Reproductive allometry in Malaysian rain forest trees: Biomechanics versus optimal allocation

Summary Quantitative predictions of reproductive allometry in iteroparous plants may be derived from two bodies of theory: biomechanics and optimal allocation theory. Biomechanical theory predicts allometric scaling exponents between reproductive (R) and vegetative (V) biomass in the range of 0.44–1.33, while very general models of life history evolution predictR–V exponents > 1 in all cases. These predictions are examined in light of allometric patterns of flower and fruit production in 32 species of Malaysian rain forest trees. Among these species the mean estimatedR–V exponents are in the range 1.8–2.0 for staminate flower, pistillate flower and fruit production. This range of exponent values provides unambiguous support for some of the general predictions of optimal allocation models, but not for biomechanical theory. Optimal allocation models also predict a positive relationship between species size andR–V slope and a positive relationship between species size andR–V intercept parameters. The latter, but not the former prediction is supported by the data.R–V allometries in sexes of dioecious species were also found to differ in intercept, though not slope, reflecting smaller sizes at reproductive onset in staminate trees. Further critical examinations of reproductive allometry are encouraged as a relatively unexplored avenue for increasing the contact of theory and data in studies of life history evolution in long-lived organisms such as tropical trees.     

Reproductive patterns of selected understorey trees in the Malaysian rain forest: the sexual species

HA, C. O., SANDS, V. E., SOEPADMO, E. & JONG, K., 1988. Reproductive patterns of selected understorey trees in the Malaysian rain forest: the sexual species. Investigations were carried out on the floral, fruit and seed biology, and the embryology of representatives of the Sapindaceae and Bombacaceae which occur in the lower canopy of the lowland rain forest of Peninsular Malaysia. Pollination studies indicated cross-compatibility and outbreeding in all species examined, associated with self-compatibility in the polygamous monecious Pometia pinnata and Allophylus cobbe and, to a limited extent, in the effectively dioecious Xerospermum intermedium . In these sapindaceous species, together with the self-incompatible androdioecious Nephelium lappaceum , the structurally hermaphrodite flowers were determined embryologically to be functionally female with no anther dehiscence. The self-incompatible Durio griffithii of the Bombacaceae, and X. intermedium, N. lappaceum and P. pinnata were shown by embryological studies to form a sexual zygote. The observations on these outcrossing species are considered in relation to their potential for gene exchange and their contribution to the maintenance of species diversity in the lowland rain forest.     

Reproductive patterns of selected understorey trees in the Malaysian rain forest: the apomictic species

HA, C. O., SANDS, V. E., SOEPADMO, E. & JONG, K., 1988. Reproductive patterns of selected understorey trees in the Malaysian raia forest: the apomictic species. Garcinia is the predominant representative of the Clusiaceae in the understorey of the lowland rain forest of Peninsular Malaysia. In Pasoh Forest Reserve the dioecious species G. paruifolia was investigated since no male trees, but only trees with structurally hermaphodite flowers which later set fruit, were found there. Pollination and embryological studies indicated a condition of non-pseudogamous agamospermy, with no viable pollen grains being formed in the staminodes, and the unfertilized egg cell giving rise to the embryo. Supportive studies were made of microspore development in G. forbesi , and of embryogenesis and seed development in G. malaccensis and G. scortechinii which also appear to be agamospermous. The role of either facultative or obligate apomixis in gene fixation in these dioecious species is discussed, and the possible adaptive significance of reproductive versatility among certain rain forest trees is examined.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Pollination success in a population of dioecious rain forest trees

Pollination success in female trees was determined for a population of Neolitsea dealbata (R. Br.) Merr., a locally abundant dioecious tree pollinated by small, unspecialized insects in northern Queensland rain forest, Australia. The population consisted of a clustered group of trees with a mean male-to-female distance of 4.5 m and more isolated individuals, including females more than 90 m away from the nearest pollen source. A map of all reproductive trees was produced to determine accurate male-to-female distances. The size of the pollen source available to females was defined as a function of the distance to the nearest ten male trees and their sizes (male neighbourhood index). The rate of pollen movement to females was measured by counting pollen tubes (and the number of tubes per style) in female trees 6 days after the commencement of population flowering. The pollination rate decreased steeply to less than half when the nearest male was only 6.5 m away. Although pollen reached a female 330 m away from the nearest pollen source, only 10% of receptive flowers had been pollinated. The short flowering period (2–3 weeks) combined with the the slow rate of pollen movement means that a large proportion of flowers in isolated trees are unpollinated, confirming an earlier finding that isolated females set fewer fruits than gregarious females. The reliability of pollen transfer to females was determined by quantifying insects and their pollen loads trapped at female trees with a range of male neighbourhood indices. Quantities of insects and pollen were significantly correlated with the size of the male neighbourhood index, indicating a strong density-dependent response by vectors to flowering. Pollen was also collected from insect visitors to non-flowering trees. Females with large male neighbourhood indices received more pollen than non-flowering trees with equivalent male neighbourhood indices. However, when the male neighbourhood indices were small for both female and non-flowering trees, the changces of pollinators encountering female and non-flowering trees were similar, suggesting random movements of pollinators in sparse-flowering sub-populations. The dioecious breeding system, brief, synchronous flowering period, clustered population structure and random, opportunistic foraging behaviour of vectors interacted in a way that reduced reproduction in relatively isolated trees. These results demonstrate a mechanism for differential breeding success between trees in natural populations and emphasize the possible impact of logging regimes on pollen flow between trees. Large interconspecific distances in species-rich environments may have been a factor in the selection for synchronous flowering between trees in outcrossing tree species with generalist insect pollinators.     

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.     

