Reproductive Biology of the Distylous Manettia luteo-rubra (Rubiaceae)

The reproductive biology of Manettia luteo-rubra was studied in the coastal montane Atlantic rain forest of southeastern Brazil. This Rubiaceae is a perennial vine that flowers all the year round, but has a flowering peak during the dry season. It presents reciprocal herkogamy, thus the plants are morphologically distylous. The morphs occur in a 1:1 ratio, and pollen diameter and corolla length vary between short and long-styled flowers. Manettia luteo-rubra displays typical heterostylous self-incompatibility and sets almost no fruits from self- or intramorph pollinations. The flowers are tubular, red with yellow lobes, and odourless. Anthesis is asynchronous, and the flowers last about four days. Concentration of sugars in nectar is similar in both morphs, ca. 24%, this concentration being typical for hummingbird flowers. Three species of hummingbirds are the major pollinators of the flowers of M. luteo-rubra at the study site: the hermits Phaethornis eurynome and P. squalidus, and the trochiline Thalurania glaucopis. Three species of Heliconius butterflies act as minor pollinators. Both morphs of M. luteo-rubra exhibit natural fruit-set of about 80%, this reproductive output being maintained throughout the year by the pollinators' constancy to the flowers.     

The missing stink: sulphur compounds can mediate a shift between fly and wasp pollination systems

The radiation of the angiosperms is often attributed to repeated evolutionary shifts between different pollinators, as this process drives diversification of floral forms and can lead to reproductive isolation. Floral scent is an important functional trait in many pollination systems but has seldom been implicated as a key mechanism in pollinator transitions. In this study, we suggest a role for sulphur compounds in mediating a shift between specialized carrion-fly and pompilid-wasp pollination systems in Eucomis (Hyacinthaceae). Flowers of closely related Eucomis species pollinated by carrion flies or pompilid wasps have very similar greenish-white flowers, but differ markedly in floral scent chemistry (determined by GC–MS analysis of headspace extracts). Comparison of the floral colours of the four Eucomis species in the visual systems of flies and wasps suggests that colour plays little role in pollinator discrimination. Nectar properties and morphology also do not differ strongly between fly- and wasp-pollinated flowers. By comparing floral scent bouquets and experimentally manipulating the scent of plants in the field, we demonstrate that shifts between wasp and fly pollination in these four congeners can depend on the production or suppression of sulphur compounds (dimethyl disulphide and dimethyl trisulphide) in the fragrance bouquet. This suggests that mutations affecting the production of particular scent compounds could precipitate shifts between pollinators, independently of floral morphology, colour or nectar properties.     

茜草科二型花柱植物滇丁香传粉精确性研究

异型花柱是一种受遗传因素控制的花型多态性现象,其适应意义在于提高不同花型间传粉精确性,从而促进异交。为检验二型花柱植物滇丁香（Luculia pinceana）是如何促进异交并确定有效传粉者的类型,实验统计自然种群中长、短柱型的植株数量,测量花部特征,统计自然种群植株柱头上的型间花粉落置的数量和比例,比较型内、型间花粉的萌发率以及花粉管长度,比较不同人工授粉处理结实情况,观察传粉昆虫类型并测量虫体特征。结果表明（1）滇丁香自然居群中长、短柱型植株的数量没有显著性差异。长柱型的筒深、筒直径、开口直径、雄蕊长和柱头长均显著小于短柱型,而长柱型的叶片长和宽、雌蕊长、花药体积、花粉粒体积、柱头厚均显著大于短柱型。（2）自然种群植株柱头上型间花粉所占比例显著低于型内花粉的比例。（3）型间授粉的花粉萌发率以及花粉管长度均大于型内授粉和混合授粉。（4）型间授粉的结实率显著高于型内授粉的结实率,型内授粉具有一定的结实。（5）夜行性长喙条背天蛾（Cechenena lineosa）是滇丁香的有效传粉者,其吻长和滇丁香的花筒深相适应。本研究表明二型花柱植物滇丁香主要是通过与其花部特征相匹配的长喙天蛾实现有效的传粉,通过促进型间花粉的萌发和花粉管的伸长而有利于型间授粉结实,提高传粉精确性。     

Dioecy and wind pollination in Pernettya rigida (Ericaceae) of the Juan Fernández Islands

