Field of Dreams: Restitution of Pollinator Services in Restored Bird‐Pollinated Plant Populations

Ecosystem functionality is an increasingly important objective of ecological restoration. Despite this, a few studies have rigorously assessed reproductive functionality within restored plant populations, and it is largely assumed that pollinators follow restoration of plant communities—“build it and they will come.” Here, we applied an ecological genetic approach to determine the impact of spatial separation on mating in Banksia menziesii (Proteaceae), a dominant bird‐pollinated species of Banksia woodlands of Western Australia. All plants at three post‐mining restored sites (n = 72 [13 years old], n = 21 [8 years old], and n = 20 [9 years old]), as well as a sample from an adjacent natural reference site (n = 42), were genotyped at nine microsatellite loci. Seed set, mating system parameters, realized pollen dispersal through the assignment of paternity to seed, and avian pollinator species composition, abundance and behavior, were assessed. All patches displayed equivalent heterozygosity (H_e = 0.53–0.59) and very weak genetic divergence (F_ST ≤ 0.01). Seed of plants within restored sites showed complete outcrossing and relatively high seed set, 26% of which were sired by pollen donors located beyond the local patch. Similar abundance and movement of nectar‐feeding birds was observed in restored and natural sites, despite lower bird species diversity in the restored site, where a smaller, less aggressive species was dominant. Our results demonstrate the restitution of wide outcrossing in these restored Banksia patches within an active mine‐site, and suggest that restored bird‐pollinated Banksia populations are resilient to human impacts, due largely to their generalist pollinator requirements and highly‐mobile avian pollinators.     

Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads

Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self‐compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self‐compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.     

Does community‐level floral abundance affect the pollination success of a rewardless orchid,Calanthe reflexa Maxim.?

Decreases in pollinator abundance may particularly constrain plants that lack floral rewards, since they are poor competitors for pollinators in the plant community. Here, we documented the pollination ecology of a rewardless orchid, Calanthe reflexa Maxim., and examined effects of forest understory degradation by deer browsing on pollination success of the species in the light of a change in the abundance of neighboring flowering plants in 2010 and 2011. Bombus species were the only pollinators at each site and the flowering phenology of C. reflexa did not overlap with that of other rewarding plants. Pollinator visit rates (assessed by time‐lapse photography), and pollinia removal rate were higher in the undegraded understory site than the degraded site in both years, while the fruit set ratio did not differ between the sites in 2011. Coverage by neighboring flowering plants was extremely low in the degraded site. Our results suggest that, although its flowering phenology and consequently lower interspecific competition of C. reflexa with rewarding plants for attracting bumblebees, neighboring flowering plants may play an important role for maintaining the visitation frequency of bumblebees of C. reflexa and contribute to its pollination success.     

Individual flowering phenology shapes plant–pollinator interactions across ecological scales affecting plant reproduction

The balance of pollination competition and facilitation among co-flowering plants and abiotic resource availability can modify plant species and individual reproduction. Floral resource succession and spatial heterogeneity modulate plant–pollinator interactions across ecological scales (individual plant, local assemblage, and interaction network of agroecological infrastructure across the farm). Intraspecific variation in flowering phenology can modulate the precise level of spatio-temporal heterogeneity in floral resources, pollen donor density, and pollinator interactions that a plant individual is exposed to, thereby affecting reproduction. We tested how abiotic resources and multi-scale plant–pollinator interactions affected individual plant seed set modulated by intraspecific variation in flowering phenology and spatio-temporal floral heterogeneity arising from agroecological infrastructure. We transplanted two focal insect-pollinated plant species (Cyanus segetum and Centaurea jacea, n = 288) into agroecological infrastructure (10 sown wildflower and six legume–grass strips) across a farm-scale experiment (125 ha). We applied an individual-based phenologically explicit approach to match precisely the flowering period of plant individuals to the concomitant level of spatio-temporal heterogeneity in plant–pollinator interactions, potential pollen donors, floral resources, and abiotic conditions (temperature, water, and nitrogen). Individual plant attractiveness, assemblage floral density, and conspecific pollen donor density (C. jacea) improved seed set. Network linkage density increased focal species seed set and modified the effect of local assemblage richness and abundance on C. segetum. Mutual dependence on pollinators in networks increased C. segetum seed set, while C. jacea seed set was greatest where both specialization on pollinators and mutual dependence was high. Abiotic conditions were of little or no importance to seed set. Intra- and interspecific plant–pollinator interactions respond to spatio-temporal heterogeneity arising from agroecological management affecting wild plant species reproduction. The interplay of pollinator interactions within and between ecological scales affecting seed set implies a co-occurrence of pollinator-mediated facilitative and competitive interactions among plant species and individuals.     

