Pollen limitation in three sympatric species ofVaccinium (Ericaceae) in the Upper Ardennes, Belgium

The reproductive success of three co-flowering species ofVaccinium (V. myrtillus, V. vitis-idaea andV. uliginosum) was studied in one heathland of the Upper Ardennes, Belgium, during three years (1988–1990). The purpose was to examine whether pollen limitation, flower position and flowering phenology may influence patterns of fruit set in these three sympatric species. I quantified fruit and seed set following supplementary hand-pollinations and compared this to natural fruit set. On the same plants, I also quantified fruit and seed set in relation to the spatial position of the flowers on the ramet and their temporal sequence of blooming. Hand-pollination had no significant effect on fruit set inV. vitis-idaea andV. uliginosum, but significantly increased seed number per fruit in 1989. InV. myrtillus both fruit and seed set were increased by supplementary pollination, but significantly in only one year. Analyses of position effects revealed that the fruits in the lower positions in the ramet did not mature preferentially and did not contain more seeds inV. uliginosum and inV. vitis-idaea. Flowering phenology also had no significant effect for these species. InV. myrtillus hand-pollinated flowers showed a seasonal decline in seed number, and control (naturally pollinated) flowers showed a seasonal increase in fruit set. Fruit and seed set appear to be pollen-limited rather than resource-limited inV. uliginosum. In the two other species, unfavorable weather (frosts) could be a more important cause of low fruit and seed set.     

Pollen and water limitation in Astragalus scaphoides, a plant that flowers in alternate years

Mast seeding is common in plant populations, but its causes have rarely been tested experimentally. We tested mechanisms of alternate-year flowering and fruit set in an iteroparous, bee-pollinated, herbaceous plant, Astragalus scaphoides, in semi-arid sagebrush steppe. Patterns of reproduction from 1986 to 1999 indicated that spring precipitation was a cue for synchronous flowering, and that increased pollination in high-flowering years was a fitness advantage of synchrony. We tested these patterns by adding supplemental water and pollen to plants in high- and low-flowering sites and years. Supplemental water had no effect on flowering or seed set, so water is not a proximate cue for reproduction, though it could be important over longer (>3 year) time scales. Supplemental pollination increased fruit set in low- but not high-flowering years, indicating that synchronous flowering increases pollination success. Many shorter-term studies also report increased fruit set after pollen supplementation, but not after resource addition. This pattern may reflect the fact that plants can store and reallocate resources, but not pollen, across multiple years. For animal-pollinated herbs such as these, uniting theories about pollination ecology and mast seeding may promote an understanding of the mechanisms that determine patterns of reproduction over time.     

北黄花菜的开花物候及传粉特性研究

北黄花菜是百合科萱草属的优秀观赏植物材料,也是进行品种创新的极有价值的野生种质资源。该研究对野生环境中北黄花菜的开花物候与传粉特性进行了连续三年的追踪观测,探讨其开花进程和传粉机制。结果表明:(1)北黄花菜种群的开花物候只有1个高峰期;种群始花时间为5月末,盛花期为6月中旬,开花时间在30 d以上,单株花期为6～12 d;开花同步性指数(Si)分别为0.36、0.27和0.21,种群内个体开花时间重叠程度较低。(2)北黄花菜的开花数量与坐果数呈显著正相关(P<0.05),坐果数与单株花期长度呈极显著正相关(P<0.01),单株花期长度与始花时间呈显著负相关(P<0.05),表明北黄花菜开花数量越多,其坐果数越高。(3)北黄花菜的访花昆虫分属于4目10科共10种;各访花昆虫的访花时间和行为区别较大;主要的传粉昆虫有4种,其活动范围大,活动特点与北黄花菜午后开放的规律相匹配,能保证传粉效果;部分访花昆虫虽然活动范围小,但可以帮助植物进行自花授粉;两类访花昆虫的共同作用,使得北黄花菜的自然结实得到较大保障。上述结果可为北黄花菜的引种栽培和资源创新发挥实际作用。     

