• Background and Aims Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0·2–2·5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan.
• Methods The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance.
• Key Results Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity.
• Conclusions This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.
Cool-temperate trees; crown allometry; crown architecture; height growth; Indonesia; saplings; plasticity; tropical trees
Background and Aims
Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation.
A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening.
Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth.
Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.
Advance regeneration; canopy gap; biomechanics; gravitropism; shade tolerance; Fagus sylvatica; Acer pseudoplatanus
Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-009-1304-2) contains supplementary material, which is available to authorised users.
Diversity-functioning relationships; Leaf damage; Mines; Multitrophic interactions; Plant–animal interactions
Background and Aims
In rain forests, sapling survival is highly dependent on the regulation of trunk slenderness (height/diameter ratio): shade-intolerant species have to grow in height as fast as possible to reach the canopy but also have to withstand mechanical loadings (wind and their own weight) to avoid buckling. Recent studies suggest that mechanosensing is essential to control tree dimensions and stability-related morphogenesis. Differences in species slenderness have been observed among rainforest trees; the present study thus investigates whether species with different slenderness and growth habits exhibit differences in mechanosensitivity.
Recent studies have led to a model of mechanosensing (sum-of-strains model) that predicts a quantitative relationship between the applied sum of longitudinal strains and the plant's responses in the case of a single bending. Saplings of five different neotropical species (Eperua falcata, E. grandiflora, Tachigali melinonii, Symphonia globulifera and Bauhinia guianensis) were subjected to a regimen of controlled mechanical loading phases (bending) alternating with still phases over a period of 2 months. Mechanical loading was controlled in terms of strains and the five species were subjected to the same range of sum of strains. The application of the sum-of-strain model led to a dose–response curve for each species. Dose–response curves were then compared between tested species.
The model of mechanosensing (sum-of-strain model) applied in the case of multiple bending as long as the bending frequency was low. A comparison of dose–response curves for each species demonstrated differences in the stimulus threshold, suggesting two groups of responses among the species. Interestingly, the liana species B. guianensis exhibited a higher threshold than other Leguminosae species tested.
This study provides a conceptual framework to study variability in plant mechanosensing and demonstrated interspecific variability in mechanosensing.
Mechanosensing; interspecific variability; trees; lianas; rain forest; neotropical species; bending; biomechanics; Bauhinia; Eperua; Symphonia; Tachigali
Background and Aims
The volume of tree stems is made up of three components: solid wood, gas and water. These components have important consequences for the construction costs, strength and stability of trees. Here, the importance of stem components for sapling growth and survival in the field was investigated, and then these stem components were related to two important life history axes of variation: the light requirements for regeneration and the adult stature of the species.
Stem fractions of wood, gas and water were determined for saplings and adults of respectively 30 and 58 Bolivian tropical moist-forest species. Sapling height growth and survival were monitored for 2 years in the field as indicators of sapling performance.
Sapling stems consisted of 26 % wood (range 7–36 % for species), 59 % water (range 49–88 %), and 15 % gas (range 0–38 %) per unit volume. The wood fraction was the only determinant of sapling performance and was correlated with increased survival and decreased growth rate across species. The wood fraction decreased with light requirements of the species, probably because a high wood fraction protects shade-tolerant species against pathogens and falling debris. The gas fraction increased with the light requirements and adult stature of the species; probably as an aid in realizing a rapid height growth and accessing the canopy in the case of light-demanding species, and for rapidly attaining stability and a large reproductive size in the case of tall species. The water fraction was not correlated with the life history variation of tree species, probably because it leads to increased stem loading and decreased stability.
The wood fraction might partially explain the growth–survival trade-off that has been found across tropical tree species. The wood and gas fractions are closely related to the regeneration light requirements of the species. Tall species have a high gas fraction, probably not only because gas is a cheap filler, but also because it might lead to an increased stability of these tall trees.
Adult stature; biomechanics; Bolivia; shade tolerance; second moment of area; tropical rain forest; wood density; wood specific gravity
The senescence and loss of photosynthetic and support structures is a nearly universal aspect of tree life history, and can be a major source of disturbance in forest understoreys, but the ability of falling canopy debris in determining the stature and composition of understorey communities seems not to have been documented. In this study, we show that senescent fronds of the palm Iriartea deltoidea cause substantial disturbance in tropical forest sapling communities. This disturbance influences the species composition of the canopy and subcanopy by acting as an ecological filter, favouring sapling species with characteristics conducive to recovery after physical damage. The scale of this dominance suggests that falling I. deltoidea debris may be influencing sapling community structure and species composition in Amazonian rainforests over very large spatial scales.
