Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

Radial and vertical variation of wood nutrients in Bornean tropical forest trees

Nitrogen, phosphorus, potassium, calcium, and magnesium concentrations in woody tissue are poorly documented, but are necessary for understanding whole-tree nutrient use and storage. Here, we report how wood macronutrient concentrations vary radially and along the length of a tree for 10 tropical tree species in Sabah, Malaysia. Bark nutrient concentrations were consistently high: 2.9–13.7 times greater than heartwood depending on the nutrient. In contrast, within the wood both the radial (sapwood vs. heartwood) and vertical (trunk bottom vs. trunk middle) variation was modest. Higher concentrations in sapwood relative to heartwood provide empirical support for wood nutrient resorption during sapwood senescence. Dipterocarp species showed resorption rates of 25.3 ± 7.1% (nitrogen), 62.7 ± 11.9% (phosphorus), and 56.2 ± 12.5% (potassium), respectively, while non-dipterocarp species showed no evidence of nutrient resorption in wood. This suggests that while dipterocarps have lower wood nutrient concentrations, this family is able to compensate for this by using wood nutrient resorption as an efficient nutrient conservation mechanism. In contrast to other nutrients, calcium and magnesium tended to accumulate in heartwood. Wood density (WD) showed little vertical variation along the trunk. Across the species (WD range of 0.33 to 0.94 mg/cm³), WD was negatively correlated with wood P and K concentration and positively correlated with wood Ca concentration. As our study showed exceptionally high nutrient concentrations in the bark, debarking and leaving the bark of the harvested trees on site during logging operations could substantially contribute to maintaining nutrients within forest ecosystems.     

Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees

This study examined the linkage between xylem vulnerability, stomatal response to leaf water potential (Ψ_L), and loss of leaf turgor in eight species of seasonally dry tropical forest trees. In order to maximize the potential variation in these traits species that exhibit a range of leaf habits and phenologies were selected. It was found that in all species stomatal conductance was responsive to Ψ_L over a narrow range of water potentials, and that Ψ_L inducing 50% stomatal closure was correlated with both the Ψ_L inducing a 20% loss of xylem hydraulic conductivity and leaf water potential at turgor loss in all species. In contrast, there was no correlation between the water potential causing a 50% loss of conductivity in the stem xylem, and the water potential at stomatal closure (Ψ_SC) amongst species. It was concluded that although both leaf and xylem characters are correlated with the response of stomata to Ψ_L, there is considerable flexibility in this linkage. The range of responses is discussed in terms of the differing leaf‐loss strategies exhibited by these species.     

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.     

Fire survival of lowland tropical rain forest trees in relation to stem diameter and topographic position

The objective of this study was to relate patterns in forest structure, tree species diversity, and tree species composition to stem diameters and topography in unburned, once burned and twice burned lowland dipterocarp rain forests in East Kalimantan, Indonesia. To do this four unburned old growth forests were compared with three forests that burned once (1997/1998) and three forests that burned twice (1982/1983 and 1997/1998). Fire resulted in a strong reduction of climax tree density which was negatively related to tree diameter. However, a disproportionate reduction in small diameter understorey climax tree species occurred only after repeated fires. Climax tree species in both burned forest types were most common in swamps, river valleys and on lower slopes, while their density was much lower on places higher along hillsides. In unburned forest the opposite was observed, with climax tree density increasing steadily from swamp and river valleys to upper slopes and ridges. In contrast to climax trees, pioneer trees were abundant throughout the burned forest, with highest numbers on hill sides and ridges. Our results indicate that both diameter and topographic position of trees strongly affect their fire survival chances in tropical lowland forests.     

Plant functional types broadly describe water use strategies in the Caatinga,a seasonally dry tropical forest in northeast Brazil

1. In seasonally dry tropical forests, plant functional type can be classified as deciduous low wood density, deciduous high wood density, or evergreen high wood density species. While deciduousness is often associated with drought‐avoidance and low wood density is often associated with tissue water storage, the degree to which these functional types may correspond to diverging and unique water use strategies has not been extensively tested.
2. We examined (a) tolerance to water stress, measured by predawn and mid‐day leaf water potential; (b) water use efficiency, measured via foliar δ¹³C; and (c) access to soil water, measured via stem water δ¹⁸O.
3. We found that deciduous low wood density species maintain high leaf water potential and low water use efficiency. Deciduous high wood density species have lower leaf water potential and variable water use efficiency. Both groups rely on shallow soil water. Evergreen high wood density species have low leaf water potential, higher water use efficiency, and access alternative water sources. These findings indicate that deciduous low wood density species are drought avoiders, with a specialized strategy for storing root and stem water. Deciduous high wood density species are moderately drought tolerant, and evergreen high wood density species are the most drought tolerant group.
4. Synthesis. Our results broadly support the plant functional type framework as a way to understand water use strategies, but also highlight species‐level differences.

