The pollination of Cymbidiella flabellata (Orchidaceae) in Madagascar: A system operated by sphecid wasps

Interactions between the terrestrial endemic Cymbidiella flabellata (Thou.) Rolfe (Orchidaceae) and flower-visiting insects were observed in a marsh on the east coast of Madagascar. The plant acted by deceit. Two large sphecid wasp species, females of Sceliphron fuscum Klug and males of Sphex torridus Sm. (Hymenoptera, Sphecidae), made frequent visits to the showy, ca. 5 cm wide, nectarless and fragrant flowers. During a visit the wasps alighted on the large, hinged labellum which they depressed while forcing their heads towards the basal cavity below the column. When the wasps withdrew they pressed powerfully down on the labellum and brushed their heads backwards against the floral sexual organs removing the pollinarium with the frons or vertex – from which position pollination took place. The wasps on average visited only two flowers consecutively and about 55% of the flowers in the orchid population received visits. About six times more pollinaria were dispersed by pollen vectors than reached stigmas and a high proportion of the flowers were aborted due to lack of pollination. Data suggest that S. fuscum , a spider hunting and mud-daubing species, is a principal pollinator. C. flabellata is interpreted as playing the role of a generalized food-flower mimic which is specialized to moist habitats where females of S. fuscum collect mud for their nests.     

草豆蔻传粉生物学的研究

比较研究了花柱卷曲性植物草豆蔻 Alpinia hainanensis两种表型(上举型和下垂型)的传粉生物学特性。结果表明:草豆蔻的花柱卷曲运动节律与其他已报道的山姜属Alpinia植物基本一致,而下垂型花的花柱卷曲运动明显滞后于上举型花约2 h。草豆蔻的花柱卷曲运动中存在一些不同步的现象,甚至在同一花序上的两朵花的花柱运动节律亦会表现出不一致的现象,但上举型花的花药开裂时间却严格同步,都发生在同类型个体的柱头全部位于花药上方之后进行。草豆蔻上举型花的花蜜分泌量、单花花粉量、花粉/胚珠比率(P/O)均明显比下垂型花多,而两种表型的胚珠数、花蜜糖含量、氨基酸含量无显著差异。在草豆蔻的单花期内不同时间段进行人工授粉,上举型花在其柱头位于花药下方时进行人工授粉,其结实率明显比柱头位于花药上方时人工授粉的处理高,下垂型花则没有显著差异。在自然居群中,草豆蔻的主要传粉者是无垫蜂Ameglla sp.和两种木蜂Xylocopa sp.,但存在传粉者不足而影响结实率的现象。完全套袋、去雄和去雌(去柱头)处理的均不结实,表明草豆蔻中不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象。而人工自交和异交均具有较高的结实率,表明草豆蔻为自交亲和植物。草豆蔻的繁育系统是具花柱卷曲性运动的异花授粉的交配系统。     

Flowering phenology and reproductive biology of Drosera anglica (Droseraceae)

Prompted by the sparse knowledge of the reproductive biology of carnivorous plants, compared with studies of their trapping habits, we investigated the flowering phenology and pollination biology of Drosera anglica Huds. in two fens in mid-western Canada. Seed set and germination were used to compare the effectiveness of a series of pollination treatments, including single insect visits to virgin flowers. Flowers opened during mid-morning but closed by early afternoon, and exhibited pseudo-cleistogamic behaviour in cool, overcast weather. D. anglica was found to be self-compatible, and able to self-pollinate and self-fertilize. Geitonogamy was an uncommon mode of self-reproduction because plants typically possessed a lone inflorescence upon which a single, short-lived flower opened, a few days before the next bud reached anthesis. Insect visits to the fragrance-lacking, nectarless flowers, chiefly by flies (Diptera: Syrphidae), were infrequent (one visit per 1 h 40 min of observation), and the low frequency of seed set and low numbers of seeds per fruit in pollination treatments involving insects, suggest the species does not rely on insects to effect pollination. Self-pollination, with or without the aid of a vector (insects, wind) was as effective as natural pollination; ultimately, autogamy is chiefly responsible for natural seed set. Thus, the species exhibits characteristics of facultative autogamy.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 417–426.     

