Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Habitat fragmentation,EFN‐bearing trees and ant communities: Ecological cascades in Atlantic Forest of northeastern Brazil

Habitat fragmentation has a marked impact on the functional composition of tropical forest tree assemblages, and such change is likely to cascade through other trophic levels. Here, we investigate how habitat fragmentation affects extrafloral nectary (EFN)‐bearing plants and ant functional groups known to attend EFNs in a fragmented landscape of the Atlantic Forest. Extrafloral nectary‐bearing trees were identified in 50 0.1‐ha plots located in forest fragments, edge and interior patches. Ants were surveyed in 30 1‐m² litter samples in each of 17 forest fragments and in forest interior. Extrafloral nectary‐bearing plants accounted for 19.9% of individuals and 10.5% of species and included both pioneer and shade‐tolerant species similarly rich in the three habitat types. However, shade‐tolerant individuals accounted for >80% of EFN‐bearing plants in forest interior, compared with 2% in forest edge and 29% in fragments. Forest edge and fragment plots had a third fewer EFN‐bearing individuals and species compared with forest interior. This appeared to have important implications for local ant communities as the density of EFN‐bearing trees was the most important variable explaining the species richness of arboreal dominant ants. Our results show that plant loser–winner replacements promoted by forest fragmentation can cascade through higher trophic levels, with implications for forest dynamics and biodiversity conservation.     

Ecological filtering by a dominant herb selects for shade tolerance in the tree seedling community of coastal dune forest

The regeneration niche is commonly partitioned along a gradient from shade-tolerant to shade-intolerant species to explain plant community assembly in forests. We examined the shade tolerance of tree seedlings in a subtropical coastal forest to determine whether the ecological filtering effect of a dominant, synchronously monocarpic herb (Isoglossa woodii) selects for species at either end of the light response continuum during the herb’s vegetative and reproductive phases. Photosynthetic characteristics of seedlings of 20 common tree species and the herb were measured. Seedlings were grown in the greenhouse at 12–14% irradiance, and their light compensation points measured using an open-flow gas exchange system. The light compensation points for the tree species were low, falling within a narrow range from 2.1 ± 0.8 μmol m⁻² s⁻¹ in Celtis africana to 6.4 ± 0.7 μmol m⁻² s⁻¹ in Allophylus natalensis, indicating general shade tolerance, consistent with a high and narrow range of apparent quantum yield among species (0.078 ± 0.002 mol CO₂ mol⁻¹ photon). Rates of dark respiration were significantly lower in a generalist pioneer species (Acacia karroo) than in a forest pioneer (C. africana), or in late successional phase forest species. We argue that the general shade tolerance, and phenotypic clustering of shade tolerance, in many tree species from several families in this system, is a result of ecological filtering by the prevailing low light levels beneath the I. woodii understorey, which excludes most light-demanding species from the seedling community.     

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade‐tolerance

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Continuous and Fluctuating Leaf Phenology of Adults and Seedlings of a Shade‐tolerant Emergent Tree,Dipterocarpus sublamellatus,in Malaysia

Continuous leaf phenology, with sequential production of metamers, is usually found in pioneer species, but is rare in shade‐tolerant species. Because of the nature of ‘continuous’ growth, continuous phenology has seldom been related to meteorological factors. We studied the leaf demography of seedlings and tall trees of a shade‐tolerant emergent species, Dipterocarpus sublamellatus (Dipterocarpaceae), in Malaysia. Although leaf phenology of D. sublamellatus was classified as continuous, the rates of leaf production and leaf fall were not constant and greatly fluctuated during the census period. Seasonal patterns of leaf production and fall were not synchronized across tall trees in most combinations or between seedlings and tall trees. Mean number of leaves produced per month and its seasonal fluctuation were greater in tall trees than in seedlings. Moreover, relationships between leaf phenology and meteorological factors (monthly rainfall, net radiation, maximum and minimum temperature, and minimum relative humidity) differed among trees. This suggests that endogenous rhythms, i.e., differences in allocation of resources to growth and storage among trees, are important to fluctuations in leaf phenology. These patterns are in contrast to patterns found in various pioneer species with continuous phenology, in which rates of leaf production and fall are generally constant. Different patterns might be caused by differences in allocation to growth and storage between pioneer and shade‐tolerant species, and might be related to their regeneration strategies.     

