trees, which might disguise any climate effect. Thus, there were species differences in response to climate for each life form, and the tree-ring width of Cleyera japonica at the northern distribution limit was more limited by low temperatures compared with co-dominating species. It is suggested that growth of Cleyera japonica is increased by global warming at the latitudinal ecotone.     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Photosynthesis-nitrogen relationship in a Hinoki cypress (Chamaecyparis obtusa) canopy: a comparison with Japanese cedar (Cryptomeria japonica)

The relationship between light-saturated photosynthetic capacity (P _max) and leaf nitrogen (N) content was investigated for one year in a 15-year-old Chamaecyparis obtusa canopy and was compared with a Cryptomeria japonica canopy previously described. The linear regression between P _max and leaf N content tended to converge toward a single line segment from July to January and in May for C. obtusa. The slope of the linear regression between P _max and leaf N content of C. obtusa was gentler than that of C. japonica. The smaller regression coefficient of C. obtusa may reflect species differences in nitrogen nutrition requirements between C. obtusa and C. japonica. A pronounced decrease in the slope of the linear regression lines due to low temperature was observed in February and March. During this period, P _max of C. obtusa declined more than that of C. japonica suggesting that C. obtusa is less tolerant to low temperatures than C. japonica.     

Crown Spread Patterns for Five Deciduous Broad-leaved Woody Species: Ecological Significance of the Retention Patterns of Larger Branches

Patterns of crown spread and branch retention of two shade-intolerantspecies (Betula platyphyllaandB. maximowicziana) were comparedwith three more tolerant species (Quercus mongolica,Acer sieboldianumandMagnoliaobovata). Branching height (height of the lowest living branch)rose more rapidly with age for the twoBetulaspecies than forthe shade tolerant species. Branching heights ofA. sieboldianumandM.obovatawere similar, irrespective of tree height and age, andlarger trees tended to produce wider crowns than theBetulaspecieswhen trees of similar height were compared. In all species,the branch basal area (cross-sectional area of a branch at itsbase) and the leaf area per branch generally increased as thebranch position on a stem became lower. Therefore, retaininglarger branches contributed significantly to the support ofa larger leaf area per tree. The number of larger branches (branchbasal area >80 cm²) for bothBetulaspecies was significantlysmaller than that of the shade tolerant species. The branchretention pattern ofBetulaspecies was probably a consequenceof intolerance of the leaves to shade. The decline ofBetulaspecieswith forest succession is likely to occur through their inabilityto retain branches with a large base area in closed forests.Copyright1997 Annals of Botany Company Shade tolerance; crown spread; branch retention; branch size; broad-leaved woody species; leaf area index per tree     

Biological spectrum and other structural functional attributes of the vegetation of Kumaun Himalaya

Spectra on life form, leaf size, leaf persistence, flowering season, and shade tolerance of trees in different vegetation types occurring within the north-western catchment of the river Gola in Kumaun Himalaya are presented. The flora of Quercus leucotrichophora, Quercus lanuginosa and Quercus floribunda forests is phanerophytic, that of Pinus roxburghii and mixed forests therophytic. The grassland vegetation is characterized by the largest percentage of hemicryptophytes. The flora of the whole area, is therophytic. The biological spectrum for the entire Kumaun Himalaya is characterized as therohemigeophytic. Among the various altitudinal zones, the tropical has a preponderance of phanerophytes, the temperate and the alpine of hemicryptophytes.The observations on leaf size indicate that with the exception of Pinus roxburghii forest, in all vegetation types, the species with microphylls are greater in number. In this region, the vegetation expression is evergreen, although the tree flora has a considerable content of deciduous elements. In all the forests, the flowering period in most of the trees is vernal.On the basis of relative density, the greatest proportions of adult trees in the Pinus roxburghii and Quercus lanuginosa forests are shade intolerant, while in the mixed and Quercus leucotrichophora forests maximum trees are intermediate in shade tolerance. With the exception of the Pinus roxburghii forest, all the forests exhibit the dominance of trees which are shade tolerant at the seedling stage. On the basis of relative density, all forest types, except for Pinus roxburghii forest, have 74.5 to 100% trees with the potentiality of vegetative reproduction.Financial support from the Indian Space Research Organisation, Banglore and University Grants Commission, New Delhi is gratefully acknowledged.     

Leaf growth dynamics of two congener gymnosperm tree species reflect the heterogeneity of light intensities given in their natural ecological niche

Chamaecyparis obtusa var. formosana and Chamaecyparis formosensis are congener gymnosperm tree species native to Taiwan cloud forests; occupying different niches there. While the seedlings of C. formosensis occur predominantly under bright conditions in large forest gaps, seedlings of C. obtusa var. formosana are mainly found below the canopy of mature forests or in small gaps. It is well known that congener species occupying different niches typically differ in several ecophysiological and morphological traits, but the differences in growth dynamics of such species are still totally unclear, as the diurnal growth dynamics of gymnosperm leaves have not been investigated before. Modern methods of digital image sequence processing were used in this study to analyse the leaf growth dynamics of the two species. We found that both species show similar base–tip gradients and pronounced diurnal growth rhythms with maxima in the evening. Differences between the two species concerning their growth dynamics correlated closely with their ecological amplitudes and abundances. Chamaecyparis obtusa var. formosana grew faster than C. formosensis in low light intensity, typical for closed-canopy situations, and reacted quickly by increasing or decreasing growth rate when light intensity changed within a range typically found below small canopy gaps. In contrast to this, C. formosensis grew better in light intensities typical for open vegetation situations, but reacted slower towards changes of light intensity. Based on those results, the hypothesis can be developed that fluctuations of leaf growth dynamics reflect heterogeneities of the light environment within the niche occupied by a given species.     

