Permian polar forests: deciduousness and environmental variation

Forests are expected to expand into northern polar latitudes in the next century. However, the impact of forests at high latitudes on climate and terrestrial biogeochemical cycling is poorly understood because such forests cannot be studied in the modern. This study presents forestry and geochemical analyses of three in situ fossil forests from Late Permian strata of Antarctica, which grew at polar latitudes. Stem size measurements and stump spacing measurements indicate significant differences in forest density and canopy structure that are related to the local depositional setting. For forests closest to fluvial systems, tree density appears to decrease as the forests mature, which is the opposite trend of self‐thinning observed in modern forests. We speculate that a combination of tree mortality and high disturbance created low‐density mature forests without understory vegetation near Late Permian river systems. Stable carbon isotopes measured from permineralized wood in these forests demonstrate two important points: (i) recently developed techniques of high‐resolution carbon isotope studies of wood and mummified wood can be applied to permineralized wood, for which much of the original organic matter has been lost and (ii) that the fossil trees maintained a deciduous habit at polar latitudes during the Late Permian. The combination of paleobotanical, sedimentologic, and paleoforestry techniques provides an unrivaled examination of the function of polar forests in deep time; and the carbon isotope geochemistry supplements this work with subannual records of carbon fixation that allows for the quantitative analysis of deciduous versus evergreen habits and environmental parameters, for example, relative humidity.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.     

Factors limiting the distribution of deciduous broadleaved trees in warm-temperate mountainous riparian forests

We examined differences in the responses of deciduous and evergreen broadleaved trees to fluvial disturbance and light environment near a river-facing forest edge in order to identify the factors limiting the distribution of deciduous trees, which are important components of warm-temperate mountainous riparian forests. Deciduous trees tended to be distributed on lower terraces of less than 2-m relative elevation from the water level, which corresponded with sites suffering from strong bank erosion due to high flood frequency. On the other hand, evergreen trees showed an opposite trend of distribution, indicating that high flood frequency associated with strong erosion-dominated soil disturbances might be a constraint for their establishment and/or survival. Furthermore, crowns of deciduous trees tended to be in the canopy layer at the forest edge, with fewer individuals observed beneath the canopy of evergreen trees, even at the forest edge. In contrast, evergreen trees were concentrated in the understory of the forest interior. These observations suggested that the light environment is the predominant factor affecting the establishment and survival of deciduous trees in warm-temperate mountainous riparian forests; however, the river-facing forest edge beneath the evergreen canopy is not a suitable habitat for deciduous trees. We concluded that erosion-dominated soil disturbance on lower terraces provides suitable habitats for deciduous trees by facilitating their reaching the canopy layer by limiting the establishment and/or survival of evergreen trees.

     

山东植物区系的演变和来源

1　现代植物区系山东省位于我国东部、黄河下游 ,北濒渤海 ,东临黄海 ,地理范围介于北纬 34°2 3′～38°2 4′,东经 1 1 4°4 8′～ 1 2 2°4 3′之间。全省总面积为 1 5.72万 km2 ,占全国总面积的 1 .6%。属暖温带季风气候 ,沿海比较湿润 ,地带性植被主要是暖温带落叶阔叶林和松、栎类针阔叶混交林。山东省在中国植物区系的分区地位隶属于泛北极植物区、中国 -日本森林植物亚区、华北植物地区 [1 ]。据最近研究统计 ,现有野生维管植物 1 47科、61 4属 ,约 1 547种 (包括变种 ,下同 )。其中蕨类植物 2 4科 39属 1 0 5种 ,裸子植物 3科 3属 …     

Fire in tropical dry forests: corticolous lichens as indicators of recent ecological changes in Thailand

The distribution of lichens in lowland deciduous and evergreen forests in Thailand is used to interpret recent changes in the distribution of these forests. The role of fire in changing the forest structure, microclimate and species content is discussed. Characteristic corticolous lichen communities of dry deciduous and moist evergreen forests are described, as well as changes in the composition of the flora following fire events. Where frequent fires have altered the forest rates of change in forest type are suggested using lichen data from randomly selected trees in forest plots, and growth rates of sampled species in quadrats. The disjunct nature of the lichen floras in lowland deciduous and evergreen forests is discussed, their origin and use in interpreting changes in forest types in monsoon climates over long periods of time.     