Disturbance Level Determines the Regeneration of Commercial Tree Species in the Eastern Amazon

The effects of reduced‐impact logging (RIL) on the regeneration of commercial tree species were investigated, as long‐term timber yields depend partly on the availability of seedlings in a managed forest. On four occasions during a 20‐month period in the Tapajós National Forest (Eastern Amazon, Brazil), seven commercial tree species were assessed as follows: the long‐lived pioneers Bagassa guianensis and Jacaranda copaia; the partially shade‐tolerant Hymenaea courbaril, Dipteryx odorata, and Carapa guianensis; and the totally shade‐tolerant Symphonia globulifera and Manilkara huberi. In 2439 10 × 10 m plots, all individuals < 20 cm diameter at breast height (dbh) were assessed over three intervals, before, during, and after the forest being logged. Before logging, the density of seedlings and saplings of the seven species did not change. Logged trees were spatially aggregated, with 9.2 percent of the plots being heavily impacted by logging. After logging, the recruitment rate increased more than the mortality rate, so that post‐harvesting densities of seedlings and saplings increased. The increase in density was concentrated in logged plots with more disturbances. It is concluded that post‐harvesting heterogeneity of micro‐environments created by RIL may be an important component to be taken into account for sustainable forest management and conservation of commercial species.     

Climate warming and precipitation redistribution modify tree–grass interactions and tree species establishment in a warm‐temperate savanna

Savanna tree–grass interactions may be particularly sensitive to climate change. Establishment of two tree canopy dominants, post oak (Quercus stellata) and eastern redcedar (Juniperus virginiana), grown with the dominant C₄ perennial grass (Schizachyrium scoparium) in southern oak savanna of the United States were evaluated under four climatic scenarios for 6 years. Tree–grass interactions were examined with and without warming (+1.5 °C) in combination with a long‐term mean rainfall treatment and a modified rainfall regime that redistributed 40% of summer rainfall to spring and fall, intensifying summer drought. The aim was to determine: (1) the relative growth response of these species, (2) potential shifts in the balance of tree–grass interactions, and (3) the trajectory of juniper encroachment into savannas, under these anticipated climatic conditions. Precipitation redistribution reduced relative growth rate (RGR) of trees grown with grass. Warming increased growth of J. virginiana and strongly reduced Q. stellata survival. Tiller numbers of S. scoparium plants were unaffected by warming, but the number of reproductive tillers was increasingly suppressed by intensified drought each year. Growth rates of J. virginiana and Q. stellata were suppressed by grass presence early, but in subsequent years were higher when grown with grass. Quercus stellata had overall reduced RGR, but enhanced survival when grown with grass, while survival of J. virginiana remained near 100% in all treatments. Once trees surpassed a threshold height of 1.1 m, both tiller number and survival of S. scoparium plants were drastically reduced by the presence of J. virginiana, but not Q. stellata. Juniperus virginiana was the only savanna dominant in which neither survival nor final aboveground mass were adversely affected by the climate scenario of warming and intensified summer drought. These responses indicate that climate warming and altered precipitation patterns will further accelerate juniper encroachment and woody thickening in a warm‐temperate oak savanna.     

Variation in wood density,wood water content,stem growth and mortality among twenty‐seven tree species in a tropical rainforest on Borneo Island

