Fallow deer impacts on Wakatipu beech forest

This study uses data from forty-nine 20 m × 20 m permanent plots measured in 1976, 1982, 1989 and 1997-2002 in Wakatipu Forest, western Otago. We relate changes in red (Nothofagus fusca), silver (Nothofagus menziesii) and mountain beech (Nothofagus solandri var. cliffortioides) forest vegetation to the presence of fallow deer (Dama dama). Vegetation composition is likely to have been altered prior to plot establishment, and results show that there was little change in vegetation composition during the study. There are some signs of fallow deer reducing silver beech seedling abundance at some plots, but most beech stands appear to be in the late phase of development and undergoing self thinning, so are probably not immediately vulnerable to suppression of canopy regeneration. In the future, widespread disturbance is likely to release plants in the understorey from competition for light and nutrients, and at that stage fallow deer browsing of beech species may be able to alter successional pathways more than has occurred over the past two decades. Seedling and sapling density of the palatable Griselinia littoralis (broadleaf) was probably insufficient to ensure successful regeneration. Comparisons of seedling densities with other mixed beech forests throughout New Zealand suggest that fallow deer density will need to be nearly zero in Wakatipu Forest before regeneration of all palatable subcanopy hardwood species is assured.     

The impact of beech thickets on biodiversity

Beech bark disease has dramatically altered hardwood forest structure and composition across northeastern North America. Extensive overstory mortality has resulted in prolific root-sprouting in some stands leading to the development of understory thickets of clonal small-stemmed beech. Beech thickets may impact local forest biodiversity, but this has not been adequately evaluated. We hypothesized significant differences in diversity of groundcover flora, craneflies, amphibians, and small mammals between plots with and without beech thickets. Paired plots were established in uneven-aged northern hardwood forest stands with no recent management history at two sites in the Adirondack Mountains of New York State. Groundcover plants, terrestrial craneflies, amphibians and small mammals were sampled on twenty paired plots. Discriminant analysis showed a significant difference between thicket and non-thicket (control) areas; significant variables in plot type separation were beech sapling abundance, leaf litter depth, and coarse woody debris volume. Groundcover plant cover, richness, and diversity were significantly lower in thicket compared to non-thicket plots, while beech sapling density explained 17–38 % in groundcover plant species diversity. There were no significant differences between the diversity of cranefly, amphibian and small mammal communities of each plot type. Beech thickets are important determinants of local biodiversity.     

Analysis of natural Fagus sylvatica L. s.l. regeneration in low elevation stands located in the central part of the Evros region in northeastern Greece: Is sprout origin regeneration significant for species maintenance?

Abstract

The aim of this study was (a) to analyse natural Fagus sylvatica L. s.l. regeneration in low elevation stands located in the central part of Evros region in northeastern Greece in relation to factors such as site productivity, type of silvicultural treatment and regeneration origin and (b) to determine whether or not sprout origin regeneration is significant for the maintenance of beech stands. One hundred and twenty plots (3 m × 3 m) were established in areas where thinnings and regeneration fellings had taken place, and in two site productivity regimes. In each plot, the number of beech seedlings, saplings, and stump or root sprouts (regeneration plants) with a height of up to 3 m was counted, and their height measured. Our findings indicate that even though reproduction by seed is the predominant regeneration mechanism in our stands, in the medium productivity sites sprouts represent a significant percentage of the total number of plants. Moreover, sprouting functions as a mechanism of maintenance of beech stands in medium productivity sites. During the regeneration fellings practised in this area, it is advisable to cut the seedlings and saplings of the advanced regeneration in order to supplement the seed-derived plants with seedling sprouts.     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than environmental factors.     

Variation of plant diversity in a temperate unmanaged forest in northern Spain: behind the environmental and spatial explanation

The persistence of seedlings in the forest understorey is of major importance for the maintenance and regeneration of canopy trees in several forested ecosystems. In the present study, we examine the small-scale spatial pattern of a mixed beech and oak seedling–sapling bank in two areas of an unmanaged temperate deciduous forest with different environmental conditions. We used environmental, biotic and spatial variables to establish the main factors that explain the spatial pattern of these seedling–sapling banks at different scales. The stand structure in both areas was similar, but while in plot A beech dominated the canopy, plot B was dominated by oaks. In both areas, established beech individuals showed a clear reverse J-shaped distribution, whereas established oaks showed a unimodal distribution with only a few young individuals. Seedlings of beech and oak were distributed in aggregates, whereas beech saplings had a random distribution. At broader scales, the abundance of seedlings and saplings is affected by the environment as well as by inter-species competition, while at finer scales the spatial pattern is mainly influenced by stochastic processes, probably related to seed predation and establishment. The structure of the seedling–sapling bank indicates an advantage of beech over oak as far as regeneration is concerned. Beech seedlings and saplings tolerate the stress induced by the canopy and the understorey and persist for many years, while oak seedlings decline in a few years. Therefore, if current conditions persist, after canopy opening beech seedlings and saplings can grow rapidly into the canopy and the stands will move towards beech dominance.     