The ecology, silviculture and biogeography of mahogany (Swietenia macrophylla): a critical review of the evidence

Recent studies of Swietenia macrophylla King argue that natural regeneration of this highly valued tree is episodic and occurs only after infrequent catastrophic disturbance. An analysis of 14 population size-distributions in natural forests shows that population structure varies across this species' range. In forests that are in successional transition from open deciduous forest to evergreen forest there is often prolific regeneration. We show that in this type of forest in southern Pará (Brazil) patches of even-sized trees are small. Large-scale disturbance may be necessary for regeneration only in evergreen forests where little light penetrates through the canopy. The transition zone between evergreen and deciduous forest in Pará is moving south, and we propose that mahogany populations to the north are in the process of competitive exclusion. Sustainable harvesting of mahogany must take account of the successional stage of the forest.     

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.     

Geographical and climatic gradients of evergreen versus deciduous broad‐leaved tree species in subtropical China: Implications for the definition of the mixed forest

Understanding climatic influences on the proportion of evergreen versus deciduous broad‐leaved tree species in forests is of crucial importance when predicting the impact of climate change on broad‐leaved forests. Here, we quantified the geographical distribution of evergreen versus deciduous broad‐leaved tree species in subtropical China. The Relative Importance Value index (RIV ) was used to examine regional patterns in tree species dominance and was related to three key climatic variables: mean annual temperature (MAT ), minimum temperature of the coldest month (MinT), and mean annual precipitation (MAP ). We found the RIV of evergreen species to decrease with latitude at a lapse rate of 10% per degree between 23.5 and 25°N, 1% per degree at 25–29.1°N, and 15% per degree at 29.1–34°N. The RIV of evergreen species increased with: MinT at a lapse rate of 10% per °C between ?4.5 and 2.5°C and 2% per °C at 2.5–10.5°C; MAP at a lapse rate of 10% per 100 mm between 900 and 1,600 mm and 4% per 100 mm between 1,600 and 2,250 mm. All selected climatic variables cumulatively explained 71% of the geographical variation in dominance of evergreen and deciduous broad‐leaved tree species and the climatic variables, ranked in order of decreasing effects were as follows: MinT > MAP  > MAT . We further proposed that the latitudinal limit of evergreen and deciduous broad‐leaved mixed forests was 29.1–32°N, corresponding with MAT of 11–18.1°C, MinT of ?2.5 to 2.51°C, and MAP of 1,000–1,630 mm. This study is the first quantitative assessment of climatic correlates with the evergreenness and deciduousness of broad‐leaved forests in subtropical China and underscores that extreme cold temperature is the most important climatic determinant of evergreen and deciduous broad‐leaved tree species’ distributions, a finding that confirms earlier qualitative studies. Our findings also offer new insight into the definition and distribution of the mixed forest and an accurate assessment of vulnerability of mixed forests to future climate change.     

Californian mixed-conifer forests under unmanaged fire regimes in the Sierra San Pedro Mártir, Baja California, Mexico

