Survival of Flower-visiting Chrysomelids during Non General-flowering Periods in Bornean Dipterocarp Forests

In SE Asian rain forests, general flowering, a community-wide synchronous flowering, occurs at irregular and supra-annual intervals. During general flowering periods (GFP), most Dipterocarpaceae and many other trees flower profusely, while flowering plants are scant between GFP. During flowerless periods, anthophilous animals that depend on floral resources for food may suffer food shortages and subsequently decrease in abundance. Flower-visiting chrysomelid adults are major pollinators for some canopy tree species that flower during GFP. Although such chrysomelids feed on flower petals, the means by which they survive flowerless periods remains unknown. We determined the abundance of flower-visiting chrysomelids in GFP and non-GFP through light trap samples and examined the effects of the presence of young leaves and flowers of dipterocarps on local abundance, and feeding preferences of flower-visiting chrysomelids. We found no clear tendency that the chrysomelid species number and the abundance during GFP were consistently higher than those during non-GFP. Chrysomelid adults were more abundant on trees with many young leaves or flowers than on trees lacking young leaves and flowers. At least a few flower-visiting chrysomelids were observed feeding on young dipterocarp leaves and visiting young leaves and flowers of non-dipterocarps in the canopy during non-GFP. All results consistently suggest that chrysomelids are able to survive flowerless periods by feeding on the young leaves of dipterocarps and on the young leaves and flowers of non-dipterocarps; through this alternate feeding, chrysomelid populations are maintained at sufficient levels to function as effective pollinators of trees that flower during GFP.     

Spatiotemporal Variability in the Reproductive Success of the Continually Flowering Shrub Dillenia suffruticosa in Borneo

Continually flowering plants bloom continuously throughout the year, as often seen in plants distributed along the roadsides or in the understory layers in Southeast Asia's tropical rain forests. Dillenia suffruticosa (Griff. ex Hook. f. & Thomson) Martelli (Dilleniaceae) is one such continually flowering shrub that flowers during periods of community‐wide mass flowering, general flowering (GF), and non‐GF. During irregularly occurring GF periods, when species of all forest layers flower synchronously for several months, some pollinators migrate to the canopy layer, where GF promotes the pollination success of participating plants. Continually flowering plants share the available pollinator community with GF plants, and the reproductive success of continually flowering plants may be affected during the GF period. To assess the effects of GF on the reproductive success of a diverse range of continually flowering plants, we examined the differences in pollinator density and reproductive success between GF and non‐GF periods in D. suffruticosa at four different research sites. Although the seed set differed among the four research sites, pollinator density and fruit set did not differ between GF and non‐GF periods or research sites. Our results suggest that the reproductive success of D. suffruticosa was maintained at an approximately constant level, regardless of the flowering phenology of the canopy layer or other vegetation components.     

Generalized food-deceptive orchid species flower earlier and occur at lower altitudes than rewarding ones

Aims Food-deceptive pollination, in which plants do not offer any food reward to their pollinators, is common within the Orchidaceae. As food-deceptive orchids are poorer competitors for pollinator visitation than rewarding orchids, their occurrence in a given habitat may be more constrained than that of rewarding orchids. In particular, the success of deceptive orchids strongly relies on several biotic factors such as interactions with co-flowering rewarding species and pollinators, which may vary with altitude and over time. Our study compares generalized food-deceptive (i.e. excluding sexually deceptive) and rewarding orchids to test whether (i) deceptive orchids flower earlier compared to their rewarding counterparts and whether (ii) the relative occurrence of deceptive orchids decreases with increasing altitude.Methods To compare the flowering phenology of rewarding and deceptive orchids, we analysed data compiled from the literature at the species level over the occidental Palaearctic area. Since flowering phenology can be constrained by the latitudinal distribution of the species and by their phylogenetic relationships, we accounted for these factors in our analysis. To compare the altitudinal distribution of rewarding and deceptive orchids, we used field observations made over the entire Swiss territory and over two Swiss mountain ranges.Important findings We found that deceptive orchid species start flowering earlier than rewarding orchids do, which is in accordance with the hypotheses of exploitation of naive pollinators and/or avoidance of competition with rewarding co-occurring species. Also, the relative frequency of deceptive orchids decreases with altitude, suggesting that deception may be less profitable at high compared to low altitude.     