Abstract Interspecific variation among wood density (WD), wood water content (WWC), tree mortality and diameter at breast height (d.b.h.) increment was examined for 27 tree species (from 13 families), based on a 9‐year interval data obtained from a permanent 1‐ha forest plot setup for long‐term studies of tree dynamics in Kuala Belong rainforest, Brunei, on Borneo Island. The species were also categorized into three adult stature groups of understorey (maximum height ≤15–20 m tall, n = 14), midcanopy (maximum height, 20–30 m tall, n = 8) and canopy/emergent (>maximum height, >30 m tall, n = 5) tree species. All measured traits varied appreciably among species. Tree WD varied between 0.3 and 0.8 g cm⁻³, and exhibited the least coefficient of variation (14.7%). D.b.h. increment was low, averaging 1.05 (95% confidence limits: 0.57–2.13) mm year⁻¹ and was attributed to predominance of understorey species in the sampled plot. Overall, annual mortality was also low, averaging 2.73% per year. The three adult stature groups differed significantly in d.b.h. increment and WWC but not in tree mortality and WD. Across species and especially more so when phylogenetic effect is minimized, WD was negatively related to tree mortality and d.b.h. increment, while a positive trend was observed between d.b.h. increment and tree mortality. A negative trend was also detected between maximum plant height and WWC, which was interpreted as a consequence of increased evaporative demand and use of xylem stored water by taller trees in order to compensate for hydraulic limitations to water transport induced by frictional resistance. No doubt, the traits chosen may vary spatially, but the consistent interspecific patterns observed in this study among coexisting species of differing adult stature reflect ‘vertical’ niche differentiation and may help to explain population regulation in a multispecies ecosystem like tropical rainforest.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Hyperspectral remote detection of niche partitioning among canopy trees driven by blowdown gap disturbances in the Central Amazon

Advanced recruitment and neutral processes play important roles in determining tree species composition in tropical forest canopy gaps, with few gaps experiencing clear secondary successional processes. However, most studies are limited to the relatively limited spatial scales provided by forest inventory plots, and investigations over the entire range of gap size are needed to better understand how ecological processes vary with tree mortality events. This study employed a landscape approach to test the hypothesis that tree species composition and forest structural attributes differ between large blowdown gaps and relatively undisturbed primary forest. Spectral mixture analysis on hyperspectral satellite imagery was employed to direct field sampling to widely distributed sites, and blowdown plots were compared with undisturbed primary forest plots. Tree species composition and forest structural attributes differed markedly between gap and non-gap sites, providing evidence of niche partitioning in response to disturbance across the region. Large gaps were dominated by classic Neotropical pioneer genera such as Cecropia and Vismia, and average tree size was significantly smaller. Mean wood density of trees recovering in large gaps (0.55 g cm⁻³) was significantly lower than in primary forest plots (0.71 g cm⁻³), a difference similar to that found when comparing less dynamic (i.e., tree recruitment, growth, and mortality) Central Amazon forests with more dynamic Western Amazon forests. Based on results, we hypothesize that the importance of neutral processes weaken, and niche processes strengthen, in determining community assembly along a gradient in gap size and tree mortality intensity. Over evolutionary time scales, pervasive dispersal among colonizers could result in the loss of tree diversity in the pioneer guild through competitive exclusion. Results also underscore the importance of considering disturbance processes across the landscape when addressing forest carbon balance.     

Survival strategy of the endangered tree Acer catalpifolium Rehd., based on 13C fractionation

We conducted a field investigation and evaluation of ¹³C natural abundance to determine the growth habit and propagation strategy of Acer catalpifolium Rehd., a tree species native to China that is highly endangered. The results showed that A. catalpifolium is a K‐selected strategist and pioneer species. Its narrow ecological range limits its geographical distribution, and poor fecundity limits its population size. The analysis of ¹³C natural abundance showed that A. catalpifolium does not use organic matter for reproduction when its stand volume is less than 1.08 × 10⁶ cm³ or it is less than 18.6 m tall, but it does use this strategy when it has a sufficient 1.08 × 10⁶ cm³ stand volume or more or is taller than 18.6 m. If environmental conditions are not conducive (e.g., severe human disturbance, cliff edges, or fierce interspecific competition) to the continued growth of the tree, A. catalpifolium may allocate organic matter for reproduction. Human disturbance seems to promote the population expansion of A. catalpifolium. We provide our suggestions for the promotion and protection of A. catalpifolium as a species.     

Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.     

