Tree form in a mixed dipterocarp forest in Indonesian Borneo

The form of tropical trees was studied with reference to the production structure of the component individuals of a tropical rain forest stand in Sebulu, East Kalimantan in Indonesian Borneo, since the production structure as a physical or bio-economical basis of tree form still remains obscure in tropical rain forests. The pipe model theory successfully explained the crown shapes of different trees, and its parameter, designated as specific pipe length, suggested an increase in the cost of leaf mass growth with an increase in crown size. A mathematical model consisting of exponential functions of aboveground height was applied for describing stem form, and its properties were examined through changes in its coefficients and by adopting an assumption of the geometrical similarity of individual stem form as a criterion for comparing differences in stem form among individual trees. Furthermore, the cost of buttersses was discussed using the relation between bole- and buttress weight calculated from the mathematical model.     

The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance

Dipterocarps dominate the canopy of lowland tropical rain forest in South‐east Asia. Seedlings of these species form diverse assemblages on the forest floor where low irradiance severely limits their growth. Further growth depends largely upon the increased irradiance that can occur with the creation of canopy gaps. However, the response of dipterocarp seedlings to increased irradiance and their subsequent establishment in the canopy may be influenced by the availability of other resources, such as nutrient availability. We investigated the influence of nitrogen supply on aspects of the photosynthetic physiology and growth of seedlings of four dipterocarp species (Shorea leprosula, Shorea johorensis, Shorea oleosa and Dryobalanops lanceolata) growing under low irradiance, during transfer from low to high irradiance, and during subsequent growth at high irradiance. All four species increased growth and photosynthetic capacity in response to N‐supply at high irradiances but not at low irradiance approximating that which can be expected to occur in the forest understorey. When seedlings grown at low irradiances and varying N‐supply were exposed to a large increase in irradiance, all species showed some degree of initial photodamage (measured through chlorophyll fluorescence), the extent of which was similar between species but differed markedly depending on the pre‐exposure growth irradiance and N‐supply. Greater photodamage occurred in seedlings grown at lower compared with higher N‐supply and irradiance. Despite these initial difference in the extent of this photodamage, all seedlings demonstrated a similar capacity to recover from damage. However, the alterations in the photosynthetic physiology of leaves during this recovery differed between species and depended on N‐supply. Under high N‐supply all species apart from S. oleosa increased photosynthetic capacity per unit chlorophyll following exposure to high irradiance by increasing photosynthetic capacity per unit leaf area while, under low N‐supply, an increase in photosynthetic capacity per unit leaf only occurred in D. lanceolata. Our results suggest that variations in N‐availability may have a much greater impact on the relative competitiveness of dipterocarp seedlings during the regenerative phase following canopy gap formation than physiological differences between seedlings. Our results demonstrate a potentially significant role for N‐availability in the regeneration dynamics and distribution of canopy‐dominating dipterocarp species.     

Aboveground biomass of tropical rain forest stands in Indonesian Borneo

Aboveground plant biomass was examined in a tall virgin tropical lowland evergreen rain forest dominated by Dipterocarpaceae in Sebulu, East Kalimantan, Indonesia, with special reference to the gap-, building- and mature phases of the forest growth cycle. From the records of dimensions of sample trees examined by the stratified clip technique and DBH inventory data of trees in a study plot, the biomass of larger trees (DBH 4.5 cm) was estimated by the allometric correlation method. The biomass of smaller plants (DBH < 4.5 cm) was estimated by harvesting the plants in small quadrat plots. Although large differences were found between aboveground-biomass-estimates in different patches of different growth stages, the aboveground biomass in a 1.0 ha plot was 509 t/ha, and the one-sided LAI was 7.3 ha/ha. These values seem to result from the tall forest architecture with huge emergent trees (over 70 m high) and a moderate packing of plant mass indicated by the basal area value of 38.8 m²/ha for trees with DBH 4.5 cm.This study was financed through a grant to H. Ogawa from the Overseas Scientific Research Funds of the Ministry of Education, Science, and Culture, Tokyo. Sponsorship from the Lembaga Ilumu Pengetahuan Indonesia (LIPI), Jakarta, the Lembaga Biologi National (LBN), Bogor, and the Herbarium Bogoriense, Bogor is gratefully acknowledged. We are also grateful to Drs Soetiyati, M. Rifai, K. Kartawinata, and the staffs of P. T. Kutai Timber Indonesia for their kind support, Dr K. Ogino for his advice and cooperation in field work, and Dr H. Kataoka for providing us with the geological map of Samarinda Province.     