浙江古田山亚热带常绿阔叶林开花物候:气候因素、系统发育关系和功能性状的影响

植物的开花物候受气候因素、植物系统发育关系和功能性状的影响。然而当前植物开花物候研究中未见同时考虑这3个因素的报道。为了解它们相互之间的影响, 本研究利用中国东部地区浙江省古田山国家级自然保护区亚热带常绿阔叶林24 ha大样地(GTS; 118°03′50′′-118°11′12.2′′ E, 29°10′19′′-29°17′41′′ N)设置的130个种子雨收集器5年的开花数据检验这3个因素对开花的影响。结果表明, 古田山植物的开花高峰期集中在5月, 群落开花格局明显受温度和降雨的影响。利用植物DNA条形码数据研究发现, 植物间系统发育关系对古田山植物开花时间有显著影响, 亲缘关系近的物种开花时间更相近。植物的平均开花时间受最大树高的影响, 但不受传粉方式、花色、种子质量和扩散方式的影响。该研究结果说明气候因素、植物系统发育关系和功能性状都可能影响植物开花物候格局, 同时考虑这3个因素能够帮助我们更好地理解开花物候格局。     

An Assemblage of Hummingbird-pollinated Flowers in a Montane Forest in Southeastern Brazil

Relationships between ornithophilous flowers and hummingbirds have been little studied in southern South America, where hummingbird species richness is low. We studied an ornithophilous flower assemblage and the hummingbird pollinators in a montane forest in southeastern Brazil. Twenty-three native hummingbird-pollinated plant species in 21 genera and 14 families were observed. Bromeliaceae, Fabaceae, Gesneriaceae, and Lobeliaceae are represented by more than one species within the assemblage. Flower shapes vary from narrow tube to bowl-shape, but tubular flowers prevail. The variety of flower shapes and sizes results in diverse pollen placement on the body parts of hummingbird visitors, although pollen is deposited mostly on the bill. Sugar concentration in nectar averages 22.1%, and nectar volume per flower averages 16.9 μl. The plant populations bloom for one month to year-round, and their flowering approaches the steady-state pattern. Four flower subsets may be defined within the assemblage, each subset related to the bill size and foraging habits of the most frequent bird visitor. Of the six species of hummingbirds recorded at the study site, four are common and largely resident. The four hummingbirds differ in bill size, body mass, and favoured foraging sites, attributes which reflect their favoured flower subsets. One hermit and one trochiline hummingbird share most of the flower species they use, these two birds being the major pollinators within the flower assemblage. This montane forest community may be viewed as medium-rich in ornithophilous flower species and poor in hummingbird species.     

Pollinator coupling can induce synchronized flowering in different plant species

Synchronous and intermittent plant reproduction has been identified widely in diverse biomes. While synchronous flowering is normally observed within the same species, different species also flower in synchrony. A well-known example of interspecific synchrony is “general flowering" in tropical rain forests of Southeast Asia. Environmental factors, such as low temperature and drought, have been considered as major trigger of general flowering. However, environmental cues are not enough to explain general flowering because some trees do not flower even when they encounter favorable environmental cues. We propose alternative explanation of general flowering; “pollinator coupling”. When species flower synchronously, the elevated pollen and nectar resource may attract increased numbers of generalist pollinators, with a concomitant enhancement of pollination success (facilitation). However, under these circumstances, plants of different species may compete with one another for limited pollinator services, resulting in declines in pollination success for individual species (competition). Here, we present a model describing resource dynamics of individual trees serviced by generalist pollinators. We analyze combinations of conditions under which plants reproduce intermittently with synchronization within species, and/or (sometimes) between different species. We show that plants synchronize flowering when the number of pollinators attracted to an area increases at an accelerating rate with increasing numbers of flowers. In this case, facilitation of flowering by different species exceeds the negative influence of interspecific plant competition. We demonstrate mathematically that co-flowering of different species occurs under a much narrower range of circumstances than intraspecific co-flowering.     