The effect of flower position on male and female reproductive success in a deceptively pollinated tropical orchid

In many plants, including orchids, differential fruit set along the inflorescence has been attributed to pollinator behaviour. For instance, the pollinator, moving up the inflorescence, becomes satiated with the resources and leaves before visiting the upper flowers. Consequently, the pollinators do not visit flowers as frequently higher up the inflorescence. Alternatively, flower size may vary along the inflorescence, making pollination ineffective as flowers decrease in size. I tested for the presence of differential pollination along the inflorescence in a pollinator-limited tropical epiphyte, Lepanthes rupestris Stimson, and determined the likely cause of the observed pattern. As this species has inflorescences with sequential flowering, pollinator behaviour, moving up the inflorescence as in synchronous multiflowering inflorescences, can be discounted as an explanation for differential fruit set. Fruit set is shown to be more frequent at the base of the inflorescence, but male reproductive success through pollinarium removal is basically independent of flower position. Moreover, cross-pollination by hand at variable flower positions along the inflorescence results in equal fruit set, suggesting that resources are not limiting and cannot explain the cause of differential fruit production along the inflorescence in natural populations. Furthermore, flower size is shown to diminish along the inflorescence, suggesting that the pollinator(s) may be ineffective at depositing the pollinarium in the smaller higher flowers. Consequently, pollinator behaviour and its interaction with flower size, and not resource limitation, is likely to be the main cause of differential fruit set along the inflorescence in L. rupestris .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 405–410.     

INFLORESCENCE SIZE AND FRUIT DISTRIBUTION AMONG INDIVIDUALS IN THREE ORCHID SPECIES

Fruit distribution among individuals in populations of three orchid species was analyzed. Fruit set was low in the three species. The distribution of fruits in the non-autogamous species was skewed to the right, with high variance: mean ratios, and a moderate to high proportion of fruitless plants. Larger inflorescence size did not necessarily result in an increase in percent fruit set but tended to increase the probability to produce at least one fruit. Pollinator-mediated selection on inflorescence size through fruit production may be limited by the low overall level of visitation and the resulting uncertainty of pollination at the individual level.     

Notes on the pollination biology of <Emphasis Type="Italic">Notylia nemorosa</Emphasis> (Orchidaceae): do pollinators necessarily promote cross pollination?

The pollination biology of Notylia nemorosa was elucidated from field and ex situ observations. Field observations were carried out in Minas Gerais, southeastern Brazil, where this orchid is pollinated by males of Eulaema nigrita(Euglossini). Plants cultivated in Campinas (S?o Paulo State, southeastern Brazil) were eventually pollinated by males of Eulaema nigrita and Euglossa melanothricha(Euglossini). In both cases, Euglossini males collect aromatic compounds produced on the surface of the lip, fix the pollinaria on the dorsal surface of the labrum and perform pollination. This species is protandrous. Recently opened flowers present their stigmatic surfaces blocked and thus flowers act only as pollen donors. After 2-3 days the viscidium dries out and pollinaria cannot be removed anymore. Then, a narrow slit opens in the stigmatic cavity, allowing pollen deposition. Experiments performed with cultivated plants suggest that this orchid species is strongly, though not completely self-incompatible. Pollination experiments showed that bees tend to stay several minutes in each inflorescence. This behavior may promote some degree of geitonogamous pollination because flowers in male and female phases coexist in the same inflorescence. The combination of self-incompatibility with this specific pollinator behavior may explain the rarity of fruits in several neotropical reward-offering Orchidaceae.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Specialization for pollination by beetles and wasps: the role of lollipop hairs and fragrance in Satyrium microrrhynchum (Orchidaceae)