Are mismatches the norm? Timing of flowering,fruiting, dispersal and germination and their fitness effects in Frangula alnus (Rhamnaceae)

The close morphological and temporal links between phases of plant growth and reproduction call for integrated studies incorporating several reproductive phases from flowering to recruitment, and associated plant‐animal interactions. Phenological strategies, as well as plastic phenological response to climate change, incorporate complex interactions between developmental constraints, pollination and seed dispersal. Relationships between reproductive phenology and components of fitness were studied for two years in the north‐temperate, self‐incompatible, insect‐pollinated, and bird‐dispersed shrub Frangula alnus (Rhamnaceae). Fruit set, dispersal, germination and juvenile survival, as well as seed mass and juvenile size were measured in relation to flowering, fruiting and germination time. The results suggest that effects of flowering and fruiting time prevailed in subsequent phases, to some extent as far as to the juvenile phase, but effects of timing were complex and had partly opposing effects on different fitness components. Early flowers had higher fruit‐set and experiments indicated that synchronous peak flowering increased fruit‐set, but later flowers had higher seed mass. Peak fruiting was not associated with peak dispersal. Late fruits derived from late flowers promoted dispersal. Juvenile recruitment was enhanced by increasing seed size. We conclude that the phenology of flowering and fruiting in F. alnus comprises several features, each with different and sometimes counteracting effects on fitness components. From a general perspective, this result implies that we should not expect to find finely tuned matches in timing specifically between flowering and pollinators, and fruiting and seed dispersing birds.     

The impacts of flowering phenology on the reproductive success of the narrow endemic Nouelia insignis Franch. (Asteraceae)

Nouelia insignis Franch. (Asteraceae) is a short, narrow endemic and endangered tree, growing with a natural population in the dry and hot valley of the Jinsha River in the southwest area of China. In this work, flowering phenology (time and duration), floral biology, visit frequency and behavior of pollinators, and pollination characteristics were studied based on investigation in the field and analysis in the laboratory with the help of a stereomicroscope, and the relationship between seed setting rate and reproductive traits, as well as the relationship between flowering time and rainfall before flowering, was tested using the method of general linear regression model. The results showed that natural population of N. insignis exhibited high flowering synchrony with relatively stable flowering duration, and the flowering time fluctuated greatly depending on the rainfall 5 months before flowering. The pollination of N. insignis required pollinators, and insect activities played a very important role in the pollination process. However, lack of the pollinators was not a limitation for reproductive fitness in N. insignis, although the number of pollinators was small and the frequency of visits was low. In addition, no pollen limitation was found during pollination. The average seed setting rate of N. insignis in the natural condition was only 1.52%–3.73%, and it was generally affected by changes in flowering phenology between years and had a higher seed set in early flowering year. The annual variation of seed set might be related to the annual variations of stamen and pistil functions, such as changes of pollen viability and stigma receptivity, which were closely related to flowering time. The results of this study are of value for further conservation actions on natural population of this threatened endemic plant.     

Pollen limitation and pollinator preferences in Scorzonera hispanica

The plant life cycle is often affected by animal–plant interactions. In insect‐pollinated plants, interaction with pollinators is very important. When pollen transfer due to a lower abundance of pollinators limits seed production, selection pressures on plant traits related to plant attraction to pollinators might occur, e.g. on flowering phenology, height or number of flowerheads. Landscape changes (e.g. habitat fragmentation or changed habitat conditions) may cause plant–pollinator systems to lose balance and consequently affect population dynamics of many plant species. We studied the relationship between measured plant traits, environmental variables and pollinator preferences in Scorzonera hispanica (Asteraceae), a rare perennial, allogamous herb of open grasslands. We estimated the pollen limitation by comparing seed set of supplemental‐pollinated plants with that of open‐pollinated ones. Pollinators selected plants based on position within the locality (isolated plants close to trees) rather than on their traits. In spite of a high proportion of undeveloped seeds on the plants, we demonstrated that they are not pollen limited. Instead, seed set and weight of seeds was correlated with plant size traits (height and flowerhead number), with larger plants producing more and larger seeds. This suggests that the studied plants are likely resource limited. Overall, the results suggest that pollinators are not a selection factor in this system, in contrast to studies on various plant species, including self‐compatible species of the Asteraceae. The lack of any effect of pollinators in the system may be caused by a strong negative effect of ungulate herbivores, which could play a decisive role in functioning of the system.     

