Declining insolation induces synchronous flowering of <Emphasis Type="Italic">Montanoa</Emphasis> and <Emphasis Type="Italic">Simsia</Emphasis> (Asteraceae) between Mexico and the Equator

We analyze the latitudinal shift in the onset of synchronous flowering in the woody genera Montanoa and Simsia (Asteraceae) between Mexico (28° N) and the Equator, where it cannot be caused by declining day length. Synchronous flowering of >100 Montanoa quadrangularis trees was observed during two consecutive years near Cali, Colombia (4° N). Analysis of herbarium specimens yielded flowering periods for 21 Montanoa species and 18 Simsia species between 4 and 28° N. Daily insolation is a function of day length and the angle at which the sun’s rays strike the earth. Between Mexico and Colombia (4° N), the maximum of insolation gradually shifts from the summer solstice to the autumn equinox. In parallel, flowering of Montanoa and Simsia starts progressively later between July and November, during the period of declining insolation. Near the Equator, there are two periods of declining insolation, and correspondingly, two flowering periods. Thus, at all tropical latitudes, flowering time of Montanoa and Simsia is highly correlated with declining insolation. The seasonal decline in daily insolation, rather than in photoperiod, apparently induces synchronous flowering of Montanoa and Simsia at the same time each year.     

Synchronous flowering of the rubber tree (Hevea brasiliensis) induced by high solar radiation intensity

How tropical trees flower synchronously near the equator in the absence of significant day length variation or other meteorological cues has long been a puzzle. The rubber tree (Hevea brasiliensis) is used as a model to investigate this phenomenon. The annual cycle of solar radiation intensity is shown to correspond closely with the flowering of the rubber tree planted near the equator and in the subtropics. Unlike in temperate regions, where incoming solar radiation (insolation) is dependent on both day length and radiation intensity, insolation at the equator is due entirely to the latter. Insolation at the upper atmosphere peaks twice a year during the spring and autumn equinoxes, but the actual solar radiation that reaches the ground is attenuated to varying extents in different localities. The rubber tree shows one or two flowering seasons a year (with major and minor seasons in the latter) in accordance with the solar radiation intensity received. High solar radiation intensity, and in particular bright sunshine (as distinct from prolonged diffuse radiation), induces synchronous anthesis and blooming in Hevea around the time of the equinoxes. The same mechanism may be operational in other tropical tree species.     

Constraints on flowering phenology in a phryganic (East Mediterranean shrub) community

The phenological pattern of flowering at the community level was studied in a Greek phryganic ecosystem near Athens for 4 consecutive years. Flowering is strongly seasonal: 80% of the insect-pollinated flora, which consists of 133 species, blooms between February and June. There is a variably expressed secondary flowering period in autumn. The pollinating fauna follows a strongly correlated pattern of abundance. Two types of plants were distinguished: pauciflorous species bearing <10 flowers that are large compared to the plant body, and multiflorous species with many small flowers. Pauciflorous species flower in the winter half of the year, while multiflorous species flower mainly in the summer half. The mean flower life spans are 9 and 3 days, respectively. The duration of flowering (DF) for each species is 55 days on average, which is long compared to other communities. The DF shows year-to-year variations, concomitant with the vicissitudes of the climate. The start of flowering of a species is statistically correlated with the temperature in the previous month, not with rainfall; its end date of flowering only partly compensates for the time gained or lost. DF is maximal in winter. The average flower life span of species flowering at any given date varies strongly and independently of the average DF. We tested the hypothesis that flowering phenology is set by phylogenetic and life form constraints. This could not be corroborated for phylogeny, evidently because of the overriding influence of the mediterranean climate, and probably also for biogeographical reasons. In contrast, life forms and multiflorous and pauciflorous species show strong differences. Many (51) of the species are therophytes; we tested the hypothesis that because of their annual habit they would be more dependent on pollination than perennials. Thus we anticipated that therophytic species would be differentiated from perennials in their flowering phenologies. This is not corroborated. We therefore conclude that the seed bank plays a role that is analogous to that of a perennial plant body.     