Eight fungal species known to produce wood pigmentation were tested for reaction to various moisture contents in two hardwood species. Fungal pigmentation by Trametes versicolor and Xylaria polymorpha was stimulated at low water concentrations in both Acer saccharum (sugar maple) and Fagus grandifolia (American beech), while Inonotus hispidus and Polyporus squamosus were stimulated above 22-28% and 34-38% moisture content in beech and in sugar maple respectively. Fomes fomentarius and Polyporus brumalis produced maximum pigmentation in beech at 26 - 41% and in sugar maple at 59 - 96% moisture content. The pink staining Scytalidium cuboideum pigmented both wood species at above 35% moisture content. This research indicates that controlling the moisture content values of wood substrates can stimulate the intensity of pigmentation of specific fungi when spalting wood for decorative and commercial purpose.
Fungal melanin; Pigment; Moisture content; Spalting
Canopy structure, which can be defined as the sum of the sizes, shapes and relative placements of the tree crowns in a forest stand, is central to all aspects of forest ecology. But there is no accepted method for deriving canopy structure from the sizes, species and biomechanical properties of the individual trees in a stand. Any such method must capture the fact that trees are highly plastic in their growth, forming tessellating crown shapes that fill all or most of the canopy space.
We introduce a new, simple and rapidly-implemented model–the Ideal Tree Distribution, ITD–with tree form (height allometry and crown shape), growth plasticity, and space-filling, at its core. The ITD predicts the canopy status (in or out of canopy), crown depth, and total and exposed crown area of the trees in a stand, given their species, sizes and potential crown shapes. We use maximum likelihood methods, in conjunction with data from over 100,000 trees taken from forests across the coterminous US, to estimate ITD model parameters for 250 North American tree species. With only two free parameters per species–one aggregate parameter to describe crown shape, and one parameter to set the so-called depth bias–the model captures between-species patterns in average canopy status, crown radius, and crown depth, and within-species means of these metrics vs stem diameter. The model also predicts much of the variation in these metrics for a tree of a given species and size, resulting solely from deterministic responses to variation in stand structure.
This new model, with parameters for US tree species, opens up new possibilities for understanding and modeling forest dynamics at local and regional scales, and may provide a new way to interpret remote sensing data of forest canopies, including LIDAR and aerial photography.
It is generally believed that asymmetric competition for light plays a predominant role in determining the course of succession by increasing size inequalities between plants. Size-related growth is the product of size-related light capture and light-use efficiency (LUE). We have used a canopy model to calculate light capture and photosynthetic rates of pioneer species in sequential vegetation stages of a young secondary forest stand. Growth of the same saplings was followed in time as succession proceeded. Photosynthetic rate per unit plant mass (Pmass: mol C g−1 day−1), a proxy for plant growth, was calculated as the product of light capture efficiency [Φmass: mol photosynthetic photon flux density (PPFD) g−1 day−1] and LUE (mol C mol PPFD−1). Species showed different morphologies and photosynthetic characteristics, but their light-capturing and light-use efficiencies, and thus Pmass, did not differ much. This was also observed in the field: plant growth was not size-asymmetric. The size hierarchy that was present from the very early beginning of succession remained for at least the first 5 years. We conclude, therefore, that in slow-growing regenerating vegetation stands, the importance of asymmetric competition for light and growth can be much less than is often assumed.
Canopy model; Light capture; Photosynthesis; Regeneration; Tropical forest
The maintenance in the long run of a positive carbon balance under very low irradiance is a prerequisite for survival of tree seedlings below the canopy or in small gaps in a tropical rainforest. To provide a quantitative basis for this assumption, experiments were carried out to determine whether construction cost (CC) and payback time for leaves and support structures, as well as leaf life span (i) differ among species and (ii) display an irradiance-elicited plasticity. Experiments were also conducted to determine whether leaf life span correlates to CC and payback time and is close to the optimal longevity derived from an optimization model. Saplings from 13 tropical tree species were grown under three levels of irradiance. Specific-CC was computed, as well as CC scaled to leaf area at the metamer level. Photosynthesis was recorded over the leaf life span. Payback time was derived from CC and a simple photosynthesis model. Specific-CC displayed only little interspecific variability and irradiance-elicited plasticity, in contrast to CC scaled to leaf area. Leaf life span ranged from 4 months to >26 months among species, and was longest in seedlings grown under lowest irradiance. It was always much longer than payback time, even under the lowest irradiance. Leaves were shed when their photosynthesis had reached very low values, in contrast to what was predicted by an optimality model. The species ranking for the different traits was stable across irradiance treatments. The two pioneer species always displayed the smallest CC, leaf life span, and payback time. All species displayed a similar large irradiance-elicited plasticity.