Long‐term tree fern dynamics linked to disturbance and shade tolerance

Question: Do New Zealand tree ferns have recognizable shade tolerance niches? Location: Lowland temperate rain forest of New Zealand (41°20′S, 174°58′E). Methods: Growth, death and recruitment of five tree fern species were estimated from a 38‐year record of stem heights, collected within a 2.25‐ha block of forest, and electron transport rates (ETR) of photosystem II of fronds were measured. Results: Two species of Cyathea were comparatively common (603 and 351 stems in total) and two were comparatively rare (155 and 17 stems in total) on the site. The common species had lower rates of growth, recruitment and mortality than the rare species, had skewed age distributions typical of shade‐tolerant species and were probably recruited soon after a catastrophic earthquake in 1855. The two rare species were failing to recruit under closed forests; their age distributions indicated that all had regenerated long after the earthquake. ETR were higher for faster‐growing than for the shade‐tolerant species. A tree fern that regenerates vegetatively from aerial buds, Dicksonia squarrosa, was common on the site (361 stems in total). Its age distribution suggested it was relatively shade tolerant, but its mortality and recruitment rates were much higher than those of the two shade‐tolerating Cyathea species, suggesting that this multi‐stemmed species functions differently from the monopodial Cyathea species. Conclusions: New Zealand Cyathea tree ferns occupy distinct niches along a shade tolerance spectrum and their relative abundances are strongly influenced by disturbance history. The study provides evidence that tree fern species differ strongly in their responses to canopy disturbance and are not ecologically equivalent.     

Techniques for seedling production of two native grasses: new perspectives for Brazilian Cerrado restoration

There is a major need for restoration of nonforest ecosystems in the Brazilian Cerrado, and planting seedlings is promising. However, there is no currently available information on nontree species production in nurseries. Grasses are dominant in the Cerrado herbaceous layer; therefore, we tested whether tree seedling production techniques can be used for two native grasses widely spread across the Cerrado ecoregion (Schizachyrium sanguineum and Loudetiopsis chrysothrix). We performed a factorial experiment with three levels: (1) seedbed substrate, (2) container volume, and (3) final substrate. Seeds were germinated in seedbeds with native soil or sand and, after emergence, transferred to 30, 55, or 290 cm³ containers filled with native soil, commercial substrate, or a mixture thereof. We evaluated seedling survival, height relative growth rate (RGR), tillering, and dry biomass. Survival rates after transfer from seedbeds to containers were greater than 80% for both species. Generally, species had greater height RGR, tillering, and biomass in the 290 cm³ containers than in the other containers, and the presence of native soil in the seedbeds, as well as in the containers, fostered seedling growth. However, the use of native soil is not recommended for large‐scale production, because its acquisition could degrade Cerrado areas. The commercial substrate improvement could be an alternative to increase seedling growth, since grasses survived well and presented greater growth in the largest container. Our study demonstrated that the techniques currently used in commercial nurseries for growing trees are suitable for both grasses, and substrates and container size drive seedling growth.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Diversity and spatial clustering of shade trees affect cacao yield and pathogen pressure in Costa Rican agroforests

The importance of the spatial organisation of individuals in explaining species coexistence within a community is widely recognised. However, few analyses of spatial structure have been performed on tropical agroforests.The main objective of this study was to highlight the links between spatial organisation of shade trees on the one hand, and shade tree species richness and cacao yield on the other, using data from 29 cacao agroforests in Costa Rica.A method of spatial statistics, Ripley's K-function, was used to analyse the spatial organisation of shade and cacao trees in the study plots. For each stand, the X and Y coordinates of ≥2.5-m-tall trees were recorded. In each plot we also assessed shade tree species richness and cacao yield (with total number of pods = number of pods damaged by frosty pod rot + number of healthy pods).Three types of stands were identified: the first was characterised by significant clustering of shade trees, the highest shade tree species richness (S = 6), and the highest number of damaged pods (139 pods ha^?1 year^?1). The second type was characterised by random spatial organisation of shade trees. The third type showed a trend towards regular organisation. Species richness of shade trees did not differ significantly between the last two types (S = 4 for both), nor did the number of damaged pods (56 pods ha^?1 year^?1 and 67 pods ha^?1 year^?1 respectively).Although the trends were not statistically significant for all the variables in our data set, the clustered spatial structure appears to favour a synergy between environmental (tree species richness), and provisioning (cacao production) services.     

