The pollination ecology ofBletilla striata (Orchidaceae)

Anthecological observations of an orchidBletilla striata were carried out in Kobe, Japan. The purplish pink flowers were visited by 26 species of insects belonging to Hymenoptera, Diptera and Lepidoptera, although the flower did not offer food rewards of any kind. Candidates for pollination were seven species of aculeate Hymenoptera with a body size that well fitted the space in between the column and labellum. The most effective candidate was the male of the longhorn beeTetralonia nipponensis, judging from its abundance and legitimate intrafloral behavior. The female ofT. nipponensis and the other six species also contributed to promote out-crossing, but to a lesser extent than the maleT. nipponensis, since they occasionally exhibited illegitimate intrafloral behaviors for pollination and also, their flower visitation rate was lower.     

Leaves of the Orchid Twayblade (Listera ovata) Contain a Mannose-Specific Lectin

A new lectin was isolated from leaves of the twayblade (Listera ovata). It is a dimeric protein built up of two subunits of M_r 12,500. This lectin, which is the first to be isolated from a species of the family Orchidaceae, exhibits exclusive specificity towards mannose.     

The pollination ecology of Actaea spicata (Ranunculaceae)

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

The pollination ecology of Echeandia macrocarpa (Liliaceae)

Field research and observation of the breeding system of the salvadoran extension of theEcheandia macrocarpa complex indicate an entirely mellittophilous syndrome, withBombus ephippiatus Say workers as the primary pollen vectors. Foraging behavior coupled with the floral morphology/ phenology of the host plant suggests a trend towards obligatory out-crossing. The foraging behavior ofB. ephippiatus workers on the host plant and on flowers of co-blooming species is reported to elucidate the dynamic processes that determine the present co-evolutionary status ofE. macrocarpa and its chief pollinator.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

The pollination ecology of buzz-pollinated Rhexia virginica (Melastomataceae)

We examined the function of floral traits associated with buzz pollination through studies of Rhexia virginica (Melastomataceae) in the Muskoka region of Ontario, Canada. Controlled pollinations demonstrated that the species is self-compatible, but dependent on insects for pollen transfer. Bumble bees made 82 and 90% of observed insect visits to R. virginica in 1996 and 1997, respectively, and effectively buzzed flowers. Buzz pollination did not appear to be highly "specialized" since various species of bumble bee were capable of pollination, and pollen transfer efficiency appeared to be relatively low. Experimental manipulations provided little support for the hypothesis that the yellow color of melastome anthers mimics abundant pollen, thereby deceiving pollinators to visit regardless of whether most pollen has been removed. Fruit set averaged 52.6% among populations, owing largely to infrequent pollinator visits and pollen limitation. Flowers of R. virginica were infertile after a single day of anthesis, but petals were subsequently maintained for 1-2 d and stamens underwent a color change from bright yellow to red. Second-day flowers may function to increase floral display size and hence fertility, without a concomitant increase in pollen discounting. Studies of bumble bee foraging behavior and correlates of seed set provided indirect support for this hypothesis.     

芳香石豆兰的繁殖生态学

芳香石豆兰(Bulbophyllum ambrosia)是分布于我国南方石灰岩山热带季节性雨林中的一种常见附生兰科植物, 附着生长于岩石或树干上, 野外监测发现其自然不结实。对芳香石豆兰繁殖生态学的研究表明, 不同于石豆兰属其他具有蝇类传粉综合征的种类, 其花具有香味和花蜜, 中华蜜蜂(Apis cerana cerana)是芳香石豆兰唯一的有效传粉者, 具有较高的访问频率, 自然种群中有较高的花粉块移出率和柱头花粉块沉降率, 在中华蜜蜂的传粉过程中, 芳香石豆兰铰链结构的唇瓣起着重要作用。人工授粉试验表明, 芳香石豆兰自交授粉不结实, 异交授粉结实率达90%以上, 芳香石豆兰为自交不亲和的繁育系统。芳香石豆兰具有较强的克隆生长能力, 同一个体花期内有大量的花同时开放, 其传粉者中华蜜蜂平均单次访花(4.29 ± 0.40)朵( n= 66), 造成了严重的同株异花传粉。芳香石豆兰附生于石灰岩山顶岩石或树干上, 种群密度较低, 使得不同个体之间花粉交流困难, 自交不亲和的特性导致了芳香石豆兰自然不结实。本研究结果为开展芳香石豆兰的有效保护提供了依据。     

