Selective nectar robbing in a gynodioecious plant (Glechoma longituba) enhances female advantage

Nectar robbing not only affects the reproductive fitness of the plant but it may also potentially affect the pollination dynamics of the associated coflowering individuals. In this study, we established that the nectar robber Xylocopa sinensis robs nectar only from the hermaphrodite ramets of the gynodioecious plant Glechoma longituba but not from the female ramets. In populations with high rates of nectar robbing, this results in hermaphrodite ramets having reduced seed set whereas the female ramets have a slightly increased seed set. We hypothesize that selective nectar robbing confers an advantage to female individuals and thus ensures their maintenance in gynodioecious populations. Results of controlled experiments indicated that the reduction in the amounts of nectar available occasioned by nectar robbing resulted in some legitimate pollinators switching to visiting flowers on female rather than hermaphrodite ramets. This resulted in lower pollination rates and seed set for hermaphrodites and higher pollination rates and seed set for females. This study presents a previously unreported mechanism causing female advantage in gynodioecious plants.     

Consequences of nectar robbing for the fitness of a threatened plant species

The effect of nectar robbing on plant fitness is poorly understood and restricted to a few plant species. Furthermore, the available studies generally evaluate the effects of nectar robbing on female fitness, disregarding the male component. Here we measured the effects of the nectar-robbing bumblebees on male (measured as pollen analogue flow distance) and female (measured as seed production) reproductive success in the insect-dependent Polygala vayredae, a narrow endemic species from the pre-Pyrenees (Spain). Intense nectar robbing by bumblebees significantly reduced the nectar available to legitimate pollinators in the studied population, and this reduction affected both male and female fitness. Significant differences were observed in fluorescent dye dispersion between robbed and non-robbed flowers within the population. Fluorescent dyes from non-robbed flowers were dispersed to larger distances and over a larger number of flowers when compared with robbed ones. Moreover, significant differences were observed in both fruit set and seed ovule ratios between the two groups, with non-robbed flowers presenting higher reproductive outcomes. However, no effect on seed weight was detected among treatments. The data obtained suggest that in this species, nectar robbing has important indirect and negative effects on plant fecundity, through both male and female functions, due to a modification in the foraging behaviour of legitimate visitors.     

盗蜜对角蒿传粉者行为和生殖成功的影响

一些研究显示盗蜜对自交植物的结实和结籽没有显著影响。然而, 对于既有传粉者为其传粉实现异交又能通过自交实现生殖保障的兼性自交植物来说, 盗蜜对其生殖的影响还知之甚少。由于兼性自交植物可以自交, 盗蜜对其总体结实可能不会有显著影响, 但可能会通过影响传粉者行为而影响传粉者介导的结实。为了验证这一假说, 本研究以兼性自交的一年生角蒿(Invarvillea sinensis var. sinensis)为研究材料, 通过野外调查和控制实验, 探讨了盗蜜对传粉者介导的结实(传粉者行为)和总体结实率的影响。结果表明: 角蒿的盗蜜者和主要传粉者相同, 均为密林熊蜂(Bombus patagiatus)。熊蜂盗蜜频率平均为20.24% (范围为0-51.43%)。盗蜜对角蒿总体结实率、每果结籽数和每果种子重量没有显著影响。然而, 被盗蜜花的柱头闭合比率显著高于未被盗蜜花, 说明盗蜜影响传粉者的访花行为和传粉者介导的结实率。另外, 被盗蜜花的高度显著高于未被盗蜜花, 说明盗蜜者倾向于从较大较高的花上盗蜜。这些结果为全面认识盗蜜对植物生殖的影响提供了新的信息。     

Study on the Chemical Constituents of Comastoma pulmonarium (Turcz.) Toyohuni

Eight compounds were isolated from Comastoma pulmonarium (Gentianaceae). Theirstructures are identified as1,8-dihydroxy-3,5-dimethoxyxanthone(Ⅰ),1,8-dihydroxy-3,7-dimetho-xyxanthone(Ⅱ), 1-hydroxy-3,7,8-trimethoxyxanthone(Ⅲ),8-hydroxy-l,3,5-trimethoxyxanthone(Ⅳ), 1,3,8-trihydroxy-7-methoxyxanthone(Ⅴ), 1-O-β-D-glucopyranosyl-3,8-dihydroxy-7-methxy-xanthone(Ⅵ), oleanolic acid(Ⅶ) and swertisin(Ⅷ) by means of chemical methods andUV, IR, NMR and MS respectively. The compound Ⅵ is a new natural product. It wasnamed as comastomaside.     

Corolla stickiness prevents nectar robbing in Erica

Journal of Plant Research - Floral stickiness is a rare trait with unknown function, but it is common in the mega-diverse Cape genus Erica (Ericaceae). This study investigated the role of...     