Plant-pollinator interactions in tropical monsoon forests in Southeast Asia

Forests with different flora and vegetation types harbor different assemblages of flower visitors, and plant-pollinator interactions vary among forests. In monsoon-dominated East and Southeast Asia, there is a characteristic gradient in climate along latitude, creating a broad spectrum of forest types with potentially diverse pollinator communities. To detect a geographical pattern of plant-pollinator interactions, we investigated flowering phenology and pollinator assemblages in the least-studied forest type, i.e., tropical monsoon forest, in the Vientiane plain in Laos. Throughout the 5-year study, we observed 171 plant species blooming and detected flower visitors on 145 species. Flowering occurred throughout the year, although the number of flowering plant species peaked at the end of dry season. The dominant canopy trees, including Dipterocarpaceae, bloomed annually, in contrast to the supra-annual general flowering that occurs in Southeast Asian tropical rain forests. Among the 134 native plant species, 68 were pollinated by hymenopterans and others by lepidopterans, beetles, flies, or diverse insects. Among the observed bees, Xylocopa, megachilids, and honeybees mainly contributed to the pollination of canopy trees, whereas long-tongued Amegilla bees pollinated diverse perennials with long corolla tubes. This is the first community-level study of plant-pollinator interactions in an Asian tropical monsoon forest ecosystem.     

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.     

The genetic basis of flowering responses to seasonal cues

Plants respond to the changing seasons to initiate developmental programmes precisely at particular times of year. Flowering is the best characterized of these seasonal responses, and in temperate climates it often occurs in spring. Genetic approaches in Arabidopsis thaliana have shown how the underlying responses to changes in day length (photoperiod) or winter temperature (vernalization) are conferred and how these converge to create a robust seasonal response. Recent advances in plant genome analysis have demonstrated the diversity in these regulatory systems in many plant species, including several crops and perennials, such as poplar trees. Here, we report progress in defining the diverse genetic mechanisms that enable plants to recognize winter, spring and autumn to initiate flower development.     

Long‐term increases in tropical flowering activity across growth forms in response to rising CO2 and climate change

Mounting evidence suggests that anthropogenic global change is altering plant species composition in tropical forests. Fewer studies, however, have focused on long‐term trends in reproductive activity, in part because of the lack of data from tropical sites. Here, we analyze a 28‐year record of tropical flower phenology in response to anthropogenic climate and atmospheric change. We show that a multidecadal increase in flower activity is most strongly associated with rising atmospheric CO₂ concentrations using yearly aggregated data. Compared to significant climatic factors, CO₂ had on average an approximately three‐, four‐, or fivefold stronger effect than rainfall, solar radiation, and the Multivariate ENSO Index, respectively. Peaks in flower activity were associated with greater solar radiation and lower rainfall during El Niño years. The effect of atmospheric CO₂ on flowering has diminished over the most recent decade for lianas and canopy trees, whereas flowering of midstory trees and shrub species continued to increase with rising CO₂. Increases in flowering were accompanied by a lengthening of flowering duration for canopy and midstory trees. Understory treelets did not show increases in flowering but did show increases in duration. Given that atmospheric CO₂ will likely continue to climb over the next century, a long‐term increase in flowering activity may persist in some growth forms until checked by nutrient limitation or by climate change through rising temperatures, increasing drought frequency and/or increasing cloudiness and reduced insolation.     