The pollination mechanism in the 'Pelexia alliance' (Orchidaceae: Spiranthinae)

The biological function of the wedge–like, dorsally adhesive viscidium of the neotropical orchid genera Cyclopogon, Pelexia and Sarcoglottis (grouped in the so-called 'Pelexia alliance') is elucidated by the study of the pollination biology of three species occurring in Sao Paulo State, southeastern Brazil. Cyclopogon congestus is pollinated by the bee, Pseudoaugochloropsis graminea (Halictidae), Pelexia oestrifera by workers of Bombus (Fervidobombus) atratus (Apidae) and Sarcoglottis fasciculata by males and females of Euglossa cordata (Apidae: Euglossini). These three species offer nectar as a reward and are self-compatible, though they need pollinators to set fruits. In spite of the difference in flower sizes and in their pollinators' taxonomic groups, the pollination mechanism is essentially the same for these species. The pollinarium adheres to the ventral surface of the bee labrum. The viscidium needs to be dorsally pressed in order to liberate a glue which fixes the pollinarium to a bee. Pollination is achieved by the interaction of the orchid column and the mouthparts of the bees. Fivation to the ventral surface of the labrum is advantageous for the orchid, since it is a difficult place for the bees to clean. Another advantage is that, since the labrum is articulated, when the bees fold and close their mouthparts, the pollinarium remains protected under the bee's head, thus reducing the risks of pollen loss. Since the wedge-like, dorsally adhesive viscidiurn is a characteristic feature of Cyclopogon, Pelexia and Sarcoglottis , it is suggested that some kind of phylogenetic constraint may exist, impeding the occurrence of pollinators other than bees in these orchid genera. All other flower-visiting animals lack the labrum-like structure needed to fix the pollinarium.     

Bird pollination in South African species ofSatyrium (Orchidaceae)

Field observations showed that three South African orchid species,Satyrium carneum, S. coriifolium andS. princeps, are pollinated by sunbirds. Foraging sunbirds insert their bills into the labellum chamber of the flowers and suck nectar from the labellum spurs with their tongues. The column overarches the entrance to the labellum and pollinaria become affixed to the upper mandible of the bill. Birds often attempt to remove pollinaria by rubbing their bills against a branch, but are mostly unsuccessful due to the large plate-like viscidia which attach the pollinaria very firmly to the bill. Other modifications for bird pollination in theseSatyrium species may include the red, carmine or orange colour of the flowers, sturdy inflorescence stems used for perching and large amounts of dilute nectar in the spurs. EachSatyrium species was pollinated by several species of sunbird, and despite some differences in flowering time, occasional hybrids occur at a site whereS. carneum andS. coriifolium share the same sunbird pollinators.     

Gynostemium micromorphology and pollination in Epipactis microphylla (Orchidaceae)

Self-pollination conditions have been recorded frequently in local populations of the genus Epipactis, and structural modifications have sometimes been reported, supporting taxonomic recognition of new species. As part of a survey of gynostemium micromorphology of Italian Epipactis populations, we studied an Apennine population of Epipactis microphylla (Ehrh.) Sw. a species listed as autogamous but with residual allogamous characters. In this population we observed the tendency to perform pre-anthesis cleistogamy. However, self-pollination was not a consequence of modification in column morphology. Cryptic pollen germination occurred in anther thecae or in the clinandrium, bypassing any contact with the receptive stigma. Pollen germination started from the distal pool of the pollinium, close to the rostellum-viscidium, suggesting that the rostellum-viscidium may have a function in promoting pollen-tube growth. Germinated pollen was exposed at the anthesis.     

