Division of labour within flowers: heteranthery,a floral strategy to reconcile contrasting pollen fates

In many nectarless flowering plants, pollen serves as both the carrier of male gametes and as food for pollinators. This can generate an evolutionary conflict if the use of pollen as food by pollinators reduces the number of gametes available for cross‐fertilization. Heteranthery, the production of two or more stamen types by individual flowers reduces this conflict by allowing different stamens to specialize in ‘pollinating’ and ‘feeding’ functions. We used experimental studies of Solanum rostratum (Solanaceae) and theoretical models to investigate this ‘division of labour’ hypothesis. Flight cage experiments with pollinating bumble bees (Bombus impatiens) demonstrated that although feeding anthers are preferentially manipulated by bees, pollinating anthers export more pollen to other flowers. Evolutionary stability analysis of a model of pollination by pollen consumers indicated that heteranthery evolves when bees consume more pollen than should optimally be exchanged for visitation services, particularly when pollinators adjust their visitation according to the amount of pollen collected.     

Morphological specialization of heterantherous Rhynchanthera grandiflora (Melastomataceae) accommodates pollinator diversity

• The tropical Melastomataceae are characterized by poricidal anthers which constitute a floral filter selecting for buzz‐pollinating bees. Stamens are often dimorphic, sometimes with discernible feeding and pollinating functions. Rhynchanthera grandiflora produces nectarless flowers with four short stamens and one long stamen; all anthers feature a narrow elongation with an upwards facing pore.
• We tested pollen transfer by diverse foraging bees and viability of pollen from both stamen types. The impact of anther morphology on pollen release direction and scattering angle was studied to determine the plant's reproductive strategy.
• Medium‐sized to large bees sonicated flowers in a specific position, and the probability of pollen transfer correlated with bee size even among these legitimate visitors. Small bees acted as pollen thieves or robbers. Anther rostrum and pore morphology serve to direct and focus the pollen jet released by floral sonication towards the pollinator's body. Resulting from the ventral and dorsal positioning of the short and long stamens, respectively, the pollinator's body was widely covered with pollen. This improves the plant's chances of outcrossing, irrespective of which bee body part contacts the stigma. Consequently, R. grandiflora is also able to employ bee species of various sizes as pollen vectors.
• The strategy of spreading pollen all over the pollinator's body is rather cost‐intensive but counterbalanced by ensuring that most of the released pollen is in fact transferred to the bee. Thus, flowers of R. grandiflora illustrate how specialized morphology may serve to improve pollination by a functional group of pollinators.

     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Floral reward presentation favored the expression of male function in the pollen‐only flower Melastoma malabathricum

Abstract Animal‐pollinated plant species modulate the presentation of pollinator rewards to maximize reproductive success. In plants providing pollen as the only reward for pollinators, it is usually difficult to unravel the dual roles of reward presentation and the realization of male and female functions (pollen removal and deposition). Exploiting the two types of anther in the androecia of Melastoma malabathricum L., we examined whether the removal of pollen for reward is regulated primarily to favor male function or female function. Pollen removal by carpenter bees from the feeding and pollination anthers, as well as pollen deposition on the stigmas, were quantified during anthesis of M. malabathricum. There was no significant difference in pollen removal rates from the feeding and pollination anthers of M. malabathricum between the onset of anthesis and flower wilting. The stigmatic pollen loads exceeded the ovule number after three sonication bouts, and female function was satisfied earlier than male function. The results support the hypothesis that the presentation of pollination reward in this species is regulated primarily to favor the expression of male function, rather than female function, in agreement with the pollen‐donation hypothesis. A cooperative relationship between the feeding and pollination anthers was demonstrated in heterantherous flowers, which optimizes the balance in investments between pollinator rewards and “functional pollen” for gene transfer.     

