Linkage between seasonal gas exchange and hydraulic acclimation in the top canopy leaves of Fagus trees in a mesic forest in Japan

We investigated how leaf gas exchange and hydraulic properties acclimate to increasing evaporative demand in mature beech trees, Fagus crenata Blume and Fagus japonica Maxim., growing in their natural habitat. The measurements in the top canopy leaves were conducted using a 16-m-high scaffolding tower over two growing seasons. The daily maxima of net photosynthetic rate for the early growing season were close to the annual maximum value (11.9 mol m^–2 s^–1 in F. crenata and 7.7 mol m^–2 s^–1 in F. japonica). The daily maxima of water vapor stomatal conductance were highest in the summer, approximately 0.3 mol m^–2 s^–1 in F. crenata and 0.15 mol m^–2 s^–1 in F. japonica. From the early growing season to the summer season, the leaf-to-air vapor pressure deficit increased and the daily minima of leaf water potentials decreased. However, there was no loss of leaf turgor in the summer as a result of effective osmotic adjustment. Both the soil-to-leaf hydraulic conductance per unit leaf area and the twig hydraulic conductivity simultaneously increased in the summer, probably as a result of production of new vessels in the xylem. These results suggest that both osmotic adjustment and increased hydraulic conductance resulted in the largest diurnal maximum of stomatal conductance in the summer, resulting in the lowest relative stomatal limitation on net photosynthetic rate, although the leaf-to-air vapor pressure deficit was highest. These results indicate that even in a mesic forest, in which excessive hydraulic stress does not occur, the seasonal acclimation of hydraulic properties at both the single leaf and whole plant levels are important for plant carbon gain.     

Photosynthetic acclimation to light in juveniles of two cloud forest tree species

The photosynthetic response of juveniles of Decussocarpus rospigliosii, an emergent primary forest species and shade tolerant in its juvenile stages and Alchornea triplinervia, a gap-colonizing species of tropical cloud forest in Venezuela was studied. Daily courses of microenvironmental variables and gas exchange under contrasting light conditions (gap and understory) were carried out in their natural environment and transplanted to different light regimes (shade and sun) in the field. The photosynthetic response and some anatomical characteristics of plants from different treatments were analyzed in the laboratory. Photosynthetic rates were low for both species, and were negative during some diurnal periods, related to the low photosynthetically active radiation levels obtained at both gap (6% of total radiation) and understory (2%). A. triplinervia shows higher rates (1.5–3.0 molm^{-2 -1}) than D. rospigliosii (0.7–1.1 molm^-2s^-1). Both species showed increased photosynthetic rates when grown in gaps. A. triplinervia did not adjust its maximum photosynthetic rates to the prevailing light conditions. In contrast, D. rospigliosii responded to increased light levels. Both species showed low light compensation points when grown under total shade. There was a partial stomatal closure generally during midday in D. rospigliosii. A. triplinervia presented lower leaf conductances, transpiration rates and lesser stomatal control. Some leaf anatomical characteristics, in both species, were affected by variations in the light regime (i.e. increased leaf thickness, leaf specific weight and stomatal density). These results suggest that both species have the ability to respond to variations in their natural light environments, therefore maintaining a favorable carbon balance during the day.     

Field water relations of a wet-tropical forest tree species,Pentaclethra macroloba (Mimosaceae)

