Fine root dynamics and trace gas fluxes in two lowland tropical forest soils

Fine root dynamics have the potential to contribute significantly to ecosystem‐scale biogeochemical cycling, including the production and emission of greenhouse gases. This is particularly true in tropical forests which are often characterized as having large fine root biomass and rapid rates of root production and decomposition. We examined patterns in fine root dynamics on two soil types in a lowland moist Amazonian forest, and determined the effect of root decay on rates of C and N trace gas fluxes. Root production averaged 229 (±35) and 153 (±27) g m^?2 yr^?1 for years 1 and 2 of the study, respectively, and did not vary significantly with soil texture. Root decay was sensitive to soil texture with faster rates in the clay soil (k=?0.96 year^?1) than in the sandy loam soil (k=?0.61 year^?1), leading to greater standing stocks of dead roots in the sandy loam. Rates of nitrous oxide (N₂O) emissions were significantly greater in the clay soil (13±1 ng N cm^?2 h^?1) than in the sandy loam (1.4±0.2 ng N cm^?2 h^?1). Root mortality and decay following trenching doubled rates of N₂O emissions in the clay and tripled them in sandy loam over a 1‐year period. Trenching also increased nitric oxide fluxes, which were greater in the sandy loam than in the clay. We used trenching (clay only) and a mass balance approach to estimate the root contribution to soil respiration. In clay soil root respiration was 264–380 g C m^?2 yr^?1, accounting for 24% to 35% of the total soil CO₂ efflux. Estimates were similar using both approaches. In sandy loam, root respiration rates were slightly higher and more variable (521±206 g C m² yr^?1) and contributed 35% of the total soil respiration. Our results show that soil heterotrophs strongly dominate soil respiration in this forest, regardless of soil texture. Our results also suggest that fine root mortality and decomposition associated with disturbance and land‐use change can contribute significantly to increased rates of nitrogen trace gas emissions.     

Leaf nutrients in relation to stature and life form in tropical rain forest

Abstract. To document the relationship between a plant's position in the canopy and its leaf nutrient content, leaf nitrogen and phosphorus were determined for 30 species growing in mature evergreen lowland rain forest at La Selva Biological Station, Costa Rica. Species that grow either in the understory, midstory, or the canopy were selected. Species were further separated into three life forms: self-supporting monocots, self-supporting dicots, and climbers. Mass-based nutrient concentrations were expected to decrease with stature, as has been reported in studies of other forests. In fact, mass-based nitrogen and phosphorus did not vary significantly among the three adult-stature classes, although area-based values differed greatly: canopy plants averaged 60 % more nitrogen and 90 % more phosphorus per unit leaf area than understory plants. Differences in leaf characteristics were evident among the three life forms. Most notably, area-based phosphorus and leaf specific mass were lowest in climbers, intermediate in self-supporting dicots, and highest in self-supporting monocots. These results support the characterization of climbers as investing in inexpensive structures, perhaps in order to gain competitive advantage in light capture by allocating resources to maximize elongation rates.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Arthropod faunal similarity of Bornean rain forest trees

ABSTRACT. 1. Arthropod samples, collected by insecticide fogging ten Bornean lowland rain forest trees, were sorted to species and used to study some of the determinants of faunal similarity of these trees.
2. Similarity indices were calculated between pairs of trees for guilds of beetles, beetle families and for different orders and other families of insects.
3. Multiple regression analyses of five independent variables of the trees (taxonomic similarity, distance apart, vertical canopy overlap, similarity in tree height, epiphytic load) were carried out against these similarity indices.
4. In eight of the eighteen insect groups analysed, taxonomic similarity of the trees is the most important variable affecting their species composition, accounting for a maximum of 29.4% of the variation.
5. For the Homoptera, Gryllidae, Anthicidae, Chrysomeloidea and scavengers, distance between the trees and the similarity of the trees'epiphyte load have a greater effect on their faunal similarities than taxonomic similarity of the trees.
6. The maximum variation accounted for by variables with significant T ratios was 38.6% in the Anthicidae. In four groups, the Formicidae, Heteroptera, Galerucinae and Corylophidae, none of the variables was significant.
7. These data may support the view that for many insect groups there is less host-specificity in the tropics than in temperate regions.     

