The role of remnant forest trees in tropical secondary succession

This paper emphasizes the contribution of remnant trees to the establishment of woody species during succession on abandoned fields and pastures in the Mexican rain forest area, Los Tuxtlas, Veracruz. Remnant trees, original large forest trees left in the clearings by traditional farmers will become natural perching sites for both passing and resident birds. Frugivorous birds drop or regurgitate seeds and fruits which fall under the canopies of remnant trees during their stay, thus contributing to an accumulation of species, which make these remnant trees into ‘regeneration nuclei’. The species transported into these sites belong chiefly to older stages of successional development and reach these otherwise isolated areas, counteracting the depauperization of tropical land, brought about by both intensive and extensive clearing. In a study of seven remnant trees, 29 woody species and two climbers were found, 86% of which are bird dispersed. The total number of species per tree varied from 6 to 15 and was higher under remnant trees with fruits attractive to birds. Floristic variations of the understorey as detected by detrended correspondence analysis was correlated with the relative amount of shade-tolerant primary and late secondary trees versus light dependent pioneers and early successional trees.     

A new species of Kalanchoe (Crassulaceae) from northeast tropical Africa

The new species Kalanchoe subrosulata from deciduous bushlands in southwestern Somalia and northeastern Kenya is described and illustrated.     

Studies in Garcinia, dioecious tropical forest trees: agamospermy

RICHARDS, A. J., 1990. Studies in Garcinia , dioecious tropical forest trees: agamospermy . In Garcinia , agamospermy is known or suspected for ten species. Most are facultative agamosperms, with males occurring, but males are probably lacking in G. mangostana . In G. mangostana and G. hombroniana , adventitious embryony occurs autonomously, and haploid parthenogenesis may also occur rarely. In G. parvifolia , it is reported previously that gametophytic agamospermy occurs, and adventitious embryos are suppressed. Autonomous endospermy is found in G. mangostana and G. parvifolia . In G. hombroniana , endosperm probably only develops after PEN fertilization. In G. hombroniana , some proembryos are formed precociously, and further proembryo formation is inhibited by sexual embryos. Asexual proembryos tend to occur in large ovules in small ovaries, and sexual embryos tend to occur in small ovules in large ovaries. It is considered that facultative agamospermy renders the genus Garcinia particularly suitable for the development of new types of fruit. Agamospermy in a dioecious genus, adventitious embryony in the absence of fertilization and/or pseudogamy, and the co-occurrence of gametophytic and sporophytic agamospermies in the same genus are unusual phenomena, which are discussed in the context of the lifestyle of a tropical forest tree. For G. hombroniana, n = 24, and for G. cowa, n = 26 are reported.     

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

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

Coordination of leaf and stem water transport properties in tropical forest trees

Stomatal regulation of transpiration constrains leaf water potential (Ψ_L) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the stems to which they are attached. We evaluated a suite of leaf and stem functional traits governing water relations in individuals of 11 lowland tropical forest tree species to determine the manner in which the traits were coordinated with stem xylem vulnerability to embolism. Stomatal regulation of Ψ_L was associated with minimum values of water potential in branches (Ψ_br) whose functional significance was similar across species. Minimum values of Ψ_br coincided with the bulk sapwood tissue osmotic potential at zero turgor derived from pressure–volume curves and with the transition from a linear to exponential increase in xylem embolism with increasing sapwood water deficits. Branch xylem pressure corresponding to 50% loss of hydraulic conductivity (P ₅₀) declined linearly with daily minimum Ψ_br in a manner that caused the difference between Ψ_br and P ₅₀ to increase from 0.4 MPa in the species with the least negative Ψ_br to 1.2 MPa in the species with the most negative Ψ_br. Both branch P ₅₀ and minimum Ψ_br increased linearly with sapwood capacitance (C) such that the difference between Ψ_br and P ₅₀, an estimate of the safety margin for avoiding runaway embolism, decreased with increasing sapwood C. The results implied a trade-off between maximizing water transport and minimizing the risk of xylem embolism, suggesting a prominent role for the buffering effect of C in preserving the integrity of xylem water transport. At the whole-tree level, discharge and recharge of internal C appeared to generate variations in apparent leaf-specific conductance to which stomata respond dynamically.     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

