Solar radiation and ENSO predict fruiting phenology patterns in a 15‐year record from Kibale National Park,Uganda

Fruiting, flowering, and leaf set patterns influence many aspects of tropical forest communities, but there are few long‐term studies examining potential drivers of these patterns, particularly in Africa. We evaluated a 15‐year dataset of tree phenology in Kibale National Park, Uganda, to identify abiotic predictors of fruit phenological patterns and discuss our findings in light of climate change. We quantified fruiting for 326 trees from 43 species and evaluated these patterns in relation to solar radiance, rainfall, and monthly temperature. We used time‐lagged variables based on seasonality in linear regression models to assess the effect of abiotic variables on the proportion of fruiting trees. Annual fruiting varied over 3.8‐fold, and inter‐annual variation in fruiting is associated with the extent of fruiting in the peak period, not variation in time of fruit set. While temperature and rainfall showed positive effects on fruiting, solar radiance in the two‐year period encompassing a given year and the previous year was the strongest predictor of fruiting. As solar irradiance was the strongest predictor of fruiting, the projected increase in rainfall associated with climate change, and coincident increase in cloud cover suggest that climate change will lead to a decrease in fruiting. ENSO in the prior 24‐month period was also significantly associated with annual ripe fruit production, and ENSO is also affected by climate change. Predicting changes in phenology demands understanding inter‐annual variation in fruit dynamics in light of potential abiotic drivers, patterns that will only emerge with long‐term data.     

Disparity in Onset Timing and Frequency of Flowering and Fruiting Events in Two Bornean Peat‐Swamp Forests

The timing and frequency of flowering and fruiting events are key tropical forest characteristics that have substantial influence on fauna. Although our understanding of geographic variation in habitat‐wide timing and frequency of flowering and fruiting is advancing, corresponding information for individual tree species is limited. Thus, we compared climate and reproductive phenology of 16 tree species over 70 mo at two Bornean tropical peat‐swamp forest sites. We found significant inter‐site correlations in rainfall and temperature, and only small absolute temperature differences. In both sites, most species exhibited within‐site synchrony in flowering and fruiting onset. Broad‐scale flowering and fruiting onset frequency classifications showed high congruence between sites. Significant correlations in flowering and fruiting onset timing between sites were found for only 19 and 17 percent of the species, respectively. This remained the case when applying 1‐ and 2‐month lag periods for both sites, with neither site consistently lagging behind. Significant differences in the exact frequency of new flowering and fruiting events were detected for 44 and 58 percent of species, respectively, and no significant relationships between the onset timing synchrony and exact frequency of new reproductive events were found for either flowers or fruit. We conclude that inter‐site climatic and ecological similarities do not necessarily lead to high inter‐site synchrony in either onset timing or exact frequency of tree reproductive events. Potential reasons for this are discussed, as are the implications for understanding tropical forest ecology and improving forest restoration project seed collections.     

Global patterns in fruiting seasons

Aim   To identify geographical and climatic correlates of the timing of fruit production in fleshy fruited plant communities.
Location   Global.
Methods   We searched the literature for studies documenting monthly variation in the number of fleshy fruited species bearing ripe fruits in plant communities (i.e. fruit phenologies). From these data, we used circular vector algebra to characterize seasonal peaks in fruit production (mean date, as an angle) and the length of fruiting seasons (as a circular standard deviation). Generalized linear models and circular correlations were used to assess whether latitudinal patterns in fruit phenologies could be explained by variation in temperature, precipitation and actual evapotranspiration (AET).
Results   Dates of peak fruit production and the length of fruiting seasons showed consistent differences with latitude. Annual peaks in fruit production occurred 1 to 3 months after the summer solstice at high-latitude sites in both hemispheres. Fruiting seasonality increased with latitude, indicating that fruiting seasons were longer in the tropics and shorter toward the poles. AET was the best climatic predictor of fruit phenologies. Annual peaks in fruit production were positively associated with annual peaks in AET and temperature, while fruiting seasons were shorter in areas with pronounced annual variation in AET.
Main conclusions   Global patterns in fruiting seasons are associated with global variation in climate. Across the globe, fleshy fruits are produced after annual periods of elevated water–energy availability. Fruiting seasonality is also more pronounced in areas with strongly seasonal water–energy inputs. Therefore, the timing of reproduction in fleshy fruited plant communities appears to be determined, at least in part, by spatial and temporal variation in energy supplies needed to subsidise plant reproduction.     