Genetic variation in flowering phenology and avoidance of seed predation in native populations of Ulex europaeus

The genetic variation in flowering phenology may be an important component of a species’ capacity to colonize new environments. In native populations of the invasive species Ulex europaeus, flowering phenology has been shown to be bimodal and related to seed predation. The aim of the present study was to determine if this bimodality has a genetic basis, and to investigate whether the polymorphism in flowering phenology is genetically linked to seed predation, pod production and growth patterns. We set up an experiment raising maternal families in a common garden. Based on mixed analyses of variance and correlations among maternal family means, we found genetic differences between the two main flowering types and confirmed that they reduced seed predation in two different ways: escape in time or predator satiation. We suggest that this polymorphism in strategy may facilitate maintain high genetic diversity for flowering phenology and related life‐history traits in native populations of this species, hence providing high evolutionary potential for these traits in invaded areas.     

西双版纳地区鞘花的繁殖物候特征及影响因素

桑寄生植物作为关键种和关键性食物资源在生态系统中扮演着重要角色,其鞘花的繁殖物候特征不仅会影响自身的繁殖适合度而且还会影响依赖于其获取食物资源的动物。为了解鞘花的繁殖物候特征及其影响因素,探究其与寄主植物和种子散布者之间的相互作用关系。该研究以西双版纳地区分布的鞘花为对象,通过定期观测鞘花和其寄主植物木荷的繁殖物候,测量它们的生物学特性和温湿度等环境因子,并分析鞘花在个体水平和种群水平上的繁殖物候特征以及寄主植物和温湿度对其繁殖物候的影响。结果表明:(1)鞘花的开花物候属于集中大量开花模式,整个种群的花期和果期的持续时间分别约为20 d和72 d,花期和果期的同步性指数都较高,6月中旬鞘花果实被取食的数量和速率最大,之后逐渐降低。(2)鞘花的始花期与木荷的始花期相关性较高,花期和果期与木荷的繁殖物候基本重叠。(3)每月开花和果熟的个体数量与同期和前一个月的平均温度和相对湿度的相关关系均不显著。总之,桑寄生植物的繁殖物候特征可能受到较多因素的影响,若想全面了解半寄生植物的繁殖物候特征,就要综合考虑生物和非生物等多种因子的共同作用。     

金沙江干热河谷滇榄仁开花物候与繁育系统

通过野外定点观测,并利用套袋实验、解剖实验以及花粉形态特征观察与活力检测、柱头可受性检测、花粉/胚珠比以及异交指数等,研究了金沙江干热河谷滇榄仁自然群体花部形态特征与开花进程、繁育系统特征以及传粉媒介与环境的适应性等内容。结果表明,滇榄仁始花期在每年4月上旬,群体花期持续时间为30d左右,开花同步性高,呈典型的"集中开花模式"。单花开放时,柱头先伸长和膨大,约4d后雄蕊伸出,具有雌雄异熟的特征,一定程度上避免了自交的发生。单花开放进程可划分为花蕾期、柱头伸长期、雄蕊始露期、盛开期、盛开后期、花谢期等6个时期。除了花蕾期外,整个开花期的花粉均具有活力。花粉活力与柱头可受性之间避开了最适授粉期,但也有部分重叠。滇榄仁可能同时具有风媒和虫媒传粉。繁育系统以异交为主,部分自交亲和,需要传粉者。滇榄仁花部特征、开花物候和繁育系统为其适应干热河谷恶劣气候环境提供了一定的生殖保障和进化潜力。良好的群落环境有助于促进滇榄仁异花传粉,产生更多有效的种子,从而促进林下自然更新。     

Effects of multiple competitors for pollination on bumblebee foraging patterns and Mimulus ringens reproductive success

When co‐occurring plant species overlap in flowering phenology they may compete for the service of shared pollinators. Competition for pollination may lower plant reproductive success by reducing the number of pollinator probes or by decreasing the quality of pollen transport to or from a focal species. Pair‐wise interactions between plants sharing pollinators have been well documented. However, relatively few studies have examined interactions for pollination among three or more plant species, and little is known about how the outcomes and mechanisms of competition for pollination may vary with competitor species composition. To better understand how the dynamics of competition for pollination may be influenced by changes in the number of competitors, we manipulated the presence of two competitors, Lythrum salicaria and Lobelia siphilitica, and quantified reproductive success for a third species, Mimulus ringens. Patterns of pollinator preference and interspecific transitions in mixed‐species arrays were significantly influenced by the species composition of competitor plants present. Both pair‐wise and three‐species competition treatments led to a similar ～ 40% reduction in Mimulus ringens seed set. However, the patterns of pollinator foraging we observed suggest that the relative importance of different mechanisms of competition for pollination may vary with the identity and number of competitors present. This variation in mechanisms of competition for pollination may be especially important in diverse plant communities where many species interact through shared pollinators.     