population dynamics of some tropical trees that yield non-timber forest products

The population biology ofAquilaria malaccensis, one source of gharu, and Cinnamomum mollissimum, one source of wild cinnamon, was studied in a 50 ha permanent plot of primary rain forest in Malaysia. Median diameter growth rates of 0.22 cm yr^-1 and 0.1 cm yr^-1 should not be prohibitive of economic exploitation, and suggest that the trees could be grown commercially. However, the natural densities were between 2 and 3 trees over 1 cm d.b.h. per ha, which is roughly the median for all trees in the plot, would preclude economic exploitation of these natural populations. The economics of harvesting natural populations is considered in a preliminary fashion by allowing favorable assumptions of quantity and quality of production. The price likely to be fetched from either a first time extraction (on the order of US$10.00 per ha) or from sustained production (on the order of USThe population biology ofAquilaria malaccensis, one source of gharu, and Cinnamomum mollissimum, one source of wild cinnamon, was studied in a 50 ha permanent plot of primary rain forest in Malaysia. Median diameter growth rates of 0.22 cm yr^-1 and 0.1 cm yr^-1 should not be prohibitive of economic exploitation, and suggest that the trees could be grown commercially. However, the natural densities were between 2 and 3 trees over 1 cm d.b.h. per ha, which is roughly the median for all trees in the plot, would preclude economic exploitation of these natural populations. The economics of harvesting natural populations is considered in a preliminary fashion by allowing favorable assumptions of quantity and quality of production. The price likely to be fetched from either a first time extraction (on the order of US10.00 per ha) or from sustained production (on the order of US10.00 per ha) or from sustained production (on the order of US0.10 per ha per yr), are too small to be of interest as single-product schemes, and are negligible compared to the extraction of multi-species crops of timber. However, it is possible that by combining multiple products under a “High Diversity Forestry” scheme, one could increase the density of harvestable products, reduce the unit cost of labor and improve the economic portrait. La poblacion dinamica de algunos arboles tropicales. La población biológica deAquilaria malaccensis, un original de gharu, yCinnamomum mollissimum, un original del cinnamon salvage, ha sido estudiado en 50 ha en un terreno primario permanente de lluvia tropical en. Con un índice de crecimiento de medio dimetro 0.22 cm yr^-1 0.1 cm yr^-1 no deberia prohibirse la explotación económica, y se segiuere que los árboles pueden ser cultivados comercialmente. De cualquier manera, la densidad natural siendo aproximadamente entre 2 y 3 árboles per encima de 1 cm dbh per ha, lo cúal es un promedio de todos los árboles en el terreno, esto representa un impedimento a la explotacion económica de la población natural. La economía de la población natural de recolección de la cosecha está considerada como una moad preliminar por permitirse favorablemente la supuesta cantidad y calidad de productión. Los precios problemente alcanzen, bien sea, la primera vex de su extracción sobre el pedido de US10.00 per ha o desde una continua productión ininterrumpida, (sobre un pedido US10.00 per ha o desde una continua productión ininterrumpida, (sobre un pedido US0.10 per ha per yr), siendo demassiado peque?o para ser de interés tal proyecto comoproducto-unico, y son insignificantes comparados con la extracción de multiples especies de semillas de madera. De caulquier manera esposible que sepueda combinar los multiple productos en el proyecto de “Alta Diversidad Forestal,” uno puede la un puede incrementar la densidad de la productión de la cosecha, reduciendo el costo de mano de obra por unidad y mejorando la imagen económica.     