POLLINATION AND SEED SET IN DIERVILLA LONICERA (CAPRIFOLIACEAE): TEMPORAL PATTERNS OF FLOWER AND OVULE DEPLOYMENT

Ovule development in Diervilla lonicera (Caprifoliaceae) is limited by insufficient pollination early in the blooming period and during extensive rainy periods. Production of flowers is skewed in time; an initial burst of flowering is followed by a long period of sparse flower production. Ovule number per flower increases through the blooming period. I discuss the interactions of fruit and seed set, ovule number, and bumble bee pollinator visitation patterns. When certain flowers have a higher a priori probability of successful pollination, it may be advantageous for plants to put more ovules in those flowers. Selective ovule deployment may be a general adaptive phenomenon that has received little attention.     

Pollination ecology of Ruellia angustiflora (Ness) Lindau ex Rambo (Acanthaceae) in the Serra da Bodoquena,Mato Grosso do Sul,Brazil

In Brazil, studies focusing on reproductive biology and aspects of pollination in the genus Ruellia have demonstrated common characteristics such as pollination by hummingbird, the predominance of self-fertility and spontaneous self-pollination. The objectives of this study were to describe the floral biology, nectar production, the reproductive system, the effect of pollination intensity and flowering phenology of Ruellia angustiflora, as well as keeping a record of the hummingbirds that visit the flowers of this species. Data collection was performed in a stretch of the Salobrinha stream in the Serra da Bodoquena, Mato Grosso do Sul, from August 2005 to July 2006. R. angustiflora flowered throughout most of the study period; it has diurnal anthesis and has increased nectar production in the morning hours with decreasing production rates during the day. The results of the reproduction experiments indicate self-fertility and pollen limitation in R. angustiflora. Among experiments with different pollination intensity there was no significant difference in fruit formation. Four hummingbird species were observed visiting the flowers of R. angustiflora, and Phaethornis pretrei was the most frequent followed by females of Thalurania furcata. According to its behavior and frequency of visits, P. pretrei can be considered the main pollinator of R. angustiflora in the study area. The experiments on flowers treated with different pollen loads indicate that most successful fruit formation in this species is obtained in flowers that received greater pollen loads on their stigmas. Besides, data suggest that major success in fruit set could be reached with more than three visits of P. pretrei.     

Early blooming's challenges: extended flowering season, diverse pollinator assemblage and the reproductive success of Gynodioecious Daphne laureola

BACKGROUND AND AIMS: The scarcity and unpredictability of active pollinators during late winter in temperate areas tends to favour extended flowering seasons and increased floral longevity in early blooming species, which are usually pollinated by diverse sets of insects. Daphne laureola is a gynodioecious woody perennial that flowers from January to April in southern Spain, a period characterized by cold temperatures, frequent rains and irregular snowfalls. METHODS: Pollinators were excluded at four different times during the flowering season in order to determine the effect of decreased exposure to pollinators on fruit set in female and hermaphrodite individuals. The role of nocturnal and diurnal pollination on reproductive success in each gender was simultaneously evaluated by selective exclusion. KEY RESULTS: A 50 % reduction in the flowering period decreased fruit set of females by 50 %, whereas the corresponding decrease in self-compatible hermaphrodites was only approx. 25 %. Day-active hymenopterans and lepidopterans were infrequent visitors, and nocturnal pollinators were inefficient, suggesting that pollen beetles, Meligethes elongatus, were the main pollinators of D. laureola in the study region. CONCLUSIONS: Beetles were less abundant in pollenless females, although discrimination did not apparently result in pollination limitation of female reproduction. A preference of beetles for sunny locations emphasized the relevance of abiotic conditions for pollination of this early blooming shrub.     