Changes in first flowering dates and flowering duration of 232 plant species on the island of Guernsey

Climate change has affected plant phenology; increasing temperatures are associated with advancing first flowering dates. The impact on flowering duration, however, has rarely been studied. In this study, we analysed first flowering dates and flowering durations from a 27 year dataset of weekly flower observations on 232 plant species from the island of Guernsey in the English Channel. The aim of this study was to explore variation in trends and relationships between first flowering dates, flowering duration and temperature. We specifically looked for evidence that traits, such as life forms and phylogenetic groups, explained variation in sensitivity of first flowering and flowering duration among species. Overall trends revealed significantly earlier flowering over time, by an average of 5.2 days decade^?1 since 1985. A highly significant shortening of flowering duration was observed by an average of 10 days decade^?1. Correlations between first flowering, flowering duration and year varied between different species, traits and flowering periods. Significant differences among traits were observed for first flowering and to a lesser degree for flowering duration. Overall, in comparison to first flowering, more species had significant trends in flowering duration. Temperature relationships revealed large differences in strength and direction of response. 55% of the species revealed a significant negative relationship of first flowering dates and temperature. In contrast, only 19% of flowering durations had a significant negative temperature relationship. The advance in first flowering date together with a shortening of flowering duration suggests potentially serious impacts on pollinators, which might pose a major threat to biodiversity, agriculture and horticulture. Human health, in terms of pollen allergies, however, might benefit from a shortening of specific plant pollen seasons.     

Flowering phenology in Solidago altissima: adaptive strategies against temporal variation in temperature

The evolution of flowering phenology is the result of a trade-off that balances many factors, including growth, reproductive capacity, and temporal overlap with pollinators. When there is large temporal variation in temperature, particularly in the onset of frost, the optimum flowering strategy will vary from year to year. In Duluth, MN, USA, the end of the growing season can vary by more than 30 days. In this study, we observed flowering phenology and pollinator abundance on 15 genotypes of Solidago altissima in Duluth, MN. We predicted that temporal variation in temperature would lead to a range of flowering strategies in the S. altissima population; some genotypes flower early and in synchrony, some ‘hedge their bets’ by flowering over a range of dates, and others have an intermediate strategy. Our results indicate that genotypes vary in mean flowering date and duration of flowering and, for the two observed years, pollinator abundance was highest for early-flowering genotypes.     

Flowering and Fruiting Phenology of Magnolia hypoleuca

Abstract The floral biology of Magnolia hypoleuca , a tall tree species widely distributed in Japan, was studied in a deciduous broad-leaved forest in Hokkaido near the northern limit of distribution of the species. The flower is large, protogynous and nectarless. Its petals close after the female stage and again after the male stage. The mean duration of flowering of an individual flower is 3–4 days, while the flowering period of an individual tree lasts up to 40 days. The main pollinators of the flowers were beetles. Success of pollination is assumed to be affected by weather conditions at anthesis. Average fruit sets (ratios of fruits to flowers) were 13–25%. Hand-pollination of 44 flowers produced a significantly higher fruit set (28 fruits) than control flowers (33/133). These results suggest that the fruit set is limited by pollination. The frequency of infertile ovules was higher in controls than in hand-pollinated fruits, indicating that the seed set was also limited by pollination. We propose a mimicry hypothesis in which the non-rewarding female-stage flower mimics the rewarding male-stage flower to explain the short life-span of an individual flower, the long flowering period of an individual tree, and the coexistence of flowers of various stages on a single tree.     

Flowering Patterns in a Seasonal Tropical Lowland Forest in Western Amazonia

The phenological behavior of many tropical plant species is highly dependent on rainfall, but these plants may also respond to changes in photoperiod. Without a better knowledge of the proportion of species responding to different factors, it is difficult to predict how global climate change may affect natural ecosystem processes. The aim of this study was to describe flowering patterns for more than 100 species in Tinigua Park, Colombia, and to propose which factors may trigger flower production ( e.g. , rainfall, temperature, cloud cover, and photoperiod). Data gathered in 5.6 km of phenological transects during 4 yr and complementary information indicated that the vast majority of species showed intraspecific synchronization, and annual production was the most common pattern, followed by episodic frequency. The annual patterns were common in tree species, while episodic patterns were common in lianas. Simple and multiple regression analyses suggested several aspects of photoperiod as the most likely triggers for flowering in most species. However, the fact that many of these species produce flowers in different periods each year, suggests that the proportion of species responding to photoperiodic cues is less that 23 percent in this community. The flowering times of taxonomically related species seldom showed significantly staggered distributions; however they do not necessarily occur at the same time, suggesting that flowering patterns are not strongly constrained by phylogeny.     