Heteranthery as a solution to the demand for pollen as food and for pollination – Legitimate flower visitors reject flowers without feeding anthers

• Heteranthery, the presence of feeding and pollinating anthers in the same flower, seems to mediate the evolutionary dilemma for plants to protect their gametes and yet provide food for pollinators. This study aims to elucidate the role of heteranthery in the buzz‐pollinated Senna reniformis.
• The fecundity of pollen from long‐, medium‐ and short‐sized anthers was determined by hand cross‐pollination experiments, and the quantity, size, ornamentation and viability of pollen of different anthers were compared. Rates of flower rejection by bees were measured in anther removal experiments to assess the preferences of flower visitors for feeding or pollinating anthers.
• Large bees, which were the effective pollinators of self‐incompatible S. reniformis, avoided flowers without short feeding anthers, but not those without medium or long anthers. Illegitimate small and medium‐sized bees were unresponsive to anther exclusion experiments. Long anthers deposited pollen on the back and short anthers on the venter of large bees. Pollen from long anthers had higher in vitro viability and higher fruit and seed set after cross‐pollination than pollen from other sized anthers.
• Short anthers produce feeding pollen to effective pollinators and long anthers are related to pollination of S. reniformis. Bee behaviour and size was found to directly influence the role of anthers in the ‘division of labour’. Only large bee pollinators that carry the pollinating pollen from long anthers in ‘safe sites’ associated short anthers with the presence of food. In the absence of these larger bee pollinators, the role of heteranthery in S. reniformis would be strongly compromised and its function would be lost.

     

Intrafloral differentiation of stamens in heterantherous fowers

Flowers that have heteromorphic stamens (heterantherous flowers) have intrigued many researchers ever since the phenomenon was discovered in the 19th century. The morphological differentiation in androecia has been suggested as a reflection of "labor division" in pollination in which one type of stamens attracts pollina-tors and satisfies their demand for pollen as food and the other satisfies the plant's need for safe gamete dispersal. The extent and patterns of stamen differentiation differ notably among taxa with heterantherous flowers. Seven species with heteromorphic stamens in three genera were sampled from Leguminosae and Melastomataceae, and the morphological difference of androecia, pollen content, pollen histochemistry and viability, pollen micro-morphology, as well as the main pollinators were examined and compared. Pollen number differs significantly between stamen sets of the same flower in most species investigated, and a correlation of pollen number and anther size was substantiated. Higher pollen viabilities were found in the long (pollinating) stamens of Senna alata (L.) Roxb. and S. bicapsularis (L.) Roxb. Dimorphic pollen exine ornamentation is reported here for the first time in Fordiophytonfaberi Stapf. The height of stigma and anther tips of the long stamens in natural conditions was proved to be highly correlated, supporting the hypothesis that they contact similar areas of the pollinator's body.     

塔落岩黄芪主要传粉蜂的传粉效率研究

塔落岩黄芪 (Hedysarum laeve)是毛乌素沙地重要豆科植物, 在我国西部生态系统恢复与重建中起着重要作用, 但其自交率非常低, 必须依靠传粉昆虫授粉才能结实。野生传粉蜂是其主要传粉者。为了明确塔落岩黄芪的主要传粉蜂组成及优势传粉蜂的传粉效率, 2004–2006年, 我们设置了4个2 m×2 m样方观测访花昆虫的组成并对主要传粉蜂的访花频率、花粉移出率和柱头花粉沉降数目及单花停留时间进行了观测。结果表明白脸条蜂(Anthophora albifronella)、海切叶蜂(Megachile maritima)和散熊蜂(Bombus sporsdicus)是塔落岩黄芪的主要传粉蜂, 其中白脸条蜂在数量和访花频率上占有明显优势, 但其花粉移出率比海切叶蜂和散熊蜂的低, 3种蜂的柱头花粉沉降数目没有显著区别。通过对这3种蜂的花粉移出率、柱头花粉沉降数目和访花频率综合分析后, 我们认为白脸条蜂是塔落岩黄芪最有效的传粉蜂。     

Evidence for reductions in floral attractants with increased selfing rates in two heterandrous species

Although theoretical models predict low allocation to attractive structures with increased selfing in animal-pollinated plants, empirical measurement of the reproductive costs and benefits is complicated. Here, floral sex allocation was compared in two nectarless heterandrous species with different mating systems: Monochoria korsakowii (Pontederiaceae), which has moderate outcrossing rates, and Monochoria vaginalis, a predominant selfer. In both species, mirror-image flowers have one large dark-purple anther and five small yellow anthers. Experimental evidence is provided for functional differences between the two sets of anthers using data on pollinator visitation, pollen removal and deposition, and seed set after hand pollinations. Flower manipulations in bee-pollinated M. korsakowii demonstrated different functions of the two sets of anthers: the yellow (feeding) anthers function to attract pollinators, but have similar pollen performance to the purple (pollinating) anthers. Furthermore, a disproportional reduction in pollen production of the feeding anthers in the selfing species was found. This differential allocation between feeding and pollinating anthers in heterandrous species has not been recognized before. The finding of reduced allocation to attractive structures with an increase in the rate of self-fertilization supports the theory of sex allocation.     