影响不同林型天然红松混交林林隙更新的土壤特征因子

采用野外调查、样品采集和统计分析等相结合的方法,对小兴安岭天然红松混交林3种不同林型(椴树红松混交林(TP)、枫桦红松混交林(BP)、云冷杉红松混交林(PAP))的林隙及其邻近郁闭林分的土壤特征因子和树木更新的相关性进行了研究,旨在阐明林隙土壤特征因子对树木更新的影响,从而为小兴安岭天然红松混交林植被更新、退化生态系统的恢复和可持续经营提供基础数据和实践参考。结果表明:郁闭林分土壤有机质、全氮质量分数显著高于3种不同林型的林隙。有效磷和速效钾含量在BP内与其他林型之间差异显著。3种林型林隙内p H值均略高于其郁闭林分,但与其差异均不显著。3种林型林隙内更新总密度、幼树更新密度与郁闭林分差异显著(P0.05),PAP林隙中更新总密度和幼树更新密度最高。BP林隙面积与更新密度相关不显著,乔木幼苗、幼树更新密度与有机质(r=-0.400,r=-0.475)、全氮均呈显著负相关(r=-0.519,r=-0.603)。TP林隙内全氮与乔木幼苗更新密度呈正相关(r=0.092),而与乔木幼树更新密度呈显著负相关(r=-0.585)。PAP林隙内全氮与乔木幼苗、幼树更新密度均呈负相关。郁闭林分幼苗更新密度分别与有机质、全氮、速效钾、p H值、脲酶和蛋白酶呈负相关。主成分分析表明,全氮是影响林隙和郁闭林分树木更新的关键因素。     

Influence of site conditions on the survival of Fagus sylvatica seedlings in an old‐growth beech forest

Abstract. Following a full mast production in autumn 1995 in the old‐growth beech forest of la Tillaie (France), cupules, live and dead 1‐yr old seedlings were counted in September 1996 in 40 plots, representing a wide range of ecological conditions. The influence of geomorphology and forest stage on mast production and on seedling establishment and survival was studied. Maximum seedling abundance did not occur in areas with the largest cupule abundance, but this result was not statistically significant. The presence of a shallow sandstone table under Fontainebleau sand, where beech is far from its ecological optimum, positively influenced the production of mast but negatively influenced seedling survival. Contrary to expectation, the early biostatic rather than the late biostatic phase seemed to be the best stage for beech regeneration. Gaps also provide good environmental conditions for seedling establishment. Humus and light conditions, water availability and competition between beech individuals may explain the results. Light was the main factor influencing mast production in mature stages, and drought was an important factor causing seedling mortality. This study points out the importance of seedling survival for regeneration patterns in beech forests.     

Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

Understory vegetation response to mountain pine beetle disturbance in northern Colorado lodgepole pine forests

Understory plants are an important element of forests, having a considerable influence on ecosystem functioning and canopy-tree development following disturbance. Recent bark beetle outbreaks across western North American forests have caused extensive canopy mortality, creating new growing conditions that provide the opportunity for changes within the intact understory. Over a five-year period following peak mountain pine beetle (MPB) activity across lodgepole pine-dominated forests in Rocky Mountain National Park, Colorado, we measured the changes in plant diversity, cover, and dominance by lifeform and quantified tree regeneration rates. Average species richness and diversity increased, but overall plant cover did not change. Graminoids appeared to benefit the most, increasing in average cover, richness, and relative dominance. The rise in graminoid dominance was largely at the expense of shrubs, which showed little ability to benefit from overstory mortality within the first years following attack. Most plant responses were positively related to the total tree basal area lost since the peak of the outbreak, suggesting that increased resource availability following tree death may benefit understory plants. However, a negative relationship between several understory variables and tree sapling density provides evidence that understory plants compete with saplings for the newly available resources. Tree seedling density nearly doubled over the duration of the study, indicating a strong regeneration pulse. Among species, lodgepole pine displayed the greatest tree seedling establishment. This is one of the first studies to use repeated measurements to describe this often-overlooked component of forest change associated with MPB disturbance.