Exposed nectar presentation is a key trait in flowers specialized for pollination by short-tongued insects. We investigated the pollination of Satyrium microrrhynchum, a rare South African orchid in which nectar is secreted as droplets on long floral hairs ("lollipop hairs") at the mouth of a shallow labellum. Our observations indicate that this orchid is pollinated specifically by two insect species: a cetoniid beetle (Atrichelaphinus tigrina) and a pompilid wasp (Hemipepsis hilaris). Both insects have short mouthparts and remove nectar from the hairs with sweeping motions of their mouthparts. Pollinaria become attached to the upper surface of their heads while they feed on the nectar. Beetles damage the hairs while feeding, which may explain the positive relationship between hair damage and pollination success in plants of S. microrrhynchum from populations where beetles were common. The orchid has cryptic green-yellow flowers with spectral reflectance similar to that of its leaves. The fragrance from plants in three populations, analyzed using gas chromatography coupled to mass spectrometry, was dominated by various terpenoids; linalool was the most abundant. Plants in different populations emitted similar compounds, but eugenol and derivatives of this compound were found in only one of the three populations. In an electrophysiological study (gas chromatography coupled to electroantennography), using antennae of A. tigrina, clear signals were elicited by some of the floral scent compounds.     

Evidence for widespread pollen limitation of fruiting success in Cape wildflowers

We used spot checks of stigmatic pollen deposition and hand-pollination experiments to test whether fruit production in Cape wildflower populations is limited by pollen availability. Natural levels of stigmatic pollen deposition were very low (median = 30.0% of flowers) in populations of 33 orchid species. We found similarly low levels of fruit set (median = 32% of flowers per plant) in six Orchidaceae and four Amaryllidaceae species. Experimental hand pollination at the whole plant level caused significant increases in fruit production in 11 of the 12 study populations. These results indicate that pollen limitation of fruit set may occur frequently among some plant families in the Cape flora. Received: 15 April 1996 / Accepted: 10 November 1996     

Reproductive biology of Cyrtopodium punctatum in situ: implications for conservation of an endangered Florida orchid

Cyrtopodium punctatum (Linnaeus) Lindley is an endangered epiphytic orchid restricted in the USA to southern Florida. This species has been extensively collected from the wild since the early 1900s, and today only a few plants remain in protected areas. As part of a conservation plan, a reproductive biology study was conducted to better understand the ecology of this species in Florida. Cyrtopodium punctatum relies on a deceit pollination system using aromatic compounds to attract pollinators. Nine aromatic compounds were identified as components of the fragrance of C. punctatum inflorescences, including two compounds that are known to be Euglossine bee attractants. However, this group of bees is not native to Florida. Of the four bee species observed to visit C. punctatum flowers in the present study, carpenter bees ( Xylocopa spp.) are likely to be the main pollinators. Pollination experiments demonstrated that C. puntatum is self-compatible, but requires a pollinator and thus does not exhibit spontaneous autogamy. In addition, the rates of fruit set were significantly higher for flowers that were outcrossed (xenogamy) than for those that were self-crossed. Thus, the species has evolved a degree of incompatibility. Examples of natural pollination and fruit set were observed during the present study (2007–2008), but the rates of reproduction were modest as a consequence of the low plant numbers and possible changes in insect densities as a result of anthropogenic influences.     

