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Most canopy species in lowland tropical rain forests in Southeast Asia, represented by Dipterocarpaceae, undergo mast reproduction synchronously at community level during a general flowering event. Such events occur at irregular intervals of 2–10 years. Some species do not necessarily participate in every synchronous mast reproduction, however. This may be due to a lack of carbohydrate resources in the trees for masting. We tested the hypothesis that interspecific differences in the time required to store assimilates in trees for seed production are due to the frequency of masting and/or seed size in each species. We examined the relationship between reproductive frequency and the carbon accumulation period necessary for seed production, and between the seed size and the period, using radiocarbon analysis in 18 dipterocarp canopy species. The mean carbon accumulation period was 0.84 years before seed maturation in all species studied. The carbon accumulation period did not have any significant correlation with reproductive frequency or seed size, both of which varied widely across the species studied. Our results show that for seed production, dipterocarp masting species do not use carbon assimilates stored for a period between the masting years, but instead use recent photosynthates produced primarily in a masting year, regardless of the masting interval or seed size of each species. These findings suggest that storage of carbohydrate resources is not a limiting factor in the masting of dipterocarps, and that accumulation and allocation of other resources is important as a precondition for participation in general flowering.

1. Our understanding of the factors that shape reproductive phenology in Southeast Asian tropical forests comes almost exclusively from studies of everwet, general flowering (GF) forests. Therefore, we examined the phenology of an evergreen dipterocarp forest in a climate exhibiting seasonal drought in Southeast Asia, with the aim of evaluating what evidence it brings to bear on the hypothesis that reproductive phenology in contemporary dipterocarp forests is the product of evolutionarily conserved responses to seasonal drought. 2. We hypothesized that (1) dipterocarp forests that experience seasonal drought would exhibit annual reproduction and seasonality similar to that of forests with similar climates in other parts of the world, (2) within species, flowering frequency would be climate-dependent, not a fixed trait, and (3) climatic events that cue flowering in everwet forests would also cue flowering in seasonally dry forests. 3. From 2001 to 2009, we monitored flowering and fruiting monthly for 1,344 trees (>300 spp.) in a forest in Khao Chong (KC), Thailand that experiences an annual 2-4 month dry season, assessing frequency, duration, seasonality and synchrony of reproduction. We also examined phenological records for trees in three dipterocarp-dominated forests in everwet climates. 4. Reproductive phenology of the KC forest was more similar to tropical forests with similar rainfall seasonality in other parts of the world than it was to dipterocarpdominated forests in everwet regions of Southeast Asia. The forest exhibited annual reproduction, with peak flowering occurring at the end of the dry season and peak fruiting occurring early in the wet season. Approximately half of all species and individuals reproduced annually, including several species that are known to exhibit \"general flowering\" in everwet climates. Short dry spells appeared to cue flowering for early flowering species, while later flowering species may have responded to either low precipitation or temperature, consistent with data from everwet forests. GF behaviour also showed significant phylogenetic signal. 5. Synthesis. Our results support the hypothesis that the phenological behaviour of both seasonal and everwet dipterocarp forests may have initially evolved in response to seasonal drought and that the general flowering phenomenon is a product of evolutionary conservatism.

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 canopy-emergent 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.

Climatic factors have commonly been attributed as the trigger of general flowering, a unique community-level mass flowering phenomenon involving most dipterocarp species that forms the foundation of Southeast Asian tropical rainforests. This intriguing flowering event is often succeeded by mast fruiting, which provides a temporary yet substantial burst of food resources for animals, particularly frugivores. However, the physiological mechanism that triggers general flowering, particularly in dipterocarp species, is not well understood largely due to its irregular and unpredictable occurrences in the tall and dense forests. To shed light on this mechanism, we employed ecological transcriptomic analyses on an RNA-seq dataset of a general flowering species, Shorea curtisii (Dipterocarpaceae), sequenced from leaves and buds collected at multiple vegetative and flowering phenological stages. We assembled 64,219 unigenes from the transcriptome of which 1,730 and 3,559 were differentially expressed in the leaf and the bud, respectively. Differentially expressed unigene clusters were found to be enriched with homologs of Arabidopsis thaliana genes associated with response to biotic and abiotic stresses, nutrient level, and hormonal treatments. When combined with rainfall data, our transcriptome data reveals that the trees were responding to a brief period of drought prior to the elevated expression of key floral promoters and followed by differential expression of unigenes that indicates physiological changes associated with the transition from vegetative to reproductive stages. Our study is timely for a representative general flowering dipterocarp species that occurs in forests that are under the constant threat of deforestation and climate change as it pinpoints important climate sensitive and flowering-related homologs and offers a glimpse into the cascade of gene expression before and after the onset of floral initiation.