Summary The water relations of Pentaclethra macroloba (Willd.) Kuntze, a dominant, shade-tolerant, tree species in the Atlantic lowlands of Costa Rica, were examined within the forest canopy. Pressure-volume curves and diurnal courses of stomatal conductance and leaf water potential were measured in order to assess differences in water relations between understory, mid-canopy and canopy leaves. Leaves in the canopy had the smallest pinnules but the largest stomatal frequencies and stomatal conductances of the three forest levels. Osmotic potentials at full turgidity decreased with height in the forest; in the canopy and midcanopy they were reduced relative to those in the understory just enough to balance the gravitational component of water potential. Consequently, maximum turgor pressures were similar for leaves from all three canopy levels. Bulk tissue elastic modulus increased with height in the canopy. Leaf water potentials were lowest in the canopy and highest in the understory, even when the gravitational component was added to mid-canopy and canopy values. As a result, minimum turgor pressures were also lowest in the canopy compared to those at lesser heights, and approached zero in full sunlight on clear days.Osmotic potentials at each canopy level were similar for both wet and dry season samples dates suggesting that seasonal osmotic adjustment does not occur. Despite lowered predawn water potentials during the dry season, turgor was maintained in the understory by reduced stomatal conductances.     

3个温带阔叶树种根系长期分解速率研究

根系分解对土壤碳固持和养分长期有效性具有重要意义,但目前对根系的长期分解模式仍知之甚少。比较3个温带阔叶树种不同直径根系7 a分解动态,可为生态过程模型提供数据支撑。在帽儿山生态站采用分解袋法对白桦（Betula platyphylla）、春榆（Ulmus davidiana var.japonica）、水曲柳（Fraxinus mandshurica）的5个直径等级（< 1 mm）、（1-2 mm）、（2-5 mm）、（5-10 mm）、（10-20 mm）根系进行了7 a野外分解实验。重复测量方差分析表明：时间、树种、直径与树种交互作用、树种与时间交互作用显著影响根系质量保持率。根系质量保持率随时间呈指数下降趋势,7 a间根系经历早期快速分解和后期慢速分解两个阶段,实验结束时根系仍剩余相当部分的初始质量（24%-56%）。利用Olson指数衰减模型估计各树种根系分解系数发现：白桦根系分解系数与根直径间具有显著的线性正相关关系,水曲柳具有显著对数正相关关系,春榆的关系不显著。3个树种短期分解系数均高估根系分解速率,而且不能完全代表长期分解系数的种内、种间差异。研究结果对理解根系长期分解速率随直径变化模式提供了数据支撑。     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Drought during canopy development has lasting effect on annual carbon balance in a deciduous temperate forest

* Climate change projections predict an intensifying hydrologic cycle and an increasing frequency of droughts, yet quantitative understanding of the effects on ecosystem carbon exchange remains limited. * Here, the effect of contrasting precipitation and soil moisture dynamics were evaluated on forest carbon exchange using 2 yr of eddy covariance and microclimate data from a 50-yr-old mixed oak woodland in northern Ohio, USA. * The stand accumulated 40% less carbon in a year with drought between bud-break and full leaf expansion (354 +/- 81 g C m(-2) yr(-1) in 2004 and 252 +/- 45 g C m(-2) yr(-1) in 2005). This was caused by greater suppression of gross ecosystem productivity (GEP; 16% = 200 g) than of ecosystem respiration (ER; 11% = 100 g) by drought. Suppressed GEP was traced to lower leaf area, lower apparent quantum yield and lower canopy conductance. The moisture sensitivity of ER may have been mediated by GEP. * The results highlight the vulnerability of the ecosystem to even a moderate drought, when it affects a critical aspect of development. Although the drought was preceded by rain, the storage capacity of the soil seemed limited to 1-2 wk, and therefore droughts longer than this are likely to impair productivity in the region.     

Depth distribution and composition of seed banks under different tree layers in a managed temperate forest ecosystem