The effect of wild radish floral morphology on pollination efficiency by four taxa of pollinators

The effects of floral morphology on rates of pollen removal and deposition by different pollinators in generalist plant species are not well known. We studied pollination dynamics in wild radish, Raphanus raphanistrum, a plant visited by four groups of pollinators: honey bees, small native bees, butterflies, and syrphyd flies. The effects of anther position and other factors on pollen removal during single visits by all four pollinator taxa were measured. Flowers with high anther exsertion (i.e., anthers placed higher above the opening of the corolla tube) tended to have the highest numbers of pollen grains removed, but this effect was strongest for honey bees and butterflies. For all pollinator taxa, pollen removal increased with the number of pollen grains available on a flower and whowed a positive, decelerating relationship with the duration of the visit. The effects of stigma position and other factors on pollen deposition during single visits by honey bees and butterflies were also studied. The nectar-feeding butterflies had a higher pollination efficiency (percentage of pollen grains removed from anthers that were subsequently deposited on a stigma) than the nectar- and pollen-feeding honey bees. Flowers with intermediate stigma exsertion had the highest numbers of pollen grains deposited on their stigmas by butterflies, but stigma exsertion had no effect on deposition by honey bees. For both butterflies and honey bees, pollen deposition on the recipient flower increased with the amount of pollen removed from the donor flower, and there was a positive, decelerating relationship between deposition and time spent at the flower; these results are analogous to those for pollen removal. The effects of anther and stigma exsertion on pollen removal and denosition did not fit predictions based on patterns of floral correlations, but results for morphology, pollen availability, time spent per visit, and pollinator efficiency are in broad agreement with previous studies, suggesting the possible emergence of some general rules of pollen transfer.     

Pollination ecology of Campanula species on Mt Olympos, Greece

Nine Campanula species occurring along the elevation gradient of Mt Olympos were studied regarding their pollination ecology. The main issues considered were 1) the relative importance of various insect taxa as Campanula pollinators, 2) the patterns of pollinators' size and activity as a function of altitude, 3) the effect of pollinator exclusion on floral longevity, and 4) the extent to which the morphological difference of C. versicolor from the other Campanula species on Mt Olympos is expressed in its pollinating fauna. The vast majority of Campanula pollinators were solitary bees. Andrenidae and Megachilidae bees (mainly Chelostoma campanulorum) dominated the pollinating fauna of most species, Melittidae and bumblebees were the commonest pollinators of high altitude species. Campanula versicolor differs from the other Campanula species in that its corolla is not bell-shaped but flat. Mainly Apis mellifera, syrphid flies, and carpenter bees, unlike all other Campanula species on Mt Olympos pollinated it. At the species level, rather large altitudinal differences of Campanula populations did not result into large diversification of their pollinating fauna. The insect visitation rate to flowers decreased with altitude. When pollinators were excluded, the floral longevity of the species examined increased three to five times. Neither flower phase (male of female) was consistently favoured in the absence of pollinators. The pollen loads of the different insect taxa (Apis mellifera included) were of variable purity. The majority of Megachilidae bees carried pollen loads of high purity. Pollen loads from insects visiting Campanula species at high altitudes did not differ significantly in their purity from those visiting lowland species. The distribution of Campanula pollinators' body size along the altitudinal gradient exhibited a U-shaped pattern. No relationship was found between insect-pollinator body size and corolla size of Campanula species.     

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.     