Phenology of neotropical pepper plants (Piperaceae) and their association with their main dispersers, two short-tailed fruit bats, Carollia perspicillata and C. castanea (Phyllostomidae)

To relate differences in phenological strategies of a group of closely related plants to biotic (pollinators, dispersers) and abiotic (water, light) factors, we studied leafing, flowering, and fruiting phenology of 12 species of Piper (Piperaceae) in a neotropical lowland forest in Panama for 28 months. We asked how Piper may partition time and vertebrate frugivores to minimize possible competition for dispersal agents. Based on habitat preferences and physiological characteristics we discriminate between forest Piper species (eight species) and gap Piper species (four species). Forest Piper species flowered synchronously mostly at the end of the dry season. Gap Piper species had broader or multiple flowering peaks distributed throughout the year with a trend towards the wet season. Both groups of Piper species showed continuous fruit production. Fruiting peaks of forest Piper species were short and staggered. Gap Piper species had extended fruiting seasons with multiple or broad peaks. Both groups of Piper species also differed in their time of ripening and disperser spectrum. Forest Piper species ripened in late afternoon and had a narrow spectrum consisting mainly of two species of frugivorous bats: Carollia perspicillata and C. castanea (Phyllostomidae). Fruits of gap Piper species, in contrast, ripened early in the morning and were eaten by a broader range of diurnal and nocturnal visitors, including bats, birds, and ants. We conclude that the differences in flowering phenology of forest and gap Piper species are primarily caused by abiotic factors, particularly the availability of water and light, whereas differences in fruiting patterns are mostly influenced by biotic factors. The staggered fruiting pattern of forest Piper species may reflect competition for a limited spectrum of dispersers. The long and overlapping fruiting periods of gap Piper species are associated with a larger spectrum of dispersers and may be a strategy to overcome the difficulty of seed dispersal into spatially unpredictable germination sites with suitable light conditions.     

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.     

Impact of flower and seed predators on seed-set in two Banksia shrubs

A non-systemic Insecticide was used to exclude flower and seed predators from Banksia ericifolia and B. oblongifolia Inflorescenses. In B. ericifolia this treatment doubled the number of inflorescences that set seeds, and increased by 40% the number of seeds produced per Inflorescence. Insecticide treatment did not alter either of these components of seed-set in B. oblongifolia partly because the experiment began too late to exclude flower predators In this species. The results obtained for B. ericifolia support the hypothesis that flower and seed predators explain, in part, the low incidence of seed-bearing cones and the low seed:flower ratios reported for many Banksia species. Each Banksia species supports two guilds of Inflorescence predators. Two Lepidopteran species feed on young flowers; one of these causes tunnel damage to the rachis. Each Banksia species also supports five Lepidopteran and two Curcullonid seed-predators. About 40% of these herbivorous Insects are common to both shrubs.     

Effectiveness of short‐tongued bees as pollinators of apparently ornithophilous New Zealand mistletoes

Abstract The flowers of two species of threatened New Zealand mistletoes (Peraxilla tetrapetala and Peraxilla colensoi, Loranthaceae) have explosive buds that do not open unless force is applied by birds or two species of native short‐tongued bees. Opened flowers are visited by a variety of birds and insects. Although both species of Peraxilla conform to a pollination syndrome of ornithophily, bees may be effective alternative pollinators. We investigated the effectiveness of bees and birds as pollinators of P. colensoi at one site and P. tetrapetala at two sites in the South Island. Bees and other insects outnumbered birds as flower visitors at all three sites. By excluding birds with wire cages, we showed that two bee species regularly open flowers of P. tetrapetala, but only rarely open flowers of P. colensoi. Few pollen grains were deposited when either birds or bees opened buds, so opening buds was not by itself sufficient for adequate pollination. Instead, pollen continued to accumulate over the next 6 or 7 days, even inside cages that excluded birds. Both populations of P. tetrapetala were regularly pollen‐limited, but in different ways. At Ohau, opened flowers gained enough pollen to produce seeds, but many buds were not opened and hence failed to set seed. In contrast, at Craigieburn, nearly all buds were opened, but many of these did not receive enough pollen. These results demonstrate that native bees can partially replace birds as pollinators of mistletoes, despite their apparent ornithophilous syndrome. Ongoing reductions in New Zealand forest bird numbers means that the service bees provide may be important for the long‐term future of these plants.     