Bateman’s principle and plant reproduction: The role of pollen limitation in fruit and seed set

Bateman’s principle states that male fitness is usually limited by the number of matings achieved, while female fitness is usually limited by the resources available for reproduction. When applied to flowering plants this principle leads to the expectation that pollen limitation of fruit and seed set will be uncommon. However, if male searching for mates (including pollen dissemination via external agents) is not sufficiently successful, then the reproductive success of both sexes (or both sex functions in hermaphroditic plants) will be limited by number of matings rather than by resources, and Bateman’s principle cannot be expected to apply. Limitation of female success due to inadequate pollen receipt appears to be a common phenomenon in plants. Using published data on 258 species in which fecundity was reported for natural pollination and hand pollination with outcross pollen, I found significant pollen limitation at some times or in some sites in 159 of the 258 species (62%). When experiments were performed multiple times within a growing season, or in multiple sites or years, the statistical significance of pollen limitation commonly varied among times, sites or years, indicating that the pollination environment is not constant. There is some indication that, across species, supplemental pollen leads to increased fruit set more often than increased seed set within fruits, pointing to the importance of gamete packaging strategies in plant reproduction. Species that are highly self-incompatible obtain a greater benefit relative to natural pollination from artificial application of excess outcross pollen than do self-compatible species. This suggests that inadequate pollen receipt is a primary cause of low fecundity rates in perennial plants, which are often self-incompatible. Because flowering plants often allocate considerable resources to pollinator attraction, both export and receipt of pollen could be limited primarily by resource investment in floral advertisement and rewards. But whatever investment is made is attraction, pollinator behavioral stochasticity usually produces wide variation among flowers in reproductive success through both male and female functions. In such circumstances the optimal deployment of resources among megaspores, microspores, and pollinator attraction may often require more flowers or more ovules per flower than will usually be fertilized, in order to benefit from chance fluctuations that bring in large number of pollen grains. Maximizing seed set for the entire plant in a stochastic pollination environment might thus entail a packaging strategy for flower number or ovule number per flower that makes pollen limitation of fruit or seed set likely. Pollen availability may limit female success in individual flowers, entire plants (in a season or over a lifetime), or populations. The appropriate level must be distinguished depending on the nature of the question being addressed.     

Seasonal variation in the mating system of a selfing annual with large floral displays

Background and Aims Flowering plants display considerable variation in mating system, specifically the relative frequency of cross- and self-fertilization. The majority of estimates of outcrossing rate do not account for temporal variation, particularly during the flowering season. Here, we investigated seasonal variation in mating and fertility in Incarvillea sinensis (Bignoniaceae), an annual with showy, insect-pollinated, ‘one-day’ flowers capable of delayed selfing. We examined the influence of several biotic and abiotic environmental factors on day-to-day variation in fruit set, seed set and patterns of mating.Methods We recorded daily flower number and pollinator abundance in nine 3 × 3-m patches in a population at Mu Us Sand land, Inner Mongolia, China. From marked flowers we collected data on daily fruit and seed set and estimated outcrossing rate and biparental inbreeding using six microsatellite loci and 172 open-pollinated families throughout the flowering period.Key Results Flower density increased significantly over most of the 50-d flowering season, but was associated with a decline in levels of pollinator service by bees, particularly on windy days. Fruit and seed set declined over time, especially during the latter third of the flowering period. Multilocus estimates of outcrossing rate were obtained using two methods (the programs MLTR and BORICE) and both indicated high selfing rates of ∼80 %. There was evidence for a significant increase in levels of selfing as the flowering season progressed and pollinator visitation declined. Biparental inbreeding also declined significantly as the flowering season progressed.Conclusions Temporal variation in outcrossing rates may be a common feature of the mating biology of annual, insect-pollinated plants of harsh environments but our study is the first to examine seasonal mating-system dynamics in this context. Despite having large flowers and showy floral displays, I. sinensis attracted relatively few pollinators. Delayed selfing by corolla dragging largely explains the occurrence of mixed mating in I. sinensis, and this mode of self-fertilization probably functions to promote reproductive assurance when pollinator service is limited by windy environmental conditions and competition from co-occurring flowering plants.     

Natural selection on floral morphology can be influenced by climate

Climate has the potential to influence evolution, but how it influences the strength or direction of natural selection is largely unknown. We quantified the strength of selection on four floral traits of the subalpine herb Ipomopsis sp. in 10 years that differed in precipitation, causing extreme temporal variation in the date of snowmelt in the Colorado Rocky Mountains. The chosen floral traits were under selection by hummingbird and hawkmoth pollinators, with hawkmoth abundance highly variable across years. Selection for flower length showed environmental sensitivity, with stronger selection in years with later snowmelt, as higher water resources can allow translation of pollination success into fitness based on seed production. Selection on corolla width also varied across years, favouring narrower corolla tubes in two unusual years with hawkmoths, and wider corollas in another late snowmelt year. Our results illustrate how changes in climate could alter natural selection even when the primary selective agent is not directly influenced.     