     

Herbivory and plant competition reduce mountain beech seedling growth and establishment in New Zealand

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently. Nomenclature: Beever et al. (1992); Parsons et al. (1995); Edgar and Connor (2000); and Brownsey and Smith-Dodsworth (2000). Raukaua simplex is described by Mitchell et al. (1997). Coprosma “taylorae” is referred to by Eagle (1986) and Halocarpus biformis, Phyllocladus alpinus, Podocarpus hallii and Podocarpus nivalis by Wilson and Galloway (1993).     

Ecological consequences of a massive flowering event of bamboo (Chusquea culeou) in a temperate forest of Patagonia,Argentina

Question: What changes occur as a consequence of the massive flowering and senescence of the dominant understory species of bamboo, Chusquea culeou (E. Desvaux)? In this study, we documented some of the ecological consequences of this rare event that occurred in 2001, the previous flowering having occurred more than 60 years ago. Location: Nothofagus temperate forest, Patagonia, Argentina. Methods: We assessed changes in environmental variables and bamboo biomass post‐flowering in an old‐growth southern beech forest. In addition, we monitored the demography of emergent Ch. culeou seedling and Nothofagus nervosa saplings, comparing non‐flowered (live understory) and flowered (senescent understory) patches within the forest matrix. Results: Bamboo flowering dramatically increased light availability in the forest understory but, surprisingly, other environmental changes were not observed. Bamboo seedlings emerged in both patch types, and experienced gradual but modest mortality through time. Bamboo dieback promoted higher survivorship and an increment in biomass, height, number of leaves and buds in the saplings of Nothofagus nervosa. Conclusion: The high density of bamboo seedlings 5 years after the flowering event and the independence of emergence from environmental variables suggest that understory regeneration is a gradual process that is not strongly regulated by initial seedling density or resource limitation. In contrast, microenvironmental conditions created after the flowering event significantly increased Nothofagus sapling growth and survival. These results suggest that overstory forest regeneration could be enhanced in this temperate forest in the first years after this infrequent bamboo flowering event.     

Inferring controls on the epidemiology of beech bark disease from spatial patterning of disease organisms

• 1 Spatial pattern in the distribution and abundance of organisms is an emergent property of collective rates of reproduction, survival and movement of individuals in a heterogeneous environment.
• 2 The form, intensity and scale of spatial patterning can be used to test hypotheses regarding the relative importance of candidate processes to population dynamics.
• 3 Using 84 plots across eastern North America, we studied populations of two associated plant parasites, the invasive felted beech scale Cryptococcus fagisuga Lind. and the native Neonectria fungi, which together cause beech bark disease (BBD).
• 4 We evaluated spatial patterns at the scales of trees within stands, stands within the forest and forests within the landscape to examine four hypothetically important factors in the ecology of the disease: (i) local contagion within stands; (ii) regional contagion, or among patch infection–reinfection dynamics; (iii) variation in host susceptibility linked to genetic and/or environmental heterogeneity; and (iv) climate effects on population growth of BBD organisms.
• 5 Analyses revealed an unexpected lack of spatial aggregation in BBD populations among trees, stands and forests. This implies that propagule pressure is generally sufficiently high throughout the infested region of North America such that neither trees nor stands are spared from the disease by dispersal limitations of the disease agents. Furthermore, variation in tree and stand level susceptibility has minimal impact on BBD dynamics and climate is not a conspicuous driver of abundance within the core range of BBD.

     

The distribution pattern of herbivory in a beech canopy

Abstract. (1) The spatial distribution pattern of herbivory by phyllophagous canopy arthropods in a 90-year-old beech stand was analysed.
(2) Two main patterns of spatial distribution were recorded. The leaf mining moths Phyllonorycter maestingella and Stigmella hemargyrella and the adult weevils Rhynchaenus fagi and Phyllobius argentatus mainly attacked the under-story and the latter species especially the low canopy in the central part of the stand. Mines of R. fagi chiefly occurred in the top of the overstory canopy or at the fringe of the stand. However, comparable distribution patterns are only expected in mature beech stands composed of distinct overstory and understory layers.
(3) The effect of the actual spatial distribution pattern of herbivory on photosynthesis, productivity, and transpiration of beech in the research stand is discussed. It is suggested that in some cases the shade leaves of the understory canopy may cushion the effect of herbivory in the beech stand.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