Pernetttya rigida is endemic to the Juan Fernández archipelago. Although all flowers are complete, with seemingly fertile stamens and pistils, differences in fruit production and detailed field, anatomical, and morphological studies indicate they are functionally unisexual, and the species is consequently dioecious. A comparison of 15 features demonstrated sigdicant differences between the sexes. The populational sex ratio is 1:1. Nectaries located between the filaments produce small amounts of floral nectar with similar sugar composition in both sexes. There are =33,357 (or = 133,429) pollen tetrads (or pollen grains)/male flower and =109 ovules/female flower. No tetrads in either hand self-pollinated or open pollinated male flowers showed any pollen germination. Tetrads on open- and hand-pollinated female flowers germinate. Female flowers do not show autogamy or apomixis. During more than 80 hours of field observation, we recorded only seven floral visitors (representing three insect species). In spite of this, openpollinated female flowers have abundant fruit and seed-set. Thus, we conclude that pollen is transferred abiotically and the ever-present wind over the exposed ridges of the islands is the likely dispersal agent. A number of anemophilous features, such as dry pollen and exposed habitat, support this conclusion. Thus, dioecy and anemophily have evolved independently, in situ , in this species in this remote locality. Preservation of habitat and elimination of competitive invasives are the primary conservation challenges.     

A preliminary study on pollination biology of Omphalogramma souliei Franch. (Primulaceae), a species endemic to China

The pollination biology of Omphalogramma souliei Franch., a species endemic to southwestern China, was investigated. Floral phenology, flower visitors, pollen/ovule ratio, attractants and rewards to the visitors were observed, measured, and recorded. Bagging experiments to exclude pollinators were carried out in the wild for two years. Our results revealed five important aspects of the reproductive biology of Omphalogramma souliei. 1) The pollen-ovule ratio was 1748±233. The breeding system was self-compatible, with facultative xenogamy. 2) The pollination syndrome is entomophily, and this species could not be pollinated by wind if the pollinators were unavailable. 3) Six insect species were observed visiting the flowers of Omphalogramma souliei in the wild, of them, three species of hymenoptera, Lasioglossum sp., Heriades parvula Cockerell and Micrapis florae Fabricius, are the principal floral visitors and effective pollinators. 4) The visual attractants to the visitors are floral color and shape, the large yellow anthers, and the rewards for visitors are pollen and nectar. 5) Cleistogamy may also occur, since the anthers of some flowers dehisced before opening of the corolla. The results of floral biology and pollination characters suggest that xenogamy predominated and autogamy played an assistant role in the evolution of reproduction and breeding system of O. souliei.     

Pollinator activity and pollination success of Medicago sativa L. in a natural and a managed population

Medicago sativa L. is an important cash crop in the arid region of northwest China. Pollinator activity is an essential aspect of pollination success, but the relationships between pollinator visitation rate and seed set still need further study of M. sativa. We investigated the following characteristics of M. sativa in natural and managed populations: floral traits, pollinator activity, and breeding system. Our results indicated the management could affect the number of flowers produced; however, there was no detectable effect on the seed set per flower. We found the percentage of seeds among pollinated flowers in the managed population was significantly higher than that in the natural population. Moreover, the increase in the proportion of pollinated flowers could significantly increase seed set per flower, and pollinator visitation rate was the important limiting factor for seed set in both populations. Andrena lebedevi Popov was found to be the most frequent pollinator in both populations. Outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. Our study suggested that proper management (artificial selection) could promote pollination success of M. sativa.     

深山含笑传粉生物学研究

为明确深山含笑(Michelia maudiae)的传粉生物学特性,该文以西华师范大学校园内的深山含笑为研究材料,采取野外观察法记录了深山含笑的开花动态、访花者及访花频率;用游标卡尺法,测定了深山含笑的异交指数(OCI);用醋酸洋红法,测定了花粉胚珠比(P/O);用套袋和人工授粉法,测定了深山含笑的繁育系统。结果表明:(1)深山含笑为早春开花植物,其种群花期为2~3月,持续30d左右,单花花期5~6d;在单花开放初期,最内层的花瓣未绽开,是适应潮湿或多雨的环境而保护花药与柱头免受雨水的冲刷,提高雌雄性适合度,确保繁殖成功的一种策略;(2)深山含笑的有效传粉者为蜜蜂,访花时间集中在晴朗天气的中午;(3)深山含笑的异交指数(OCI)等于5,花粉胚珠比(P/O)为2 933±50;(4)深山含笑为雄蕊先熟,柱头可授性在开花第2~3天达到最高;(5)套袋和人工授粉表明深山含笑为兼性自交和异交授粉,其中异交授粉的座果率和结籽率显著高于自交授粉。早春开花的深山含笑,其传粉者为单一的蜜蜂,繁育系统为异交,部分自交亲和,结果为引种栽培、良种选育等提供依据更好地利用和保护此资源奠定基础。     