Relationship between flowering time and fruit set of the entomophilous alpine shrub,Rhododendron aureum (Ericaceae), inhabiting snow patches

Flowering phenology, fruit set, and pollinator frequency of the ericaceous alpine dwarf shrub, Rhododendron aureum, were compared among three quadrats arranged along a snowmelt gradient on a slope. Bagging and self-pollination experiments showed that R. aureum was physiologically self-compatible, but pollinator visitation highly enhanced fruit set. Depending on the lateness of snowmelt, flowering time varied from mid-June to late July among quadrats. Pollinator visitation increased as the flowering season progressed, and fruit set was significantly higher in the quadrat with latest snowmelt. Emasculation experiments suggested that later flowering might promote cross-pollination. Thus, later flowering was advantageous for effective pollination. On the other hand, later-flowering plants often failed to set fruits because of the onset of autumn frost and snow before fruit maturation.     

Reproductive biology of the andromonoecious Cucumis melo subsp. agrestis (Cucurbitaceae)

Background and Aims

Cucumis melo subsp. agrestis (Cucurbitaceae) is cultivated in many African regions for its edible kernels used as a soup thickener. The plant, an annual, andromonoecious, trailing-vine species, is of high social, cultural and economic value for local communities. In order to improve the yield of this crop, the first step and our aim were to elucidate its breeding system.

Methods

Eight experimental pollination treatments were performed during three growing seasons to assess spontaneous selfing, self-compatibility and effects of pollen source (hermaphroditic vs. male flowers). Pollination success was determined by pollen tube growth and reproductive success was assessed by fruit, seed and seedling numbers and characteristics. The pollinator guild was surveyed and the pollination distance determined both by direct observations and by indirect fluorescent dye dispersal.

Key Results

The species is probably pollinated by several Hymenoptera, principally by Hypotrigona para. Pollinator flight distances varied from 25 to 69 cm. No evidence for apomixis or spontaneous self-pollination in the absence of insect visitors was found. The self-fertility index (SFI = 0) indicated a total dependence on pollinators for reproductive success. The effects of hand pollination on fruit set, seed number and seedling fitness differed among years. Pollen tube growth and reproductive success did not differ between self- and cross-pollinations. Accordingly, a high self-compatibility index for the fruit set (SCI = 1·00) and the seed number (SCI = 0·98) and a low inbreeding depression at all developmental stages (cumulative δ = 0·126) suggest a high selfing ability. Finally, pollen origin had no effect on fruit and seed sets.

Conclusions

This andromonoecious species has the potential for a mixed mating system with high dependence on insect-mediated pollination. The selfing rate through geitonogamy should be important.     

Significance of intra-inflorescence variation on flowering time of a spring ephemeral, <Emphasis Type="Italic">Gagea lutea</Emphasis> (Liliaceae), under seasonal fluctuations of pollinator and light availabilities

I studied the relationship between seed-set patterns within inflorescences and temporal variations in light and pollinator availabilities for 2 years in the spring ephemeral species Gagea lutea in a deciduous forest. Timing of canopy closure and seasonal trend of pollinator frequency did not synchronize with the annual fluctuation in flowering phenology. In the early snowmelt year, seed-set success reflected the seasonal pollinator abundance from early to middle flowering periods. In the late snowmelt year, however, seed-set rates were independent of pollinator activity and decreased with canopy closing even after hand pollination. The restricted seed production by defoliation and the increase in seed-set rates at the forest edge suggested that seed production was supported by current photosynthetic carbon gain. Thus, annual fluctuations of reproductive success can explain the variation in flowering phenology within a population although seasonal light deterioration would serve as a selective force for flowering in the early season.     