     

Interactions between body size,abundance, seasonality,and phenology in forest beetles

Body size correlates with a large number of species traits, and these relationships have frequently been used to explain patterns in populations, communities, and ecosystems. However, diverging patterns occur, and there is a need for more data on different taxa at different scales. Using a large dataset of 155,418 individual beetles from 588 species collected over 13 years of sampling in Norway, we have explored whether body size predicts abundance, seasonality, and phenology in insects. Seasonality is estimated here by flight activity period length and phenology by peak activity. We develop several methods to estimate these traits from low‐resolution sampling data. The relationship between abundance and body size was significant and as expected; the smaller species were more abundant. However, smaller species tended to fly for longer periods of the summer and peaked in midsummer, while larger species were restricted to shorter temporal windows. Further analysis of repeated sampling from a single location suggested that smaller species had increased flight period lengths in warmer years, but larger species showed the opposite pattern. The results 1) indicate that smaller species are likely to be disproportionately valuable in ecological interactions, and 2) provide potential insights into the traits influencing the vulnerability of some larger species to disturbances and climate change.     

Growth and mortality of trees in Amazonian tropical rain forest in Ecuador

Abstract. Structural changes are analysed in four samples representing 4 ha, two line transects and two hectare plots, of Amazonian tropical lowland rain forest in northern Ecuador. Only trees with a DBH ≥ 10 cm were included. A sample of floodplain forest in Añangu represents the largest turnover found in tropical forests (stand half-life = 23 yr). The line transect and hectare plot both of tierra firme forest in Añangu have the same turnover (37 yr) and were balanced for death and in-growth of both individuals and wood (basal area). The 1-ha tierra firme sample in Cuyabeno had a turnover of 67 yr and was in a growing phase. The floodplain line transect in Añangu was in a phase of structural breakdown. However, the floodplain line transect had the largest growth of basal area per tree (23.4 cm²/yr). The tierra firme samples had a growth of 9.6, 10.1, and 13.6 cm²/yr. Most of the dead trees fell with some uprooting in three of the four samples. However, no significant difference in the distribution of mode of death was found between the four samples. Death was independent of topography and the dead trees were randomly distributed. As the trees grow up they occupy more space and larger trees (DBH ≥ 15 cm) become more uniformly distributed, whereas smaller trees (DBH ≤ 15 cm) were randomly distributed. Our study confirms that plots of 1 ha are not sufficient to include representative samples of different stages of forest structure.     

Morphological and physiological differentiation of seedlings between dry and wet habitats in a tropical dry forest

A common observation in tropical dry forests is the habitat preference of tree species along spatial soil water gradients. This pattern of habitat partitioning might be a result of species differentiation in their strategy for using water, along with competing functions such as maximizing water exploitation and tolerating soil water stress. We tested whether species from drier soil conditions exhibited a tolerance strategy compared with that of wet-habitat species. In a comparison of 12 morphophysiological traits in seedlings of 10 closely related dry and wet-habitat species pairs, we explored what trade-offs guide differentiation between habitats and species. Contrary to our expectations, dry-habitat species showed mostly traits associated with an exploitation strategy (higher carbon assimilation capacity, specific leaf area and leaf-specific conductivity and lower water-use efficiency). Strikingly, dry-habitat species tended to retain their leaves longer during drought. Additionally, we detected multiple strategies to live within each habitat, in part due to variation of strategies among lineages, as well as functional differentiation along the water storage capacity-stem density (xylem safety) trade-off. Our results suggest that fundamental trade-offs guide functional niche differentiation among tree species expressed both within and between soil water habitats in a tropical dry forest.     