Low temperature influence on flowering in Citrus. The separation of inductive and bud dormancy releasing effects

The flowering response of Owari Satsuma mandarin ( Citrus unshiu Marc) to low temperature treatments has been determined using potted trees and in vitro bud cultures. In potted trees the chilling treatments released bud dormancy and enhanced both sprouting and flowering, but these two responses could not be separated. However, bud cultures showed no dormancy, and a specific effect of low temperature on flower induction was demonstrated. Low temperature appears to have a dual effect, releasing bud dormancy and inducing flowering. Potential flower buds have a deeper dormancy than vegetative buds, and the first stages of flower initiation seem to occur before the winter rest period.     

Pollination biology in a lowland dipterocarp forest inSarawak, Malaysia. I. Characteristics of the plant-pollinator communityin a lowland dipterocarp forest

Flowerings and flower visitors were observed continuously in alowland dipterocarp forest in Sarawak, Malaysia, for 53 mo in1992-1996. Flower visitors of 270 plant species were observed orcollected, and pollinators were assessed by observing body contact tostigmas and anthers. We recognized 12 categories of pollination systems.Among them, plants pollinated by social bees included the largest numberof species (32%) and were followed by beetle-pollinated species(20%). Pollination systems were significantly related with somefloral characters (flowering time of day, reward, and floral shape), butnot with floral color. Based on the relationships between pollinatorsand floral characters, we described pollination syndromes found in alowland dipterocarp forest. The dominance of social bees and beetlesamong pollinators is discussed in relation to the general floweringobserved in dipterocarp forests of West Malesia. In spite of high plantspecies diversity and consequent low population densities of lowlanddipterocarp forests, long-distance-specific pollinators were uncommoncompared with theNeotropics.     

浙江低山地区多用途植物无患子的开花物候特征

无患子(Sapindus mukorossi Gaertn.)是我国长江以南地区传统的重要绿化树种,其果皮富含皂苷,种仁富含油脂,是国家林业局审定的新型木本油料树种之一。为研究多用途树种无患子在浙江低山地区的开花特征,2012年和2013年连续两年对位于浙江省天台县9年生无患子人工林在群体、个体、花序和单花水平进行开花物候观测和比较,并运用开花日期、相对开花强度和同步性等指数研究了无患子开花物候特征。观察结果显示:无患子花呈浅黄白色,花的类型有雄花和两性花,没有发现雌花。雄花较两性花大(花径分别为5.09 mm和3.72 mm),雄蕊多为8枚,个别7或9、10枚;雌蕊退化仅留下浅绿色凸起。两性花花萼稍抱拢,花药藏于花被片下,雄蕊大多8枚,极少数7枚或9枚,柱头高于花药并伸出花苞,子房一般具3室,极少数4室或仅2室。无患子2012年的开花进程略早于2013年,其花期集中在在5月中旬至6月上旬,单花从花蕾膨大到花朵凋谢一般为8—9 d。在2012年和2013年,无患子在群体、个体和花序水平的花期约为30 d、20 d、11 d和28 d、19 d、13 d。个体水平的开花振幅均呈单峰曲线,年际间相似性较高;开花同步性在个体水平同步性较高(同步指数为0.868),表现出一种大量、集中的开花式样;相对开花强度在单株间分布范围相对宽泛,但主要分布在30%—40%,在年际间和年际内均呈现极显著差异。花期同步指数在两年的变异范围分别是0.81—0.97和0.70—0.98,不同单株开花同步性在年际内差异极显著,但在年际间差异性则不显著。由此可见,无患子的生殖资源分配存在明显的时空差异,较长的花期可以减少非法花粉的干扰、保持其种群基因多样性,遗传因子是决定无患子开花物候的主要因素,生态环境对无患子开花物候的影响还需进一步研究,本研究以期为探索无患子开花的主要限制因子奠定基础。     