Identification of major quantitative trait loci underlying floral pollination syndrome divergence in Penstemon

Distinct floral pollination syndromes have emerged multiple times during the diversification of flowering plants. For example, in western North America, a hummingbird pollination syndrome has evolved more than 100 times, generally from within insect-pollinated lineages. The hummingbird syndrome is characterized by a suite of floral traits that attracts and facilitates pollen movement by hummingbirds, while at the same time discourages bee visitation. These floral traits generally include large nectar volume, red flower colour, elongated and narrow corolla tubes and reproductive organs that are exerted from the corolla. A handful of studies have examined the genetic architecture of hummingbird pollination syndrome evolution. These studies find that mutations of relatively large effect often explain increased nectar volume and transition to red flower colour. In addition, they suggest that adaptive suites of floral traits may often exhibit a high degree of genetic linkage, which could facilitate their fixation during pollination syndrome evolution. Here, we explore these emerging generalities by investigating the genetic basis of floral pollination syndrome divergence between two related Penstemon species with different pollination syndromes—bee-pollinated P. neomexicanus and closely related hummingbird-pollinated P. barbatus. In an F₂ mapping population derived from a cross between these two species, we characterized the effect size of genetic loci underlying floral trait divergence associated with the transition to bird pollination, as well as correlation structure of floral trait variation. We find the effect sizes of quantitative trait loci for adaptive floral traits are in line with patterns observed in previous studies, and find strong evidence that suites of floral traits are genetically linked. This linkage may be due to genetic proximity or pleiotropic effects of single causative loci. Interestingly, our data suggest that the evolution of floral traits critical for hummingbird pollination was not constrained by negative pleiotropy at loci that show co-localization for multiple traits.     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

Reproductive isolation and pollination success of rewarding Galearis diantha and non-rewarding Ponerorchis chusua (Orchidaceae)

Background and Aims

Increasing evidence challenges the conventional perception that orchids are the most distinct example of floral diversification due to floral or prezygotic isolation. Regarding the relationship between co-flowering plants, rewarding and non-rewarding orchids in particular, few studies have investigated whether non-rewarding plants affect the pollination success of rewarding plants. Here, floral isolation and mutual effects between the rewarding orchid Galearis diantha and the non-rewarding orchid Ponerorchis chusua were investigated.

Methods

Flowering phenological traits were monitored by noting the opening and wilting dates of the chosen individual plants. The pollinator pool and pollinator behaviour were assessed from field observations. Key morphological traits of the flowers and pollinators were measured directly in the field. Pollinator limitation and interspecific compatibility were evaluated by hand-pollination experiments. Fruit set was surveyed in monospecific and heterospecific plots.

Key Results

The species had overlapping peak flowering periods. Pollinators of both species displayed a certain degree of constancy in visiting each species, but they also visited other flowers before landing on the focal orchids. A substantial difference in spur size between the species resulted in the deposition of pollen on different regions of the body of the shared pollinator. Hand-pollination experiments revealed that fruit set was strongly pollinator-limited in both species. No significant difference in fruit set was found between monospecific plots and heterospecific plots.

Conclusions

A combination of mechanical isolation and incomplete ethological isolation eliminates the possibility of pollen transfer between the species. These results do not support either the facilitation or competition hypothesis regarding the effect of nearby rewarding flowers on non-rewarding plants. The absence of a significant effect of non-rewarding P. chusua on rewarding G. diantha can be ascribed to low levels of overlap between the pollinator pools of two species.     

Fruit regulates seasonal expression of flowering genes in alternate-bearing 'Moncada' mandarin

Background and Aims

The presence of fruit has been widely reported to act as an inhibitor of flowering in fruit trees. This study is an investigation into the effect of fruit load on flowering of ‘Moncada’ mandarin and on the expression of putative orthologues of genes involved in flowering pathways to provide insight into the molecular mechanisms underlying alternate bearing in citrus.

Methods

The relationship between fruit load and flowering intensity was examined first. Defruiting experiments were further conducted to demonstrate the causal effect of fruit removal upon flowering. Finally, the activity of flowering-related genes was investigated to determine the extent to which their seasonal expression is affected by fruit yield.