Carbon balance; construction cost; functional diversity; leaf life span; payback time; photosynthesis; tropical rainforest
Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (Nmass) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (Narea). In evergreen conifers, LMA and Narea were less powerful predictors of area-based photosynthetic rate (Amax(area)) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, Amax(mass) was related to Nmass but not to LMA, whereas LMA became a significant codeterminant of Amax(mass) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-012-2279-y) contains supplementary material, which is available to authorized users.
Plant functional types; Leaf plasticity; Shade acclimation; Evergreen leaves; Leaf mass-per-area
Background and Aims
In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest.
The relationship between seedling survival and seedling relative growth rate in mass (RGRM) or height (RGRH) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes.
Within each species, both RGRM and RGRH usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGRM and RGRH in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG.
The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.
Coexistence; gap; gap size; habitat selection; habitat width; light; niche partitioning; relative growth rate; seed size; successional status
Background and Aims
The light availability on a temperate, deciduous-forest floor varies greatly, reflecting the seasonal leaf dynamics of the canopy trees. The growth and/or reproductive activity of understorey plants should be influenced by the length of the high-irradiance period from snowmelt to canopy closure. The aim of the present study was to clarify how spring-blooming species regulate the translocation of photosynthetic products to current reproduction and storage organs during a growing season in accordance with the changing light conditions.
Growth pattern, net photosynthetic rate, seed production, and shoot and flower production in the next year of Trillium apetalon were compared between natural and experimentally shaded conditions. Furthermore, translocation of current photosynthetic products within plants was assessed by a labelled carbon-chase experiment.
During the high-irradiance period, plants showed high photosynthetic ability, in which current products were initially used for shoot growth, then reserved in the rhizome. Carbon translocation to developing fruit occurred after canopy closure, but this was very small due to low photosynthetic rates under the darker conditions. The shading treatment in the early season advanced the time of carbon translocation to fruit, but reduced seed production in the current year and flower production of the next year.
Carbon translocation to the storage organ had priority over seed production under high-irradiance conditions. A shortened bright period due to early canopy closure effectively restricts carbon assimilation, which greatly reduces subsequent reproductive output owing to low photosynthetic products for fruit development and small carbon storage for future reproduction. As populations of this species are maintained by seedling recruitment, acceleration of canopy closure timing may influence the maintenance and dynamics of populations.
13C labelling; canopy closure; carbon translocation; deciduous forest; light availability; photosynthesis; spring-bloomer; Trillium apetalon
Due to a long history of intensive forest exploitation, few European beech (Fagus sylvatica L.) old-growth forests have been preserved in Europe.
Material and Methods
We studied two beech forest reserves in southern Slovenia. We examined the structural characteristics of the two forest reserves based on data from sample plots and complete inventory obtained from four previous forest management plans. To gain a better understanding of disturbance dynamics, we used aerial imagery to study the characteristics of canopy gaps over an 11-year period in the Kopa forest reserve and a 20-year period in the Gorjanci forest reserve.
The results suggest that these forests are structurally heterogeneous over small spatial scales. Gap size analysis showed that gaps smaller than 500 m2 are the dominant driving force of stand development. The percentage of forest area in canopy gaps ranged from 3.2 to 4.5% in the Kopa forest reserve and from 9.1 to 10.6% in the Gorjanci forest reserve. These forests exhibit relatively high annual rates of coverage by newly established (0.15 and 0.25%) and closed (0.08 and 0.16%) canopy gaps. New gap formation is dependant on senescent trees located throughout the reserve.
We conclude that these stands are not even-sized, but rather unevenly structured. This is due to the fact that the disturbance regime is characterized by low intensity, small-scale disturbances.