In the present work we examined the composition and distribution across three soil layers of the buried soil seed bank under three different overstory types (Fagus sylvatica, Quercus robur, Pinus sylvestris) and in logging areas in a 4383-ha forest in central Belgium. The objectives were: (1) to investigate whether species composition and species richness of soil seed banks are affected by different forest stands; (2) to examine how abundant are habitat-specific forest species in seed banks under different planted tree layers. The study was carried out in stands which are replicated, managed in the same way (even-aged high forest), and growing on the same soil type with the same land-use history. In the investigated area, the seed bank did show significant differences under oak, beech, pine and in logging areas, respectively in terms of size, composition and depth occurrence. All species and layers taken together, the seed bank size ranked as follows: oakwood > beechwood > logging area > pinewood. The same pattern was found for forest species. Seed numbers of Betula pendula, Calluna vulgaris, Dryopteris dilatata and Rubus fruticosus were significantly higher under the beech canopy. Carex remota, Impatiens parviflora and Lotus sp. showed a significantly denser seed bank in logging areas, while Digitalis purpurea seeds were significantly more abundant in soils under the oak canopy. The fact that the seed bank of an originally homogeneous forest varies under different planted stands highlights that a long period of canopy conversion can affect the composition and depth of buried seeds.     

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.     

Comparative recruitment patterns of two non-pioneer canopy tree species in French Guiana

Summary A comparative study was conducted on the recruitment patterns of two non-pioneer tree species, one dispersed by arboreal mammals and birds (Virola michelii, Myristicaceae) and the other by rodents (Moronobea coccinea, Clusiaceae). These species differ in fruiting phenology, seed size, dispersal distance, germination time and seed nutrient exhaustion. In both species, establishment patterns were consistent with the escape hypothesis and the Janzen-Connell model. Virola seeds need not be buried to survive and germinate, and may produce a seedling carpet beneath the parent. Moronobea seedlings only establish from seeds buried by scatterhoarding rodents in the surrounding understory. One-year survival of Virola seedlings was 47.8% and was greater >10 m than < 10 m from the largest parent tree. In contrast, survival of Moronobea seedlings was 56% 3 years after seed dispersal. Survival of juveniles was greater in gaps than in the understory for Virola but not for Moronobea. Moronobea survival was greater than Virola survival in both microhabitats. Both species establish in the understory, yet both grew faster in gaps. Virola appeared to be more gap-dependent than Moronobea which may persist several years in the understory until a gap occurs. Virola and Moronobea illustrate two intermediate recruitment patterns along an hypothetical continuum of nonpioneer species replacement (Bazzaz and Pickett 1980; Swaine and Whitmore 1988).     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

Herbivory and seedling performance in a fragmented temperate forest of Chile

Forest fragmentation alters plant-animal interactions, including herbivory. Relying manipulative experiments, we test if the reduction in insect herbivory associated with forest fragmentation translates into increased seedling growth and survival of three tree common species (Aristotelia chilensis, Cryptocarya alba and Persea lingue) in forest fragments and continuous forests in coastal Maulino forest, central Chile. Furthermore, we test if after protecting seedlings from herbivorous insects, plant performance is increased regardless of forest fragmentation. Nursery grown seedlings were transplanted into four forest fragments and a continuous forest during 2002. Insects, important herbivores in this forest, were excluded from half the seedlings by repeated applications of insecticides. Compared to continuous forests, in forest fragments, herbivory was reduced in all three species, seedling growth was greater in A. chilensis and C. alba but not in P. lingue, and survivorship was unaffected by herbivory or fragmentation in all three species. Protecting seedlings from insects reduced herbivory in the continuous forest to similar levels attained in the forest fragments. No change in herbivory results from by protecting seedlings in forest fragments. These results confirm that insects are important herbivores in the Maulino forest and also support the hypothesis that fragmentation can have strong indirect effects on plant communities as mediated through trophic interactions.     