The effect of different anthropogenic disturbances on litterfall of a dominant pioneer rain forest tree in Gabon

Tropical rain forests worldwide are affected by anthropogenic disturbances, and secondary forests that develop afterwards are often dominated by pioneer tree species, but the consequences of different anthropogenic disturbances on nutrient cycling are poorly understood. Because most nutrient cycling in tropical rain forests occurs through litterfall and in the soil organic layer, we measured litterfall of a widespread and dominant pioneer tree, okoume (Aucoumea klaineana, Burseraceae) in Gabon, in one forest previously disturbed by logging and in another by agriculture. Litterfall of okoume trees, measured over 14 months, was 18.2 Mg ha⁻¹ year⁻¹ in the formerly logged forest, which was 72.6% greater than in the forest more recently disturbed by agriculture (10.6 Mg ha⁻¹ year⁻¹). Litter decomposition rates were more rapid in the formerly logged forest, and this may explain why litter thickness was 32% lower in the formerly logged forest, despite the greater litterfall within it. Our results highlight that two widespread anthropogenic disturbances in Gabonese rain forests have significantly different consequences on litterfall of a dominant tree and therefore are likely to have different effects on nutrient cycling and forest ecosystem function.     

Leaf attributes and tree growth in a tropical dry forest

Questions: How are leaf attributes and relative growth rate (RGR) of the dominant tree species of tropical deciduous forest (TDF) affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional attributes with each other? Can leaf attributes singly or in combination predict the growth rate of tree species of TDF? Location: Sonebhadra district of Uttar Pradesh, India. Methods: Eight leaf attributes, specific leaf area (SLA); leaf carbon concentration (LCC); leaf nitrogen concentration (LNC); leaf phosphorus concentration (LPC); chlorophyll concentration (Chl), mass‐based stomatal conductance (Gs_mass); mass based photosynthetic rate (A_mass); intrinsic water use efficiency (WUEi); and relative growth rate (RGR), of six dominant tree species of a dry tropical forest on four sites were analysed for species, site and season effects over a 2‐year period. Step‐wise multiple regression was performed for predicting RGR from mean values of SMC and leaf attributes. Path analysis was used to determine which leaf attributes influence RGR directly and which indirectly. Results: Species differed significantly in terms of all leaf attributes and RGR. The response of species varied across sites and seasons. The attributes were positively interrelated, except for WUEi, which was negatively related to all other attributes. The positive correlation was strongest between Gs_mass and A_mass and the negative correlation was strongest between Gs_mass and WUEi. Differences in RGR due to site were not significant when soil moisture was controlled, but differences due to season remained significant. The attributes showed plasticity across moisture gradients, which differed among attributes and species. Gs_mass was the most plastic attribute. Among the six species, Terminalia tomentosa exhibited the greatest plasticity in six functional attributes. In the step‐wise multiple regression, A_mass, SLA and Chl among leaf attributes and SMC among environmental factors influenced the RGR of tree species. Path analysis indicated the importance of SLA, LNC, Chl and A_mass in determining RGR. Conclusion: A _mass, SMC, SLA and Chl in combination can be used to predict RGR but could explain only three‐quarters of the variability in RGR, indicating that other traits/factors, not studied here, are also important in modulating growth of tropical trees. RGR of tree species in the dry tropical environment is determined by soil moisture, whereas the response of mature trees of different species is modulated by alterations in key functional attributes such as SLA, LNC and Chl.     

Recovery from clearing,cyclone and fire in rain forests of Tonga,South Pacific: Vegetation dynamics 1995–2005

Abstract In late 2001 a category 3 cyclone impacted forest plots that were established in Tonga in 1995, and additionally, one plot was accidentally burned by an escaped land‐clearing fire. Subsequent surveys provide observations of 10 years of forest dynamics in this poorly studied region, and the first reported observations of large interannual variation in juvenile (seedling and sapling) abundance in the western tropical Pacific. The severely disturbed (burned) plot was initially colonized by a non‐native early pioneer, Carica papaya L., but 3.5 years later a native pioneer, Macaranga harveyana (Muell. Arg.) Muell. Arg., was the most abundant tree species. The seedling layer included some long‐lived pioneers and shade‐tolerant species. Two mature forest plots affected only by the cyclone changed very little over a decade. Late‐successional shade‐tolerant species that dominated the overstory were also abundant as seedlings and saplings. This is in contrast with a 30‐ to 40‐year‐old, formerly cultivated, secondary forest plot that still shows no recruitment of late‐successional dominants, in spite of the proximity of remnant forest patches. This study suggests differing pathways of succession following shifting cultivation versus cyclone and fire disturbances in Tonga. Land use legacies appear to have a long‐lasting effect on community composition.