Climbing plants are a key component of rainforests, but mechanistic approaches to their distribution and abundance are scarce. In a southern temperate rainforest, we addressed whether the dominance of climbing plants across light environments is associated with the expression of ecophysiological traits. In mature forest and canopy gaps, we measured leaf size, specific leaf area, photosynthetic rate, and dark respiration in six of the most abundant woody vines. Mean values of traits and their phenotypic change (%) between mature forest and canopy gaps were predictor variables. Leaf size and specific leaf area were not significantly associated with climbing plant dominance. Variation in gas-exchange traits between mature forest and canopy gaps explained, at least partly, the dominance of climbers in this forest. A greater increase in photosynthetic rate and a lower increase in dark respiration rate when canopy openings occur were related to the success of climbing plant species. Dominant climbers showed a strategy of maximizing exploitation of resource availability but minimizing metabolic costs. Results may reflect phenotypic plasticity or genetic differentiation in ecophysiological traits between light environments. It is suggested that the dominant climbers in this temperate rainforest would be able to cope with forest clearings due to human activities.
A series of growth experiments and observations on natural populations have been carried out on dipterocarp species of contrasting ecology growing in artificial gaps and the forest understorey. These studies have demonstrated that although differences exist between species in photosynthetic and growth responses to the high-light environment, competition for light in canopy gaps is highly asymmetrical and tends to reinforce any pre-existing dominance hierarchy. We propose that differences in seedling persistence in forest canopy shade are highly influenced by species-specific biotic and abiotic interactions. Our experiments suggest that as seedlings, dipterocarp species trade off traits which enhance persistence and growth in shade against those that enhance their ability to exploit gaps. Less competitive species survive for progressively longer periods of time after a gregarious fruiting event. This leads to significant shifts with time in the number of species present in the seedling bank and hence in the importance of interspecific competition in determining which species dominates regrowth in gaps. We propose that this special case of dispersal limitation is more likely to account for coexistence of dipterocarp species than differences in growth responses to gaps of different size, with stochastic and environmental variables interacting to determine species distribution and abundance.
1.Insect herbivory can strongly affect ecosystem processes, and its relationship with plant diversity is a central topic in biodiversity–functioning research. However, very little is known about this relationship from complex ecosystems dominated by long-lived individuals, such as forests, especially over gradients of high plant diversity.
2.We analysed insect herbivory on saplings of 10 tree and shrub species across 27 forest stands differing in age and tree species richness in an extraordinarily diverse subtropical forest ecosystem in China. We tested whether plant species richness significantly influences folivory in these highly diverse forests or whether other factors play a more important role at such high levels of phytodiversity.
3.Leaf damage was assessed on 58 297 leaves of 1284 saplings at the end of the rainy season in 2008, together with structural and abiotic stand characteristics.
4.Species-specific mean damage of leaf area ranged from 3% to 16%. Herbivory increased with plant species richness even after accounting for potentially confounding effects of stand characteristics, of which stand age-related aspects most clearly covaried with herbivory. Intraspecific density dependence or other abiotic factors did not significantly influence overall herbivory across forest stands.
5.Synthesis.The positive herbivory–plant diversity relationship indicates that effects related to hypotheses of resource concentration, according to which a reduction in damage by specialized herbivores might be expected as host plant concentration decreases with increasing plant diversity, do not seem to be major determinants for overall herbivory levels in our phytodiverse subtropical forest ecosystem. We discuss the potential role of host specificity of dominant herbivores, which are often expected to show a high degree of specialization in many (sub)tropical forests. In the forest system we studied, a much higher impact of polyphagous species than traditionally assumed might explain the observed patterns, as these species can profit from a broad dietary mix provided by high plant diversity. Further testing is needed to experimentally verify this assumption.
BEF China; biodiversity; ecosystem functioning; Gutianshan; resource concentration; succession; trophic interactions; Zhejiang
Savannahs are a mixture of trees and grasses often occurring as alternate states to closed forests. Savannah fires are frequent where grass productivity is high in the wet season. Fires help maintain grassy vegetation where the climate is suitable for woodlands or forests. Saplings in savannahs are particularly vulnerable to topkill of above-ground biomass. Larger trees are more fire-resistant and suffer little damage when burnt. Recruitment to large mature tree size classes depends on sapling growth rates to fire-resistant sizes and the time between fires. Carbon dioxide (CO2) can influence the growth rate of juvenile plants, thereby affecting tree recruitment and the conversion of open savannahs to woodlands. Trees have increased in many savannahs throughout the world, whereas some humid savannahs are being invaded by forests. CO2 has been implicated in this woody increase but attribution to global drivers has been controversial where changes in grazing and fire have also occurred. We report on diverse tests of the magnitude of CO2 effects on both ancient and modern ecosystems with a particular focus on African savannahs. Large increases in trees of mesic savannahs in the region cannot easily be explained by land use change but are consistent with experimental and simulation studies of CO2 effects. Changes in arid savannahs seem less obviously linked to CO2 effects and may be driven more by overgrazing. Large-scale shifts in the tree–grass balance in the past and the future need to be better understood. They not only have major impacts on the ecology of grassy ecosystems but also on Earth–atmosphere linkages and the global carbon cycle in ways that are still being discovered.