Delayed inducible resistance against a leaf-chewing insect in four deciduous tree species

Summary Both mechanical damage to mountain birch foliage and rearing of moth larvae on the trees reduced the growth of Epirrita autumnata larvae reared on these trees in the following year. The effects of physical damage and some other cues from insects were additive. On bird cherry the performance of Epirrita larvae was equal on untreated trees and on trees artificially defoliated in the previous year, but larval growth was reduced on previously insect-damaged branches. With mountain ash just physical damage per se reduced the performance of Epirrita larvae. On Salix phylicifolia there were no significant differences in the growth or survival of Epirrita on untreated control bushes and on bushes with partial larval damage during the previous year. Among untreated control trees the growth and survivorship of Epirrita were higher on fast-growing willow and bird cherry than on the slow-growing mountain birch. Mountain birch and mountain ash, the two deciduous tree species adapted to nutrient-poor soils, showed delayed inducible resistance triggered by defoliation (artificial or insect-made). This supports the hypothesis that delayed inducible resistance may be a passive response due to nutrient-stress caused by defoliation. On the other hand, the additional increase in the resistance of mountain birch triggered by specific cues from insects suggests that this response may be an evolved defense against leaf-eating insects.     

Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

Herb layer response to broadleaf tree species with different leaf litter quality and canopy structure in temperate forests

Question: How do broadleaf tree species affect humus characteristics, herb layer composition and species diversity through their leaf litter quality and canopy structure? Location: Mixed broadleaf forests in Brandenburg, NE Germany. Methods: We studied the herb and tree layer composition in 129 undisturbed stands using a 10‐degree cover‐abundance and percentage scale, respectively. The main floristic gradients were extracted by non‐metric multidimensional scaling. Effects of tree species on the herb layer were analysed with partial Spearman rank correlation. We assessed affinities for specific tree species using indicator species analysis. Results: Both beech and oak influenced herb layer composition mainly through their litter quality, which resulted in deep Ol and Of horizons, respectively. The less dense canopy of oak, in contrast to the dense beech canopy, enhanced species diversity in favour of indifferent herb species (species not closely tied to forests). Lime was correlated with a distinct floristic gradient, but a direct effect on the herb layer cannot be proven with the available data. Effects of hornbeam were less pronounced. Conclusions: The relationship between the tree and herb layer must be partly attributed to pH differences. However, tree species effects on humus characteristics and on light flux to the ground were largely responsible as well. The results suggest that tree species can influence herb layer composition and diversity, but the missing correlation with lime and hornbeam raise questions requiring further detailed investigation.     

Thermal acclimation of photosynthesis: a comparison of boreal and temperate tree species along a latitudinal transect

Common gardens were established along a ～900 km latitudinal transect to examine factors limiting geographical distributions of boreal and temperate tree species in eastern North America. Boreal representatives were trembling aspen (Populus tremuloides Michx.) and paper birch (Betula papyrifera Marsh.), while temperate species were eastern cottonwood (Populus deltoides Bartr ex. Marsh var. deltoides) and sweetgum (Liquidambar styraciflua L.). The species were compared with respect to adjustments of leaf photosynthetic metabolism along the transect, with emphasis on temperature sensitivities of the maximum rate of ribulose bisphosphate (RuBP) carboxylation (E_V) and regeneration (E_J). During leaf development, the average air temperature (T_growth) differed between the coolest and warmest gardens by 12 °C. Evidence of photosynthetic thermal acclimation (metabolic shifts compensating for differences in T_growth) was generally lacking in all species. Namely, neither E_V nor E_J was positively related to T_growth. Correspondingly, the optimum temperature (T_opt) of ambient photosynthesis (A_sat) did not vary significantly with T_growth. Modest variation in T_opt was explained by the combination of E_V plus the slope and curvature of the parabolic temperature response of mesophyll conductance (g_m). All in all, species differed little in photosynthetic responses to climate. Furthermore, the adaptive importance of photosynthetic thermal acclimation was overshadowed by g_m's influence on A_sat's temperature response.     