Effects of conspecific and heterospecific floral density on the pollination of two related rewarding orchids

Variation in within-population floral density can affect interactions between plants and pollinators, resulting in variable pollen export for plants. We investigated the effects of conspecific and heterospecific floral densities on pollination success both of two related, self-compatible, nectar-rewarding orchid species in Ireland, Spiranthes romanzoffiana (rare and listed as endangered) and its congener, S. spiralis (more abundant and not of conservation concern). Floral densities, insect visitation rates, and orchid pollen transport were recorded in multiple quadrats in four populations of both orchid species over their flowering season. We found that conspecific and heterospecific co-flowering plant density affected pollination in both orchid species. For S. romanzoffiana, higher heterospecific density increased pollen removal. For S. spiralis, higher conspecific visitation increased pollen removal and increased heterospecific density decreased pollen deposition. In addition, increased conspecific density increased pollen deposition in both species. This study shows that plants may interact to facilitate or compete for different components of the pollination process, namely; pollinator attraction, pollen removal and deposition. Such interactions have immediate consequences for endangered plant species, as increases in both conspecific and heterospecific coflowering density may ameliorate the negative effects of rarity on pollination, hence overall reproductive success.     

Comparative Studies on Pollination Biology of Mosla hangchouensis and M.chinensis (Labiatae)

Mosla hangchouensis Matsuda and M. chinensis Maxim. are sibling species. The former is endemic to the coastal areas of Zhejiang Province, and the latter occupies southern China. Their morphological differences were associated with their different breeding systems indicated by pollen-ovule ratio (P/O). M. hangchouensis was predominantly outcrossing with P/O ranging from 756 to 6163 due to the variation among populations. Floral characters also showed that M. hangchouensis is an outcrosser: corolla tube 5～10 mm long, 3~5 mm wide, bright red: style 6～15 mm long; stigma 1.5～2.0 mm horizontally away from anthers. On the contrary, M. chinensis was predominantly a selfer with a low P/O ratio (P/O= 110). Its floral parameters contrasted sharply with M. hangchouensis: corolla tube 3.0 mm long, 1.0 ～ 1.5 mm wide, pale colored; style 2.5 mm long. stigma close to or touching anthers. The pollinators of M. hangchouensis most frequently seen are: Bombus trifascitus Smith., Scolia spp., Apis cerana Fabricius, Hylaeus sp. and Lasioglossum spp. These pollinators began to visit flowers at about 8:00 am, when flower buds were ready to open. When insects entered corolla tube for honey and (or) pollen, the stigmata touched the insects body hairs to which pollen grains adhered, and pollen grains were transferred to the stigmata. Meanwhile the hairs collected pollen from anthers under the upper lips of corolla as the insects thrusted their bodies forward. Bumble bees and honey bees were considered more efficient pollinators because they were hair-rich and their body size fitted corolla tube fairly well. Due to the'short flight distance of pollinators between two visits (average flight distances of honey bees and solitary bees were 0.52 m and 0.32 m respectively), pollen dispersal was probably restricted. Pollen vectors were important for the reproductive success of M. hangchouensis because 84.3 % of its seed-set depended on the pollinators' visitation. Although insects did pay visits to M. chinensis occasionally, such visitation was insignificant in seed-set due to cleistogamy. Discussions are made on the evolutionary significance of breeding system divergence between M. hangchouensis and M. chinensis, and the contribution of studies on pollination biology to conservation biology. Further studies are suggested for more thorough understanding of pollination biology of M. hangchouensis.     

Colored floral organs influence pollinator behavior and pollen transfer in Commelina communis (Commelinaceae)

Visual floral guides such as colored anthers, lines, dots, and UV-absorption patterns on petals are commonly observed in insect-pollinated angiosperms. Floral guides that are known to enhance foraging efficiency of visitors on flowers thus promote return visits (foraging facilitation hypothesis, which predicts that visitors will discriminate against flowers with inferior floral guides). In this study, we experimentally examined the hypothesis that floral guides also prevent pollen-theft behavior by floral visitors (theft prevention hypothesis), which has rarely been tested. Nectarless flowers of Commelina communis have three types of brightly colored floral organs: large blue petals, rewarding yellow anthers, and nonrewarding yellow anthers. Colored floral organs were removed artificially from plants in two natural populations of C. communis. Removal of the nonrewarding yellow anthers diminished hoverings in front of flowers and tended to reduce the number of total floral visitor landings, supporting the foraging facilitation hypothesis. Additionally, removal of the rewarding yellow anther decreased the frequency of legitimate landings on flowers and the legitimate landing-to-total landing ratio, which is consistent with the theft prevention hypothesis. The nonrewarding anthers and the rewarding yellow anthers were shown to play an important role in increasing visitor landings and orienting floral visitors toward a landing point appropriate for pollination, respectively. We also showed that the absence of yellow anthers decreased both pollen dispatch from brown anthers and receipt by stigmas in C. communis. These findings support both the foraging facilitation hypothesis and the theft prevention hypothesis.     