Predicting the effects of nectar robbing on plant reproduction: implications of pollen limitation and plant mating system

The outcome of species interactions is often difficult to predict, depending on the organisms involved and the ecological context. Nectar robbers remove nectar from flowers, often without providing pollination service, and their effects on plant reproduction vary in strength and direction. In two case studies and a meta-analysis, we tested the importance of pollen limitation and plant mating system in predicting the impacts of nectar robbing on female plant reproduction. We predicted that nectar robbing would have the strongest effects on species requiring pollinators to set seed and pollen limited for seed production. Our predictions were partially supported. In the first study, natural nectar robbing was associated with lower seed production in Delphinium nuttallianum, a self-compatible but non-autogamously selfing, pollen-limited perennial, and experimental nectar robbing reduced seed set relative to unrobbed plants. The second study involved Linaria vulgaris, a self-incompatible perennial that is generally not pollen limited. Natural levels of nectar robbing generally had little effect on estimates of female reproduction in L. vulgaris, while experimental nectar robbing reduced seed set per fruit but not percentage of fruit set. A meta-analysis revealed that nectar robbing had strong negative effects on pollen-limited and self-incompatible plants, as predicted. Our results suggest that pollination biology and plant mating system must be considered to understand and predict the ecological outcome of both mutualistic and antagonistic plant-animal interactions.     

位置效应对喉毛花（龙胆科）的花特征和繁殖成功的影响

同一个植株上不同位置花的特征和繁殖成功通常表现出较大的差异,这种差异是源于不同位置花间的资源竞争还是植株本身发育过程中的结构效应仍需进一步阐明。作者以青藏高原高寒草甸中常见的一年生自交植物——喉毛花（Comastoma pulmonarium）为研究对象,通过控制植株的资源状态比较植株上不同位置花的特征和繁殖成功,探讨花序内资源竞争和结构效应的影响。结果发现,与其它位置的花相比,位于植株顶端的首先开放的花具有较大花高度和花直径,并产生了更多的花粉和胚珠 但蕾期去除顶花后,其它位置的花高度和花直径与对照植株上同位置的花相比并没有显著变化,表明喉毛花不同位置的花特征变异是由植株发育过程中的结构效应引起,改变植株的资源状态不会改变喉毛花的花特征。喉毛花顶果的种子数量、重量和萌发率均高于其它位置果实的种子 但去除顶花后,其它位置果实的种子数量与对照植株相同位置的果实相比,均呈现增加的趋势,表明喉毛花顶果以外其它不同位置果实的种子发育期间存在资源限制 改变植株的资源状态后,植株的可用资源会在不同位置的果实间重新分配,这对后期发育的果实具有一定的保障作用。研究结果表明,喉毛花植株上不同位置间的花特征和繁殖成功差异分别由结构效应和资源竞争引起,与其它的研究结果存在分歧,其原因可能与喉毛花一年生的生活史特征和资源有限的高原环境有关,但仍需要进一步的研究证明。     

Social learning: nectar robbing spreads socially in bumble bees

Social transmission of learned behaviour is well documented in vertebrates but much less so among invertebrates. New research shows that nectar robbing can spread socially among bumble bees, even in the absence of nectar-robbing models.     

Novel nectar robbing negatively affects reproduction in Digitalis purpurea

With many plant–pollinator interactions undergoing change as species’ distributions shift, we require a better understanding of how the addition of new interacting partners can affect plant reproduction. One such group of floral visitors, nectar robbers, can deplete plants of nectar rewards without contributing to pollination. The addition of nectar robbing to the floral visitor assemblage could therefore have costs to the plant´s reproductive output. We focus on a recent plant colonist, Digitalis purpurea, a plant that in its native range is rarely robbed, but experiences intense nectar robbing in areas it has been introduced to. Here, we test the costs to reproduction following experimental nectar robbing. To identify any changes in the behavior of the principal pollinators in response to nectar robbing, we measured visitation rates, visit duration, proportion of flowers visited, and rate of rejection of inflorescences. To find the effects of robbing on fitness, we used proxies for female and male components of reproductive output, by measuring the seeds produced per fruit and the pollen export, respectively. Nectar robbing significantly reduced the rate of visitation and lengths of visits by bumblebees. Additionally, bumblebees visited a lower proportion of flowers on an inflorescence that had robbed flowers. We found that flowers in the robbed treatment produced significantly fewer seeds per fruit on average but did not export fewer pollen grains. Our finding that robbing leads to reduced seed production could be due to fewer and shorter visits to flowers leading to less effective pollination. We discuss the potential consequences of new pollinator environments, such as exposure to nectar robbing, for plant reproduction.     

