1. In a unique phenomenon restricted to the ever wet forests of Southeast Asia, hundreds of species from dozens of plant families reproduce synchronously at irregular, multi-year intervals. The ...proximate environmental cues that synchronize these general flowering events have not been evaluated systematically because there have been no long-term, high temporal resolution, species-level records from the region. 2. We present 13 years of weekly flowering records for five Shorea species as well as daily temperature and rainfall records from the Pasoh Forest Reserve, Peninsular Malaysia. We constructed models to evaluate hypothesized relationships between flowering and cool temperature, drought, and additive and synergistic effects of cool temperature and drought for each species. Model parameters include periods of time for floral cue accumulation and flower development and temperature and/or rainfall thresholds required for floral initiation. Parameters estimated using flowering observations from 2001 to 2011 were used to forecast flowering for 2011-2014. 3. We show that drought and cool temperatures acting synergistically best explain the timing of flowering events for all Shorea species in the section Mutica and forecast the largest general flowering event accurately. Periods estimated for signal accumulation ranged from 54 to 90 days among species. Periods estimated for flowers to develop ranged between 43 and 96 days and closely followed the interspecific sequence of flowering in the Shorea species. Drought and temperature thresholds also varied among species, with Shorea maxwelliana requiring the most severe drought and Shorea leprosula the lowest temperatures. 4. Synthesis. Our results indicate that cool temperatures and low rainfall occurring on seasonal time-scales of about 2-3 months rather than brief cold snaps or brief droughts best explain general flowering in Shorea species at the Pasoh Forest Reserve. Low rainfall is equally likely in winter (December-February) and summer (July–August) and cool temperatures are most likely in winter at Pasoh, which explains why general flowering events are restricted to spring and fall, with more frequent and stronger flowering in spring. In addition, species-specific sensitivity to environmental cues suggests that future climate change will have differential impacts on the frequency of reproduction, with potential consequences for regeneration of these dominant species of lowland tropical forests.
Aims: With the aim of understanding why some of the world's forests exhibit higher tree beta diversity values than others, we asked: (1) what is the contribution of environmentally related variation ...versus pure spatial and local stochastic variation to tree beta diversity assessed at the forest plot scale; (2) at what resolution are these beta-diversity components more apparent; and (3) what determines the variation in tree beta diversity observed across regions/continents? Location: World-wide. Methods: We compiled an unprecedented data set of 10 large-scale stem-mapping forest plots differing in latitude, tree species richness and topographic variability. We assessed the tree beta diversity found within each forest plot separately. The non-directional variation in tree species composition among cells of the plot was our measure of beta diversity. We compared the beta diversity of each plot with the value expected under a null model. We also apportioned the beta diversity into four components: pure topographic, spatially structured topographic, pure spatial and unexplained. We used linear mixed models to interpret the variation of beta diversity values across the plots. Results: Total tree beta diversity within a forest plot decreased with increasing cell size, and increased with tree species richness and the amount of topographic variability of the plot. The topography-related component of beta diversity was correlated with the amount of topographic variability but was unrelated to its species richness. The unexplained variation was correlated with the beta diversity expected under the null model and with species richness. Main conclusions: Because different components of beta diversity have different determinants, comparisons of tree beta diversity across regions should quantify not only overall variation in species composition but also its components. Global-scale patterns in tree beta diversity are largely coupled with changes in gamma richness due to the relationship between the latter and the variation generated by local stochastic assembly processes.
1 The rapid growth rates of light-demanding tree species have been attributed in part to their low-density, low-cost stems. We evaluated the influence of light and biomass support costs on growth ...rates of trees 8-25 cm in diameter at breast height (d.b.h.) among 21 species differing in wood density in two aseasonal rain forests. 2 Measurements of crown width, tree height, d.b.h. and wood density (p) were used to estimate the stem biomass (Ms) of a standard-sized tree (17 m tall and$16 m^2$in crown area), i.e. the cost in stem biomass of supporting a given sized crown at a given height. 3 The species showed a three-fold range in support cost, which was highly correlated with wood density ($M_s \propto p^{0.77}$, r2= 0.72 for the log-transformed relationship). This relationship is due to the high interspecific variation in wood density and the fact that the stem diameter of the standard-sized tree increased only slightly with decreasing wood density, i.e. light-wooded species did not compensate for their lighter, weaker wood by substantially increasing stem thickness. 4 Mean growth rate per species showed a 10-fold range and increased with the fraction of trees at least partly in gaps (gap fraction), the reciprocal of support cost$(1/M_s)$, and the reciprocal of wood density$(1/\rho)$. The relationship between mean growth rate and$1/M_s$was particularly strong when one outlier was excluded (r2= 0.88) and among the Dipterocarpaceae$(r^2 = 0.89)$. 5 Log(mortality rate), as determined for all trees per species$\geq 1 cm$d.b.h., increased linearly with$1/M_s$,$1/\rho$and gap fraction. 6 These results suggest an important role for wood density and support costs in the classic tradeoff between rapid growth and increased risks of damage and death.