林分光照空间异质性对水曲柳更新的影响

 光照是森林更新动态中极为关键的环境因子。本研究在东北天然次生林中选择样地,应用地统计学理论结合实验生态学方法,定量分析林分光照空间异质性特征,同时进行林下播种水曲柳（Fraxinus mandshurica）的更新实验,探测更新格局特征。通过空间关联性分析,研究探讨是否林分光照的空间变异对水曲柳更新有重要影响作用。研究结果表明： 1）生长季林分平均光照仅为全光照的4.2%～4.6%,在空间分布上呈现非常明显的异质性特征,空间自相关变异（62.5%～78.2%）主要体现在10.9～12.4 m的尺度上。2）林分光照不同的空间样点上,水曲柳更新幼苗发生数量和秋季幼苗存活数量均存在显著差异,光照相对强的微生境中,更新苗发生数量较多且存活率高,而在光照较弱的微生境中则更新较差,更新具有十分明显的空间格局。3）更新格局与林分光照空间异质性特征紧密关联。在光照空间异质性程度较高、空间变异较复杂的条件下,更新相应呈现出较复杂的空间格局,自相关的变异表现在较小尺度范围（2.12～6.97 m）。在光照空间异质性程度较低、空间变异复杂性较小的条件下,更新格局的复杂程度明显变小,空间自相关变异（>83%）表现在较大的尺度上（30 m）,对更新格局起主要的影响作用,随机变异的影响很小（<17%）。这些研究结果说明,林分光照环境在水曲柳更新中的作用十分关键,林分光照的空间异质性对更新格局起决定性作用。     

Influence of overstory composition on understory colonization by native species in plantations on a degraded tropical site

Abstract. Patterns of understory colonization by native and naturalized trees and shrubs were evaluated in 4.5-year-old plantations of three exotic tree species, Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala, on a degraded coastal grassland site with reference to overstory composition and understory environmental conditions. 19 secondary forest species were established in the plantation understories (with a total area of 0.52 ha), while no natural regeneration occurred in unplanted, though protected, control areas. The majority of these species (90 %) and the total seedling population (97 %) were zoochorous, indicating the importance of frugivorous bats and particularly birds as facilitators of secondary forest species colonization. Understory species richness and seedling densities were affected significantly by overstory composition, the most abundant regeneration occurring beneath Leucaena and least under Casuarina. Understory colonization rates within mixed-species stands were intermediate between those of single-species stands of the trees comprising their overstories. Significant negative correlations were found between understory species richness and seedling density, and forest floor depth and dry mass, especially for small-seeded ornithochorous species. Higher colonization rates near the peripheries of plantation plots relative to plot interiors were due in part to roosting site preferences by frugivores, particularly bats. The study results indicate that overstory species selection can exert a significant influence on subsequent patterns of colonization by secondary forest species and is an important consideration in the design of plantations for ‘catalyzing’ succession on deforested, degraded sites.     

Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.     

Bioclimate and growth history affect beech lifespan in the Italian Alps and Apennines

When site factors reduce growth rates, tree lifespan tends to increase. This study investigates processes leading to such inverse relationship in Fagus sylvatica stands distributed along two elevation gradients, with an emphasis on climatic response, suppression periods, and growth trends. Dendrochronological records from old‐growth beech populations sampled at different elevations within two different bioclimatic regions (Alps vs. Apennines), were used to investigate factors that control tree lifespan. Differences between old‐growth (12) and nearby managed (15) stands were used to assess effects of silvicultural practices on maximum age. Logging reduced tree lifespan not only by removing older trees, but also by reducing the number of years beech individuals spent in the shaded understory. Tree lifespan and growth rates were affected by climate (spring–summer temperature) and were inversely related to one another along elevation gradients. The greatest lifespan was observed in old‐growth high‐mountain populations, and was related not only to slower growth due to a shorter growing season, but also to multidecadal periods of growth suppression during the initial development stages in the understory (i.e., slower growth rates at the youngest cambial ages). Past unfavorable climatic periods (in this case, the Little Ice Age) also helped increase tree lifespan. Using a linear model, we estimated a reduction in beech lifespan of 23 ± 5 years for each degree of warming. Basal area increment of trees with the maximum observed lifespan showed an increasing trend over time. Because growth of old (>300 years) trees has increased in the Alps, while it has recently declined in the Apennines, different bioclimatic regions can have opposite responses to global climatic change. In the next decades, if warming continues, beech lifespan could be reduced in the Alps by faster growth and in the Apennines by drought‐induced mortality.     

Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.