Determinants of mortality across a tropical lowland rainforest community

Tree mortality is an important process determining forest dynamics. However, in species‐rich tropical forests it is largely unknown, how species differ in their response of mortality to resource availability and individual condition. We use a hierarchical Bayesian approach to quantify the impact of light availability, tree size and past growth on mortality of 284 woody species in a 50‐ha long‐term forest census plot in Barro Colorado Island, Panama. Light reaching each individual tree was estimated from yearly vertical censuses of vegetation density. Across the community, 78% of the species showed increasing mortality with increasing light. Considering species individually, just 29 species showed a significant response to light, all with increasing mortality at high light. Past growth had a significant impact on all but three species, with higher growth leading to lower mortality. For the majority of species, mortality decreased sharply with diameter in saplings, then levelled off or increased slightly towards the maximum diameter of the species. Diameter had the biggest impact on mortality, followed by past growth and finally light availability. Our results challenge many previous reports of higher mortality in shade, and we suggest that it is crucial to control for size effects when assessing the impact of environmental conditions on mortality.     

Diversity and distribution of ground-dwelling ants in a lowland rainforest in southeast Cameroon

Detailed information on the ground-dwelling ant diversity and distribution in the rainforests of the Congo Basin is lacking so far. A new ant species list from a continuous mixed secondary lowland rainforest on the northern periphery of the Dja Biosphere Reserve in southeast Cameroon is presented. We investigated the effect of vegetation type on ant species density, activity and composition. Ants were collected by pitfall trapping in seven vegetation types in each of three seasons during 12 months in 2003 and 2004–2005. Our site in Cameroon had a total observed ant diversity of 145 species. Vegetation type significantly influenced the ground-dwelling and -foraging (arboreal) ant assemblages. Differences in species density, activity and composition between vegetation types were explained not only by soil type (hydromorphous – terra firme), but also by developmental stage of vegetation types. The highly inundated Raphia-swamp had the lowest ant species density and activity. Older secondary forests still had a different species composition than near primary forest even after 25 years of recovery after anthropogenic disturbance. Finally, the high ant species number captured by our pitfalls, the absence of the invasive exotic ant Wasmannia auropunctata and the high Dorylus (Anomma) army ant diversity confirm the importance of the Cameroonian rainforest for biodiversity conservation programs. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Received 4 October 2006; revised 4 June 2007; accepted 2 July 2007.     

Blind population genetics survey of tropical rainforest trees

Rainforest tree species can be difficult to identify outside of their period of reproduction. Vascular tissues from Carapa spp. individuals were collected during a short field trip in French Guiana and analysed in the laboratory with nuclear and chloroplast markers. Using a Bayesian approach, > 90% of the samples could be assigned to one of two distinct clusters corresponding to previously described species, making it possible to estimate the genetic structure of each species and to identify cases of introgression. We argue that this blind procedure represents a first-choice rather than a fallback option whenever related taxa are investigated.     

Viable soil seed banks in disturbed lowland tropical rainforest sites in North Queensland

Four 0.125 m² soil samples, 50 mm deep, from each of eight tropical lowland rainforest sites ranging from primary and secondary forests to degraded Imperata grasslands were exposed to shade-house conditions. Germinations of soil seeds were monitored over 28 weeks in April-October. The viable seed banks in the primary forest (592 m^-2) and the secondary forest sites resulting from one cycle of clearing and abandonment (1300–1400 m^-2) were composed predominantly of secondary species. The viable seed banks were small in Imperata grassland sites that had been burnt regularly (200–400 m^-2) and very large (2000–3000 m^-2) in sites that had not been burnt for five years or more. The seed banks of the burnt and unburnt Imperata grassland sites were composed mostly of agricultural weeds whilst seeds of secondary rainforest trees were rare or absent. Similarities between the species composition of soil seed banks and regrowth vegetation types suggest that soil seed composition is important in determining the initial floristic composition following disturbance and that lack of seed of secondary rainforest trees is one factor which restricts re-growth and secondary succession on degraded Imperata grasslands. Species such as Eucalyptus intermedia R. T. Bak., E. pellita F. Muell. and Imperata cylindrica (L.)Beauv. which can establish and deflect the normal rainforest secondary succession were not present in the soil seed bank, and these species appear to rely on fresh seed inputs, lignotubers or rhizomes to colonize and regenerate after disturbance.     