Fine root biomass under light gap openings in an Amazon rain forest

Summary Belowground processes in light gap openings are poorly understood, particularly in tropical forests. Fine roots in three zones of light gap openings and adjacent intact forest were regularly measured in buried bags and surface litter envelopes for 2 years. Fine root biomass does not vary significantly within gaps for either buried bags or for surface litter envelopes. When entire gaps are compared without regard for within gap zones, root growth into both surface litter and buried bags is significantly different between gaps, with highest rates of fine root biomass accumulation in the smallest gap. These results suggest that the aboveground within-gap zones do not result in a congruent pattern of below-ground zonation. Gap size, decomposition of the fallen tree, and pre-gap fine root growth rates should be considered to determine fine root growth patterns following the formation of light gap openings.     

Guild structure of arthropods from Bornean rain forest trees

ABSTRACT. 1. 23275 arthropods collected by insecticide fogging from ten Bornean lowland rain forest trees were sorted to approximately 3000 species and assigned to guilds using two sets of criteria.
2. The rank order for proportions of guilds of species but not individuals is similar in tropical and temperate canopy samples.
3. Misplacement in the guild assignments of a few species can cause important differences in the proportions of certain guilds. This can seriously affect the results of comparisons of different samples, and views on proportional constancy of guilds of species on different trees.
4. Guilds of arthropod species such as chewers, suckers, all phytophages, predators, ants and tourists, occur in constant proportions in samples from different tropical trees and this constancy of proportion is similar in samples from groups of'closely related'and'distantly related'trees.
5. The guild concept of community structure and problems in assigning species to guilds are considered with respect to arthropod samples. The conclusions on constancy of proportion for guilds of species are tempered by remarks on the problems of guild composition.     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Plurality of tree species responses to drought perturbation in Bornean tropical rain forest

Drought perturbation driven by the El Niño Southern Oscillation (ENSO) is a principal stochastic variable determining the dynamics of lowland rain forest in S.E. Asia. Mortality, recruitment and stem growth rates at Danum in Sabah (Malaysian Borneo) were recorded in two 4-ha plots (trees ≥ 10 cm gbh) for two periods, 1986–1996 and 1996–2001. Mortality and growth were also recorded in a sample of subplots for small trees (10 to <50 cm gbh) in two sub-periods, 1996–1999 and 1999–2001. Dynamics variables were employed to build indices of drought response for each of the 34 most abundant plot-level species (22 at the subplot level), these being interval-weighted percentage changes between periods and sub-periods. A significant yet complex effect of the strong 1997/1998 drought at the forest community level was shown by randomization procedures followed by multiple hypothesis testing. Despite a general resistance of the forest to drought, large and significant differences in short-term responses were apparent for several species. Using a diagrammatic form of stability analysis, different species showed immediate or lagged effects, high or low degrees of resilience or even oscillatory dynamics. In the context of the local topographic gradient, species’ responses define the newly termed perturbation response niche. The largest responses, particularly for recruitment and growth, were among the small trees, many of which are members of understorey taxa. The results bring with them a novel approach to understanding community dynamics: the kaleidoscopic complexity of idiosyncratic responses to stochastic perturbations suggests that plurality, rather than neutrality, of responses may be essential to understanding these tropical forests. The basis to the various responses lies with the mechanisms of tree-soil water relations which are physiologically predictable: the timing and intensity of the next drought, however, is not. To date, environmental stochasticity has been insufficiently incorporated into models of tropical forest dynamics, a step that might considerably improve the reality of theories about these globally important ecosystems.     