A Simple model of the leaf daily water potential dynamics of some forest tree species

A simple model was developed to characterize the daily water potential dynamics (Ψ_x of sun and shade leaves of three forest tree species (Quercus cerris, Acer campestre andCarpinus betulus) under anticyclonic weather types. Input data used for this-model were the vapour pressure deficit (d) and the soil moisture content (w.). The model is usable for the calculation of the actual Ψ_x-values with a probable error 0.18 –0.28 MPa and limits the maximum and minimum Ψ_x-values which may occur with the particular tree species. The model makes it possible to establish for each species the regime, determined byd andw, at which the water potential of the leaves reacts most sensitively to the changes of the environmental parameters.     

Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

Effects of the common cormorant,phalacrocorax carbo,on evergreen forests in two nest sites at Lake Biwa,Japan

The effects of the habitation of the common cormorant on vegetation were evaluated in two colonies in Lake Biwa, central Japan. The state of the vegetation of each colony was classified arbitrarily into the following three types according to the appearance of the tree, shrub and herb layers; dense tree layer (Type 1), sparse tree layer and herb layer (Type 2), and sparse tree layer with dense herb layer (Type 3). The density of cormorant nesting was almost similar among the three types, except for part of Type 2. Trees, especially large ones on which cormorants nested, were found to be heavily damaged, and there were few seedlings and saplings found under the canopy trees.Chamaecyparis obtusa andCryptomeria japonica were more heavily damaged than other evergreen and deciduous trees. In the herb layer, only a few species such asReynoutria japonica orPhytolacca americana were abundant. These results suggest that the habitation of cormorants could simplify the structure and the species composition of forests.     

Influence of shade tolerance and development stage on the allometry of ten temperate tree species

Allometry studies the change in scale between two dimensions of an organism. The metabolic theory of ecology predicts invariant allometric scaling exponents, while empirical studies evidenced inter- and intra-specific variations. This work aimed at identifying the sources of variations of the allometric exponents at both inter- and intra-specific levels using stem analysis from 9,363 trees for ten Eastern Canada species with a large shade-tolerance gradient. Specifically, the yearly allometric exponents, α _v,DBH [volume (v) and diameter at breast height (DBH)], β _v,h [v and height (h)], and γ _h,DBH (h and DBH) were modelled as a function of tree age for each species. α _v,DBH, and γ _h,DBH increased with tree age and then reached a plateau ranging from 2.45 to 3.12 for α _v,DBH, and 0.874–1.48 for γ _h,DBH. Pine species presented a local maximum. No effect of tree age on β _v,h was found for conifers, while it increased until a plateau ranging from 3.71 to 5.16 for broadleaves. The influence of shade tolerance on the growth trajectories was then explored. In the juvenile stage, α _v,DBH, and γ _h,DBH increased with shade tolerance while β _v,h was shade-tolerance independent. In the mature stage, β _v,h increased with shade tolerance, whereas γ _h,DBH decreased and α _v,DBH was shade-tolerance independent. The interaction between development stage and shade tolerance for allometric exponents demonstrates the importance of the changing functional requirements of trees for resource allocation at both the inter- and intra-specific level. These results indicate the need to also integrate specific functional traits, growth strategies and allocation, in allometric theoretical frameworks.     

Fungal Community Analyses of Endophytic Fungi from Two Oak Species,Quercus mongolica and Quercus serrata,in Korea

Fungal endophytes have been recorded in various plant species with a richness of diversity, and their presence plays an essential role in host plant protection against biotic and abiotic stresses. This study applied the Illumina MiSeq sequencing platform based on the amplification of fungal ribosomal ITS2 region to analyze fungal endophytic communities of two oak species (Quercus mongolica and Q. serrata) with different oak wilt disease susceptibilities in Korea. The results showed a total of 230,768 sequencing reads were obtained and clustered at a 97% similarity threshold into 709 operational taxonomic units (OTUs). The OTUs of Q. serrata were higher than that of Q. mongolica with the number of 617 OTUs and 512 OTUs, respectively. Shannon index also showed that Q. serrata had a significantly higher level of fungal diversity than Q. mongolica. Total of OTUs were assigned into 5 fungal phyla, 17 classes, 60 orders, 133 families, 195 genera, and 280 species. Ascomycota was the dominant phylum with 75.11% relative abundance, followed by Basidiomycota with 5.28%. Leptosillia, Aureobasidium and Acanthostigma were the most abundant genera detected in Q. serrata with the average relative abundance of 2.85, 2.76, and 2.19%, respectively. On the other hand, Peltaster, Cladosporium and Monochaetia were the most common genera detected in Q. mongolica with the average relative abundance of 4.83, 3.03, and 2.87%, respectively. Our results indicated that fungal endophytic communities were significantly different between two oak species and these differences could influence responses of host trees to oak wilt disease caused by Raffaelea quercus-mongolicae.     