早春短命植物黑鳞顶冰花的花粉呈现时序及其适应性

花粉的时序呈现是指植物花药按一定次序释放花粉的现象, 被认为是对传粉者访问频率的一种适应。在传粉者充足的环境中, 植物通过限制花粉1次被移出的数量, 使花粉供体能作为多个父本, 从而提高雄性适合度。该文从开花习性、花部特征、传粉者及繁育系统等方面对早春短命植物黑鳞顶冰花(Gagea nigra)的花粉呈现时序及其适应性进行研究, 结果表明, 黑鳞顶冰花单花期约5-7天; 白天开放, 晚上闭合, 花药次序开裂, 呈拉链式散粉, 散粉期4-6天。黑鳞顶冰花以异交为主, 部分自交亲和。蝇类和食蚜蝇为主要传粉者, 访花频率为(0.141±0.078) flower∙h^-1。在雄蕊的时序散粉过程中, 雌蕊持续生长, 经历了从低于雄蕊到等高、再到高于雄蕊阶段。在等高阶段, 单花早晚的开闭, 使得雌雄蕊紧靠在一起, 促成了自动自花授粉。在传粉者缺乏的环境中, 黑鳞顶冰花的花粉时序呈现延长了散粉期, 在等待传粉者和分摊风险方面具重要作用。这种花粉渐次呈现的策略, 在新疆的早春开花植物中可能广泛存在。     

蝶形花亚科植物花部适应机制与传粉系统

蝶形花亚科(Papilionoideae)植物种类丰富、繁育系统多样,与其传粉者关系密切,主要表现在花部适应与传粉系统的形成。蝶形花形态复杂,为完全花,花萼、花瓣5基数,花萼呈箭头状,相邻花瓣螺旋轮生,旗瓣较大位于最外边,两片翼瓣紧贴旗瓣着生并包裹两片合生的龙骨瓣,龙骨瓣内包裹着雌、雄蕊,雌蕊位于正中央,雄蕊轮生,构成二体雄蕊(多数9+1,少数5+5)。对蝶形花亚科植物的花部特征、传粉功能群、酬物与传粉系统构建进行了回顾,重点论述了蝶形花形态和化学组成与传粉系统的进化关系以及花粉呈现机制。其中,泛化的传粉系统以蜂媒传粉为主,同时存在鸟媒传粉、蝙蝠传粉和松鼠传粉等方式,花部结构和传粉者的相互选择和相互适应推动了传粉系统的演化。花粉释放是体现蝶形花植物与其传粉功能群相互作用的重要方面,二者协同进化形成了以下4种花粉呈现机制:弹花机制、活塞机制、瓣膜机制和毛刷机制。蝶形花的花部特征与传粉功能群的相互作用形成相应的传粉系统,而传粉系统是执行传粉功能的重要集合体,各个构件的相互协调和适应保证了蝶形花亚科植物传粉过程的顺利完成。     

Effects of local density on pollination and reproduction in Delphinium nuttallianum and Aconitum columbianum (Ranunculaceae)

Plant populations vary in density both naturally and as a consequence of anthropogenic impacts. Density in turn can influence pollination by animals. For example, plants in dense populations might enjoy more frequent visitation if pollinators forage most efficiently in such populations. We explored effects of plant density on pollination and seed set in the larkspur Delphinium nuttallianum and monkshood Aconitum columbianum. At our site in the Colorado Rocky Mountains, flowers of D. nuttallianum are pollinated primarily by queen bumble bees, solitary bees, and hummingbirds, whereas those of A. columbianum are pollinated primarily by queen and worker bumble bees. We found that the quantity of pollination service to both species (pollinator visitation rate and pollen deposition) was at best weakly related to density. In contrast, seed set declined by approximately one-third in sparse populations relative to nearby dense populations. This decline may stem from the receipt of low-quality pollen, for example, inbred pollen. Alternatively, sparsity may indicate poor environmental conditions that lower seed set for reasons unrelated to pollination. Our results demonstrate the value of simultaneously exploring pollinator behavior, pollen receipt, and seed set in attempting to understand how the population context influences plant reproductive success.     