Pollination of Androdioecious Xerospermum intermedium Radlk. (Sapindaceae) in a rain forest

Xerospermum intermedium, a fairly common understorey species in the West Malesian lowland dipterocarp forest is androdioecious, with delayed self-compatibility. It flowers annually, and has an extended flowering period, with individuals flowering somewhat asynchronously, presenting small flowers with minimal visual lures. The flowers are visited by an unrestricted array of apparently imprecise opportunistic feeders, predominated by trigonid bees and butterflies. Most of these visitors exhibit low fidelity and forage opportunistically on numerous competing tree species. Presentation of nectar in alternating rhythms between the male and hermaphrodite trees appears to induce pollinator movement between them. Out of such alternating visits between the sexes, a low level of short-range inter-tree pollen transfer by trigonids and butterflies seems to occur. Despite the elaborate system which promotes pollinator movement between the sexes, this species has retained a low level of self-compatibility. The physical barriers to pollinator movement in the understorey, and high interspecific competition for pollinators by a large number of tree species, appear therefore to make it difficult to maintain an obligate outbreeding system. Hence, androdioecism may be one of the explanations for the survival of X. intermedium.     

杧果花期延迟技术研究进展

杧果Mangifera indica花期大多集中在2～3月,易受低温阴雨天气影响,导致其授粉受精不良、落花落果,并加重白粉病和炭疽病的危害。本文简要介绍低温阴雨天气对杧果花期的影响,并详细阐述生产上推迟花期的常用措施,如回缩、重修剪、摘花穗、抹花、应用植物生长调节剂、采后灌水、培养二次秋梢或利用冬梢结果等,为杧果开花调控及花期管理提供参考。     

Community-level flowering phenology and fruit set: Comparative study of 25 woody species in a secondary forest in Japan

We investigated the flowering phenology, pollinator visitation, and fruit set of 25 animal-pollinated woody species in a warm temperate secondary forest in Japan. Various species flowered sequentially from February to October. The principal pollinators were bumblebees, honey-bees, flies and/or beetles and birds; bumblebees and flies/beetles pollinated most trees. The duration of flowering was shorter for species that bloomed in the middle of the season than it was for species that bloomed earlier or later in the season. The timing of flowering was more synchronous within species that had a shorter flowering duration; this was also detected when phylogenetically independent contrasts were calculated. This could be important for the effective pollination of species with a short flowering duration because such species bloom sequentially over a short period of less than 1month around May. Fruit set was related not to pollinator type, sex expression, flowering sequence (in order of the date of peak flowering) or flowering duration, but to the relative abundance of the species in the forest. This correlation was detected for fly- and beetle-pollinated species but not for bumblebee-pollinated species. Thus, relatively rare plant species with opportunistic pollinators might experience limited fruit set because of insufficient pollinator services. Bagging experiments conducted on eight hermaphrodite species revealed that the fruit set of bagged flowers was nearly zero, lower than that of control flowers. These results indicate the importance of pollinators for successful reproduction and thus for the coexistence of plants in this secondary forest.     

The pollination ecology of Actaea spicata (Ranunculaceae)

Pollination ecology of Actaea spicata L. (Ranunculaceae) was studied at localities ranging from deciduous to spruce forest. The species is self–incompatible and possibly facultatively apomictic. No anemogamy seems to occur. In deciduous forest, the plant is primarily pollinated by pollen–foraging and mate–searching Byturus ochraceus (Scriba) (Coleoptera). It flowers during a period between emergence of the beetle and onset of anthesis of the prime pollen source and only larval host plant, Geum ur–banum L. (Rosaceae). The beetles sit on the suitably long anther filaments or on top of the stigma. Floral morphology promotes pollination–performing movements by the beetle. The complex floral fragrance seems to induce close–range behaviour, i.e. alighting and/or foraging. Flies become abundant visitors only shortly before anthesis is over, and their contribution is small. Large syrphids are relatively effective as pollinators, since they grasp the stamens near the base and often touch the stigma with pollen–dusted parts of their bodies. Due to lack of its food–plant, B. ochraceus is absent at spruce forest localities and northwards. At these localities flowering is delayed until the main fly appearance, and the flowers are heavily pollinated by several syrphids and muscids. At three deciduous–forest localities a second flowering took place with new, juxtaposed inflorescences on separate stalks. The distribution of such inflorescences within the population along with the distribution and abundance of pollinators among the two generations of flowers suggest that these inflorescences occur as a response to low rate of pollination in the first anthesis.     