vegetation fires; woody thickening; bush encroachment; C4 grass
Four isolates of Chlorociboria aeruginascens were tested for possible stimulatory effects when grown on malt agar media containing wood additives. The addition of any of the four types of test wood (Acer saccharum, Populus tremuloides, spalted P. tremuloides, and Ailanthus altissima), stimulated colony growth and xylindein production in C. aeruginascens. Addition of any amount of wood produced more growth than no wood additions, while ground wood produced more growth than chopped wood. Of the wood types tested, A. saccharum wood stimulated all four isolates, while spalted Populus tremuloides stimulated three of the four isolates. High glucose and sucrose amounts may be partially responsible for the greater stimulatory affect of some woods over others. The development of this simple and reliable method for growth and pigment stimulation of C. aeruginascens in laboratory conditions will allow for further development of this fungus for decorative and commercial use.
Chlorociboria; Green Pigment; Spalting; Xylindein
We hypothesised that the decomposition rates of leaf litter will increase along a gradient of decreasing fraction of the European beech (Fagus sylvatica) and increasing tree species diversity in the generally beech-dominated Central European temperate deciduous forests due to an increase in litter quality. We studied the decomposition of leaf litter including its lignin fraction in monospecific (pure beech) stands and in stands with up to five tree genera (Acer spp., Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Tilia spp.) using a litterbag approach. Litter and lignin decomposition was more rapid in stand-representative litter from multispecific stands than in litter from pure beech stands. Except for beech litter, the decomposition rates of species-specific tree litter did not differ significantly among the stand types, but were most rapid in Fraxinus excelsior and slowest in beech in an interspecific comparison. Pairwise comparisons of the decomposition of beech litter with litter of the other tree species (except for Acerplatanoides) revealed a “home field advantage” of up to 20% (more rapid litter decomposition in stands with a high fraction of its own species than in stands with a different tree species composition). Decomposition of stand-representative litter mixtures displayed additive characteristics, not significantly more rapid than predicted by the decomposition of litter from the individual tree species. Leaf litter decomposition rates were positively correlated with the initial N and Ca concentrations of the litter, and negatively with the initial C:N, C:P and lignin:N ratios. The results support our hypothesis that the overall decomposition rates are mainly influenced by the chemical composition of the individual litter species. Thus, the fraction of individual tree species in the species composition seems to be more important for the litter decomposition rates than tree species diversity itself.
Biodiversity; Central Europe; Element ratio; Litter degradation; Litterbag
• Background and Aims Burial is a recurrent stress imposed upon plants of coastal dunes. Woody plants are buried on open coastal dunes and in forested areas behind active blowouts; however, little is known about the burial responses and adaptive traits of these species. The objectives of this study were: (a) to determine the growth and morphological responses to burial in sand of seven woody plant species native to central Canadian coastal dunes; and (b) to identify traits that determine burial tolerance in these species.
• Methods Field experiments were conducted to determine the responses of each species to burial. Saplings were exposed to burial treatments of 0, 10, 25, 50 and 75 % of their height. Burial responses were evaluated based on regressions of total biomass, height, adventitious root production and percentage allocation to shoot, root and adventitious root biomass on percentage burial.
• Key Results
Pinus strobus and Picea glauca lacked burial tolerance. In response to the burial gradient, these species showed a strong linear decline in total biomass, minimal adventitious root production that peaked at moderate levels (25–50 % burial) and no change in allocation to shoots vs. roots. The tolerant species Juniperus virginiana, Thuja occidentalis and Picea mariana showed a quadratic response to burial, with little change in biomass up to 50 % burial, but a large decline at 75 %. These species produced abundant adventitious roots up to 50 % burial, but did not alter allocation patterns over the range of burial levels. Populus balsamifera and Salix cordata were stimulated by burial. These species showed linear increases in biomass with increasing burial, produced copious adventitious roots across the gradient and showed a clear shift in allocation to vertical shoot growth and adventitious root production at the expense of the original roots under high burial conditions.