Reproductive biology and pollination of Govenia utriculata: A syrphid fly orchid pollinated through a pollen-deceptive mechanism

The reproductive biology and pollination mechanisms of Govenia utriculata (Sw.) Lindl. were studied in a mesophytic semideciduous forest at Serra do Japi, south-eastern Brazil. The floral visitors and pollination mechanisms were recorded, and experimental pollinations were carried out to determine the breeding system of this species. Populations of G. utriculata growing at Serra do Japi are exclusively visited and pollinated by two species of hoverflies in the genus Salpingogaster (Diptera: Syrphidae) that are attracted by deceit to the flowers of this orchid species. The lip apex and the column base present small brownish and yellow to orange spots that mimic pollen clusters. Govenia utriculata is self-compatible, but pollinator dependent. Natural fruit set was low (10%), but similar to that of other non-obligatorily autogamous sympatric orchid species that occur at Serra do Japi and of other fly-pollinated orchid species pollinated through deceptive mechanisms.     

Floral scent composition predicts bee pollination system in five butterfly bush (Buddleja,Scrophulariaceae) species

Traditionally, plant–pollinator interactions have been interpreted as pollination syndrome. However, the validity of pollination syndrome has been widely doubted in modern studies of pollination ecology. The pollination ecology of five Asian Buddleja species, B. asiatica, B. crispa, B. forrestii, B. macrostachya and B. myriantha, in the Sino‐Himalayan region in Asia, flowering in different local seasons, with scented inflorescences were investigated during 2011 and 2012. These five species exhibited diverse floral traits, with narrow and long corolla tubes and concealed nectar. According to their floral morphology, larger bees and Lepidoptera were expected to be the major pollinators. However, field observations showed that only larger bees (honeybee/bumblebee) were the primary pollinators, ranging from 77.95% to 97.90% of total visits. In this study, floral scents of each species were also analysed using coupled gas chromatography and mass spectrometry (GC‐MS). Although the five Buddleja species emitted differentiated floral scent compositions, our results showed that floral scents of the five species are dominated by substances that can serve as attractive signals to bees, including species‐specific scent compounds and principal compounds with larger relative amounts. This suggests that floral scent compositions are closely associated with the principal pollinator assemblages in these five species. Therefore, we conclude that floral scent compositions rather than floral morphology traits should be used to interpret plant–pollinator interactions in these Asian Buddleja species.     

Pollination biology of Harrisia portoricensis (Cactaceae), an endangered Caribbean species

Traits associated with self-pollination are common features of island plant communities. In this work, we studied the pollination biology and the breeding system of Harrisia portoricensis, an island columnar cactus, to test for the presence of inbreeding and inbreeding depression. For H. portoricensis, which bears flowers with typical outcrossing morphology, the results from 322 h of direct observations and videotaping showed that visits to flowers by animals were uncommon. Controlled pollinations demonstrated that H. portoricensis has a partially self-compatible breeding system that it is not autogamous and thus requires an external mechanism for the movement of pollen to set fruit. We detected differences in seed size, seed mass, germination success, and multiplicative fitness estimates between self- and cross-pollination treatments. We found that progeny resulting from natural and self-pollination treatments showed signs of inbreeding depression compared with progeny resulting from cross-pollination; however, the magnitude of the inbreeding depression was less than 50%. Our combined results suggest that for this species an endogamous breeding system should be favored by natural selection.     

Highly efficient pollination by bumblebees ensures seed production in Pedicularis lachnoglossa (Orobanchaceae), an early‐flowering Himalayan plant

Abstract Pedicularis (Orobranchaceae) is a common high altitude genus of the Himalayas that may be affected by pollination limitation. Using Pedicularis lachnoglossa from Yulong (Jade Dragon) Snow Mountain in Lijiang (Yunnan Province, southwest China), we investigated the effects of high altitude habitats on the process of pollination and seed production. Floral biology, pollinator foraging behavior, breeding system, and pollination efficiency were examined using observation and exclusionary techniques. Pedicularis lachnoglossa was found to be entomophilous and exclusively pollinated by Bombus friseanus and B. yunnanicola. Our results indicated that pollination limitation in P. lachnoglossa was not significant. Under open pollination, approximately 80% of flowers were successfully pollinated and developed to fruits, and about 38% of ovules developed to mature seeds. Bumblebee pollination is highly precise and efficient in P. lachnoglossa, because its flowering phenology and floral characters enhance the foraging of bumblebees on flowers. This study supports that animal pollination plays a crucial role in the outbreeding of the early flowering Pedicularis. The evolution of floral specification in Pedicularis has the advantages of adaptation to bumblebee pollination in adverse high altitude habitats.     