兰科植物欺骗性传粉

植物与传粉动物的互利关系在生态系统中非常普遍。然而,有许多植物不为传粉者提供任何报酬,而是利用各种欺骗方式诱骗昆虫拜访,从而实现传粉,即欺骗性传粉。兰科是被子植物大科之一,其高度特化的繁殖器官和适应于昆虫传粉的精巧结构令人称奇。进化论创始人达尔文描述了许多兰花与昆虫精巧的传粉系统,但他忽视了欺骗性传粉的存在。事实上,近1/3的兰科植物都依赖于欺骗性传粉。欺骗性传粉可能是导致兰科植物多样性的重要原因之一。兰花利用或操作昆虫觅食、交配、产卵和栖息等行为,演化出各种各样的欺骗性传粉机制,常见的类型包括泛化的食源性欺骗、Batesian拟态、性欺骗、产卵地拟态和栖息地拟态。花的颜色、形态和气味在欺骗性传粉的成功实现中起到了重要作用。欺骗性兰花与传粉昆虫之间的演化可能是不同步的,兰花追踪昆虫的行为信号而发生分化,然而欺骗性传粉可能对昆虫造成一定的伤害,从而对昆虫也施加选择压力。由于昆虫的学习行为,欺骗性的兰花一般具有低的昆虫拜访率和结实率,其繁殖成功率受各种因素的影响。欺骗性加剧了兰花对传粉昆虫的依赖,使其具有更高的灭绝风险,传粉生物学的研究能为兰科植物的有效保护提供指导。在欺骗性传粉系统中,有报酬的伴生植物、拟态模型和其他拟态信号提供者对传粉成功有重要影响。因此,研究欺骗性传粉兰花、传粉昆虫和相关的生物和生态因子的网状进化关系具有重要理论和实践意义。     

Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.     

濒危植物独花兰的传粉生物学初步观察

独花兰(Changnienia amoena Chien)为我国特有的单种属植物,近年因生境破碎化和过度采挖,其野生资源日渐减少。迄今对独花兰极为有限的研究表明,其结实率很低或根本不结实,其传粉媒介也一直未被发现。2002年3～4月,我们对神农架2个移植居群和5个天然居群进行了传粉生物学的定点观察,发现雌性三条熊蜂(Bombus (Diversobombus)trifasciatus Smith)、仿熊蜂(Bombus (Tricornibombus) imitator Pittion)和蜜蜂均访问独花兰,但只有二条熊蜂身体粘有花粉块,是独花兰的有效传粉者。三条熊蜂的访问频率很低,在113h的观察中只有9次访问,但在一个天然居群(population 3)中曾观察到一天4次的最高访问频率;访问主要在12：00～15：00出现,但在花上停留时间很短,不超过10s。在末被授粉的情况下,独花兰花朵大约3周后自然枯萎,但受粉后3、4d内即出现～系列形态和颜色的变化,包括花梗逐渐伸长,子房在花梗逐渐停止伸长后开始膨大等,表明花梗伸长可作为结实(授粉成功)的指标。人工授粉实验表明,自花、异花受粉后花梗均伸长,而套袋隔离花的则花梗不伸长,说明独花兰是自交亲和的异交种,需要昆虫传粉。根据传粉者的访问频率、居群中果实的分布,尤其是花距内无花蜜等特征,我们认为独花兰是一种欺骗性传粉的兰花。相对于其他欺骗性传粉的兰花,独花兰的自然结实率并不很低(26．98％),这与居群规模小会提高欺骗性传粉兰花结实率这一观点吻合。花粉块的输出数远高于结实数,这说明独花兰存在一定的花粉浪费。     

Pollinator limitaton in a deceptive orchid, Pogonia japonica, on a floating peat mat

Deceptive orchid species that offer no floral rewards to pollinators often experience pollinator limitation because they. The breeding system and fruit set were investigated in order to examine pollinator limitation for a population of a deceptive orchid, Pogonia japonica , on a floating peat mat in Mizorogaike Pond, western Japan. Fruit sets for outcross-pollinated, self-pollinated and pollinator-excluded flowers were 75%, 80%, and 0%, respectively. Thus, this species was self-compatible but neither autogamous nor apogamous. The fruit set for open-pollinated flowers was 22.9% and 17.4% in 1994 and 1995, suggesting that pollinator limitation did occur in the field. Larger flowers were more likely to develop into fruit than smaller ones in open pollination (36% vs 8%). Thus, flower size seems to affect fruiting of this species. No effective visits by major pollinators such as bees, which could carry pollinia, were observed during 16.5 h of observation during daytime. Instead, thrips were often found moving on gynostemium or pollinia and circumstantial evidence that suggests thrips may partly transfer pollen in pieces out of a granular pollinium of P. japonica was obtained. Pogonia japonica should come to use thrips as a pollinator to supplement low fruit set under limited flower visitation by major pollinators.     