Aim This study appraises historical fire regimes for Californian mixed‐conifer forests of the Sierra San Pedro Mártir (SSPM). The SSPM represents the last remaining mixed‐conifer forest along the Pacific coast still subject to uncontrolled, periodic ground fire. Location The SSPM is a north–south trending fault bound range, centred on 31°N latitude, 100 km SE of Ensenada, Baja California. Methods We surveyed forests for composition, population structure, and historical dynamics both spatially and temporally over the past 65 years using repeat aerial photographs and ground sampling. Fire perimeter history was reconstructed based on time‐series aerial photographs dating from 1942 to 1991 and interpretable back to 1925. A total of 256 1‐ha sites randomly selected from aerial photographs were examined along a chronosequence for density and cover of canopy trees, density of snags and downed logs, and cover of non‐conifer trees and shrubs. Twenty‐four stands were sampled on‐the‐ground by a point‐centred quarter method which yielded data on tree density, basal area, frequency, importance value, and shrub and herb cover. Results Forests experience moderately intense understory fires that range in size to 6400 ha, as well as numerous smaller, low intensity burns with low cumulative spatial extent. SSPM forests average 25–45% cover and 65–145 trees per ha. Sapling densities were two to three times that of overstory trees. Size‐age distributions of trees ≥ 4 cm dbh indicate multi‐age stands with steady‐state dynamics. Stands are similar to Californian mixed conifer forests prior to the imposition of fire suppression policy. Livestock grazing does not appear to be suppressing conifer regeneration. Main conclusions Our spatially‐based reconstruction shows the open forest structure in SSPM to be a product of infrequent, intense surface fires with fire rotation periods of 52 years, rather than frequent, low intensity fires at intervals of 4–20 years proposed from California fire‐scar dendrochronology (FSD) studies. Ground fires in SSPM were intense enough to kill pole‐size trees and a significant number of overstory trees. We attribute long fire intervals to the gradual build‐up of subcontinuous shrub cover, conifer recruitment and litter accumulation. Differences from photo interpretation and FSD estimates are due to assumptions made with respect to site‐based (point) sampling of fire, and nonfractal fire intensities along fire size frequency distributions. Fire return intervals determined by FSD give undue importance to local burns which collectively use up little fuel, cover little area, and have little demographic impact on forests.     

Physiological ecology of Mesozoic polar forests in a high CO2 environment

Fossils show that coniferous forests extended into polar regions during the Mesozoic, a time when models and independent paleo-CO2 indicators suggest that the atmospheric CO2 concentration was at least double that of the present day. Consequently, such polar forests would have experienced high CO2 interacting with an extreme variation in light. Here we describe an experiment investigating this plant-environment interaction for extant tree species that were important components of polar forests, and give results from the first year of treatment. Specifically, we tested the hypotheses that growth in elevated CO2 (1) stimulates photosynthesis; (2) reduces photoinhibition during the polar summer; and (3) reduces respiration of above- and below-ground plant organs. Our results indicate that CO2 fertilization generally does not affect photosynthesis under continuous daylight characteristic of the polar summer but does increase it when the period of illumination is shorter. Growth in elevated CO2 did not alter the potential for photoinhibition. CO2 enrichment significantly reduced leaf and root respiration rates by 50 and 25 %, respectively, in a range of evergreen taxa. Incorporating these observed CO2 effects into numerical simulations using a process-based model of coniferous forest growth indicates that a high paleo-CO2 concentration would have increased the productivity of Cretaceous conifer forests in northern Alaska. This results from decreased respiratory costs that more than compensate for the absence of high CO2-high temperature interactions during the polar summer. The longer-term effects of CO2 enrichment on seasonal changes in the above- and below-ground carbon balance of trees are discussed.     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Species diversity of Chinese beech forests in relation to warmth and climatic disturbances

The trends in the occurrence of climatic disturbances in the ChineseFagus range are described, and the relationship between woody species diversity and climatic factors in eight old-growth Chinese beech forests is characterized. In the ChineseFagus range that lies in the humid mountains of southern China, wind storms and heavy rain frequency increase towards the eastern coast. Thunderstorm frequency increases southwards. Snowfall frequency increases northwards. Glaze storm frequency peaks in the center near Lake Dongtian, but much higher in the east than in the west. Hailstorm frequency also peaks in the center. The forests sampled in this study are widely separated. Their canopies consist of either deciduous broad-leaved trees or a mixture of evergreen and deciduous broad-leaved trees. Their species diversity increases towards warmer sites and towards the east. The importance of the evergreen trees in relation to warmth and minimum temperature increases southwards. Our analysis suggests that wind storms and heavy rains enhance the species diversity of Chinese beech forests. Cold disturbances such as glaze and snow diminish the diversity and canopy dominance of evergreen broad-leaved trees but favor deciduous broad-leaved trees, especially beech. The annual precipitation received by the forests in this study varies from 1400–2550 mm. This is not correlated with diversity, however, probably because all of these forests grow in humid conditions with sufficient water being supplied by precipitation throughout the year.     