Habitat associations of Sterculiaceae trees in a Bornean rain forest plot

Questions: 1. Are trees in a Bornean tropical rain forest associated with a particular habitat? 2. Does the strength of habitat association with the species‐specific optimal habitat increase with tree size? Location : A 52‐ha plot in a mixed dipterocarp forest in a heterogeneous landscape at the Lambir Hills National Park, Sarawak, East Malaysia. Methods : Ten species from the Sterculiaceae were chosen as representative of all species in the plot, on the assumption that competition among closely related species is more stringent than that among more distantly related taxa. Their habitat associations were tested using data from a 52‐ha plot by a torus‐translation test. Results : The torus‐translation test showed that eight out of the ten species examined had significant association with at least one habitat. We could not find negative species‐habitat associations for rare species, probably due to their small sample sizes. Among four species small trees were less strongly associated with habitat than large trees, implying competitive exclusion of trees in suboptimal habitats. The other four species showed the opposite pattern, possibly owing to the smaller sample size of large trees. A habitat had a maximum of three species with which it was significantly positively associated. Conclusions : For a species to survive in population equilibrium in a landscape, habitats in which ‘source’ subpopulations can be sustained without subsidy from adjacent habitats are essential. Competition is most severe among related species whose source subpopulations share the same habitat. On the evidence of source subpopulations identified by positive species‐habitat association, species‐habitat association reduces the number of confamilial competitors. Our results therefore indicate that edaphic niche specialization contributes to coexistence of species of Sterculiaceae in the plot, consistent with the expectations of equilibrium hypotheses.     

Tree species distributions across five habitats in a Bornean rain forest

We examined the distribution of tree species across five habitats in 69 small plots within a single watershed of the Gunung Palung National Park, West Kalimantan, Indonesia (GPNP). The spatially complex distribution and close proximity of habitats provided an opportunity to test habitat specificity of tree species across strong environmental gradients, in a situation where dispersal into ‘inappropriate’ habitat should not be a rare event. Habitat had a weak influence on community structure, although species diversity was lower in the alluvium and peat habitats. Association tests based on two randomization models (spatially independent and explicit) were used to examine habitat distribution of 55 ‘common’ and 142 ‘frequent’ taxa. The general patterns were similar in the two models but the interpretation of specific patterns depended greatly on assumptions about dispersal ability. A majority (67%) of the common species was significantly associated with a single habitat, while few were restricted to one habitat. A small proportion (16%) of the species appear to be habitat generalists. The peat habitat had the most profound effect on species distribution. Overall, a large amount of variation was found in the degree of habitat specificity, even within speciose groups. No obvious evolutionary or ecological correlates with degree of habitat specificity were found. These results suggest that a mixture of stochastic and deterministic processes determine species distribution even across strong environmental gradients.     

Fine root mass in relation to soil N supply in a cool temperate forest

Soil inorganic nitrogen supply and fine root mass in the top layers of mineral soil (0–5 and 5–10cm) were investigated at upper and lower sites of a cool temperate forest where Fagus crenata and Quercus crispula dominate. At both sites, soil inorganic nitrogen supply was greatest in the 0–5cm layer. The predominant forms of soil inorganic nitrogen supply were NH₄⁺-N at the upper site and NO₃^–-N at the lower site. Fine roots were concentrated in the 0–5cm layer at the upper site, but not at the lower site. The form of supplied soil inorganic nitrogen supply can be important in determining the vertical distribution of fine roots.     

Tree diversity,composition, forest structure and aboveground biomass dynamics after single and repeated fire in a Bornean rain forest