Phenology of Ficus variegata in a seasonal wet tropical forest at Cape Tribulation, Australia

Abstract. We studied the phenology of 198 mature trees of the dioecious fig Ficus variegata Blume (Moraceae) in a seasonally wet tropical rain forest at Cape Tribulation, Australia, from March 1988 to February 1993. Leaf production was highly seasonal and correlated with rainfall. Trees were annually deciduous, with a pronounced leaf drop and a pulse of new growth during the August-September drought. At the population level, figs were produced continually throughout the study but there were pronounced annual cycles in fig abundance. Figs were least abundant during the early dry period (June-September) and most abundant from the late dry season (October-November) through the wet season (December-April). The annual peak in reproduction actually reflected two staggered peaks arising from gender differences in fig phenology. In this dioecious species, female and male trees initiated their maximal fig crops at different times and flowering was to some extent synchronized within sexes. Fig production in the female (seed-producing) trees was typically confined to the wet season. Male (wasp-producing) trees were less synchronized than female trees but reached a peak level of fig production in the months prior to the onset of female fig production. Male trees were also more likely to produce figs continually. Asynchrony among male fig crops during the dry season could maintain the pollinator population under adverse conditions through within- and among-tree wasp transfers.     

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.     

Phenology of temperate trees in tropical climates

Several North American broad-leaved tree species range from the northern United States at 47°N to moist tropical montane forests in Mexico and Central America at 15–20°N. Along this gradient the average minimum temperatures of the coldest month (T _Jan), which characterize annual variation in temperature, increase from –10 to 12°C and tree phenology changes from deciduous to leaf-exchanging or evergreen in the southern range with a year-long growing season. Between 30 and 45°N, the time of bud break is highly correlated with T _Jan and bud break can be reliably predicted for the week in which mean minimum temperature rises to 7°C. Temperature-dependent deciduous phenology—and hence the validity of temperature-driven phenology models—terminates in southern North America near 30°N, where T _Jan>7°C enables growth of tropical trees and cultivation of frost-sensitive citrus fruits. In tropical climates most temperate broad-leaved species exchange old for new leaves within a few weeks in January-February, i.e., their phenology becomes similar to that of tropical leaf-exchanging species. Leaf buds of the southern ecotypes of these temperate species are therefore not winter-dormant and have no chilling requirement. As in many tropical trees, bud break of Celtis, Quercus and Fagus growing in warm climates is induced in early spring by increasing daylength. In tropical climates vegetative phenology is determined mainly by leaf longevity, seasonal variation in water stress and day length. As water stress during the dry season varies widely with soil water storage, climate-driven models cannot predict tree phenology in the tropics and tropical tree phenology does not constitute a useful indicator of global warming.     

Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows

The study of phylogenetic conservatism in alpine plant phenology is critical for predicting climate change impacts; currently we have a poor understanding of how phylogeny and climate factors interactively influence plant phenology. Therefore, we explored the influence of phylogeny and climate factors on flowering phenology in alpine meadows. For two different types of alpine plant communities, we recorded phenological data, including flowering peak, first flower budding, first flowering, first fruiting and the flowering end for 62 species over the course of 5 years (2008–2012). From sequences in two plastid regions, we constructed phylogenetic trees. We used Blomberg’s K and Pagel’s lambda to assess the phylogenetic signal in phenological traits and species’ phenological responses to climate factors. We found a significant phylogenetic signal in the date of all reproductive phenological events and in species’ phenological responses to weekly day length and temperature. The number of species in flower was strongly associated with the weekly day lengths and followed by the weekly temperature prior to phenological activity. Based on phylogenetic eigenvector regression (PVR) analysis, we found a highly shared influence of phylogeny and climate factors on alpine species flowering phenology. Our results suggest the phylogenetic conservatism in both flowering and fruiting phenology may depend on the similarity of responses to external environmental cues among close relatives.     

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.     