Pollinator limitation and position dependent fruit set in the high Andean orchidMyrosmodes cochleare (Orchidaceae)

Myrosmodes cochleare is a terrestrial orchid restricted to high elevations of the northern Andes. It is self-compatible but not autogamous. Flowering begins at the apex of the inflorescence, but flower size increases steadily towards the base. Fruit set of selfed flowers was found to increase significantly from the apex to the base of the inflorescence. Open-pollinated fruit set was significantly lower than hand-pollinated fruit set overall, but the highest levels occurred in the middle of the inflorescence. Eumenid wasps and a calliphorid fly were observed to pollinate the flowers, but visits were rare and occurred only during infrequent warm periods.     

Ecological constraints on fruit initiation by Calyptrogyne ghiesbreghtiana (Arecaceae): FLORAL HERBIVORY,POLLEN AVAILABILITY,AND VISITATION BY POLLINATING BATS

Failure of a flower to initiate fruit can be attributed to an insufficiency of resources, genetic incompatibilities, or ecological constraints on pollination. Flower visitors can increase fruit initiation by pollinating flowers, or decrease fruit initiation by damaging them. If, however, resources limit fruit initiation, variation in visitation may be relatively unimportant. Visitors to Calyptrogyne ghiesbreghtiana, a rain forest understory palm, leave a record of their activity by feeding on floral tissue. I recorded variation in inflorescence visitation, floral display, and fruit initiation during 7 mo over two field seasons. On average, bats visited 50-60% of flowers; katydids and scarab beetles visited and damaged the remaining 40-50%. Four species of bat (Phyllostomidae) captured near inflorescences were found to be carrying C. ghiesbreghtiana pollen. Approximately 18% of flowers visited by bats and not damaged by insects initiated fruit, whereas insect-damaged flowers set a negligible number. Some variation in fruit initiation was explained by nightly variation in the proportion of inflorescences at each site that were potential pollen donors (male-phase inflorescences). Visitors responded to variation in floral display; katydids damaged more flowers on taller inflorescences, and in one season bats visited a greater proportion of flowers on inflorescences with many flowers. However, floral display only explained a small amount of variation in visitation. A similar amount of variation in flower visitation was explained by the degree to which inflorescences were obstructed by surrounding vegetation. I used path analysis to summarize the overall effect of floral display on visitation by both pollinators and florivores and the effect of visitation and site sex ratio on fruit initiation. Because almost all flowers were visited by bats or damaged by katydids, the amount of bat visitation was strongly negatively correlated with katydid damage. In spite of the low fruit set from bat-visited flowers, variation in bat visitation explained 27-37% of the variation in fruit initiation.     

Phylogeny and radiation of pollination systems in DISA (Orchidaceae)

We studied the patterns of adaptive radiation in Disa, a large orchid genus in southern Africa. A cladogram for 27 species was constructed using 44 morphological characters. Pollination systems were then mapped onto the phylogeny in order to analyze pathways of floral evolution. Shifts from one pollination system to another have been a major feature of the evolutionary diversification of Disa. Unlike many plant genera that are pollinated mainly by a single group of insects, radiation in Disa has encompassed nearly all major groups of pollinating insects; in all, 19 different specialized pollination systems have been found in the 27 species included in this analysis. Another striking pattern is the repeated evolution of broadly similar pollination systems in unrelated clades. For example, butterfly-pollinated flowers have evolved twice; showy deceptive flowers pollinated by carpenter bees, twice; long-spurred flowers pollinated by long-tongued flies, four times; night-scented flowers pollinated by moths, three times; and self-pollination, three times. This suggests that a few dominant pollinator species in a region may be sufficient to generate diversification in plants through repeated floral shifts that never retrace the same pathways.     