A global assessment of forest surface albedo and its relationships with climate and atmospheric nitrogen deposition

We present a global assessment of the relationships between the short‐wave surface albedo of forests, derived from the MODIS satellite instrument product at 0.5° spatial resolution, with simulated atmospheric nitrogen deposition rates (N_dep), and climatic variables (mean annual temperature T_m and total annual precipitation P), compiled at the same spatial resolution. The analysis was performed on the following five forest plant functional types (PFTs): evergreen needle‐leaf forests (ENF); evergreen broad‐leaf forests (EBF); deciduous needle‐leaf forests (DNF); deciduous broad‐leaf forests (DBF); and mixed‐forests (MF). Generalized additive models (GAMs) were applied in the exploratory analysis to assess the functional nature of short‐wave surface albedo relations to environmental variables. The analysis showed evident correlations of albedo with environmental predictors when data were pooled across PFTs: T_m and N_dep displayed a positive relationship with forest albedo, while a negative relationship was detected with P. These correlations are primarily due to surface albedo differences between conifer and broad‐leaf species, and different species geographical distributions. However, the analysis performed within individual PFTs, strengthened by attempts to select ‘pure’ pixels in terms of species composition, showed significant correlations with annual precipitation and nitrogen deposition, pointing toward the potential effect of environmental variables on forest surface albedo at the ecosystem level. Overall, our global assessment emphasizes the importance of elucidating the ecological mechanisms that link environmental conditions and forest canopy properties for an improved parameterization of surface albedo in climate models.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Variations of leaf longevity in tropical moist forests predicted by a trait‐driven carbon optimality model

Leaf longevity (LL) varies more than 20‐fold in tropical evergreen forests, but it remains unclear how to capture these variations using predictive models. Current theories of LL that are based on carbon optimisation principles are challenging to quantitatively assess because of uncertainty across species in the ‘ageing rate:’ the rate at which leaf photosynthetic capacity declines with age. Here, we present a meta‐analysis of 49 species across temperate and tropical biomes, demonstrating that the ageing rate of photosynthetic capacity is positively correlated with the mass‐based carboxylation rate of mature leaves. We assess an improved trait‐driven carbon optimality model with in situLL data for 105 species in two Panamanian forests. We show that our model explains over 40% of the cross‐species variation in LL under contrasting light environment. Collectively, our results reveal how variation in LL emerges from carbon optimisation constrained by both leaf structural traits and abiotic environment.     

桂北岩溶区青冈栎-青檀群落植物叶片比叶面积研究

为揭示岩溶地区植物叶片比叶面积变化规律和叶片形态之间的相关关系,研究了桂北岩溶区青冈栎-青檀群落的叶片长/宽(LL/LW)、叶片厚度(LT)和比叶面积(SLA)及其之间的关系。结果表明:群落中常绿树种的LL/LW和LT显著高于落叶树种,而常绿树种的SLA显著低于落叶树种;乔木和灌木之间的LL/LW存在显著差异,但SLA和LT不存在显著差异。总体上看,SLA与LL/LW、LT之间是显著负相关关系,SLA随着LL/LW、LT的变大而逐渐减小。但LT与SLA负相关趋势比LL/LW与SLA的明显。因此,LT比LL/LW对叶片SLA的影响大,其明显影响了叶片的SLA。SLA的大小是衡量叶片获取光照能力的指标,因此岩溶区植物较厚的叶片对光照的获取能力具有较大的影响,并可导致光合作用效率的降低。     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Canopy structure of tropical and sub-tropical rain forests in relation to conifer dominance analysed with a portable LIDAR system