Forest fires remain a devastating phenomenon in the tropics that not only affect forest structure and biodiversity, but also contribute significantly to atmospheric CO2. Fire used to be extremely rare in tropical forests, leaving ample time for forests to regenerate to pre-fire conditions. In recent decades, however, tropical forest fires occur more frequently and at larger spatial scales than they used to. We studied forest structure, tree species diversity, tree species composition, and aboveground biomass during the first 7 years since fire in unburned, once burned and twice burned forest of eastern Borneo to determine the rate of recovery of these forests. We paid special attention to changes in the tree species composition during burned forest regeneration because we expect the long-term recovery of aboveground biomass and ecosystem functions in burned forests to largely depend on the successful regeneration of the pre-fire, heavy-wood, species composition. We found that forest structure (canopy openness, leaf area index, herb cover, and stem density) is strongly affected by fire but shows quick recovery. However, species composition shows no or limited recovery and aboveground biomass, which is greatly reduced by fire, continues to be low or decline up to 7 years after fire. Consequently, large amounts of the C released to the atmosphere by fire will not be recaptured by the burned forest ecosystem in the near future. We also observed that repeated fire, with an inter-fire interval of 15 years, does not necessarily lead to a huge deterioration in the regeneration potential of tropical forest. We conclude that burned forests are valuable and should be conserved and that long-term monitoring programs in secondary forests are necessary to determine their recovery rates, especially in relation to aboveground biomass accumulation.     

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.     

Variation in leaf stomatal traits of 28 tree species in relation to gas exchange along an edaphic gradient in a Bornean rain forest

? Premise of the study: Quantifying variation in functional traits associated with shifts in the species composition of plant communities along resource gradients helps identify environmental attributes important for community assembly. Stomates regulate the balance between carbon assimilation and water status in plants. If environmental attributes affecting photosynthetic water-use efficiency govern species distribution along an edaphic gradient, then adaptive variation in stomatal traits of plant species specializing on different soils should reflect belowground resource availability. ? Methods: We tested this hypothesis by quantifying stomatal trait variation in understory saplings of 28 Bornean tree species in relation to gas exchange and water-use efficiency (WUE). ? Key results: Comparisons between congeneric specialists of the more fertile, moister clay and the less fertile, well-drained sandy loam revealed little evidence of similar shifts in stomatal traits across genera, nor was stomatal pore index correlated with g(max), A(max), or WUE (A(max)/g(max) or Δ(13)C), suggesting that stomates may be overbuilt in these shaded juveniles. A(max) was higher on sandy loam, likely due to higher understory irradiance there, but there were no other significant differences in gas exchange or WUE. ? Conclusions: Despite substantial diversity in stomatal anatomy, there were few strong relationships between stomatal, photosynthetic, and WUE traits in relation to soil resources. Routine differences in water availability therefore may not exert a dominant control on the distributions of these Bornean tree species. Furthermore, the clades represented by these 12 genera may possess alternative functional designs enabling photosynthetic WUE that is sufficient to these humid, understory environments, due to whole plant-functional integration of stomatal traits with other, unmeasured traits influencing gas exchange.     

Fine root dynamics in Slovenian beech forests in relation to soil temperature and water availability

Key message

Fine root ingrowth and mortality of European beech are related to evapotranspiration, cumulative forest floor precipitation, soil temperature and water content, which are affected by forest management and gap creation.

Abstract

The ingrowth and mortality of European beech (Fagus sylvatica L.) fine roots (diameters <2 mm) were studied in relation to environmental variables describing temperature and water availability at four sites, covering a range in environmental conditions likely to be encountered in Slovenian beech forests. Minirhizotron images were used to determine fine root dynamics in a stand and gap in each of the sites for 12 periods during the 2007–2009 growing seasons. The environmental variables included air and soil temperatures, precipitation, forest floor precipitation, evapotranspiration and soil water contents. For data analysis, the daily mean values for each period for all variables were used. Fine root ingrowth and mortality were higher in the managed stand and gap compared to the old-growth stand and gap, but only significantly correlated with each other in the case of the managed stand. Forest floor precipitation and soil temperature were significant in explaining fine root ingrowth, whereas maximal evapotranspiration, soil temperature and soil water content were more important for fine root mortality. However, the correlations were weak and inconsistent among the four sites. By including site as predictor as well as environmental variables, R ² values of 0.49 and 0.55 for ingrowth and mortality, respectively, were achieved. Despite this, the relationships between the fine root dynamics and selected environmental factors appeared relatively weak and complex, especially for fine root ingrowth and might be partially related also to differences in successional stages of the forests under study.