Roots of pioneer trees in the Amazonian rain forest

. On low fluvial terraces of the Caquetá River, Araracuara region, Colombia, tree root systems were examined with regard to their structure, underground stratification and biomass distribution. Excavations of skeleton roots and microscopic observation of terminal fine roots included ten species belonging to the Cecropia, Vismia, Miconia, Goupia, Clathrotropis and Brosimum genera. Roots of particular species varied in the external features of their periderm and structure of freshly cut slash. Coarse skeleton roots differed in the shape and position of the plagiotropic branches and positively geotropic taproots or sinkers. None of the observed species could be identified as a shallow-rooted tree, in spite of the general fine root concentration in the upper soil horizon. In the two seasons of study, the terminal roots displayed prevailingly secondary anatomical structure and did not maintain much primary anatomical tissue in their apices. Neither ectomycorrhizas nor endomycorrhizas were detected in the samples. In a set of regeneration stages the amount of tree roots linearly increased with the age of growth. However, fine roots below 2 mm in diameter shared 80-90% of all roots in the upper 20 cm layer of all sample plots. In an old-growth forest, the total tree root biomass amounted to 39 ton/ha, thus being comparable to the underground biomass observed in similar tropical forests.     

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.     

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.     

Crown development in tropical rain forest trees in gaps and understorey

Emergence and establishment of Tsuga sieboldii seedlings were monitored for six years in an old-growth mixed forest in Japan, to clarify the reason why adult T. sieboldii trees are distributed only on convex slopes. T. sieboldii seedlings germinated on exposed soils without broad-leaved litter cover on both convex and concave slopes, but they died out from concave slope during the first two years after germination. Along a gradient from convex to concave slope, litter cover became higher and surface soil erosion became severer, and T. sieboldii seedlings died at the plots with severe surface soil erosion. Other environmental variables, such as relative illuminance, vegetation cover of seedling layer and soil water potential were not significantly different between convex and concave slopes. We concluded that the restricted seedling establishment of T. sieboldii to convex slope is determined by stability of surface soils and broad-leaved litter cover.     

Congruence between floristic patterns of trees and lianas in a southwest Amazonian rain forest

The congruence in floristic patterns between different life-forms of woody plants remains poorly understood in tropical rain forests. We explored whether the floristic patterns of woody plants, divided into small trees 2.5–10 cm dbh, large trees ≥10 cm dbh, and lianas ≥2.5 cm dbh were associated with each other or with patterns in soil properties, elevation, and geographical distances between sample plots. We also tested whether ecological amplitudes in relation to environmental variables differed among the plant groups. Trees and lianas were inventoried in 44 0.1-ha plots, distributed among three lowland and two submontane sites in the Madidi National Park, Bolivia. Soil samples were analysed for physico-chemical properties. Floristic differences between sites (as measured with each plant group separately) yielded significant Mantel correlations with each other, and with pH, Ca, Mg, elevation and geographical distance. Mantel correlations with edaphic distances were higher for large trees than small trees, but for Mantel correlations with geographical distance the situation was reversed. Environmental and geographical distances explained 31% of the variation in floristic differences for large trees, 22% for small trees, and 10% for lianas. The ecological amplitudes of lianas were wider than those of all trees for pH, Mg and elevation. The amplitudes of the two size classes of trees did not differ. In principal coordinates ordination, the three plant groups produced similar overall floristic patterns that were explainable by environmental factors.     

Litter breakdown and mineralization in a central African rain forest dominated by ectomycorrhizal trees

Based on litter mass and litterfall data, decomposition rates for leaveswere found to be fast (k = 3.3) and the turnover times short (3.6mo) on the low-nutrient sandy soils of Korup. Leaf litteroffour ectomycorrhizal tree species (Berlinia bracteosa, Didelotiaafricana, Microberlinia bisulcata and Tetraberliniabifoliolata) and of three non-ectomycorrhizal species(Cola verticillata, Oubanguia alata andStrephonema pseudocola) from Korup were left to decomposein 2-mm mesh bags on the forest floor in three plots ofeachof two forest types forest of low (LEM) and high (HEM) abundance ofectomycorrhizal (caesalp) trees. The litter of the ectomycorrhizal speciesdecayed at a significantly slower rate than that of thenon-ectomycorrhizal species, although the former were richer in P and Nconcentrations of the start. Disappearance rates of the litter layer showed asimilar trend. Ectomycorrhizal species immobilized less N, but mineralized moreP, than non-ectomycorrhizal species. Differences between species groupsin K, Mg and Ca mineralization were negligible. Effect of forest type was clearonly for Mg: mineralization of Mg was faster in the HEM than LEM plots, apattern repeated across all species. This difference was attributed to a muchmore prolific fine root mat in the HEM than LEM forest. The relatively fastrelease of P from the litter of the ectomycorrhizal species suggests that thematmust allow an efficient uptake to maintain P in the forest ecosystem.     

Photosynthesis and successional status of Costa Rican rain forest trees

Seven tree species from three different light environments in the wet lowland forests of Costa Rica were grown under controlled environment conditions to assess light related photosynthetic potentials. Light saturated photosynthesis rates were clearly related to light levels of the field environments. Mean saturated, net photosynthetic rates ranged from 6.8 to 11.3 to 27.7 mol m^–2 sec^–1 for plants from heavy shade, canopy light gaps and man-made clearings respectively. Light saturation of plants from clearings occurred at photosynthetic photon flux densities greater than 1000 mol m^–2 sec^–1 whereas plants from heavy shade environments became light saturated near 500 mol m^–2 sec^–1. Plants that normally occur in intermediate light environments were intermediate in light saturation levels. Mean maximum stomatal conductances ranged from 1.0 to 7.3 mm sec^–1 and followed a pattern similar to photosynthetic rates.