Ricochet pollination in Senna (Fabaceae) – petals deflect pollen jets and promote division of labour among flower structures

• Naturalists Fritz and Hermann Müller hypothesised that heteranthery often leads to a division of labour into ‘feeding’ and ‘pollinating’ stamens; the latter often being as long as the pistil so as to promote successful pollination on the bees’ back. In many buzz‐pollinated species of Senna, however, the so‐called pollinating stamens are short and not level with the stigma, raising the question of how pollen is shed on the bees’ back. Here we explore a mechanism called ‘ricochet pollination’. We test whether division of labour is achieved through the interaction between short lower stamens and strongly concave ‘deflector petals’.
• We studied the arrangement and morphology of the floral organs involved in the ricochet pollination, functioning of the flowers through artificial sonication and observed the interactions between bees and flowers in the field.
• The middle stamens are adapted to eject pollen downwards, which can be readily collected on the bee mid legs. Most of the pollen is ejected towards the deflector petal(s). Pollen from this set of stamens is more likely to contribute to pollination. The pollen grains seem to ricochet multiple times against the deflector petals to eventually reach the bee's back.
• The pollen ricochet mechanism promotes a division of labour by involving additional floral organs, such as petals, reinforcing the Müllers’ division‐of‐labour hypothesis. However, alternative, non‐multiexclusive hypotheses could be explored in genus Senna and other angiosperm species.

     

Flower-visiting bees and the breakdown of the tristylous breeding system of Eichhornia azurea (Swartz) Kunth (Pontederiaceae)

Populations of Eichhornia azurea (Swartz) Kunth (Pontederiaceae) are often composed of three floral morphs that differ in style length and anther position. Hence, E. azurea is described as tristylous. For sexual reproduction the plant requires specialized pollinators able to transfer pollen from flowers with long-, mid-, or short-level anthers to flowers with long-, mid-, or short-styles, respectively. This is because within a floral morph, self-incompatibility is the rule. Populations of E. azurea were studied from six localities in south-eastern Brazil (São Paulo and Minas Gerais). The frequencies of floral morphs, floral morphology and flower-visiting bees were recorded in each population. Ten different bee species were observed visiting plants. Four of them were considered generalist bees and not appropriate for legitimate pollination of the three floral morphs. Another three species were considered opportunists and only three species were considered to be potential pollinators. At the coastal region (Juréia), the plant population lacked the short-styled morph and the mid-styled morph and exhibited a semi-homostylous condition, in which the low-level anthers were adjacent to the stigma. This morph showed high level of self-compatibility. The disappearance of the short-styled morph from this region is likely correlated with the absence of Ancyloscelis gigas Friese, a specialized long-tongued bee that is oligolectic on this plant in other localities.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 77 , 499–507.     

Response of wild bees (Hymenoptera: Apoidea: Anthophila) to surrounding land cover in Wisconsin pickling cucumber

Cucumber (Cucumis sativus L.) is among the plants highly dependent on insect-mediated pollination, but little is known about its unmanaged pollinators. Both domestic and wild bee populations in central Wisconsin pickling cucumber fields were assessed using a combination of pan trapping and floral observations before and during bloom. Together with land cover analyses extending 2,000 m from field centers, the relationship of land cover components and bee abundance and diversity were examined. Over a 2-yr sample interval distributed among 18 experimental sites, 3,185 wild bees were collected representing >60 species. A positive association was found between both noncrop and herbaceous areas with bee abundance and diversity only during bloom. Response of bee abundance and diversity to land cover was strongest at larger buffers presumably because of the heterogeneous nature of the landscape and connectivity between crop and noncrop areas. These results are consistent with previous research that has found a weak response of wild bees to surrounding vegetation in moderately fragmented areas. A diverse community of wild bees is present within the fields of a commercial cucumber system, and there is evidence of floral visitation by unmanaged bees. This evidence emphasizes the importance of wild pollinators in fragmented landscapes and the need for additional research to investigate the effectiveness of individual species in pollen deposition.