Listera fugongensis (Orchidaceae), a new species from Yunnan, China

Listera fugongensis, a new species from Yunnan, China, is described and illustrated. The new species is probably related to Listera nipponica, but differs from it by having green flowers, petals slightly wider than lateral sepals, and a toothed lip with ciliate margins.     

The pollinium ofLoroglossum hircinum (Orchidaceae) between pollination and pollen tube emission

The structure of the massulae composing the pollinium ofLoroglossum hircinum was studied before pollination and 12 and 24 hours afterwards. The grains are grouped in tetrads closely packed in massulae. The exine is only present on the outside of the massulae. The intine consists of two layers: a compact layer surrounding the pollen grain and a looser layer surrounding the pollen grain and a looser layer surrounding the tetrad. Twelve hours after pollination, pollen volume and the space between the tetrads increase due to vacuolization. Twenty-four hours after pollination, pollen volume and tetrad spacing are higher due to vacuolization and some grains have emitted pollen tubes. Pollen growth due to vacuole formation, and the absence of common walls between adjacent tetrads lead to crumbling of the massulae. The mature pollen grain does not have apertures: the site of pollen tube emission is determined after pollination. The first grains to germinate are those in the centre of the massula. The vegetative cell nucleus is the first to enter the pollen tube; the generative cell elongates and undergoes the second haploid mitosis shortly after entering the pollen tube.     

Deceptive pollination of the Lady's Slipper Cypripedium tibeticum (Orchidaceae)

To test whether the nectarless flowers of Cypripedium tibeticum attract pollinators through mimicry like the allied species C. macranthos var. rebunense, pollination biology of C. tibeticum was investigated in western China. Although C. tibeticum was also pollinated by bumble bee queens, i.e. Bombus lepidus , B. lucorum and B. hypnorum , no special, rewarding model plants were found in the habitat. Field experimentation confirmed that the flowers were self-compatible but insects were required to transfer orchid pollen to the stigma. Both Bombus queens and workers were visitors, but queens were much more frequent than workers and only queens were effective pollinators. Floral functional morphology analysis showed that it was large queens rather than small workers that fitted well with the flowers of C. tibeticum. With the faint sweet-fruity scent, the minor floral fragrance compound, ethyl acetate, probably plays a role in attracting bumble bees by food deception. The dark flowers with the inflated, trap-like labellum are hypothesized to mimic the nest site of queens. Therefore, bumble bee queens tend to be attracted by C. tibeticum through nest site mimic combined with food deception. Considering that the co-blooming flowers of C. flavum are pollinated by the Bombus workers, and C. smithii pollinated by a queen, we suggest that using the same bumblebees with different body sizes as the pollinators is the main reproductive isolation between interfertile C. tibeticum and C. flavum, while C. tibeticum and C. smithii tend to hybridize naturally.     

Evolutionary disruption in the pollination system of Vanilla (Orchidaceae)

• Pollination of the pantropical Vanilla has been linked to melittophily and food deception.
• Here we investigated the role of flower traits on the reproduction of Neotropical Vanilla. We also studied the evolution of pollination systems in order to understand the origin of production of flower resources and the diversification of pollinators in this orchid genus. Our study was founded on data of adaptations in flower morphology, production of resources, scent release, pollinators and breeding systems of Vanilla and presenting new data on reproductive biology of V. palmarum. Data on reproductive biology of Vanilla were mapped onto a phylogeny to address our queries on the evolution of pollination systems in this genus.
• Vanilla palmarum shows a mixed mating system, with its facultative autogamous flowers being pollinated by hummingbirds. Its yellow flowers are scentless and produces nectar. Mapping of the pollination system onto trees resulted in one origin for bird pollination and at least two origins for autogamy in Vanilla. Nectar secretion has a single origin in the Neotropical thick-leafed lineage.
• Bird pollination of Vanilla is shown for the first time. The origin of ornithophily within a bee-pollinated clade is supported by flower morphology. Floral transitions to ornithophily have been favoured by the occupation of a distinct niche from that of the other thick-leafed Vanilla species. Despite its specialized pollination, V. palmarum is autogamous. A mixed mating system can promote reproductive assurance in the case of a decline in pollinator populations, or in areas where pollinator services are irregular or absent.