Effects of nectar robbing on pollinator behavior and plant reproductive success of Pitcairnia angustifolia (Bromeliaceae)

Nectar robbers may have direct and indirect effects on plant reproductive success but the presence of nectar robbing is not proof of negative fitness effects. We combined census data and field experiments to disentangle the complex effects of nectar robbing on nectar production rates, pollinator behavior, pollen export, and female reproductive success of Pitcairnia angustifolia. Under natural conditions flowers were visited by four different animal species including a robber‐like pollinator and a secondary robber. Natural levels of nectar robbing ranged from 40 to 100%. Natural variation in nectar robbing was not associated with fruit set in any year whereas seed set was weakly positively associated for 1 year only. Artificial nectar robbing did not increase nectar production or concentration, did not affect the behavior of long‐billed hummingbirds, and when faced with artificially robbed flowers, these visitors behaved as secondary nectar robbers. The number of stigmas within a patch that received pollen dye analogs and the average distance traveled by these analogs were not significantly different between robbing treatments (robbed flowers versus unrobbed flowers), but the maximum distance traveled by these pollen analogs was higher when nectar robbing was not prevented. Overall, the proportion of robbed flowers on an inflorescence had a neutral effect to a weak positive effect on the reproduction of individual plants (i.e. positive association between nectar robbing and fruit set in 2002) even when it clearly changed the behavior of its most efficient pollinator potentially increasing the frequency of nectar robbing within a plant.     

Effects of nectar robbing on nectar dynamics and bumblebee foraging strategies in Linaria vulgaris (Scrophulariaceae)

Differences in morphology among bumblebee species sharing a nectar resource may lead to variation in foraging behaviour and efficiency. Less efficient bumblebees might opportunistically switch foraging strategies from legitimate visitation to secondary robbing when hole-biting primary robbers are present. We observed various aspects of pollination and nectar robbing ecology of Linaria vulgaris in the Colorado Rocky Mountains, with emphasis on the role of bumblebee proboscis length. Bees can extract nectar from a nectar spur legitimately, by entering the front of the flower, or illegitimately, by biting or reusing holes in the spur. Although L. vulgaris flowers are apparently adapted for pollination by long-tongued bees, short-tongued bees visited them legitimately for trace amounts of nectar but switched to secondary robbing in the presence of primary robbers. Longer-tongued bees removed more nectar in less time than did shorter-tongued bees, and were less likely to switch to secondary robbing even when ∼100% of flowers had been pierced. As the proportion of robbed flowers in the population increased, the relative number of legitimate visits decreased while the relative number of robbing visits increased. Robbing decreased nectar standing crop and increased the proportion of empty flowers per inflorescence. Despite these potentially detrimental effects of robbers, differences in inflorescence use among robbers and pollinators, and the placement of holes made by primary robbers, may mitigate negative effects of nectar robbing in L. vulgaris . We discuss some of the reasons that L. vulgaris pollination ecology and growth form might temper the potentially negative effect of nectar robbing.     

Selective seed abortion affects the performance of the offspring in Bauhinia ungulata

BACKGROUND AND AIMS: Under the microgametophytic competition hypothesis, a non-random pattern of seed abortion is expected, in which only the most vigorous seeds reach maturity. In a previous study, it was found that Bauhinia ungulata (Fabaceae) exhibits a pattern of seed abortion dependent on the position of the ovule within the ovary; ovules located in the stylar half of the fruit, close to the point of entry of pollen tubes to the ovary, have a low probability of seed abortion, whereas ovules in the basal half of the fruit are aborted with a high probability. METHODS: A series of experimental fruits was generated, in which ovules from either the stylar (treatments 1 and 2) or the basal (treatments 3 and 4) half of fruits were destroyed, to evaluate whether these patterns of selective seed abortion have an effect on the vigour of the offspring in B. ungulata. KEY RESULTS: Only 53 % of the seed from control fruits germinated. Seed set in fruits from treatments 1 and 2 showed a significantly lower (33-43 %) percentage of germination; the germination of seeds from fruits in treatments 3 and 4 (49-51 %) did not differ from control seeds. In addition, it was found that the differences in vigour of the offspring are not random with respect to the position of the ovule in the pod. CONCLUSIONS: The overall performance of the seeds correlated with their likelihood of maturation. Seeds located at the basal half of the treatment fruits showed lower values of vigour than seeds located on the stylar half. The differences were more marked for early measures of fitness.     