Fruiting phenology in a tropical dry evergreen forest on the Coromandel coast of India in relation to plant life-forms,physiognomic groups,dispersal modes,and climatic constraints

The fruiting phenology of 22 woody plant species belonging to 19 families was studied with respect to life-forms, physiognomic groups and dispersal modes, for 1 year at monthly intervals, in a tropical dry evergreen forest at Oorani (12°11′N, 79°57′E) on the Coromandel coast of India. At the community level, bimodal fruiting pattern prevailed, with a major peak in the dry season and a minor one in the early rainy season. An annual fruiting pattern was observed in many species and among the studied species fruiting lasted for 2–9 months. There was no significant difference in the frequency of species at three fruiting stages across the life-form categories and many species of upper and lower canopy trees and lianas were in the ripe fruiting phase during the late dry season. Plant physiognomic groups displayed distinct seasonality in fruiting pattern. The fruit maturation period was much longer for the wet season fruiting brevi-deciduous species than evergreen and deciduous species that fruited during the dry season. The variation in timing of fruiting behaviour among zoochorous species demonstrated less seasonality and zoochorous fruits were available throughout the year. Fruiting in anemochorous species peaked during the driest months and dryness favoured the dissemination of seeds. The fruiting patterns observed in the studied tropical dry evergreen forest across various plant traits were comparable with patterns recorded in other tropical seasonal forests.     

Flowering and Fruiting Patterns in a Subtropical Rain Forest,Taiwan

Reproductive patterns of tropical and temperate plants are usually associated with climatic seasonality, such as rainfall or temperature, and with their phylogeny. It is still unclear, however, whether plant reproductive phenology is influenced by climatic factors and/or phylogeny in aseasonal subtropical regions. The plant reproductive phenology of a subtropical rain forest in northern Taiwan (24°45′ N, 121°35′ E) was monitored at weekly intervals during a 7‐yr period (2002–2009). The flowering patterns of 46 taxa and fruiting patterns of 26 taxa were examined and evaluated in relation to climatic seasonality, phylogenetic constraints, and different phenophases. Our results indicated that most of the studied species reproduced annually. Additionally, both community‐wide flowering and fruiting patterns exhibited distinct annual rhythms and varied little among years. The community flowering peak matched seasonal changes in day length, temperature, and irradiance; while the community fruiting peak coincided with an increase in bird richness and the diet‐switching of resident omnivorous birds. In addition, phylogenetically closely related species tended to reproduce during the same periods of a year. Neither the mean flowering dates nor seasonal variation in solar radiation, however, was related to the fruit development times. Our results indicate the importance of abiotic, biotic, and evolutionary factors in determining the reproductive phenology in this subtropical forest.     

High interannual variation in the diet of a tropical forest frugivore (Hylobates lar)

Frugivores must deal with seasonal changes in fruit availability and changes from year to year, as most species of tropical forest fruiting trees have considerable interannual variation in phenology and many are mast fruiters. We quantified seasonal and interannual changes in the fruit diet in a frugivore and important seed disperser, the white‐handed gibbon, Hylobates lar, in Thailand. We used 40‐d following data during April and May replicated in six consecutive years to study interannual variability in the diet and compared it with seasonal changes measured in monthly samples of the same size collected in three successive years. The 40‐d periods of following also allowed us to measure the decline in dietary similarity with time over a finer scale. We measured fruit diet similarity between replicated 5‐d periods using the percentage overlap (Renkonen's) index and Jaccard's similarity index. Seasonally, average dietary overlap between adjacent months was low, and similarity approached zero after four months. Average rate of decline in similarity exceeded 20 percent per 5‐d period. Variation in fruit species in the diet between years was high and was correlated with interannual variation in fruiting phenology. The strongest correlation occurred in the case of Nephelium melliferum, a highly preferred species that dominated the diet in good fruiting years. It is difficult to separate changes in food species preference from changes in availability from year to year. We devised a relative measure of preference that depends on the degree to which the gibbons rely on prior knowledge to find sources.     

Annual cycles are the most common reproductive strategy in African tropical tree communities

We present the first cross‐continental comparison of the flowering and fruiting phenology of tropical forests across Africa. Flowering events of 5446 trees from 196 species across 12 sites and fruiting events of 4595 trees from 191 species across 11 sites were monitored over periods of 6 to 29 years and analyzed to describe phenology at the continental level. To study phenology, we used Fourier analysis to identify the dominant cycles of flowering and fruiting for each individual tree and we identified the time of year African trees bloom and bear fruit and their relationship to local seasonality. Reproductive strategies were diverse, and no single regular cycle was found in >50% of individuals across all 12 sites. Additionally, we found annual flowering and fruiting cycles to be the most common. Sub‐annual cycles were the next most common for flowering, whereas supra‐annual patterns were the next most common for fruiting. We also identify variation in different subsets of species, with species exhibiting mainly annual cycles most common in West and West Central African tropical forests, while more species at sites in East Central and East African forests showed cycles ranging from sub‐annual to supra‐annual. Despite many trees showing strong seasonality, at most sites some flowering and fruiting occurred all year round. Environmental factors with annual cycles are likely to be important drivers of seasonal periodicity in trees across Africa, but proximate triggers are unlikely to be constant across the continent.     