1. In the absence of wind, tree height is limited by elastic instability, which occurs when a tree becomes too spindly to erect itself when bent from the vertical. To assess the extent to which trees ...approach this limit and characterize species stature in diverse tropical forests, we measured tree dimensions in tall, dense forests at Lambir Hills, National Park, Sarawak, Malaysia and Pasoh Forest Reserve, Peninsular Malaysia. From these measurements we determined characteristic adult heights and trunk diameters for 91 species. 2. Across all species, adult height scaled with adult diameter to the 2/3 power, as predicted for a column at its buckling limit. These heights were, on average, 65% of the buckling limit calculated for a cylindrical column with typical wood properties. 3. At a given diameter, the species of Lambir were 9% taller than those of Pasoh, a pattern related to the greater rainfall and density of trees at Lambir. 4. On the topographically rugged Lambir plot, large emergent trees were shorter on ridges than in hollows, whereas small, sheltered trees showed no relation between allometry and elevation. 5. Thus, trees may approach gravitational limits to height in favourable environments for growth where there are large advantages of height for light interception and trees are sheltered from wind.
In South-East Asian dipterocarp forests, many trees synchronize their reproduction at the community level, but irregularly, in a phenomenon known as general flowering (GF). Several proximate cues ...have been proposed as triggers for the synchronization of Southeast Asian GF, but the debate continues, as many studies have not considered geographical variation in climate and flora. We hypothesized that the spatial pattern of GF forests is explained by previously proposed climatic cues if there are common cues for GF among regions. During the study, GF episodes occurred every year, but the spatial occurrence varied considerably from just a few forests to the whole of Peninsular Malaysia. In 2001, 2002 and 2005, minor and major GF occurred widely throughout Peninsular Malaysia (GF2001, GF2002, and GF2005), and the geographical patterns of GF varied between the episodes. In the three regional-scale GF episodes, most major events occurred in regions where prolonged drought (PD) had been recorded prior, and significant associations between GF scores and PD were found in GF2001 and GF2002. However, the frequency of PD was higher than that of GF throughout the peninsula. In contrast, low temperature (LT) was observed during the study period only before GF2002 and GF2005, but there was no clear spatial relationship between GF and LT in the regional-scale episodes. There was also no evidence that last GF condition influenced the magnitude of GF. Thus, our results suggest that PD would be essential to trigger regional-scale GF in the peninsula, but also that PD does not fully explain the spatial and temporal patterns of GF. The coarse relationships between GF and the proposed climatic cues may be due to the geographical variation in proximate cues for GF, and the climatic and floristic geographical variations should be considered to understand the proximate factors of GF.