Realized tolerance to nectar robbing: compensation to floral enemies in Ipomopsis aggregata

Background and Aims

Although the ecological and evolutionary consequences of foliar herbivory are well understood, how plants cope with floral damage is less well explored. Here the concept of tolerance, typically studied within the context of plant defence to foliar herbivores and pathogens, is extended to floral damage. Variation in tolerance to floral damage is examined, together with some of the mechanisms involved.

Methods

The study was conducted on Ipomopsis aggregata, which experiences floral damage and nectar removal by nectar-robbing bees. High levels of robbing can reduce seeds sired and produced by up to 50 %, an indirect effect mediated through pollinator avoidance of robbed plants. Using an experimental common garden with groups of I. aggregata, realized tolerance to robbing was measured. Realized tolerance included both genetic and environmental components of tolerance. It was hypothesized that both resource acquisition and storage traits, and traits involved in pollination would mitigate the negative effects of robbers.

Key Results

Groups of I. aggregata varied in their ability to tolerate nectar robbing. Realized tolerance was observed only through a component of male plant reproduction (pollen donation) and not through components of female plant reproduction. Some groups fully compensated for robbing while others under- or overcompensated. Evidence was found only for a pollination-related trait, flower production, associated with realized tolerance. Plants that produced more flowers and that had a higher inducibility of flower production following robbing were more able to compensate through male function.

Conclusions

Variation in realized tolerance to nectar robbing was found in I. aggregata, but only through an estimate of male reproduction, and traits associated with pollination may confer realized tolerance to robbing. By linking concepts and techniques from studies of plant–pollinator and plant–herbivore interactions, this work provides insight into the role of floral traits in pollinator attraction as well as plant defence.Key words: Compensation, herbivory, indirect effects, Ipomopsis aggregata, male reproductive success, nectar robbing, pollen donation, pollination, resistance, tolerance     

Differential effects of anthropogenic edges and gaps on the reproduction of a forest‐dwelling plant: The role of plant reproductive effort and nectar robbing by bumblebees

Forest fragmentation produces sharp changes in the spatial configuration of remnant forest patches, which include the increasing influence of patch edges on the interior of forest patches (‘edge effects’). Human activities responsible for forest fragmentation tend also to change the internal characteristics of remnant patches, for example, through the creation of gaps by selective logging. While edges and gaps can be expected to cause comparable changes in the micro‐environmental conditions of the forest, their effects on forest‐dwelling species and their interactions are not necessarily comparable. This study compares the effect of forest edges and anthropogenic gaps on the reproductive success of a self‐incompatible epiphytic plant (Mitraria coccinea), mediated by changes in its relationship with mutualists (pollinators) and antagonists (flower larcenists). Mitraria coccinea's flowers are pollinated by hummingbirds and robbed by the bumblebee Bombus dahlbomii. Edges and gaps had comparable positive effects on flower production and fruit set; however, nectar robbing was up to sevenfold higher in patches with numerous gaps and resulted in lower reproductive success (fruit set and total fruit crop). Forest fragmentation studies should therefore avoid treating forest remnants as homogeneous units (i.e. focusing exclusively on their characteristics and connectivity), without taking into account the internal heterogeneity caused by anthropogenic pressures (e.g. gaps and edge effects).     

Interactions among nectar robbing, floral herbivory, and ant protection in Linaria vulgaris

Nectar robbers are often assumed to be plant antagonists; however, empirical data show that the impacts of these animals range from negative to positive depending on the system and ecological conditions. We experimentally evaluated the combined effects of nectar robbing and ant visitation on three indices of reproductive fitness in Linaria vulgaris in the Colorado Rocky Mountains, via indirect effects on flower- and seed-eating beetles ( Brachypterolus pulicarius and Gymnaetron antirrhinni ). Nectar robbing Bombus occidentalis leave holes in the nectar spurs, effectively creating "extra-floral nectaries" that attract ants. In a paired-plant experiment, ants were significantly more abundant on robbed than on unrobbed plants. Manipulation of ant access and nectar robbing showed that ant exclusion increased beetle attack and decreased female fitness. There was a significant ant-by-robbing interaction on flower damage. Patterns in the other two indices were suggestive of ant-by-robbing interactions, but these were not statistically significant. We also found correlations between spider occupancy on some plants and the mean number of ants (marginally negative) or beetles (significantly positive). Although the effect we report in this study may be highly dependent on spatial and temporal distributions of several interacting species, we discuss its potential role in mitigating the costs of floral parasitism, and its importance to the study of nectar robbing in general.     

Extrafloral nectar increases seed removal by ants in Turnera ulmifolia

To investigate whether extrafloral nectar (EFN) increases seed dispersal in Turnera ulmifolia, we measured seed removal on plants with and without EFN. Plants producing EFN had more seeds removed than control plants, suggesting that EFN does play a role in seed dispersal. This is a novel function of EFN.