Wind pollination in high-mountain populations of Hormathophylla spinosa (Cruciferae)

In this paper we study aspects of the breeding system of Hormathophylla spinosa (Crucifcrae) to identify the factors responsible for seed production in the absence of insect pollinators. The pollinator-exclusion experiments show that H. spinosa, under natural conditions, does not produce seed by apomixis or spontaneous autogamy. H. spinosa appears to be self-incompatible but slightly geitonogamous. Thus, this plant species needs pollen vectors for reproduction. The results of the wind-exclusion experiments performed during two different years in two populations of H. spinosa support the hypothesis that the wind acts as a pollen vector; flowers excluded from the wind had a lower fruit set and female fertility than flowers excluded from all pollinator insects (winged and wingless). This generalist pollination system (insects and wind) permits this species to colonize and sustain viable populations in high mountains, where this species is the only woody shrub living above 3000 m above sea level in the Sierra Nevada.     

Rapid floral senescence following male function and breeding systems of some tropical orchids

No comparative study of floral senescence following male function among a range of tropical orchid genera has previously been undertaken. The timing and pattern of floral senescence and occurrence of fruit formation were studied following self-, geitonogamous and cross-pollination in 14 epiphytic and two terrestrial orchid species to determine their breeding system and assess the occurrence of floral abscission following pollinaria removal. Both pollination and pollinaria removal caused rapid floral senescence, and the pattern and timing of the floral changes were the same in all treatments. Six Dendrobium species and Pelatantheria insectifera were self-incompatible (SI) and eight other species, including one terrestrial species, were self-compatible (SC). Capsules produced from outcrossing in four SC species, Phalaenopsis cornu-cervi, Eria pubescens, Cleisostoma appendiculatum and Arundina graminifolia, were larger and heavier than those produced after selfing. Reductions in flower life span following pollinaria removal were positively correlated with flower size and longevity of unpollinated flowers but not with position in the inflorescence or nature of the breeding system. Rapid flower senescence following pollinaria removal reported here suggests that it may be widespread in tropical species. The significant association of the response with size of flowers and inflorescences among the species studied suggests that the cost of flower maintenance outweighs the benefit of remaining open for female function after pollinaria have been removed. Both SC and SI species were found among tropical orchids, but variation in capsule size following self- and cross-pollination indicates that there may be a reduction in seed production following selfing, even in SC species, and that fruit formation alone should not be taken as reliable evidence of full self-compatibility.     

濒危植物独花兰的传粉生物学初步观察

独花兰(Changnienia amoena Chien)为我国特有的单种属植物,近年因生境破碎化和过度采挖,其野生资源日渐减少.迄今对独花兰极为有限的研究表明,其结实率很低或根本不结实,其传粉媒介也一直未被发现.2002年3～4月,我们对神农架2个移植居群和5个天然居群进行了传粉生物学的定点观察,发现雌性三条熊蜂(Bombus(Diversobombus)trifasciatus Smith)、仿熊蜂(Bombus(Tricornibombus)imitator Pittion)和蜜蜂均访问独花兰,但只有三条熊蜂身体粘有花粉块,是独花兰的有效传粉者.三条熊蜂的访问频率很低,在113 h的观察中只有9次访问,但在一个天然居群(population 3)中曾观察到一天4次的最高访问频率;访问主要在12:00～15:00出现,但在花上停留时间很短,不超过10 s.在未被授粉的情况下,独花兰花朵大约3周后自然枯萎,但受粉后3、4 d内即出现一系列形态和颜色的变化,包括花梗逐渐伸长,子房在花梗逐渐停止伸长后开始膨大等,表明花梗伸长可作为结实(授粉成功)的指标.人工授粉实验表明,自花、异花受粉后花梗均伸长,而套袋隔离花的则花梗不伸长,说明独花?兰是自交亲和的异交种,需要昆虫传粉.根据传粉者的访问频率、居群中果实的分布,尤其是花距内无花蜜等特征,我们认为独花兰是种欺骗性传粉的兰花.相对于其他欺骗性传粉的兰花,独花兰的自然结实率并不很低(26.98%),这与居群规模小会提高欺骗性传粉兰花结实率这一观点吻合.花粉块的输出数远高于结实数,这说明独花兰存在一定的花粉浪费.