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 | 587 Journal of Ecology CHEN Et al.

Abstract. The importance of lianas through time and their effect on tree reproduction are evaluated for the first time in a Southeast Asian Dipterocarp forest. We quantified flower and seed production by lianas and trees for 13 years, assessed liana loads in the crowns of all trees larger than 30 cm in diameter at breast height (1.3 m) in 2002 and 2014, and assessed levels of reproduction for the same trees during a strong general flowering event in 2014 for the 50-ha forest dynamics plot at the Pasoh Forest Reserve, Malaysia. General flowering refers to synchronous reproduction by hundreds of plant species at irregular, multiyear intervals and only occurs in Southeast Asian Dipterocarp forests. Overall, lianas were present in 50% of tree crowns and comprised 31% of flower production and 46% of seed production. Lianas reduced growth, survival, and reproduction by their host trees. Lianas were less frequent in canopyemergent trees, Dipterocarps comprised a disproportionately large proportion of canopy emergents, and, as a consequence, lianas were less frequent in Dipterocarps than in trees from other plant families. Lianas infested the crowns of significantly fewer trees in 2014 (47.9%) than in 2002 (52.3%); however, the decrease was restricted to trees with the lightest liana loads and sample sizes and statistical power were enormous. Lianas comprised a stable proportion of flower production and a highly variable proportion of seed production from 2002 through 2013. We conclude lianas have a huge impact on trees in this forest and were a stable component of the forest between 2002 and 2014. The emergent habit and associated ability to avoid lianas might contribute to the success of the Dipterocarpaceae.

Summary 1The 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. 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.

Spatial and temporal variation in seedling dynamics was assessed using records of community-wide seedling demography collected with identical monitoring methods at four tropical lowland forests in Panama, Malaysia, Ecuador and French Guiana for periods of between 3 and 10 y. At each site, the fates of between 8617 and 391 777 seedlings were followed through annual censuses of the 370-1008 1-m 2 seedling plots. Within-site spatial and inter-annual variation in density, recruitment, growth and mortality was compared with among-site variability using Bayesian hierarchical modelling to determine the generality of each site's patterns and potential for meaningful comparisons among sites. The Malaysian forest, which experiences community-wide masting, was the most variable in both seedling density and recruitment. However, density varied year-to-year at all sites (CV amongyearsatsite = 8-43%), driven largely by high variability in recruitment rates (CV = 40-117%). At all sites, recruitment was more variable than mortality (CV= 5-64%) or growth (CV= 12-51%). Increases in mortality rates lagged 1 y behind large recruitment events. Within-site spatial variation and inter-annual differences were greater than differences among site averages in all rates, emphasizing the value of long-term comparative studies when generalizing how spatial and temporal variation drive patterns of recruitment in tropical forests.

Bamboo forests, due to rapid growth and short harvest rotation, play an important role in carbon cycling and local economic development. Accurate estimation of bamboo forest aboveground biomass (AGB) has garnered increasing attention during the past two decades. However, remote sensing-based AGB estimation for bamboo forests is challenging due to poor understanding of the mechanisms between bamboo forest growth characteristics and remote sensing data. The objective of this research is to examine the remote sensing characteristics of on-year and off-year bamboo forests at different dates and their AGB estimation performance. This research used multiple Sentinel-2 data to explore AGB estimation of bamboo forests in Zhejiang Province, China, by taking into account the unique characteristics of on-year and off-year bamboo forest growth features. Combining field survey data and Sentinel-2 spectral responses (spectral bands and vegetation indices) and textural images, random forest was used to identify key variables for AGB estimation. The results show that (1) the on-year and off-year bamboo forests have considerably different spectral signatures, especially in the wavelengths between red edge 2 and near-infrared wavelength (NIR2) (740-865 nm), making it possible to separate on-year and off-year bamboo forests; (2) on-year bamboo forests have similar spectral signatures although AGB increases from as small as 40 Mgha −1 to as high as 90 Mgha −1 , implying that optical sensor data cannot effectively model on-year bamboo AGB; (3) off-year bamboo AGB has significant relationships with red and shortwave infrared (SWIR) spectral bands in the April image and with red edge 2 in the July image, but the AGB saturation problem yields poor estimation accuracy; (4) stratification considerably improved off-year bamboo AGB estimation but not on-year, non-stratification using the April image is recommended; and (5) Sentinel-2 data cannot solve the bamboo AGB data saturation problem when AGB is greater than 70 Mgha −1 , similar to other optical sensor data such as Landsat. More research should be conducted in the future to integrate multiple sources-remotely sensed data (e.g., lidar, optical sensor data) and ancillary data (e.g., soil, topography)-into AGB modeling to improve the estimation. The use of very high spatial resolution images that can effectively extract tree density information may improve bamboo AGB estimation and yield new insights.

The family Dipterocarpaceae includes 470 tree species from 13 genera in South and South-East Asian tropical forests (Ashton 1982). Many dipterocarp species in aseasonal lowland rain forests of western Malesia flower synchronously during masting (or general flowering) events, which usually occur at irregular intervals of 2–10 y (Ashton et al. 1988). Very few individuals flower at other times, and successful recruitment of seedlings is limited to those masting events (Ashton et al. 1988, Curran et al. 1999).

1. Organisms of all species must balance their allocation to growth, survival and recruitment. Among tree species, evolution has resulted in different life-history strategies for partitioning resources to these key demographic processes.

Many species of Dipterocarpaceae and other plant families reproduce synchronously at irregular, multi-year intervals in Southeast Asian forests. These community-wide general flowering events are thought to facilitate seed survival through satiation of generalist seed predators. During a general flowering event, closely related Shorea species (Dipterocarpaceae) stagger their flowering times by several weeks, which may minimize cross pollination and interspecific competition for pollinators. Generalist, pre-dispersal seed predators might also track flowering hosts and influence predator satiation. We addressed the question of whether pre-dispersal seed predation differed between early and late flowering Shorea species by monitoring flowering, fruiting and seed predation intensity over two general flowering events at the Pasoh Research Forest, Malaysia. Pre-dispersal insect seed predators killed up to 63 percent of developing seeds, with Nanophyes shoreae, a weevil that feeds on immature seeds being the most important predator for all Shorea species. This weevil caused significantly greater pre-dispersal seed predation in earlier flowering species. Long larval development time precluded oviposition by adults that emerged from the earliest flowering Shorea on the final flowering Shorea. In contrast, larvae of weevils that feed on mature seeds before seed dispersal (Alcidodes spp.), appeared in seeds of all Shorea species almost simultaneously. We conclude that general flowering events have the potential to satiate post-dispersal seed predators and pre-dispersal seed predators of mature fruit, but are less effective at satiating pre-dispersal predators of immature fruit attacking early flowering species.Abstract in Malay is available with online material.