Stable nitrogen isotope patterns of trees and soils altered by long-term nitrogen and phosphorus addition to a lowland tropical rainforest

Foliar nitrogen (N) isotope ratios (δ¹⁵N) are used as a proxy for N-cycling processes, including the “openness” of the N cycle and the use of distinct N sources, but there is little experimental support for such proxies in lowland tropical forest. To address this, we examined the δ¹⁵N values of soluble soil N and canopy foliage of four tree species after 13 years of factorial N and P addition to a mature lowland rainforest. We hypothesized that N addition would lead to ¹⁵N-enriched soil N forms due to fractionating losses, whereas P addition would reduce N losses as the plants and microbes adjusted their stoichiometric demands. Chronic N addition increased the concentration and δ¹⁵N value of soil nitrate and δ¹⁵N in live and senesced leaves in two of four tree species, but did not affect ammonium or dissolved organic N. Phosphorus addition significantly increased foliar δ¹⁵N in one tree species and elicited significant N × P interactions in two others due to a reduction in foliar δ¹⁵N enrichment under N and P co-addition. Isotope mixing models indicated that three of four tree species increased their use of nitrate relative to ammonium following N addition, supporting the expectation that tropical trees use the most available form of mineral N. Previous observations that anthropogenic N deposition in this tropical region have led to increasing foliar δ¹⁵N values over decadal time-scales is now mechanistically linked to greater usage of ¹⁵N-enriched nitrate.     

Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A_max, stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A_max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.     

Leaf hydraulic architecture correlates with regeneration irradiance in tropical rainforest trees

The leaf hydraulic conductance (K(leaf)) is a major determinant of plant water transport capacity. Here, we measured K(leaf), and its basis in the resistances of leaf components, for fully illuminated leaves of five tree species that regenerate in deep shade, and five that regenerate in gaps or clearings, in Panamanian lowland tropical rainforest. We also determined coordination with stomatal characters and leaf mass per area. K(leaf) varied 10-fold across species, and was 3-fold higher in sun- than in shade-establishing species. On average, 12% of leaf hydraulic resistance (= 1/K(leaf)) was located in the petiole, 25% in the major veins, 25% in the minor veins, and 39% outside the xylem. Sun-establishing species had a higher proportion of leaf resistance in the xylem. Across species, component resistances correlated linearly with total leaf resistance. K(leaf) correlated tightly with indices of stomatal pore area, indicating a coordination of liquid- and vapor-phase conductances shifted relative to that of temperate woody species. Leaf hydraulic properties are integrally linked in the complex of traits that define differences in water use and carbon economy across habitats and vegetation zones.     

Evaluating the annual nature of juvenile rings in Bolivian tropical rainforest trees

Knowledge on juvenile tree growth is crucial to understand how trees reach the canopy in tropical forests. However, long-term data on juvenile tree growth are usually unavailable. Annual tree rings provide growth information for the entire life of trees and their analysis has become more popular in tropical forest regions over the past decades. Nonetheless, tree ring studies mainly deal with adult rings as the annual character of juvenile rings has been questioned. We evaluated whether juvenile tree rings can be used for three Bolivian rainforest species. First, we characterized the rings of juvenile and adult trees anatomically. We then evaluated the annual nature of tree rings by a combination of three indirect methods: evaluation of synchronous growth patterns in the tree- ring series, ¹⁴C bomb peak dating and correlations with rainfall. Our results indicate that rings of juvenile and adult trees are defined by similar ring-boundary elements. We built juvenile tree-ring chronologies and verified the ring age of several samples using ¹⁴C bomb peak dating. We found that ring width was correlated with rainfall in all species, but in different ways. In all, the chronology, rainfall correlations and ¹⁴C dating suggest that rings in our study species are formed annually.     

Ecological distribution of homobaric and heterobaric leaves in tree species of Malaysian lowland tropical rainforest

Tree species can generally be classified into two groups, heterobaric and homobaric leafed species, according to whether bundle-sheath extensions (BSEs) are found in the leaf (heterobaric leaf) or not (homobaric leaf). In this study, we study whether the leaf type is related to the growth environment and/or life form type, even in a tropical rain forest, where most trees have evergreen leaves that are generally homobaric. Accordingly, we investigated the distribution of leaf morphological differences across different life forms of 250 tree species in 45 families in a tropical rainforest. In total, 151 species (60%) in 36 families had homobaric leaves, and 99 species (40%) in 21 families had heterobaric leaves. We found that the proportion of heterobaric and homobaric leaf species differed clearly across taxonomic groups and life form types, which were divided into five life form types by their mature tree heights (understory, subcanopy, canopy, and emergent species) and as canopy gap species. Most understory (94%) and subcanopy (83%) species such as Annonaceae had homobaric leaves. In contrast, heterobaric leaf trees appeared more frequently in the canopy species (43%), the emergent species (96%) (such as Dipterocarpaceae), and the canopy gap species (62%). Our results suggest that tree species in the tropical rainforest adapt to spatial differences in the environmental conditions experienced at the mature height of each tree species, such as light intensity and vapor pressure difference, by having differing leaf types (heterobaric or homobaric) because these types potentially have different physiological and/or mechanical functions.     