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.     

The effect of canopy gaps on growth and morphology of seedlings of rain forest species

Summary Growth and morphology of seedlings of ten tropical rain forest species were studied at Los Tuxtlas, Mexico. Seedlings were grown in three environmental conditions: the shaded forest understorey (FU, receiving 0.9–2.3% of the daily photosynthetic photon flux, PF, above the canopy), a small canopy gap of approx. 50 m² (SG, receiving 2.1–6.1% of daily PF), and a large canopy gap of approx. 500 m² (LG, receiving 38.6–53.4% of daily PF). The growth of all species was enhanced in gaps, and in LG the effect was stronger than in SG. Plants grown in LG had a sunplant morphology, with a high root-shoot ratio (R/S), a high specific leaf weight (SLW) and a low leaf area ratio (LAR). Plants grown in SG or FU showed a shade-plant morphology, with a low R/S, a low SLW and a high LAR. Growth responses varied from species unable to grow in the shade but with strong growth in the sun, to species with relatively high growth rates in both shade and sun conditions. Shade tolerant species were able to grow in the shade because of a relatively high unit leaf rate. The pioneerCecropia had a high growth rate in LG because of a high LAR. Most species showed a complex growth response in which they resembled the shade intolerant extreme in some aspects of the response, and the shade tolerant extreme in other aspects.     

Nutrient cycling in a tropical seasonal rain forest of Xishuangbanna, Southwest China. Part 1: tree species: nutrient distribution and uptake

Tropical rain forests are characterized by large numbers of the species with diverse growth habits. The objective of the present study was to determine the distribution of nutrient content in the major trees of the tropical rain forests in Xishuangbanna. This will improve the understanding of the nutrient losses from such sites that result from harvesting and flow of nutrients within the ecosystem and lead to the development of effective and rational forest management strategies. Based on the results in this study, the distribution of nutrients among biomass components of trees varied: The ordering of major elements concentrations was K>N>Mg>Ca>P in branch, stem and root tissues but was N>K>Mg>Ca>P in leaves. The maximum amount of all nutrients per ha occurred in the stems followed by branches, roots and leaves. Of the total uptake of 6167.7 kg ha⁻¹ of all nutrients, the contribution of various nutrients was found to be N (2010.6 t ha⁻¹), P (196.3 t ha⁻¹), K (2123.8 kg ha⁻¹), Ca (832 kg ha⁻¹) and Mg (1005 kg ha⁻¹). However, comparing the nutrient uptake of other tropical and sub tropical forests, the results indicated that rates for the Xishuangbanna forests were 20–35% lower than previously reported values.     

Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil

The release of carbon as CO₂ from belowground processes accounts for about 70% of total ecosystem respiration. Insights about factors controlling soil CO₂ efflux are constrained by the challenge of apportioning sources of CO₂ between autotrophic tree roots (and mycorrhizal fungi) and heterotrophic microorganisms. In some temperate conifer forests, the reduction in soil CO₂ efflux after girdling (phloem removal) has been used to separate these sources. Girdling stops the flow of carbohydrates to the belowground portion of the ecosystem, which should slow respiration by roots and mycorrhizae while heterotrophic respiration should remain constant or be enhanced by the decomposition of newly dead roots. Therefore, the reduction in CO₂ efflux after girdling should be a conservative estimate of the belowground flux of C from trees. We tested this approach in two tropical Eucalyptus plantations. Tree canopies remained intact for more than 3 months after girdling, showing no reduction in light interception. The reduction in soil CO₂ efflux averaged 16–24% for the 3-month period after girdling. The reduction in CO₂ efflux was similar for plots with one half of the trees girdled and those with all of the trees girdled. Girdling did not reduce live fine root biomass for at least 5 months after treatment, indicating that large reserves of carbohydrates in the root systems of Eucalyptus trees maintained the roots and root respiration. Our results suggest that the girdling approach is unlikely to provide useful insights into the contribution of tree roots and heterotrophs to soil CO₂ efflux in this type of forest ecosystem.     