浙江古田山亚热带常绿阔叶林开花物候:气候因素、系统发育关系和功能性状的影响

植物的开花物候受气候因素、植物系统发育关系和功能性状的影响。然而当前植物开花物候研究中未见同时考虑这3个因素的报道。为了解它们相互之间的影响, 本研究利用中国东部地区浙江省古田山国家级自然保护区亚热带常绿阔叶林24 ha大样地(GTS; 118°03′50′′-118°11′12.2′′ E, 29°10′19′′-29°17′41′′ N)设置的130个种子雨收集器5年的开花数据检验这3个因素对开花的影响。结果表明, 古田山植物的开花高峰期集中在5月, 群落开花格局明显受温度和降雨的影响。利用植物DNA条形码数据研究发现, 植物间系统发育关系对古田山植物开花时间有显著影响, 亲缘关系近的物种开花时间更相近。植物的平均开花时间受最大树高的影响, 但不受传粉方式、花色、种子质量和扩散方式的影响。该研究结果说明气候因素、植物系统发育关系和功能性状都可能影响植物开花物候格局, 同时考虑这3个因素能够帮助我们更好地理解开花物候格局。     

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae werecompared to test in part a hypothesis that the forest understorey plays anintegral role in drought response. At Danum, Sabah, the relatively commonspecies Dimorphocalyx muricatus is associated with ridgeswhilst another species, Mallotus wrayi, occurs widely bothon ridges and lower slopes. Sets of subplots within two 4 -hapermanent plots in this lowland dipterocarp rain forest, were positioned onridges and lower slopes. Soil water potentials were recorded in1995–1997,and leaf water potentials were measured on six occasions. Soil water potentialson the ridges (–0.047 MPa) were significantly lower than onthe lower slopes (–0.012 MPa), but during the driest periodin May 1997 they fell to similarly low levels on both sites (–0.53MPa). A weighted 40-day accumulated rainfall index was developedtomodel the soil water potentials. At dry times, D.muricatus(ridge) had significantly higher pre-dawn (–0.21 v.–0.57 MPa) and mid-day (–0.59 v.–1.77 MPa) leaf water potentials than M.wrayi (mean of ridge and lower slope). Leaf osmotic potentials ofM. wrayi on the ridges were lower (–1.63MPa) than on lower slopes (–1.09 MPa), withthose for D. muricatus being intermediate (–1.29MPa): both species adjusted osmotically between wet and dry times.D. muricatus trees were more deeply rooted thanM. wrayi trees (97 v. 70cm). M. wrayi trees had greaterlateral root cross-sectional areas than D. muricatus treesalthough a greater proportion of this sectional area for D.muricatus was further down the soil profile. D.muricatus appeared to maintain relatively high water potentialsduring dry periods because of its access to deeper water supplies and thus itlargely avoided drought effects, but M. wrayi seemed to bemore affected yet tolerant of drought and was more plastic in its response. Theinteraction between water availability and topography determines these species'distributions and provides insights into how rain forests can withstandoccasional strong droughts.     

The relationship between tree biodiversity and biomass dynamics changes with tropical forest succession

Theory predicts shifts in the magnitude and direction of biodiversity effects on ecosystem function (BEF) over succession, but this theory remains largely untested. We studied the relationship between aboveground tree biomass dynamics (Δbiomass) and multiple dimensions of biodiversity over 8–16 years in eight successional rainforests. We tested whether successional changes in diversity–Δbiomass correlations reflect predictions of niche theories. Diversity–Δbiomass correlations were positive early but weak later in succession, suggesting saturation of niche space with increasing diversity. Early in succession, phylogenetic diversity and functional diversity in two leaf traits exhibited the strongest positive correlations with Δbiomass, indicating complementarity or positive selection effects. In mid‐successional stands, high biodiversity was associated with greater mortality‐driven biomass loss, i.e. negative selection effects, suggesting successional niche trade‐offs and loss of fast‐growing pioneer species. Our results demonstrate that BEF relationships are dynamic across succession, thus successional context is essential to understanding BEF in a given system.     