Geophagy in brown spider monkeys (Ateles hybridus) in a lowland tropical rainforest in Colombia

Spider monkeys and howler monkeys are the only Neotropical primates that eat soil from mineral licks. Not all species within these genera visit mineral licks, and geophagy has been restricted to populations of Ateles belzebuth belzebuth,Ateles belzebuth chamek and Alouatta seniculus in western Amazonian rainforests. With the aid of a camera trap we studied the visitation patterns of a group of brown spider monkeys (Ateles hybridus) to a mineral lick at Serrania de Las Quinchas, in Colombia. Spider monkeys visited the lick frequently throughout the year, with a monthly average of 21.7 ± 7.2 visits per 100 days of camera trapping (n = 14 months). Spider monkeys visited the mineral lick almost always on days with no rain, or very little (<3 mm) rain, suggesting that proximate environmental variables might determine spider monkeys' decisions to come to the ground at the licks. This study expands the geographical occurrence of mineral lick use by spider monkeys providing additional data for future assessments on the biogeographical correlates of mineral lick use by platyrrhines.     

Ontogenetic changes in leaf traits of tropical rainforest trees differing in juvenile light requirement

Relationships between leaf traits and the gap dependence for regeneration, and ontogenetic changes therein, were investigated in juvenile and adult tropical rainforest tree species. The juveniles of the 17 species included in the study were grown in high light, similar to the exposed crowns of the adult trees. The traits were structural, biomechanical, chemical and photosynthetic. With increasing species gap dependence, leaf mass per area (LMA) decreased only slightly in juveniles and remained constant in adults, whereas punch strength together with tissue density decreased, and photosynthetic capacity and chlorophyll increased. Contrary to what has been mostly found in evergreen tropical rainforest, the trade-off between investment in longevity and in productivity was evident at an essentially constant LMA. Of the traits pertaining to the chloroplast level, photosynthetic capacity per unit chlorophyll increased with gap dependence, but the chlorophyll a/b ratio showed no relationship. Adults had a twofold higher LMA, but leaf strength was on average only about 50% larger. Leaf tissue density, and chlorophyll and leaf N per area were also higher, whereas chlorophyll and leaf N per unit dry mass were lower. Ranking of the species, relationships between traits and with the gap dependence of the species were similar for juveniles and adults. However, the magnitudes of most ontogenetic changes were not clearly related to a species’ gap dependence. The adaptive value of the leaf traits for juveniles and adults is discussed.     

Spatial dispersion patterns of trees in a tropical rainforest in Xishuangbanna,southwest China

Ecological Research - Spatial dispersion patterns of trees at different life stages are an important aspect to investigate in understanding the mechanisms that facilitate species co-existence. In...     

Gregarious flowering in a lowland tropical rainforest: A possible response to disturbance by Cyclone Winifred

Observations in tropical lowland rainforests damaged by Cyclone Winifred on 1 February 1986 suggest that the disturbance has been followed by an unusual burst of flowering. Fifty-four species of concurrently flowering shrubs and trees were recorded in early March along a 4 km road transect in a cyclone-damaged forest. Approximately 80% of the smaller tree and shrub species observed were flowering. Twelve species which had flowered and fruited prior to the cyclone flowered again. Some species which normally flower following periods of partial or complete deciduousness in the drier months of the year flowered after being defoliated by the cyclone. Widespread defoliation of the rainforests, together with the hot, dry weather which followed the cyclone, exposed the understorey layers to abrupt increases in temperature, insolation, and water stress. It is postulated that these environmental changes may have effected the synchronization of flowering of a larger number of species than would normally occur. It is suggested that longterm changes in the species composition of the damaged forests as a result of coincidences of seed availability and regenerative space may occur.     