1 The lowland dipterocarp forests of Southeast Asia exhibit interspecifically synchronized general flowering (GF) and mast fruiting at irregular multi-year intervals of 1 to 11 years. The predator ...satiation hypothesis (PSH) posits that GF events enhance seed survival by reducing the survival, reproduction and population sizes of seed predators between GF events, and then satiating the reduced seed predator populations during GF events. 2 Three GF events of different magnitudes occurred in Pasoh Forest Reserve, Peninsular Malaysia, during 2001, 2002 and 2005. We exploited this natural experiment to test two predictions of the PSH. The first prediction was that seed survival should increase with the magnitude of the GF event. The second prediction was that seed predation should decrease with time since the previous GF event. 3 A reproductive survey of all (c. 900) dipterocarp trees 30 cm d.b.h. in a 50 ha plot showed that flowering pervasiveness (the proportion of dipterocarp species participating) was high and similar in all three GF events. However, relative flowering magnitudes (measured by an index of individual tree participation and flowering intensity in Shorea species) were 2, 5 and 8 for the 2001, 2002 and 2005 GF events, respectively. 4 The percentage of Shorea seeds surviving pre- and post-dispersal predation increased with the magnitude of GF events, which is consistent with the first prediction. 5 Pre-dispersal insect seed predators consumed 12.9%, 11.2% and 3.4% of Shorea seeds in the 2001, 2002 and 2005 GF events, respectively, which is consistent with both predictions. 6 Pre-dispersal seed predation by primates (mainly leaf monkeys) increased from 11.9% to 38.6% then fell to 9.3% in the 2001, 2002 and 2005 GF events, respectively. 7 Predator satiation occurred only at population and community levels. At the individual tree level there was no relationship between the percentage of seeds surviving pre- and post-dispersal seed predation and variation in seed crop size or seed density beneath the tree. This suggests that attempts to test the PSH on the scale of individual trees may miss key community level effects. 8 Our results suggest a more significant role of pre-dispersal seed predation in the evolution of reproductive synchrony than was recognized in the original statement of the PSH.
Importance of Demographic Niches to Tree Diversity Condit, Richard; Ashton, Peter; Bunyavejchewin, Sarayudh ...
Science (American Association for the Advancement of Science),
07/2006, Volume:
313, Issue:
5783
Journal Article
Peer reviewed
Open access
Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the ...variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.
Spatial association patterns reflect underlying mechanisms of coexistence, community structure of plant species in tropical forests. We hypothesized that if spatial associations between two species ...shift toward segregation patterns during the course of growth, deterministic mechanisms, such as interspecific competition and habitat differentiation, would prevail, whereas if no directed change in spatial associations between two species is observed and, consequently, the initial association pattern is retained through growth, the two species would experience weak interspecific competition and show no habitat differentiation. To assess the underlying mechanisms operating between confamilial species, we analysed spatial associations among 11 dipterocarp species in terms of three growth stages distinguished on the basis of dbh in the Pasoh 50-ha plot in Peninsular Malaysia. We analysed the spatial associations of all possible combinations among identical stages (165 pairs) and among different stages (330 pairs) for each pair of 11 species, except between identical species. Our previous study revealed that the 11 species could be characterized into two classes: seven fast-growing species exhibited high growth and mortality rates, spatial aggregation on a small scale, and positive habitat associations, while four slow-growing species exhibited low growth and mortality rates, spatial aggregation on a large scale, and no habitat associations except one. Spatial segregation was observed between fast-growing species (32 pairs, 17%) and between species of different classes (35 pairs, 14%), but not between slow-growing species. Throughout the growth stages, positive associations with other species were maintained for slow-growing species versus fast-growing species. In contrast, changes in initial associations toward segregation were observed more in fast-growing species. These results indicated that interspecific competition or habitat differentiation dominated for fast-growing species, while non-directed random processes dominated for slow-growing species.
General flowering (GF) is a unique phenomenon wherein, at irregular intervals, taxonomically diverse trees in Southeast Asian dipterocarp forests synchronize their reproduction at the community ...level. Triggers of GF, including drought and low minimum temperatures a few months previously has been limitedly observed across large regional scales due to lack of meteorological stations. Here, we aim to identify the climatic conditions that trigger large-scale GF in Peninsular Malaysia using satellite sensors, Tropical Rainfall Measuring Mission (TRMM) and Moderate Resolution Imaging Spectroradiometer (MODIS), to evaluate the climatic conditions of focal forests. We observed antecedent drought, low temperature and high photosynthetic radiation conditions before large-scale GF events, suggesting that large-scale GF events could be triggered by these factors. In contrast, we found higher-magnitude GF in forests where lower precipitation preceded large-scale GF events. GF magnitude was also negatively influenced by land surface temperature (LST) for a large-scale GF event. Therefore, we suggest that spatial extent of drought may be related to that of GF forests, and that the spatial pattern of LST may be related to that of GF occurrence. With significant new findings and other results that were consistent with previous research we clarified complicated environmental correlates with the GF phenomenon.
In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species ...over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16-52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha(-1) y(-1), 95% confidence intervals 0.07, 0.39 MgC ha(-1) y(-1)), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 0.09, 0.55 % y(-1)) compared with the tree community as a whole (+0.15 % y(-1)); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 0.02, 0.37 % y(-1)), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.
PDF