     

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.     

Evolution,pollination, and systematics of the tribeNeottieae (Orchidaceae)

The various classifications of the orchid tribeNeottieae are reviewed and a new classification is proposed that divides the tribe into three subtribes,Neottiinae, Limodorinae, andCephalantherinae, based primarily on characters of the column (gynostemium). A cladistic analysis illustrates that these three subtribes are more closely related to one another than either is to any other group in subfam.Neottioideae, although there are very few apomorphic characters for the tribe. Pollination biology is also discussed showing links between breeding systems and distribution. There is also a possible role between column and labellum morphology and the emergence of a deceptive pollination syndrome from one of reward.     

足茎毛兰的欺骗性传粉研究

一些兰科植物常常利用多种多样的欺骗性传粉机制吸引传粉者,唇瓣上的附属物就是其中之一.黄色被认为对昆虫具有吸引作用.在广西雅长兰科植物自治区级保护区内秋季开花的足茎毛兰(Eria coronaria)唇瓣上具有鲜明的黄色斑块,这种花部信号很可能与吸引昆虫有关系.为验证这一假设,我们在广西雅长保护区内对足茎毛兰的传粉过程进行了观察.研究发现,足茎毛兰的唯一传粉者是中华蜜蜂(Apis cerana cerana).与足茎毛兰同在秋季开花的植物主要有光叶海桐(pittosporum glabratum).光叶海桐的花中有丰富的花蜜和花粉,吸引大量中华蜜蜂访问.足茎毛兰虽然不为中华蜜蜂提供任何报酬,但其唇瓣上的黄色斑块的颜色和形态大小与光叶海桐的黄色花相似.根据中华蜜蜂在足茎毛兰和光叶海桐花上的活动情况,我们认为足茎毛兰唇瓣上鲜明的黄色斑块对中华蜜蜂有吸引作用.中华蜜蜂访花时,通常降落在唇瓣的黄色斑块上,调整身体姿势后进入花内,在退出的过程中将花粉块带出或将所携带的花粉块授到柱头上,而药帽留在原来的位置.足茎毛兰的花部构造,特别是药帽的形状与中华蜜蜂的形态以及在花上的行为十分吻合,因此中华蜜蜂的传粉效率较高.人工授粉实验表明足茎毛兰需要依赖传粉者才能完成传粉过程.足茎毛兰在研究样地的自然结实率为20.72%,接近于食源欺骗性传粉兰科植物的平均结实率(20.7%).     

Beetle pollination of Luisia teres (Orchidaceae) and implications of a geographic divergence in the pollination system

The epiphytic orchid Luisia teres on Okinawa Island, the Ryukyus, Japan, sexually attracts male Protaetia beetles for pollination through floral scent. Here, the pollination system of L. teres on the Satsuma Peninsula, Kyushu, located approximately 600 km north of Okinawa Island, is reported. The inconspicuous flowers depend on the Cetoniinae beetles Protaetia orientalis submarumorea and Pseudotorynorrhina japonica, which both usually feed on fermented sap/ripe fruits, for pollination. The flower-visiting P. orientalis submarumorea beetles included both males and females, and pseudocopulation was never observed. Pollinaria removal and pollination were achieved while beetles fed on floral nectar. Although P. orientalis submarumorea did not necessarily remove pollinaria more frequently than Ps. japonica, the pollinaria attached precisely to the central part of their clypeus, unlike on Ps. japonica. Our findings suggest that a geographic divergence in the pollination system occurs in Japanese L. teres. In the Ryukyus, the orchid adopts a specialized pollination system with only male-attractive olfactory stimuli, whereas in Kyushu it has a relatively more generalized system based on only a food reward.     