Field observed relationships between biodiversity and ecosystem functioning during secondary succession in a tropical lowland rainforest

The relationship between biodiversity and ecosystem functioning (BEF) is one of the most concerned topics in ecology. However, most of the studies have been conducted in controlled experiments in grasslands, few observational field studies have been carried out in forests. In this paper, we report variations of species diversity, functional diversity and aboveground biomass (AGB) for woody plants (trees and shrubs) along a chronosequence of four successional stages (18-year-old fallow, 30-year-old fallow, 60-year-old fallow, and old-growth forest) in a tropical lowland rainforest recovered after shifting cultivation on Hainan Island, China. Fifty randomly selected sample plots of 20 m × 20 m were investigated in each of the four successional stages. Four functional traits (specific leaf area, wood density, maximum species height and leaf dry matter content) were measured for each woody plants species and the relationships between species/functional diversity and AGB during secondary succession were explored. The results showed that both plant diversity and AGB recovered gradually with the secondary succession. AGB was positively correlated with both species and functional diversity in each stage of succession. Consistent with many controlled experimental results in grasslands, our observational field study confirms that ecosystem functioning is closely related to biodiversity during secondary succession in species rich tropical forests.     

Disturbance history mediates climate change effects on subtropical forest biomass and dynamics

The responses of forest communities to interacting anthropogenic disturbances like climate change and logging are poorly known. Subtropical forests have been heavily modified by humans and their response to climate change is poorly understood. We investigated the 9‐year change observed in a mixed conifer‐hardwood Atlantic forest mosaic that included both mature and selectively logged forest patches in subtropical South America. We used demographic monitoring data within 10 1 ha plots that were subjected to distinct management histories (plots logged until 1955, until 1987, and unlogged) to test the hypothesis that climate change affected forest structure and dynamics differentially depending on past disturbances. We determined the functional group of all species based on life‐history affinities as well as many functional traits like leaf size, specific leaf area, wood density, total height, stem slenderness, and seed size data for the 66 most abundant species. Analysis of climate data revealed that minimum temperatures and rainfall have been increasing in the last few decades of the 20th century. Floristic composition differed mainly with logging history categories, with only minor change over the nine annual census intervals. Aboveground biomass increased in all plots, but increases were higher in mature unlogged forests, which showed signs of forest growth associated with increased CO₂, temperature, and rainfall/treefall gap disturbance at the same time. Logged forests showed arrested succession as indicated by reduced abundances of Pioneers and biomass‐accumulators like Large Seeded Pioneers and Araucaria, as well as reduced functional diversity. Management actions aimed at creating regeneration opportunities for long‐lived pioneers are needed to restore community functional diversity, and ecosystem services such as increased aboveground biomass accumulation. We conclude that the effects of climate drivers on the dynamics of Brazilian mixed Atlantic forests vary with land‐use legacies, and can differ importantly from the ones prevalent in better known tropical forests.     

Nutrient fluxes via litterfall and leaf litter decomposition vary across a gradient of soil nutrient supply in a lowland tropical rain forest

The extent to which plant communities are determined by resource availability is a central theme in ecosystem science, but patterns of small-scale variation in resource availability are poorly known. Studies of carbon (C) and nutrient cycling provide insights into factors limiting tree growth and forest productivity. To investigate rates of tropical forest litter production and decomposition in relation to nutrient availability and topography in the absence of confounding large-scale variation in climate and altitude we quantified nutrient fluxes via litterfall and leaf litter decomposition within three distinct floristic associations of tropical rain forest growing along a soil fertility gradient at the Sepilok Forest Reserve (SFR), Sabah, Malaysia. The quantity and nutrient content of small litter decreased along a gradient of soil nutrient availability from alluvial forest (most fertile) through sandstone forest to heath forest (least fertile). Temporal variation in litterfall was greatest in the sandstone forest, where the amount of litter was correlated negatively with rainfall in the previous month. Mass loss and N and P release were fastest from alluvial forest litter, and slowest from heath forest litter. All litter types decomposed most rapidly in the alluvial forest. Stand-level N and P use efficiencies (ratios of litter dry mass to nutrient content) were greatest for the heath forest followed by the sandstone ridge, sandstone valley and alluvial forests, respectively. We conclude that nutrient supply limits productivity most in the heath forest and least in the alluvial forest. Nutrient supply limited productivity in sandstone forest, especially on ridge and hill top sites where nutrient limitation may be exacerbated by reduced rates of litter decomposition during dry periods. The fluxes of N and P varied significantly between the different floristic communities at SFR and these differences may contribute to small-scale variation in species composition.