Key Results

First observations and defruiting experiments indicated a significant inverse relationship between preceding fruit load and flowering intensity. Moreover, data indicated that when fruit remained on the tree from November onwards, a dramatic inhibition of flowering occurred the following spring. The study of the expression pattern of flowering-genes of on (fully loaded) and off (without fruits) trees revealed that homologues of FLOWERING LOCUS T (FT), SUPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1), APETALA1 (AP1) and LEAFY (LFY) were negatively affected by fruit load. Thus, CiFT expression showed a progressive increase in leaves from off trees through the study period, the highest differences found from December onwards (10-fold). Whereas differences in the relative expression of SOC1 only reached significance from September to mid-December, CsAP1 expression was constantly higher in those trees through the whole study period. Significant variations in CsLFY expression only were found in late February (close to 20 %). On the other hand, the expression of the homologues of TERMINAL FLOWER 1 (TFL1) and FLOWERING LOCUS C (FLC) did not appear to be related to fruit load.

Conclusions

These results suggest for the first time that fruit inhibits flowering by repressing CiFT and SOC1 expression in leaves of alternate-bearing citrus. Fruit also reduces CsAP1 expression in leaves, and the significant increase in leaf CsLFY expression from off trees in late February was associated with the onset of floral differentiation.     

Variation in pollen limitation and floral parasitism across a mating system transition in a Pacific coastal dune plant: evolutionary causes or ecological consequences?

Background and Aims Evolutionary transitions from outcrossing to self-fertilization are thought to occur because selfing provides reproductive assurance when pollinators or mates are scarce, but they could also occur via selection to reduce floral vulnerability to herbivores. This study investigated geographic covariation between floral morphology, fruit set, pollen limitation and florivory across the geographic range of Camissoniopsis cheiranthifolia, a Pacific coastal dune endemic that varies strikingly in flower size and mating system.Methods Fruit set was quantified in 75 populations, and in 41 of these floral herbivory by larvae of a specialized moth (Mompha sp.) that consumes anthers in developing buds was also quantified. Experimental pollen supplementation was performed to quantify pollen limitation in three large-flowered, outcrossing and two small-flowered, selfing populations. These parameters were also compared between large- and small-flowered phenotypes within three mixed populations.Key Results Fruit set was much lower in large-flowered populations, and also much lower among large- than small-flowered plants within populations. Pollen supplementation increased per flower seed production in large-flowered but not small-flowered populations, but fruit set was not pollen limited. Hence inadequate pollination cannot account for the low fruit set of large-flowered plants. Floral herbivory was much more frequent in large-flowered populations and correlated negatively with fruit set. However, florivores did not preferentially attack large-flowered plants in three large-flowered populations or in two of three mixed populations.Conclusions Selfing alleviated pollen limitation of seeds per fruit, but florivory better explains the marked variation in fruit set. Although florivory was more frequent in large-flowered populations, large-flowered individuals were not generally more vulnerable within populations. Rather than a causative selective factor, reduced florivory in small-flowered, selfing populations is probably an ecological consequence of mating system differentiation, with potentially significant effects on population demography and biotic interactions.     

四川黄龙沟少花鹤顶兰繁殖成功特征

少花鹤顶兰(Phaiu delavayi)是多年生、多次结实的具有克隆能力的地生兰科植物,是中国特有种。目前少花鹤顶兰开花物候、花寿命、繁殖成功等生物学特征尚不清楚,而这些资料是开展少花鹤顶兰进化和保护生物学研究的基础。利用2005-2007年四川黄龙沟少花鹤顶兰的调查数据,对少花鹤顶兰繁殖成功特征进行了研究。结果表明黄龙沟少花鹤顶兰每个花序的平均花朵数为3(1-7)朵。叶和花的形态指标在年季间无显著变异。2005-2007年3年间的开花物候没有明显的差异,花期始于6月中旬,于7月底至8月初结束,持续约6-7周,80%-90%的花在7月初的开花高峰期开放,属花集中开放模式。花寿命与是否授粉密切相关,成功授粉的花寿命比没有授粉的花寿命短。少花鹤顶兰是自交亲和的,但其结实完全依赖于两种熊蜂的传粉,不存在自动自花授粉和无融合生殖现象。柱头可授性和花粉活力可维持15 d左右。繁殖成功率年季间的变化很大,并表现出逐年下降的趋势,花粉移走率为18%-51%,自然结实率为10%-36%。繁殖成功下降的趋势可能与黄龙沟喷洒农药有关。     