• Conclusions Adventitious root production and plastic resource allocation to biomass are adaptive traits of coastal dune woody plants in central Canada, and provide a basis for assessing burial tolerance in woody plants on coastal dunes throughout the world.
Burial; coastal dunes; woody plants; adventitious roots; allocation patterns; Pinus strobus; Picea glauca; Picea mariana; Juniperus virginiana; Thuja occidentalis; Salix cordata; Populus balsamifera
Tree species differ from one another in, and display trade-offs among, a wide range of attributes, including canopy and understorey growth and mortality rates, fecundity, height and crown allometry, and crown transmissivity. But how does this variation affect the outcome of interspecific competition and hence community structure? We derive criteria for the outcome of competition among tree species competing for light, given their allometric and life-history parameters. These criteria are defined in terms of a new simple whole life-cycle measure of performance, which provides a simple way to organize and understand the many ways in which species differ. The general case, in which all parameters can differ between species, can produce coexistence, founder control or competitive exclusion: thus, competition for light need not be hierarchical as implied by R* theory. The special case in which species differ only in crown transmissivity produces neutral dynamics. The special case in which species differ in all parameters except crown transmissivity gives hierarchical competition, where the equivalent of R* is Zˆ*, the height at which trees enter the canopy in an equilibrium monoculture.
biomechanics; succession; community ecology; forest dynamics; pairwise invasibility; Sortie
In Amazonian floodplain forests, >1000 tree species grow in an environment subject to extended annual submergence which can last up to 9 months each year. Water depth can reach 10 m, fully submerging young and also adult trees, most of which reproduce during the flood season. Complete submergence occurs regularly at the seedling or sapling stage for many species that colonize low-lying positions in the flooding gradient. Here hypoxic conditions prevail close to the water surface in moving water, while anaerobic conditions are common in stagnant pools. Light intensities in the floodwater are very low.
Questions and Aims
Despite a lack of both oxygen and light imposed by submergence for several months, most leafed seedlings survive. Furthermore, underwater growth has also been observed in several species in the field and under experimental conditions. The present article assesses how these remarkable plants react to submergence and discusses physiological mechanisms and anatomical adaptations that may explain their success.
Adaptation; Amazonian floodplains; darkness; environmental stress; flooding; hypoxia; submergence tolerance; trees; underwater photosynthesis; woody species
Background and Aims
Along snowmelt gradients, the canopies of temperate alpine meadows differ strongly in their structural and biochemical properties. Here, a study is made of the effects of these canopy dissimilarities combined with the snow-induced changes in length of growing season on seasonal gross primary production (GPP).
Leaf area index (LAI) and community-aggregated values of leaf angle and leaf nitrogen content were estimated for seven alpine plant canopies distributed along a marked snowmelt gradient, and these were used as input variables in a sun–shade canopy bulk-photosynthesis model. The model was validated for plant communities of early and late snowmelt sites by measuring the instantaneous CO2 fluxes with a canopy closed-chamber technique. A sensitivity analysis was conducted to estimate the relative impact of canopy properties and environmental factors on the daily and seasonal GPP.
Carbon uptake was primarily related to the LAI and total canopy nitrogen content, but not to the leaf angle. For a given level of photosynthetically active radiation, CO2 assimilation was higher under overcast conditions. Sensitivity analysis revealed that increase of the length of the growing season had a higher effect on the seasonal GPP than a similar increase of any other factor. It was also found that the observed greater nitrogen content and larger LAI of canopies in late-snowmelt sites largely compensated for the negative impact of the reduced growing season.
The results emphasize the primary importance of snow-induced changes in length of growing season on carbon uptake in alpine temperate meadows. It was also demonstrated how using leaf-trait values of the dominants is a useful approach for modelling ecosystem carbon-cycle-related processes, particularly when continuous measurements of CO2 fluxes are technically difficult. The study thus represents an important step in addressing the challenge of using a plant functional-trait approach for biogeochemical modelling.
Alpine meadows; gross primary production; plant functional traits; snowmelt gradient; sun–shade model
Backgrounds and Aims
Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.
The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.
Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.
Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.
Masting; shoot demography; short shoot; long shoot; temporal variation; spatial variation; leaf area; shoot length; resource allocation; Betula grossa