Seasonal reversibility of acclimation to irradiance in leaves of common box (Buxus sempervirens L.) in a deciduous forest

Understorey shade plants are seasonally exposed to dramatic changes in light conditions in deciduous forests related with the dynamics of the overstorey leaf phenology. These transitions are commonly followed by changes in herb plant communities, but shade-tolerant evergreen species must be able to adapt to changing light conditions. In this work we checked the photoprotective responses of evergreen species to acclimate to the shady summer environment and reversibly de-acclimate to a more illuminated environment after leaf fall on deciduous overstoreys. For that purpose we have followed the process of light acclimation in leaves of common box (Buxus sempervirens) during the winter to spring transition, which decrease irradiance in the understorey, and conversely during the transition from summer to autumn. Four parameters indicative of the structure and degree of acclimation of the photosynthetic apparatus have been studied: chlorophyll a/b ratio which is supposed to be inversely proportional to the antenna size, α/β-carotene which increases in shade acclimated leaves and the pools of α-tocopherol and xanthophyll cycle pigments (VAZ) which are two of the main photoprotection mechanisms in plants. Among these parameters, chlorophyll a/b ratio and VAZ pool responded finely to changes in irradiance indicating that modifications in the light harvesting size and photoprotective capacity contribute to the continuous acclimation and de-acclimation of long-lived evergreen leaves.     

东北温带森林20种乔木树种叶片干旱容忍性特征

全球范围内干旱频率和强度的增加严重影响树木生长,甚至导致森林大面积死亡。压力-容积(PV)曲线能够反映树木对干旱的容忍能力,但在局域尺度上尚未确定哪个PV曲线参数具有最优指示性。通过测定东北温带森林20种主要树种(包括16种被子植物和4种裸子植物)的PV曲线性状,包括质壁分离时的相对含水量(RWC_tlp)、失膨点叶水势(TLP)、饱和含水时的叶渗透势(π₀)、细胞弹性模量(ε)、叶水容(C_leaf)及叶结构性状(比叶面积和叶密度),研究局域尺度上叶片耐旱性的最佳指示性状,并分析叶片PV性状与结构性状间的相关性。结果表明: 被子植物的RWC_tlp 显著大于裸子植物,但其C_leaf 显著小于裸子植物,这表明用RWC_tlp和C_leaf可以指示东北温带森林不同功能型树种间耐旱性的大小。在被子植物中,TLP和π₀与叶密度呈显著负相关,且均与比叶面积呈显著正相关;而ε与比叶面积呈显著负相关。然而,裸子植物PV曲线性状与叶结构性状之间呈现与被子植物完全相反的趋势。裸子植物与被子植物树种之间PV曲线性状与叶结构性状关系的差异,可能归因于二者采取不同的干旱响应和适应策略。     

Long-term persistence of aspen – a key host for many threatened species – is endangered in old-growth conservation areas in Finland

Large, dead and dying European aspens (Populus tremula L.) host many threatened species in Fennoscandian boreal forests. Large aspen trees have mostly disappeared and are being harvested from the managed forests that cover 95% of the forest area in Finland. Due to the small area protected (4.1%), the aspen-associated species may encounter major difficulties in the protected areas if aspen trees disappear due to natural forest succession. The availability of aspens was assessed in the old-growth conservation area network in eastern Finland. We mapped all the living and dead aspens in 15 protected old-growth forests. The total number of counted trees was 32 903 individuals. Current amounts of living (2.7 m³/ha) and especially dead aspens (2.8 m³/ha) in the protected areas were higher than in the surrounding managed forests (1.1 and 0.1 m³/ha for living and dead trees, respectively). However, while saplings (dbh<5 cm) occur in most of the areas (12 individuals/ha on average) they survive poorly and young aspen cohorts (5 cm <dbh<15 cm) are lacking or are very rare. The most likely reason for the poor sapling survival is high browsing pressure by the mammalian herbivores, especially the moose. The moose population has increased many times in Finland during the past decades. The poor regeneration of aspens implies that the value of the old-growth conservation areas for aspen-associated species will face a serious bottleneck within a few decades when the currently middle-aged tree cohorts disappear. If the current high browsing pressure and lack of natural disturbances continue the obligatory aspen-associated species may disappear both locally and regionally from the network of the protected areas.