Background and Aims

Globally, conifer dominance is restricted to nutient-poor habitats in colder, drier or waterlogged environments, probably due to competition with angiosperms. Analysis of canopy structure is important for understanding the mechanism of plant coexistence in relation to competition for light. Most conifers are shade intolerant, and often have narrow, deep, conical crowns. In this study it is predicted that conifer-admixed forests have less distinct upper canopies and more undulating canopy surfaces than angiosperm-dominated forests.

Methods

By using a ground-based, portable light detection and ranging (LIDAR) system, canopy structure was quantified for old-growth evergreen rainforests with varying dominance of conifers along altitudinal gradients (200–3100 m a.s.l.) on tropical and sub-tropical mountains (Mount Kinabalu, Malaysian Borneo and Yakushima Island, Japan) that have different conifer floras.

Key Results

Conifers dominated at higher elevations on both mountains (Podocarpaceae and Araucariaceae on Kinabalu and Cupressaceae and Pinaceae on Yakushima), but conifer dominance also varied with soil/substrate conditions on Kinabalu. Conifer dominance was associated with the existence of large-diameter conifers. Forests with higher conifer dominance showed a canopy height profile (CHP) more skewed towards the understorey on both Kinabalu and Yakushima. In contrast, angiosperm-dominated forests had a CHP skewed towards upper canopy, except for lowland dipterocarp forests and a sub-alpine scrub dominated by small-leaved Leptospermum recurvum (Myrtaceae) on Kinabalu. Forests with a less dense upper canopy had more undulating outer canopy surfaces. Mixed conifer–angiosperm forests on Yakushima and dipterocarp forests on Kinabalu showed similar canopy structures.

Conclusions

The results generally supported the prediction, suggesting that lower growth of angiosperm trees (except L. recurvum on Kinabalu) in cold and nutrient-poor environments results in a sparser upper canopy, which allows shade-intolerant conifers to co-occur with angiosperm trees either as emergents or as codominants in the open canopy.     

Spatial and temporal variations in photosynthetic capacity of a temperate deciduous-evergreen forest

Key message

The understory evergreen trees showed maximal photosynthetic capacity in winter, while the overstory deciduous trees showed this capacity in spring. The time lag in productive ecophysiologically active periods between deciduous overstory and evergreen understory trees in a common temperate forest was clearly related to the amount of overstory foliage.

Abstract

In temperate forests, where deciduous canopy trees and evergreen understory trees coexist, understory trees experience great variation in incident radiation corresponding to canopy dynamics represented by leaf-fall and leaf-out. It is generally thought that changes in the light environment affect understory plants’ ecophysiological traits. Thus, to project and estimate annual energy, water, and carbon exchange between forests and the atmosphere, it is necessary to investigate seasonal variation in the ecophysiological activities of both evergreen trees in the understory and deciduous trees that make up the canopy/overstory. We conducted leaf-scale gas-exchange measurements and nitrogen content analyses for six tree species along their heights throughout a complete year. Photosynthetic capacity as represented by the maximum carboxylation rate (V _cmax25) and photosynthetic nitrogen use efficiency (PNUE) of deciduous canopy trees peaked immediately after leaf-out in late May, declined and stabilised during the mid-growing season, and drastically decreased just before leaf-fall. On the other hand, the timing of lowest V _cmax25 and PNUE for evergreen understory trees coincided with that of the highest values for canopy trees. Furthermore, understory trees’ highest values appeared just before canopy tree leaf-out, when incident radiation in the understory was highest. This implies that failing to consider seasonal variation in leaf ecophysiological traits for both canopy and understory trees could lead to serious errors in estimating ecosystem productivity and energy balance for temperate forests.