Stomatal and non-stomatal limitations of photosynthesis in trees of a tropical seasonally flooded forest

Trees in the flooded forest of the Mapire River in Venezuela suffer a decrease in photosynthetic rate (A) when flood begins, which is reverted at maximum flood. Changes in A are accompanied by similar changes in stomatal conductance (g_s), and the possibility of changes in photosynthetic capacity is not ruled out. In order to understand how relative stomatal and non-stomatal limitations of photosynthesis are affected by flooding, we studied the seasonal changes in A and its response to intercellular CO₂ concentration in trees of Campsiandra laurifolia , Symmeria paniculata , Acosmium nitens and Eschweilera tenuifolia . Flooding caused in trees of C.   laurifolia and S.   paniculata a reduction in A, g_s, carboxylation efficiency and total soluble protein (TSP), whereas gas exchange in A.   nitens and E.   tenuifolia was more sensitive to drought. Under flooding, relative stomatal limitation (L_s) was on average half the highest, and relative non-stomatal limitation (L_ns) increased from the dry season to flooding. Under full flood, A, g_s and TSP regained high values. A was positively correlated to light-saturated electron transport rate, suggesting that part of the decrease in A under flooding was due to impairment of photosynthetic capacity. Under flooding, not only stomatal closure but also increased L_ns causes a reduction in photosynthesis of all four species, and a process of acclimation as flooding progresses allows gas exchange and related variables to regain high values.     

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d^–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d^–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m² increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.     

Leaf age and the timing of leaf abscission in two tropical dry forest trees

We used experimental defoliations to examine the effect of leaf age on the timing of leaf shedding in two tropical dry forest trees. Trees of the deciduous Bombacopsis quinata (bombacaceae, a.k.a. Pachira quinata) and the brevi-deciduous Astronium graveolens (anacardiaceae) were manually defoliated for three times during the rainy season. All trees started to produce a new crown of leaves 2 weeks after defoliation, and continued expanding leaves throughout the rainy season. At the transition to the dry season, the experimental groups consisted of trees with known differences in maximum leaf age. Defoliations resulted in declines in stem growth but did not affect the mineral content or water relations of the leaves subsequently produced. There was no effect of leaf age on the timing of leaf abscission in B. quinata. In A. graveolens, the initiation of leaf shedding followed in rank order, the maximum leaf age of the four treatments, but there was substantial coherence among treatments in the major period of leaf abscission such that trees completed leaf shedding at the same time. In the two species, leaf water potential (Ψ_L) and stomatal conducantce (g _S) declined with the onset of the dry season, reaching minimum values of –0.9 MPa in P. quinata and <–2.0 MPa in A. graveolens. Within each species, leaves of different age exhibited similar Ψ_L and g _S at the onset of drought, and then decreased at a similar rate as the dry season progressed. Overall, our study suggests that the environmental factors were more important than leaf age in controlling the timing of leaf shedding.     

Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ_L) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although Ψ_L did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ¹⁸O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar ¹⁸O (∆¹⁸O_l) and ¹³C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.     

Ecological implications of a flower size/number trade-off in tropical forest trees

BACKGROUND: In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. METHODOLOGY/PRINCIPAL FINDINGS: We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. CONCLUSIONS AND THEIR SIGNIFICANCE: Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.     

The distribution of forest passerine birds and Quaternary climatic change in tropical Africa

A cluster analysis of the distribution patterns of forest passerine birds in tropical Africa showed that they could be divided into three main distribution types: those of restricted distribution (144 species), those of disjunct distributions across West, Central and rarely to East Africa (65 species) and those of very wide distribution (81 species). Centres of species richness, endemism and disjunction coincide spatially, and are identified as forest refugia, where forest persisted throughout Quaternary climatic vicissitudes. These distribution patterns agree with modern interpretations of Quaternary palaeoclimatic changes, which show that glacial periods were arid and interglacials humid. Glacial periods were therefore too dry for montane forest to have spread into areas at present occupied by lowland forest, as advocated by Moreau and other supporters of ihe pluvial theory; montane forest is probably as extensive now as at any time during the Quaternary and there is no evidence of past connections between currently isolated montane forests. Bird distribution can be explained largely by slow dispersal outwards from refugia as climatic conditions allowed forest to spread, combined with a relaxing of the distinction between montane and lowland species under conditions of reduced interspecific competition.     