Spatio‐temporal patterns in pollination of deceptive Aristolochia rotunda L. (Aristolochiaceae)

• Pollination success of highly specialised flowers is susceptible to fluctuations of the pollinator fauna. Mediterranean Aristolochia rotunda has deceptive trap flowers exhibiting a highly specialised pollination system. The sole pollinators are kleptoparasitic flies in search of food. This study investigates these pollinators on a spatio‐temporal scale and the impact of weather conditions on their availability. Two potential strategies of the plants to cope with pollinator limitation, i.e. autonomous selfing and an increased floral life span, were tested.
• A total of 6156 flowers were investigated for entrapped pollinators in 10 Croatian populations. Availability of the main pollinator was correlated to meteorological data. Artificial pollination experiments were conducted and the floral life span was recorded in two populations according to pollinator availability.
• Trachysiphonella ruficeps (Chloropidae) was identified as dominant pollinator, along with less abundant species of Chloropidae, Ceratopogonidae and Milichiidae. Pollinator compositions varied among populations. Weather conditions 15–30 days before pollination had a significant effect on availability of the main pollinator. Flowers were not autonomously selfing, and the floral life span exhibited considerable plasticity depending on pollinator availability.
• A. rotunda flowers rely on insect pollen vectors. Plants are specialised on a guild of kleptoparasitic flies, rather than on a single species. Pollinator variability may result in differing selection pressures among populations. The availability/abundance of pollinators depends on weather conditions during their larval development. Flowers show a prolonged trapping flower stage that likely increases outcrossing success during periods of pollinator limitation.

     

Reproductive biology of Azadirachta indica (Meliaceae), a medicinal tree species from arid zones

Neem or the Margosa tree, Azadirachta indica (Meliaceae), is a versatile tree species common in arid zones. We investigated the reproductive biology of Neem, including phenology, floral biology, pollination ecology and the breeding system, in three natural populations in northern India. Phenological observations established that trees within a population flowered synchronously, but populations varied in the time of onset of flowering; a few trees flowered twice a year. An inflorescence bears ～ 91 bisexual, white and sweet‐scented flowers. The stigma consists of a non‐receptive apical region and a subjacent rim of receptive surface; papillae on the two regions differ in their morphology and function. The occurrence of natural pollination both by wind and insects indicates ambophily. Insect pollinators were predominantly represented by Hymenoptera and Lepidoptera; Apis spp. were the most effective pollinators. Although the occurrence of incomplete dichogamy and a pollen : ovule ratio of ～ 627 indicates the possibility of autogamy, supplemental pollinations clearly established that the trees were 100% self‐incompatible and incompatibility is regulated at the gametophytic level. Natural fruit set was low (～ 5%) and hand cross‐pollinations increased the fruit set to ～ 19%. Our study demonstrated that natural fruit set in Neem is possibly limited by insufficient amounts of xenogamous pollination and resources.     

Specialized use of pollen vectors by Caesalpinia gilliesii, a legume species with brush-type flowers

The pollination of Caesalpinia gilliesii (Fabaceae) , a legume species with long, brush-type flowers, was studied in wild populations in Argentina. A total of 341 hawkmoths (Sphingidae), belonging to 10 species, with tongues ranging from 1.5 to 13.5 cm in length, were captured by means of light traps. Hawkmoths could be classified in four significantly different tongue-length classes or guilds: short (15.35 ± 1.11 mm), medium (30.23 ± 2.24 mm), long (55.85 ± 4.98 mm) and very long (95.61 ± 13.63 mm). A total of 133 of the hawkmoths, belonging to seven species, carried pollen of Cae. gilliesii , mainly on their venter. Successful contact with anthers while drinking nectar depended upon hawkmoth tongue length. The relative frequency of pollen-carriage and the mean tongue length of each species were positively and significantly correlated. Considering a null hypothesis of nonspecialization, in which hawkmoths carrying pollen are distributed at random among individuals of the four guilds, long-tongued hawkmoths carried Cae. gilliesii pollen more frequently than would be expected by chance. In contrast, short- and medium-tongued guilds carried pollen less frequently than is predicted by random expectation. Individuals with short tongues accessed nectar from below without touching the anthers, whereas very long-tongued hawkmoths, with tongues longer than the length range of the flower reproductive organs, can probably reach nectar without touching the fertile organs. Results show that a pterotribic pattern of pollen deposition on the body of long-tongued hawkmoths is operating in this species. This suggests that brush-type blossoms are not necessarily unspecialized, despite popular belief.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 579–592.     