岷江冷杉林皆伐后次生群落结构和物种多样性的演替动态

为阐明岷江冷杉林皆伐后次生群落的演替过程,采用空间代替时间的方法,在川西米亚罗林区海拔3100—3600 m的阴坡选择岷江冷杉林皆伐后次生演替10、20、30、40和50a阶段的次生群落作为研究对象,对其群落结构和物种多样性的动态进行了研究。不同演替阶段的树木均呈显著聚集分布。按群落中优势种的重要值将该演替序列划分为3个类型:悬钩子-蔷薇灌丛、白桦阔叶林和桦木-岷江冷杉针阔混交林。随次生演替,乔木和灌木种的物种丰富度趋于增加,而草本种的物种丰富度趋于减少;乔木和灌木种的Shannon-Wiener多样性指数趋于增大,而草本的Shannon-Wiener多样性指数趋于减小;乔木、灌木和草本层的Pielou均匀度指数均趋于增大;乔木层的Simpson优势度指数趋于减小,灌木和草本层的Simpson优势度指数在演替0—40年阶段趋于增大,而在演替50a阶段趋于减小。在该演替序列中,乔木、灌木和草本层的物种组成均呈耐荫种替代非耐荫种的趋势。     

Comparative demography of three coexisting Acer species in gaps and under closed canopy

Questions: 1. Is there a trade‐off between gap dependency and shade tolerance in each of the life‐history stages of three closely related, coexisting species, Acer amoenum (Aa), A. mono (Am) and A. rufinerve (Ar)? 2. If not, what differences in life‐history traits contribute to the coexistence of these non‐pioneer species? Location: Ogawa Forest Reserve, a remnant (98 ha), species‐rich, temperate deciduous forest in central Japan (36°56’ N, 140°35’ E, 600 ‐ 660 m a.s.l.). Methods: We estimated the demographic parameters (survival, growth rate and fecundity) by stage of each species growing in gaps and under closed canopy through observations of a 6‐ha permanent plot over 12 years. Population dynamics were analysed with stage‐based matrix models including gap dynamics. Results : All of the species showed high seedling and sapling survival rates under closed canopies. However, demographic parameters for each growth stage in gaps and under closed canopies revealed inter‐specific differences and ontogenetic shifts. The trade‐off between survival in the shade and growth in gaps was detected only at the small sapling stage (height < 30 cm), and Ar had the highest growth rate both in the shade and in the gaps at most life stages. Conclusions: Inter‐specific differences and ontogenetic shifts in light requirements with life‐form differences may contribute to the coexistence of the Acer species in old‐growth forests, with Aa considered a long‐lived sub‐canopy tree, Am a long‐lived canopy tree, and Ar a short‐lived,‘gap‐phase’ sub‐canopy tree.     

Tree recruitment is determined by stand structure and shade tolerance with uncertain role of climate and water relations

1. Tree regeneration is a key process for long‐term forest dynamics, determining changes in species composition and shaping successional trajectories. While tree regeneration is a highly stochastic process, tree regeneration studies often cover narrow environmental gradients only, focusing on specific forest types or species in distinct regions. Thus, the larger‐scale effects of temperature, water availability, and stand structure on tree regeneration are poorly understood.
2. We investigated these effects in respect of tree recruitment (in‐growth) along wide environmental gradients using forest inventory data from Flanders (Belgium), northwestern Germany, and Switzerland covering more than 40 tree species. We employed generalized linear mixed models to capture the abundance of tree recruitment in response to basal area, stem density, shade casting ability of a forest stand as well as site‐specific degree‐day sum (temperature), water balance, and plant‐available water holding capacity. We grouped tree species to facilitate comparisons between species with different levels of tolerance to shade and drought.
3. Basal area and shade casting ability of the overstory had generally a negative impact on tree recruitment, but the effects differed between levels of shade tolerance of tree recruitment in all study regions. Recruitment rates of very shade‐tolerant species were positively affected by shade casting ability. Stem density and summer warmth (degree‐day sum) had similar effects on all tree species and successional strategies. Water‐related variables revealed a high degree of uncertainty and did not allow for general conclusions. All variables had similar effects independent of the varying diameter thresholds for tree recruitment in the different data sets.
4. Synthesis: Shade tolerance and stand structure are the main drivers of tree recruitment along wide environmental gradients in temperate forests. Higher temperature generally increases tree recruitment rates, but the role of water relations and drought tolerance remains uncertain for tree recruitment on cross‐regional scales.