Gap effects on leaf traits of tropical rainforest trees differing in juvenile light requirement

The relationships of 16 leaf traits and their plasticity with the dependence of tree species on gaps for regeneration (gap association index; GAI) were examined in a Neotropical rainforest. Young saplings of 24 species with varying GAI were grown under a closed canopy, in a medium-sized and in a large gap, thus capturing the full range of plasticity with respect to canopy openness. Structural, biomechanical, chemical and photosynthetic traits were measured. At the chloroplast level, the chlorophyll a/b ratio and plasticity in this variable were not related to the GAI. However, plasticity in total carotenoids per unit chlorophyll was larger in shade-tolerant species. At the leaf level, leaf mass per unit area (LMA) decreased with the GAI under the closed canopy and in the medium gap, but did not significantly decrease with the GAI in the large gap. This was a reflection of the larger plasticity in LMA and leaf thickness of gap-dependent species. The well-known opposite trends in LMA for adaptation and acclimation to high irradiance in evergreen tropical trees were thus not invariably found. Although leaf strength was dependent on LMA and thickness, plasticity in this trait was not related to the GAI. Photosynthetic capacity expressed on each basis increased with the GAI, but the large plasticity in these traits was not clearly related to the GAI. Although gap-dependent species tended to have a greater plasticity overall, as evident from a principle component analysis, leaf traits of gap-dependent species are thus not invariably more phenotypically plastic.     

Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees

Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light-wood species grew faster in diameter than heavy-wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light-wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.     

Photosynthetic rates in relation to leaf phosphorus content in pioneer versus climax tropical rainforest trees

In Guyana dense rainforest occurs on intensely weathered acid soils, low in soil phosphorus. To investigate whether low P availability limits photosynthesis of trees growing on these soils more than N does, leaf P and N content, and their relationship with the photosynthetic capacity (A _sat, mol CO₂ m^-2 s^-1) were studied for nine pioneer and climax tree species in a range of light climates. Light environment was described using hemispherical photographs. For both pioneer and climax species, leaf P content (r ²=0.71 and 0.23, respectively) is a more important determinant of A _sat than leaf N content (r ²=0.54 and 0.12, respectively). Pioneer species have a higher leaf P and N content than climax species. At similar P or N content, pioneers have a higher A _sat than climax species. The saplings studied had a relatively high A _sat, considering their low P concentration (15–30 mol P g^-1). All species studied had a constant leaf P and N concentration and photosynthetic capacity across light climates, because specific leaf mass (g m^-2) increased similarly with light availability. This acclimation to a change in light environment makes a possible limitation of A _sat by P or N independent of light environment.     

Photosynthetic responses to vapour pressure deficit in temperate and tropical evergreen rainforest trees of Australia

Rainforests occur in high precipitation areas of eastern Australia, along a gradient in seasonality of precipitation, ranging from a summer dry season in the temperate south to a winter dry season in the tropical north. The response of net photosynthesis to increasing vapour pressure deficit (VPD) was measured in a range of Australian rainforest trees from different latitudes to investigate possible differences in their response to atmospheric drought. Plants were grown in glasshouses under ambient or low VPD to determine the effect of growth VPD on the photosynthetic response. Temperate species, which experience low summer precipitation, were found to maintain maximum net photosynthesis over the measurement range of VPD (0.5–1.9 kPa). In contrast, the tropical species from climates with high summer precipitation showed large reductions in net photosynthesis with increasing VPD. Temperate species showed higher intrinsic water-use efficiencies under low VPD than the tropical species, whereas their efficiencies were similar under high VPD. Growing plants under a low VPD had little effect on either the photosynthetic response to VPD or the intrinsic water-use efficiency of the species. These different responses of gas exchange to VPD shown by the tropical and temperate rainforest species may reflect different strategies to maximise productivity in their respective climates.     

Wild pigs (Sus scrofa) mediate large-scale edge effects in a lowland tropical rainforest in Peninsular Malaysia

Edge-effects greatly extend the area of tropical forests degraded through human activities. At Pasoh, Peninsular Malaysia, it has been suggested that soil disturbance by highly abundant wild pigs (Sus scrofa), which feed in adjacent Oil Palm plantations, may have mediated the invasion of Clidemia hirta (Melastomataceae) into the diverse tropical lowland rain forest. To investigate this hypothesis, we established three 1 km transects from the forest/Oil Palm plantation boundary into the forest interior. We recorded the distribution of soil disturbance by wild pigs, C. hirta abundance, and environmental variables. These data were analyzed using a hierarchical Bayesian model that incorporated spatial auto-correlation in the environmental variables. As predicted, soil disturbance by wild pigs declined with distance from forest edge and C. hirta abundance was correlated with the level of soil disturbance. Importantly there was no effect of distance on C. hirta abundance, after controlling for the effect of soil disturbance. Clidemia hirta abundance was also correlated with the presence of canopy openings, but there was no significant association between the occurrence of canopy openings and distance from the edge. Increased levels of soil disturbance and C. hirta abundance were still detectable approximately 1 km from the edge, demonstrating the potential for exceptionally large-scale animal mediated edge effects.