兔耳兰食源性欺骗传粉的研究

兰科植物具有精巧、多样化的花部结构以及高度多样的吸引传粉者方式。作者对广西雅长兰科植物自治区级保护区内的一个兔耳兰(Cymbidium lancifolium)居群进行了连续2年的观察和研究。观察发现兔耳兰唯一的传粉者为膜翅目蜜蜂科的中华蜜蜂(Apis cerana cerana)。中华蜜蜂一般直接落在唇瓣外弯的中裂片上, 然后调整身体的方向, 进入花中, 当发现花中无蜜液等回报时, 借助于后足的蹬力退出花朵。在退出的过程中, 花粉块连同药帽会通过粘盘粘附在中华蜜蜂的胸部。中华蜜蜂在花内的停留时间为8–71 s, 平均18.3 s (N = 11)。根据观察我们推测兔耳兰可能是通过其唇瓣上无规则的紫栗色小斑点(假蜜导)来吸引中华蜜蜂为其传粉, 属于食源性欺骗方式。在传粉过程中兔耳兰的药帽与花粉团和粘盘一起粘在中华蜜蜂背部。药帽的存在能够阻止下一朵被拜访的花实现雌性功能。兔耳兰药帽高度(0.154 ± 0.032 cm) (N = 10)加上传粉昆虫胸高(2005年为0.37 ± 0.03 cm (N = 10), 2006年0.35 ± 0.04 cm (N = 7))大于传粉通道入口的高度(0.29 ± 0.04 cm) (N = 21), 支持兔耳兰可能通过药帽来减少同株异花授粉现象的推测。2005和2006年该兔耳兰居群的自然繁殖成功率分别为21.13%和21.28%。繁育系统实验证明兔耳兰是高度自交亲和物种, 自交和异交的繁殖成功率没有显著性差异, 表明该种在结实过程中未显示近交衰退。兔耳兰不存在无融合生殖和自花授粉的现象, 其结实依赖传粉者。TTC法检测结果显示兔耳兰种子活力达85.78%(N = 11), 可见种子活力不是制约兔耳兰种子萌发的主要原因。因此传粉者的密度和访问频率可能是影响兔耳兰结实的重要因素, 并最终影响兔耳兰种群的维持和扩张。     

The pollination of Habenaria rhodocheila (Orchidaceae) in South China: When butterflies take sides

Habenaria is one of the largest terrestrial genera in the family Orchidaceae. Most field studies on Habenaria species with greenish–white and nocturnal scented flowers are pollinated by nocturnal hawkmoths and settling moths. However, H. rhodocheila presents reddish flowers lacking a detectable scent and fails to fit the moth pollination syndrome. We investigated the pollinators, breeding system, and functional traits of H. rhodocheila in South China and found that two diurnal swallowtail butterflies Papilio helenus and Papilio nephelus (Papilionidae) were the effective pollinators. When butterflies foraged for nectar in the spur, the pollinia became attached between the palpi. A triangular projected median rostellar lobe was found at the entrance (sinus) of the spur of H. rhodocheila. This lobe divided the spur opening into two entrances forcing butterflies to enter their proboscides through the left or right side. When the projection of median rostellar lobe was removed, the site of pollinium attachment changed to the eyes of the butterflies, leading to a higher rate of pollinium removal but lower rate of pollinium deposition. Our quartz glass cylinder choice experiment suggested that visual rather than olfactory cues provided the major stimuli for butterflies to locate these flowers. Hand pollination experiments suggested this species was self‐compatible but pollinator‐dependent. However, the proportion of seeds with large embryos produced in self‐pollinated fruits was significantly lower than in cross‐pollinated fruits, indicating a significant inbreeding depression. Unlike many other orchid species, fruit set was higher than rates of pollinium removal, indicating a high level of pollination efficiency in a species with friable pollinia. Shifts from moth to butterfly pollination in the genus Habenaria parallel other orchid lineages providing insights into the potential for pollinator‐mediated floral trait selection.