Above-ground biomass investments and light interception of tropical forest trees and lianas early in succession

BACKGROUND AND AIMS: Crown structure and above-ground biomass investment was studied in relation to light interception of trees and lianas growing in a 6-month-old regenerating forest. METHODS: The vertical distribution of total above-ground biomass, height, diameter, stem density, leaf angles and crown depth were measured for individual plants of three short-lived pioneers (SLPs), four long-lived pioneers (LLPs) and three lianas. Daily light interception per individual Phi(d) was calculated with a canopy model. The model was then used to estimate light interception per unit of leaf mass (Phi(leaf mass)), total above-ground mass (Phi(mass)) and crown structure efficiency (E(a), the ratio of absorbed vs. available light). KEY RESULTS: The SLPs Trema and Ochroma intercepted higher amounts of light per unit leaf mass (Phi(leaf mass)) because they had shallower crowns, resulting in higher crown use efficiency (E(a)) than the other species. These SLPs (but not Cecropia) were also taller and intercepted more light per unit leaf area (Phi(area)). LLPs and lianas had considerably higher amounts of leaf mass and area per unit above-ground mass (LMR and LAR, respectively) and thus attained Phi(mass) values similar to the SLPs (Phi(mass)=Phi(area)xLAR). Lianas, which were mostly self-supporting, had light interception efficiencies similar to those of the trees. CONCLUSIONS: These results show how, due to the trade-off between crown structure and biomass allocation, SLPs, and LLPs and lianas intercept similar amount of light per unit mass which may contribute to the ability of the latter two groups to persist.     

Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (D) of xylem and soil water, soil volumetric water content (_v), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in _v was observed at 90–120 cm depth in both years. Mean soil water D during the dry season was –19 at 0–30 cm, –34 at 30–60 cm, and –50 at 90–120 cm. Average values of xylem D among the liana species ranged from –28 to –44 during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem D values (–41 to –62), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.     

Partitioning of soil water among canopy trees in a seasonally dry tropical forest

Little is known about partitioning of soil water resources in species-rich, seasonally dry tropical forests. We assessed spatial and temporal patterns of soil water utilization in several canopy tree species on Barro Colorado Island, Panama, during the 1997 dry season. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θ_v), and sap flow were measured concurrently. Evaporative fractionation near the soil surface caused soil water δD to decrease from about –15‰ at 0.1 m to –50 to –55‰ at 1.2 m depth. Groundwater sampled at the sources of nearby springs during this period yielded an average δD value of –60‰. θ_v increased sharply and nearly linearly with depth to 0.7 m, then increased more slowly between 0.7 and 1.05 m. Based on xylem δD values, water uptake in some individual plants appeared to be restricted largely to the upper 20 cm of the soil profile where θ_v dropped below 20% during the dry season. In contrast, other individuals appeared to have access to water at depths greater than 1 m where θ_v remained above 45% throughout the dry season. The depths of water sources for trees with intermediate xylem δD values were less certain because variation in soil water δD between 20 and 70 cm was relatively small. Xylem water δD was also strongly dependent on tree size (diameter at breast height), with smaller trees appearing to preferentially tap deeper sources of soil water than larger trees. This relationship appeared to be species independent. Trees able to exploit progressively deeper sources of soil water during the dry season, as indicated by increasingly negative xylem δD values, were also able to maintain constant or even increase rates of water use. Seasonal courses of water use and soil water partitioning were associated with leaf phenology. Species with the smallest seasonal variability in leaf fall were also able to tap increasingly deep sources of soil water as the dry season progressed. Comparison of xylem, soil, and groundwater δD values thus pointed to spatial and temporal partitioning of water resources among several tropical forest canopy tree species during the dry season. Received: 5 October 1998 / Accepted: 23 June 1999