Patterns of natural 15N abundance in the leaf-to-soil continuum of tropical rain forests differing in N availability on Mount Kinabalu,Borneo

We investigated the natural abundance of ¹⁵N in sun leaves and other components of tropical rain forests on altitudinal sequences of eight sites that form a gradient of soil N availability with varying ectomycorrhizal abundances on Mt. Kinabalu, Borneo. We investigated how soil N availability and ectomycorrhizal abundance related to the ¹⁵N abundance of ecosystem components. ¹⁵N values (¹⁵N abundance relative to ¹⁴N) increased consistently in the following order at each site: sun leaves, leaf litter, fine roots and from shallower organic to deeper mineral soil horizons. Enrichment (3–6 ¹⁵N) of ¹⁵N occurred at the litter–topsoil interface at all sites, and the magnitude of the enrichment correlated negatively with ¹⁵N depletion in the foliage, irrespective of ectomycorrhizal abundance. Foliar ¹⁵N values significantly positively correlated with their N concentrations. Foliar (and litter and root) ¹⁵N values correlated positively with NO₃ availability, and negatively with NH₄ availability. The two positive correlations of foliar ¹⁵N with foliar N and NO₃ availability were inconsistent with the assumption that stronger nitrification (hence a greater nitrate availability) produced a more ¹⁵N-depleted active inorganic N pool. The isotopic fractionation during the passage of N through ectomycorrhizas to plants might explain the positive correlation of foliar ¹⁵N and N concentration; however, this mechanism could not fully explain the correlation in our case because strong foliar ¹⁵N depletions occurred at the sites that lacked ectomycorrhizas. Alternatively, the positive correlation across sites reflected the tightness of N cycling. Strong nitrification and associated isotopic fractionation might have occurred at N-richer sites and the subsequent removal of NO₃ from the system could decrease isotopically `lighter' N at these sites.     

热带雨林不同生态习性树种幼苗光合作用和抗氧化酶对生长光环境的反应

于雨季研究了西双版纳热带雨林后期演替树种思茅木姜子(Litsea pierrei var.szemaois)、五桠果叶木姜子(L．alilleniifolia)和先锋树种毛果桐(Mallotus barbatus)幼苗的光合、荧光特征和抗氧化酶等对3种光水平(50％、25％和8％)的适应特性。结果表明,毛果桐的最大光合速率(Pmx)随光水平升高呈增加趋势,而两种木姜子则在50％光水平下Pmax最小,Fv／Fm的日变化表明,3个种在25％和50％光水平下发生可逆光抑制,两种木姜子较重,50％光水平下,五桠果叶木姜子受到长期光抑制．思茅木姜子和毛果桐的丙二醛(MDA)和H2O2含量,以及超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)活性随光水平升高而增加,表明光水平升高,致使植物受光胁迫的影响加剧,增强抗氧化酶系统则减轻了其伤害,两种木姜子的濒危现状可能和它们对强光生境的适应性较差有一定关系。     

Structure,composition and species diversity in an altitude-substrate matrix of rain forest tree communities on Mount Kinabalu,Borneo

We studied forest structure, composition and tree species diversity of eight plots in an environmental matrix of four altitudes (700, 1700, 2700 and 3100 m) and two types of geological substrates (ultrabasic and non-ultrabasic rocks) on Mount Kinabalu, Borneo. On both substrate series, forest stature, mean leaf area and tree species diversity (both 4.8 cm and 10 cm diameter at breast height [dbh]) decreased with altitude. The two forests on the different substrate series were similar at 700 m in structure, generic and familial composition and tree species diversity, but became dissimilar with increasing altitude. The decline in stature with altitude was steeper on the ultrabasic substrates than on the non-ultrabasic substrates, and tree species diversity was generally lower on ultrabasic substrates than on non-ultrabasic substrates at 1700 m. The forests on non-ultrabasic substrates at higher altitudes and those on ultrabasic substrates at the lower altitudes were similar in dbh versus tree height allometry, mean leaf area, and generic and familial composition at 1700 m. These contrasting patterns in forest structure and composition between the two substrate series suggested that altitudinal change was compressed on the ultrabasic substrates compared to the non-ultrabasic substrates. Tree species diversity was correlated with maximum tree height and estimated aboveground biomass, but was not with basal area, among the eight study sites. We suggest that forests with higher tree species diversity are characterized by greater biomass allocation to height growth relative to trunk diameter growth under more productive environment than forests with lower tree species diversity.     