Dipterocarpaceae, a dominant family of trees in South-East Asian tropical forests, are remarkable in that they exhibit supra-annual mass-flowering events. The flowering patterns are related to the El Niño Southern Oscillation, but the mechanism that precipitates mass-flowering is still debated. Here, we test if a cumulative-trigger model that tracks resource availability, specifically light, may better explain dipterocarp phenology than a direct-environmental-trigger mechanism. Using 11 flowering time series with an average length of 29 y and variety of candidate predictor variables (precipitation, cloud cover, minimum temperature and El Niño indices) we could not find a plausible direct-environmental-trigger (median AUCs across regions from 0.53 to 0.57 indicating near random predictions). The cumulative-trigger model based on El Niño indices showed better predictive results (AUC 0.67), which could further be improved by resetting the resource at known flowering events (AUC 0.76). Additional support for a cumulative-trigger model comes from the observation that regional differences in the time of year of peak flowering correspond to where El Niño effects are strongest. We conclude that cumulative resource tracking is an evolutionary plausible trigger mechanism that has other primary evolutionary advantages, such as predator satiation.

General flowering is a community-wide masting phenomenon, which is thus far documented only in aseasonal tropical forests in Asia. Although the canopy and emergent layers of forests in this region are dominated by species of a single family, Dipterocarpaceae, general flowering involves various plant groups. Studying proximate factors and estimating the flowering patterns of the past and future may aid our understanding of the ecological significance and evolutionary factors behind this phenomenon. Here we show that this phenomenon is most likely triggered by irregular droughts based on 10 years of observations. In the aseasonal forests of SE Asia, droughts tend to occur during transition periods from La Niña to El Niño, which results in an irregular 6-7-yr cycle involving a dry period with several droughts and a wet period without droughts. The magnitude of a flowering event also depends on the timing of droughts associated with the El Niño southern oscillation (ENSO) cycle, with the largest events occurring after an interval of several years with no flowering. Because most plant species can only reproduce successfully during large flowering events, changes in the ENSO cycle resulting from global warming, may have serious ramifications for forest regeneration in this region.

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.

1 The aim of this study was to document patterns in tree reproductive phenology in a rain forest of central Borneo and examine relationships between phenology and climatic patterns. 2 A 10-year data set (1990-2000) of monthly observations of flowering and fruit production of 171 trees (including 39 members of the Dipterocarpaceae) at Barito Ulu, Central Kalimantan, Indonesia, showed that most trees (73%) underwent reproductive activity on a supra-annual timescale. 3 There were three general flowering (GF) events, in 1991, 1994 and 1997, which were preceded by major drought periods (30-day sliding total rainfall of less than 100 mm for more than 10 days) in which at least 40% of dipterocarps and at least 18% of all other trees underwent synchronized reproductive activity; there was also a minor event in 1990. Around 1.3% of trees flowered and 3.8% produced fruit in months outside of these four events. 4 At the community level, the strongest negative correlation was found between the percentage of flowering individuals and total rainfall in the preceding 150 days. 5 Within three genera of dipterocarps examined in more detail (Dipterocarpus, Shorea and Vatica) there were clear and consistent patterns of sequential flowering with certain species flowering early in the GF events and others towards the end of these events. 6 Our results confirm the importance of large-scale climatic fluctuations (El Niño-Southern Oscillation) on plant reproductive phenology in South-east Asian tropical forests and indicate that drought may be a more important cue than low night-time temperatures.

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.