31种木本植物开花物候与系统发育的关系

植物物候通常被认为是由环境因素,如降水、温度和日照长度所决定,然而环境因素是否是物候唯一的决定因素仍然存在很大争议。谱系结构表征了植物在进化上的顺序,该发育时序是否对物候产生影响,当前仍然未知。在调查2016年春季新疆乌鲁木齐市最常见的31种木本植物的初始开花时间、败花时间和开花持续时间的基础上,通过分析植物开花物候的分布特征、开花物候在乔灌木间的差别、以及植物谱系距离与开花物候距离间的关系,试图揭示植物的开花物候和物种谱系(进化)顺序间的关系。结果表明:(1)新疆乌鲁木齐市31种木本植物的初始开花时间为4月18日±9d、败花时间为5月5日±12d、开花持续时间为(16±8)d;(2)乔木的初始开花时间和败花时间的标准差分别均低于灌木,乔木开花物候相对灌木更稳定;(3)乔木的初始开花和败花时间均显著早于灌木(P0.05),但开花持续时间在两者间未有显著性差异(P0.05);(3)31种木本植物间的初始开花时间距离、败花时间距离和开花持续时间距离均与物种谱系距离存在显著线性回归关系(P0.05)。综上可知:乔灌木在垂直空间上的分化使得木本植物的开花物候在植物生活型间存在不同。对植物的开花物候,除已被证明的降水、温度和日照长度等环境因素的影响外,物种进化顺序也可能造成了它在植物种间、时间和空间上的变异。     

Tree size and its relationship with flowering phenology and reproductive output in Wild Nutmeg trees

Reproductive strategies, sexual selection, and their relationship with the phenotype of individuals are topics widely studied in animals, but this information is less abundant for plants. Variability in flowering phenology among individuals has direct impact on their fitness, but how reproductive phenology is affected by the size of the individuals needs further study. We quantified the flowering intensity, length, and reproductive synchronization of two sympatric dioecious Wild Nutmeg tree species (Virola, Myristicaceae) in the Brazilian Atlantic forest, and analyzed its relationships with tree size. Two distinct strategies in flowering timing and intensity were found between species (annual versus biennial flowering), and among individuals in the annual flowering species (extended versus peak flowering). Only for the annual flowering species the reproductive output is related to tree size and large trees present proportionally higher flower coverage, and lower synchronization than smaller ones. Flowering is massive and highly synchronized in the biennial species. Sex ratios are not different from 1:1 in the two species, and in the two segregated reproductive subgroups in the biennial flowering species. The biennial flowering at individual level is a novelty among reproductive patterns in plants, separating the population in two reproductive subgroups. A proportional increase in the reproductive output with size exists only for the annual flowering species. A biennial flowering can allow resource storage favouring massive flowering for all the individuals diluting their relationship with size.     

Do seasonal changes in light availability influence the inverse leafing phenology of the neotropical dry forest understory shrub Bonellia nervosa (Theophrastaceae)?

In tropical dry forests most plants are deciduous during the dry season and flush leaves with the onset of the rains. In Costa Rica, the only species displaying the opposite pattern is Bonellia nervosa. To determine if seasonal changes in light availability are associated with the leaf and reproductive phenology of this species, we monitored leaf production, survival, and life span, as well as flower and fruit production from April 2000 to October 2001 in Santa Rosa National Park. Leaf flushing and flower bud production took place shortly after the autumnal equinox when day length starts to decrease. Leaves began expansion at the end of the wet season, and plants reached 70 % of their maximum leaf area at the beginning of the dry season, maintaining their foliage throughout the entire dry period. Leaf shedding occurred gradually during the first three months of the wet season. Leaf flushing and shedding showed high synchrony, with leaf numbers being related to light availability. Maximum leaf production coincided with peaks in radiation during the middle of the dry season. Decreasing day length induces highly synchronous flower bud emergence in dry forest species, but this is the first study indicating induction of leaf flushing by declining day length.     