毛茛状金莲花不同花期的花特征和访花昆虫的变化及表型选择

为了研究植物生长季内开花时间对花特征表型选择的影响,我们以青藏高原东缘高寒草地的毛茛状金莲花Trollius ranunculoides)为实验材料,在生长季内不同开花时间(花前期、花末期)测定花特征,观察访花昆虫的类群和访花频率,生长季结束后收集种子.根据昆虫访花的喜好和季节内类群与访花频率的变化,分析了不同开花时间毛茛状金莲花的花特征与昆虫的选择;并用种子产量表示雌性适合度,估计了毛茛状金莲花的花特征在不同开花时间所受的表型选择.结果表明:不同花期植物的花特征有显著差异,相应的访花昆虫的类群和频率也存在差异,不同类群昆虫访花喜好也不一样.蜂喜好花瓣和花萼较宽、花茎短和花茎数少的个体,这正符合花前期的特征,因而蜂的访花频率在花前期较高;蝇对花特征没有明显的偏好.而通过雌性适合度估计毛茛状金莲花花特征所受的表型选择则是:花前期,花茎较长和花茎数多的植株适合度大;花末期,花茎数多的植株适合度大.我们的研究表明:在植物生长季,花期的分化伴随着传粉昆虫活动的变化.不同花期,访花昆虫的变化可能对植物花特征的分化起了至关重要的作用.但是访花昆虫对花特征的选择与通过雌性适合度估计植物受到的选择不尽相同,这可能是由于其他因素造成的.     

Reproductive ecology of tropical forest trees in logged and fragmented habitats in Thailand and Costa Rica

Invertebrates mediate several important ecological processes, including pollination and seed predation, and events that affect invertebrate diversity or behaviour can potentially disrupt forest regeneration processes. This study investigates the impact of logging in Thailand and forest fragmentation in Costa Rica on the pollination and seed production of two self-incompatible forest trees. Logging in a dry deciduous dipterocarp forest in Thailand resulted in reduced densities of the common dipterocarp treeShorea siamensis and variably isolated individual trees. The number of flower visits to S. siamensis by pollinating Trigona bees was not affected by logging disturbance. However, pollinators did spend longer periods of time foraging in the canopies of isolated trees which were more prevalent in logged areas where tree density had been reduced. Consequently, at the logged site few cross-pollinations were effected and fruit set of S. siamensis was considerably lower than at nearby unlogged sites where distances between flowering conspecifics were smaller. Reduced fruit set has long-term implications for the recovery of S. siamensis populations in disturbed areas, and local population genetic structure is likely to be affected as reduced outcrossing rates among trees in disturbed regions results in relatively inbred seed. In Costa Rica forest fragmentation has restricted the once widespread tree Anacardium excelsum to forest patches located in an agriculturally-dominated landscape. As with S. siamensis, the abundance of pollinators, also Trigona bees, in the canopies of A. excelsum was largely unaffected by fragment size. Nevertheless, pollination success and seed production was positively correlated with fragment size. We propose that small bees rarely move between forest fragments and gene exchange through pollination occurs predominantly among trees within fragments and, together with likely low genetic variability in small fragments, that this contributes to the observed reduced fertilisation and seed set of A. excelsum. Thus increased tree isolation tree through selective logging or habitat fragmentation by forest clearance can result in reduced seed set due to changes in the foraging patterns of poorly mobile pollinators. Even if population sizes of the pollinators are maintained following environmental perturbation, this study shows that disturbance may disrupt pollination processes through changes in pollinator foraging behaviour. More attention needs to be focussed on changes in the behaviour of species involved in key ecological interactions following disturbance events in tropical forests.