珍稀濒危植物距瓣尾囊草(Urophysa rochkii)的开花物候和繁育系统特性

通过野外观察,运用套袋授粉和联苯胺-过氧化氢法等实验方法对距瓣尾囊草开花物候以及种群的繁育系统特性进行研究。研究结果:1)距瓣尾囊草为两性花,聚伞状花序,花期较长,从头年12月持续到次年4月,单花花期持续8—15 d,种群花期可持续4个月左右。2)距瓣尾囊草在花朵未开放前花粉没有活力,花药开裂当天花粉活力达到94.8%,第2天为90.3%,第3天为81.6%,第4天为62.9%,此后其花粉活力明显减弱;柱头可授性于散粉后第3天开始出现,散粉后4—5天最强,散粉后第8天柱头失去可授性;柱头可授性和花粉活力有5 d左右的重叠期。3)其花粉胚珠比P/O为938.5±250,按照Cruden标准划分,其有性繁育系统为兼性异交。4)按照Dafni的标准,距瓣尾囊草的OCI=4,即繁育系统为异交、部分自交亲和、需要传粉者。5)人工异花授粉结实率达80%以上,略高于自然结实率78.94%;直接套袋结实率为5.71%;去雄套袋和去雌套袋均不结实。以上结果表明,距瓣尾囊草繁育系统表现为异交为主、部分自交亲和并且不存在无融合生殖现象,其开花物候、花部特征和繁育系统为适应特殊的环境提供了一定的生殖保障,本次研究结果为距瓣尾囊草的迁地保护和种群的繁衍复壮提供理论依据。     

Severe pollen limitation in populations of the New Zealand shrub Alseuosmia macrophylla (Alseuosmiaceae) can be attributed to the loss of pollinating bird species

The loss of bird species following human colonization of New Zealand has raised concerns about the consequences for crucial ecosystem functions such as pollination. The understorey shrub Alseuosmia macrophylla (Alseuosmiaceae) exhibits characteristics typical of a bird pollination syndrome, but populations still persist in northern North Island forest remnants despite the local extinction of most endemic bird pollinators, leading to the suggestion that moths – rather than birds – may be the primary pollinators. The aim of this study was to quantify the importance of endemic birds as pollinators of A. macrophylla over several years by comparing plants on Little Barrier Island (LBI), where all extant endemic bird pollinators still occur, to plants at sites on the adjacent North Island in the Waitakere Ranges (WTK), where only one of these species remains common. Flowers on LBI were visited by endemic bellbirds (Anthornis melanura) and stitchbirds (Notiomystis cincta), while at WTK sites the most common visitors were the recently arrived silvereye (Zosterops lateralis) and the introduced honeybee (Apis mellifera), both of which acted principally as nectar robbers. Caged flowers on LBI had significantly lower fruit set than open flowers, and plants at WTK were significantly more pollen‐limited than plants on LBI. This provides evidence that the loss of endemic pollinating birds is the most likely reason for the high pollen limitation found in some North Island A. macrophylla populations, and the very low seed set of these populations could have serious implications for the long‐term persistence of this species.     

Responses of pre‐dispersal seed predators to sequential flowering of Dipterocarps in Malaysia

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.     

Pollination ecology of Isoglossa woodii,a long‐lived,synchronously monocarpic herb from coastal forests in South Africa

Synchronous monocarpy in long‐lived plants is often associated with pollination by wind, in part because infrequent mass flowering may satiate pollinators. Selfing in synchronous monocarps may provide reproductive assurance but conflict with the benefits of outcrossing, a key evolutionary driver of synchrony. We predicted that animal‐pollinated species with synchronous flowering would have unspecialised flowers and attract abundant generalised pollinators, but predictions for selfing and outcrossing frequencies were not obvious. We examined the pollination biology of Isoglossa woodii (Acanthaceae), an insect‐pollinated, monocarpic herb that flowers synchronously at 4–7‐year intervals. The most frequent visitor to I. woodii flowers was the African honeybee, Apis mellifera adansonii. Hand‐pollination failed to enhance seed production, indicating that the pollinators were not saturated. No seed was set in the absence of pollinators. Seed set was similar among selfed and outcrossed flowers, demonstrating a geitonogamous mixed‐mating strategy with no direct evidence of preferential outcrossing. Flowers contained four ovules, but most fruits only developed one seed, raising the possibility that preferential outcrossing occurs by post‐pollination processes. We argue that a number of the theoretical concerns about geitonogamous selfing as a form of reproductive assurance do not apply to a long‐lived synchronous monocarp such as I. woodii.