Seed predation by insects in tropical mangrove forests: extent and effects on seed viability and the growth of seedlings

Summary Although insects are known to be important seed predators in most terrestrial forests, their role in marine tidal (mangrove) forests has not been examined. Surveys at 12 sites in tropical Australia showed that between 3.1 and 92.7 percent of the seeds or propagules of 12 mangrove tree species had been attacked by insects. Seeds/propagules of six species (Avicennia marina, Bruguiera gymnorrhiza, B. parviflora, Heritiera littoralis, Xylocarpus australasicus and X. granatum) showed consistently high (>40%) levels of insect damage. Greater than 99% of H. littoralis seeds were attacked by insect predators. The survival and subsequent growth in height and biomass of insect-damaged and non-damaged control seeds/propagules of eight mangrove species were compared in shadehouse experiments. Mangrove species fell into 4 groups with regard to the effect of insect predators on their seeds and seedlings. Xylocarpus australasicus and X. granatum had significantly decreased survival (X 48 and 70%) and growth in height (X 61 and 96%) and biomass (X 66 and 85%). Bruguiera parviflora showed decreased survival (X 59%), but there was no effect of insects on the growth of surviving propagules. In contrast, there was no effect of insect damage on the survival of seedlings of Avicennia marina and Bruguiera exaristata, but decreased growth in height (X 22 and 25%) and biomass (X 22 and 26%). Survival and growth of seedlings of Rhizophora stylosa and Bruguiera gymnorrhiza were not affected. The influence of insect seed predators on the survival and growth of seeds of mangrove species in forests will depend on the relative abundance of seed-eating crabs and intertidal position in mangrove forests.This is Contribution No 499 from the Australian Institute of Marine Science     

The effects of light on foliar chemistry,growth and susceptibility of seedlings of a canopy tree to an attine ant

Summary Seedlings of Inga oerstediana Benth. (Mimosaceae) growing in three different light environments (the understory, tree-fall gaps and full sun) were tested for differences in chemistry (nutrients and tannins), wound-induced increases in tannins, growth, and susceptibility to leaf-cutter ants, Atta cephalotes (L.) (Formicidae: Attini). I hypothesized that seedlings of I. oerstediana would contain higher concentrations of tannins when growing in high light conditions and, therefore, would be less susceptible to leaf-cutter ants.Foliar concentrations of condensed tannins were much higher in plants growing in full sun compared to those growing in the understory. The concentrations of condensed tannins did not increase following damage. Despite higher concentrations of condensed tannins in sun foliage, leaf-cutter ants found these leaves more acceptable. The preference for sun leaves was consistent with higher concentrations of foliar nutrients. I suggest that the magnitude of the increase in condensed tannins was not great enough to override the benefits of increased concentrations of foliar nutrients. Finally, based on these results and those of others, I suggest that foraging by leaf-cutter ants may be an important factor determining patterns of succession in early successional habitats.     

Dissolved organic carbon affects soil microbial activity and nitrogen dynamics in a Mexican tropical deciduous forest

Seasonal variation of dissolved organic C (DOC) and its effects on microbial activity and N dynamics were studied during two consecutive years in soils with different organic C concentrations (hilltop and hillslope) in a tropical deciduous forest of Mexico. We found that DOC concentrations were higher at the hilltop than at the hillslope soils, and in both soils generally decreased from the dry to the rainy season during the two study years. Microbial biomass and potential C mineralization rates, as well as dissolved organic N (DON) and NH₄⁺ concentrations and net N immobilization were higher in soils with higher DOC than in soils with lower DOC. In contrast, net N immobilization and NH₄⁺ concentration were depleted in the soil with lowest DOC, whereas NO₃⁻ concentrations and net nitrification increased. Negative correlations between net nitrification and DOC concentration suggested that NH₄⁺ was transformed to NO₃⁻ by nitrifiers when the C availability was depleted. Taken together, our results suggest that available C appears to control soil microbial activity and N dynamics, and that microbial N immobilization is facilitated by active heterotrophic microorganisms stimulated by high C availability. Soil autotrophic nitrification is magnified by decreases in C availability for heterotrophic microbial activity. This study provides an experimental data set that supports the conceptual model to show and highlight that microbial dynamics and N transformations could be functionally coupled with DOC availability in the tropical deciduous forest soils. Responsible Editor: Chris Neill     

Long-term growth decline in Toona ciliata in a moist tropical forest in Bangladesh: Impact of global warming