Seasonal Patterns of Flowering and Fruiting in a Dry Tropical Forest in Jamaica1

Tropical dry forests occupy more area and are more endangered than rainforests, yet their regeneration ecology has received less study and is consequently poorly understood. We recorded the flowering and fruiting phenology of a tropical dry forest in Jamaica over a period of 26 mo within ten 15 × 15‐m plots. Community‐wide recruitment reached a maximum in the wet season, whereas no recruitment occurred during the dry season. We observed a unimodal peak in rainfall and fruit production, and the periodicity and intensity of seed production were significantly correlated with rainfall seasonality (the optimal time for germination). Flowering at the community and system levels lagged behind a significant increase and subsequent decrease in rainfall by 7 and 3 mo, respectively, indicating that the dominant factor controlling flowering periodicity is the passage of the major (4‐mo long) rainy season and changes in soil moisture conditions. Fruiting lagged behind flowering by 2 mo and a significant increase in fruiting occurred 2 mo prior to a significant increase in rainfall. At the population level, a correspondence analysis identified a major dichotomy in the patterns of flowering and fruiting between species and indicated two broad species groups based on their time of peak fruiting and the number of times they were in fruit. These were either individuals which were usually in peak fruit 1–2 mo prior to the start of the major rainy season or those that were in fruit more or less continuously throughout the year with no peak fruiting time. This study supports the view that seasonal variation in rainfall and hence soil water availability constitutes both the proximate and the ultimate cause of flowering periodicity in tropical dry forests.     

Long‐term Changes in Fruit Phenology in a West African Lowland Tropical Rain Forest are Not Explained by Rainfall

Decreases in rainfall have been proposed to have a negative impact on tropical rain forests, and West Africa is currently experiencing a decline in rainfall at the multi‐decadal scale. Here, we present analyses of a long‐term dataset on the plant fruiting status from individuals of 44 species of the tropical rain forest of Taï National Park, Côte d'Ivoire. This study includes records of 1401 individuals collected at monthly intervals for over 12 yr, 984 of which survived throughout the entire study period. The aims of this study were to: (1) quantitate inter‐annual trends in species and forest scale fruit presence; and (2) test the importance of rainfall in explaining inter‐annual fruit presence variability. Long‐term upward trends in the expected proportion of individuals with fruits were found for the majority of species, while no significant downward trends were detected, driving a significant upward trend at the community level. Peak production months of the upward trending species were not associated with the dry or wet season. Significant rainfall correlations with the total proportion of individuals showing fruit were generally negative, with only five species showing significant positive correlations. Taken together, these results suggest that the observed inter‐annual trends and variability of fruit abundance are currently not associated with rainfall. We discuss several parsimonious and complex alternative explanations.     

Resource tracking and its conservation implications for an obligate frugivore (Procnias tricarunculatus,the three‐wattled bellbird)

In Monteverde, Costa Rica, the vulnerable Three‐wattled Bellbird (Procnias tricarunculatus) feeds primarily upon the fruit of Lauraceae species during its reproductive and post‐reproductive seasons. To understand and advance appropriate conservation measures, this study identified the bellbird's foraging challenges in its search for a temporally and spatially fluctuating resource. Although there are at least 96 species of Lauraceae found in the five life zones of Monteverde, the distinct distributions of tree species both among and within life zones require the bellbirds to track seasonal fruiting across the various zones. In this 6‐year study, we monitored the fruiting of tree species and bellbird abundance in 24 study plots within its post‐reproductive life zone, the Premontane Wet forest, to identify preferred bellbird food resources and how the fruiting of these species drives the spatial distribution of the bellbird. Our research revealed phenological patterns of annual, biennial, and triennial fruiting with high levels of fruiting synchrony within several identified key fruit species. Of critical conservation importance is that no single species of Lauraceae produced a consistent food supply for bellbirds each year. Therefore, even within life zones, the bellbird's survival depends on its mobility to search for and obtain fruit, as well as the availability of fruits of multiple tree species. The conservation implications include focused attention on multiple core areas within given life zones, protection of existing forest and remnant trees, and forest restoration with plantings of multiple tree species. We suspect that other tropical frugivorous species face similar conservation challenges.