沙丘稀有种准噶尔无叶豆花部综合特征与传粉适应性

植物花部特征进化与传粉适应性一直是进化生态学领域关注的核心问题之一。以古尔班通古特沙漠自然生长的准噶尔无叶豆为对象,对其花部特征和传粉特性进行了野外观察和室内的分析研究。结果表明:种群花期历时21 d,花序花期历时7-12 d,单花花期一般3 d,若遇阴雨天气,花期可延长1-2 d,整个花期龙骨瓣一直保持闭合状态。单花10枚花药在旗瓣微张时已全部完成散粉。准噶尔无叶豆主要靠分泌花蜜、鲜艳的花色以及旗瓣基部的黄色辐射状纹理结构吸引传粉者。准噶尔无叶豆花期的有效传粉者为4种蜂类昆虫,它们的平均访花频率为(7.75±0.57)次·花^-1·d^-1,访花高峰期表现为三峰型: 13:00-14:00,16:00-17:00和19:00-20:00。准噶尔无叶豆人工套袋实验表明该种为自交亲和型,主动自交少见,生殖成功依赖传粉者。胚珠成功受精至果实完全成熟阶段存在自交衰退,柱头角质层结构和花粉刷结构是准噶尔无叶豆在进化过程中形成的减少自交,倾向异交的机制。     

姜科、闭鞘姜科植物繁育系统与传粉生物学的研究进展

综述了姜科Zingiberaceae植物繁育系统和传粉生物学方面的研究及其进展。姜科有50余属1500多种,是一个泛热带分布的、动物传粉的单子叶植物大科。姜科植物中存在着雌花两性花异株、雄花两性花同株、花柱卷曲性、雄性先熟、自交不亲和等多种性表达方式和花部机制。姜科植物重要的传粉动物包括各种蜂类、天蛾、蝴蝶、鸟类等,不同的传粉动物对应不同的花部特征。在相对有限的研究中,姜科植物展现出了一些独特的传粉和繁育机制,在豆蔻属Amomum、山姜属Alpinia等植物中发现的花柱卷曲性被认为是植物界中一种独特的促进异交的行为机制,在对黄花大苞姜Caulokaempferia coenobialis的研究中发现了植物界中一种全新的花粉滑动自花传粉机制,这些新的研究和发现丰富了我们对姜科植物传粉和繁育系统多样性的认识。本文提出了今后姜科植物繁殖生物学研究的建议:对更多未知姜科植物类群的传粉和繁育系统进行研究;从系统发育的角度开展姜科不同类群植物的传粉和繁育系统的演化研究;对花柱卷曲性这一独特的性多态现象开展全面深入的研究。     

Reproductive biology of two Himalayan alpine gingers (Roscoea spp., Zingiberaceae) in China: pollination syndrome and compensatory floral mechanisms

According to the concept of pollination syndromes, floral traits reflect specialisation to a particular pollinator or set of pollinators. However, the reproductive biology of endemic, and often specialised, plants may require increased attention as climate change accelerates worldwide. Species of Roscoea endemic to the Himalayan region have striking orchid-like flowers with long corolla tubes, suggesting pollination by long-tongued insects. Until now, the reproductive biology of species of Roscoea has been poorly documented. We investigated the floral biology, breeding system and pollination ecology of R. cautleoides and R. humeana, from Hengduan Mountains, a global biodiversity hotspot in southwest China. We also tested whether floral longevity increases pollination success. Pollination experiments showed that the two species were self-compatible and depended on insects for fruit production. Over several flowering seasons we did not observe any potential pollinators with long tongues that matched the corolla tube visiting flowers in centres of distribution. The principal pollinators observed were pollen-collecting generalist bees, with low visitation frequencies. In general, members of the ginger family are characterised by short-lived (usually 1 day) flowers, but flowers of R. cautleoides and R. humeana last 8 and 6 days, respectively. Removing stigmas decreased fruit set in both study populations. Our results suggest that the original pollinators may have been long-tongued insects that are now absent from the Chinese Himalayas because habitats have responded to climate change. However, long-lived and self-compatible flowers, coupled with the presence of generalist pollinators, are traits that have allowed these gingers to reproduce and continue to persist in the alpine habitats.