Phenological Adaptations in Ficus tikoua Exhibit Convergence with Unrelated Extra-Tropical Fig Trees

Flowering phenology is central to the ecology and evolution of most flowering plants. In highly-specific nursery pollination systems, such as that involving fig trees (Ficus species) and fig wasps (Agaonidae), any mismatch in timing has serious consequences because the plants must balance seed production with maintenance of their pollinator populations. Most fig trees are found in tropical or subtropical habitats, but the dioecious Chinese Ficus tikoua has a more northerly distribution. We monitored how its fruiting phenology has adapted in response to a highly seasonal environment. Male trees (where fig wasps reproduce) had one to three crops annually, whereas many seed-producing female trees produced only one fig crop. The timing of release of Ceratosolen fig wasps from male figs in late May and June was synchronized with the presence of receptive figs on female trees, at a time when there were few receptive figs on male trees, thereby ensuring seed set while allowing remnant pollinator populations to persist. F. tikoua phenology has converged with those of other (unrelated) northern Ficus species, but there are differences. Unlike F. carica in Europe, all F. tikoua male figs contain male flowers, and unlike F. pumila in China, but like F. carica, it is the second annual generation of adult wasps that pollinate female figs. The phenologies of all three temperate fig trees generate annual bottlenecks in the size of pollinator populations and for female F. tikoua also a shortage of fig wasps that results in many figs failing to be pollinated.     

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.     

迁地保育植物花期物候对迁入地气候变化的响应

以中国科学院武汉植物园内栽培的长果秤锤树(Sinojackia dolichocarpa C. J. Qi)、山白树(Sinowilsonia henryi Hemsl.)、夏腊梅(Sinocalycanthus chinensis Cheng et S. Y. Chang)、紫茎(Stewartia sinensis Rehd. et Wils.)和绒毛皂荚(Gleditsia vestita Chun et How ex B. G. Li) 5种迁地保育植物为对象,通过2008-2016年观察记录的初花期物候及整个花期长度的数据,研究花期的年际变化规律及其与迁入地武汉气候因子的相关性。结果显示:(1)从初花期来看,长果秤锤树的初花期每年提前1.25 d,紫茎的初花期每年推迟1.35 d,绒毛皂荚的初花期每年推迟1.22 d。(2)从花期长度来看,山白树的花期每年增加1.72 d,夏蜡梅的花期每年减少1.62 d,紫茎的花期每年增加0.32 d。(3)从花期与气候因子的相关性来看,年降水量、年平均相对湿度、 10℃有效积温、花前 10℃的有效积温是影响这5种植物初花期、花期长度的主要气候因子;不同物种间影响花期的主要气候因子有所差异。     

Community phenological patterns of a lowland tropical rainforest in north-eastern Australia

Leaf-fall, leaf-flush (newly expanded leaves), flowering and fruiting were recorded monthly between April 1982 and May 1985 in 703 individuals of 99 species of trees, shrubs and herbs in a logged lowland, complex rainforest. Weekly water availability and demand were modelled using pan evaporation, rainfall, and a range of estimated maximum soil water storage. The water balance model indicated that rainfall distribution was an inadequate measure of the seasonal availability of water for growth and reproduction. Intermittent shortages in available water were predicted between October and February, well into what is the wettest season of the year in terms of long-term rainfall averages. Adequate water was available in soil water stores through most of the August–October ‘dry season’. Leaf-fall, flowering, and fruiting all followed distinct seasonal patterns although appreciable levels of activity occurred in all months. Community leaf-fall increased rapidly following an annual low in the very wet, February–May period, to a peak in August. The rapid increase in leaf-fall activity was accompanied by pronounced depressions in flowering and leaf-flushing and coincided with the period of minimum temperatures and insolation. There was no evidence to suggest that this leaf-fall peak was accompanied or induced by drought. This ‘winter’ was followed by rising temperatures and insolation at a time when adequate soil water was usually available, and flowering activity (number of species, number of individuals, and total abundance) increased rapidly in August and peaked from September to October. The flowering peak was accompanied by a major leaf-flush. Leaf-flushing continued intermittently from October to February, a period characterized by maximum temperature, maximum insolation, and periods of predicted water stress when soil water stores were depleted during short, rainless periods. A second peak in leaf-flushing coincided with the wet in March-April. This was accompanied by a relatively small increase in leaf-fall activity. Fruiting showed a bimodal peak from October to April each year. Prediction of functional relationships between climate and species and individual behaviour from community phenological patterns was considered to be inappropriate.