Tropical forests are carbon rich ecosystems and small changes in tropical forest tree growth substantially influence the global carbon cycle. Forest monitoring studies report inconsistent growth changes in tropical forest trees over the past decades. Most of the studies highlighted changes in the forest level carbon gain, neglecting the species-specific growth changes which ultimately determine community-level responses. Tree-ring analysis can provide historical data on species-specific tree growth with annual resolution. Such studies are inadequate in Bangladesh, which is one of the most climate sensitive regions in the tropics. In this study, we investigated long-term growth rates of Toona ciliata in a moist tropical forest of Bangladesh by using tree-ring analysis. We sampled 50 trees of varying size, obtained increment cores from these trees and measured tree-ring width. Analyses of growth patterns revealed size-dependent growth increments. After correcting for the effect of tree size on tree growth (ontogenetic changes) by two different methods we found declining growth rates in T. ciliata from 1960 to 2013. Standardized ring-width index (RWI) was strongly negatively correlated with annual mean and maximum temperatures suggesting that rising temperature might cause the observed growth decline in T. ciliata. Assuming that global temperatures will rise at the current rate, the observed growth decline is assumed to continue. The analysis of stable carbon and oxygen isotopes may reveal more insight on the physiological response of this species to future climatic changes.     

Temporal and spatial partitioning of water resources among eight woody species in a Hawaiian dry forest

Lowland dry forests are unique in Hawaii for their high diversity of tree species compared with wet forests. We characterized spatial and temporal partitioning of soil water resources among seven indigenous and one invasive dry forest species to determine whether the degree of partitioning was consistent with the relatively high species richness in these forests. Patterns of water utilization were inferred from stable hydrogen isotope ratios (δD) of soil and xylem water, zones of soil water depletion, plant water status, leaf phenology, and spatial patterns of species distribution. Soil water δD values ranged from –20‰ near the surface to –48‰ at 130 cm depth. Metrosideros polymorpha, an evergreen species, and Reynoldsia sandwicensis, a drought-deciduous species, had xylem sap δD values of about –52‰, and appeared to obtain their water largely from deeper soil layers. The remaining six species had xylem δD values ranging from –33 to –42‰, and apparently obtained water from shallower soil layers. Xylem water δD values were negatively correlated with minimum annual leaf water potential and positively correlated with leaf solute content, an integrated measure of leaf water deficit. Seasonal patterns of leaf production ranged from dry season deciduous at one extreme to evergreen with near constant leaf expansion rates at the other. Species tapping water more actively from deeper soil layers tended to exhibit larger seasonality of leaf production than species relying on shallower soil water sources. Individuals of Myoporum sandwicense were more spatially isolated than would be expected by chance. Even though this species apparently extracted water primarily from shallow soil layers, as indicated by its xylem δD values, its nearly constant growth rates across all seasons may have been the result of a larger volume of soil water available per individual. The two dominant species, Diospyros sandwicensis and Nestegis sandwicensis, exhibited low leaf water potentials during the dry season and apparently drew water mostly from the upper portion of the soil profile, which may have allowed them to exploit light precipitation events more effectively than the more deeply rooted species. Character displacement in spatial and temporal patterns of soil water uptake was consistent with the relatively high diversity of woody species in Hawaiian dry forests. Received: 20 May 1999 / Accepted: 2 March 2000     

Variation in seed and seedling traits in Pithecellobium pedicellare,a tropical rain forest tree

Summary Pithecellobium pedicellare, a mimosoid legume, is a large canopy tree in the tropical rain forests of Costa Rica. We examined the pattern of variation in seed weight, germination date, hypocotyl length (stem), and rachis length (the first leaf) of the seedlings in this species. Seeds collected from widely dispersed individual trees at the La Selva Biological Station, Costa Rica, were randomly planted in blocks, and grown under controlled, indoor conditions for about 2 weeks. There-fore, we were able to quantify the effects of maternal family on mean seed weight and the effects of maternal family and microenvironment on the remaining traits examined. A significant effect of maternal family was detected for all traits. In particular, the maternal effects on germination date and seedling size traits which were consistently significant even after controlling the initial seed weight may indicate that the maternal effects reflect, at least to some extent, maternal genetic control over these traits. Despite overall strong maternal effects, the performance of maternal siblings, such as the rachis length, differed among blocks. The sensitivity of maternal siblings to the local environments may contribute to the maintenance of genetic variability in this highly outcrossing tropical species.