Orthoptera, a new order of pollinator

Background and Aims

Pollinator-mediated selection and evolution of floral traits have long fascinated evolutionary ecologists. No other plant family shows as wide a range of pollinator-linked floral forms as Orchidaceae. In spite of the large size of this model family and a long history of orchid pollination biology, the identity and specificity of most orchid pollinators remains inadequately studied, especially in the tropics where the family has undergone extensive diversification. Angraecum (Vandeae, Epidendroideae), a large genus of tropical Old World orchids renowned for their floral morphology specialized for hawkmoth pollination, has been a model system since the time of Darwin.

Methods

The pollination biology of A. cadetii, an endemic species of the islands of Mauritius and Reunion (Mascarene Islands, Indian Ocean) displaying atypical flowers for the genus (white and medium-size, but short-spurred) was investigated. Natural pollinators were observed by means of hard-disk camcorders. Pollinator-linked floral traits, namely spur length, nectar volume and concentration and scent production were also investigated. Pollinator efficiency (pollen removal and deposition) and reproductive success (fruit set) were quantified in natural field conditions weekly during the 2003, 2004 and 2005 flowering seasons (January to March).

Key Results

Angraecum cadetii is self-compatible but requires a pollinator to achieve fruit set. Only one pollinator species was observed, an undescribed species of raspy cricket (Gryllacrididae, Orthoptera). These crickets, which are nocturnal foragers, reached flowers by climbing up leaves of the orchid or jumping across from neighbouring plants and probed the most ‘fresh-looking’ flowers on each plant. Visits to flowers were relatively long (if compared with the behaviour of birds or hawkmoths), averaging 16·5 s with a maximum of 41·0 s. At the study site of La Plaine des Palmistes (Pandanus forest), 46·5 % of flowers had pollen removed and 27·5 % had pollinia deposited on stigmas. The proportion of flowers that set fruit ranged from 11·9 % to 43·4 %, depending of the sites sampled across the island.

Conclusions

Although orthopterans are well known for herbivory, this represents the first clearly supported case of orthopteran-mediated pollination in flowering plants.     

Plants,position and pollination: Planting arrangement and pollination limitation in a revegetated eucalypt woodland

The spatial arrangement of plants is a key determinant of pollination services in natural ecosystems. Despite this, plant arrangements are rarely considered an important characteristic of revegetated communities and this may be limiting successful pollination dynamics in these re‐created systems. We assessed if aggregated South Australian Blue Gum (Eucalyptus leucoxylon, F. Muell) had greater pollination success than more dispersed trees by measuring fruit set, seed production and germination in a revegetated eucalypt woodland. The proportion of buds that developed into fruit (fruit set) was similar between aggregated (five flowering Blue Gum within 30 m) and dispersed (no flowering Blue Gum within 30 m) trees (<14%). Aggregated and dispersed trees produced a similar number of seeds per fruit in 2015, when fewer trees flowered and those that flowered produced fewer flowers (ca 8 seeds/fruit). In 2016, when flowering was more abundant, aggregated trees produced more seeds per fruit than dispersed trees (12 seeds/fruit compared to 4 seeds/fruit). Despite differences in seed number, there was no difference in the quality of seed produced, indicating outcrossed pollen was received by both aggregated and dispersed trees. Although outcrossing was likely, this did not prevent pollination limitation, with the addition of outcrossed pollen increasing fruit and seed set for trees in both arrangements. Consequently, we propose that planting individuals of the same species (conspecifics) in clumps, rather than intermixing species, can increase seed set in revegetated eucalypts, particularly in good flowering years. However, aggregated plantings will be most effective if the needs and foraging behaviour of pollinators are also considered, to attract and sustain pollinators in revegetated systems, thereby minimizing the risk of pollination limitation.