Flowers, Nectar and Insect Visits: Evaluating British Plant Species for Pollinator-friendly Gardens

Twenty-four plant species native or naturalized in Britain weregrown in the Cambridge University Botanic Garden, UK and evaluatedas potential resources for nectar-foraging bees, butterfliesand hoverflies. In ten plant species a series of measurementswere made, at regular intervals from dawn to dusk, of nectarsecretion rate and standing crop, and in all species insectvisits were monitored throughout daylight hours. The study revealeddifferences between plant species in the composition of theassemblage of insect visitors, and in the magnitude and temporaldistribution of the nectar reward. In some cases we found interestingcorrelations between temperature and secretion rates or patternsof insect visits. Species that received numerous insect visitsin our study are potentially valuable forage plants that mightbe planted by gardeners to support local pollinator populations.Deep flowers whose nectar is accessible to long-tongued bumblebees(Bombus hortorum, B. pascuorum) but not to honeybees may providelong-tongued pollinators with a resource refuge relatively freefrom honeybee competition. Features that make some of thoseplant species particularly interesting to observe in the gardeninclude robbing by short-tongued bumblebees inSaponaria,baseworkingby honeybees on closed flowers inMalva sylvestris, and apparentdisplacement of bumblebees by territorial behaviour of the solitarybeeAnthidium manicatumon species ofStachys.Copyright 1999 Annalsof Botany Company Wild flowers, gardens, nectar,Silene, Lychnis, Saponaria, Stachys, Malva, Dipsacus fullonum, Lythrum salicaria,pollinators, bumblebees,Bombus,honeybees,Apis,butterflies, Lepidoptera, hoverflies, Syrphidae,Anthidium manicatum, weeds.     

Impact of flower rewards on phytophagous insects: importance of pollen and nectar for the development of the pollen beetle (<Emphasis Type="Italic">Brassicogethes aeneus</Emphasis>)

Entomophilous plants reward pollinators with provision of nutrient-rich foods such as pollen and nectar. These rewards contain compounds that are essential to insect development and can be used by pollinators as well as herbivorous insects. The pollen beetle (Brassicogethes aeneus, syn. Meligethes aeneus) whose larvae develop in oilseed rape flowers (Brassica napus) is known to feed on pollen. Previous studies already showed the importance of pollen on the development of this insect but it seems that other resource, such as nectar, could also be used. The purpose of this study was to assess the respective roles of pollen and nectar on pollen beetle development. We tested their role with behavioural and developmental experiments using flowers where the presence and absence of nectar and pollen varied. Larvae, irrespective of their instar, fed both on anthers and nectar. Nectar did not influence larval development or adult survival while pollen influenced development by increasing both larval and adult weight. However, pollen did not affect larval or adult survival nor development time. These results indicate that pollen beetle larvae are adapted to deal with various diets and can complete their development without pollen or nectar.     

Floral Nectar Secretion and Ploidy in Brassica rapa and B. napus (Brassicaceae). II. Quantified Variability of Nectary Structure and Function in Rapid-cycling Lines

Haploid, diploid and tetraploid lines ofBrassica rapaL. (syn.campestris),and allotetraploidB. napusL., were examined to determine theinfluence of ploidy on floral features, particularly nectarymorphology and anatomy, and to relate nectary structure to nectarproduction capacity. Except for haploids, all lines were rapid-cycling.Average flower dry weight, and petal length and width, werein the descending orderB. napus>B. rapa (4n) >2n>n.Pollen grains of 4nplants were larger than those of 2nplants;haploids lacked pollen. All lines developed nectaries. Typically, each flower producedtwo pairs of nectaries, of different types and nectar productioncapacity. Normally, each lateral gland was located above thebase of a short stamen, and together this pair yielded mostof a flower 's nectar carbohydrate. Each median nectary aroseat the outer junction of the bases of two adjacent long stamens.All lateral nectaries received a vascular supply of phloem alone,but median glands received reduced amounts of phloem, or lackedvasculature altogether. Most nectaries were solitary, but 14%of all flowers, and especially those of 2n B. rapa,had at leastone median and lateral gland connected. Obvious variation existed in nectary morphology between ploidylevels, between flowers of the same plant, and even within flowers.Ten forms of each nectary type were recognized. Plants producingthe most nectar carbohydrate had high frequencies of lateralnectaries which were symmetrical, unfurrowed swellings. TetraploidsofB. rapahad both the highest frequencies of furrowed lateralglands, and of isolated segments of nectarial tissue at thatposition. Even these separated nectarial outgrowths receivedphloem and produced a nectar droplet. At the median location,nectaries were commonly of two forms: peg- or fan-shaped. Lobeson median nectaries, up to four per nectary, were detected inalmost half of glands of 4nflowers examined; lobes were absentin haploids. Brassica rapa; Brassica napus; flower size; nectar production; nectary variability; petal size; ploidyphloem; pollen; rapeseed     

Are nectar-robbers mutualists or antagonists?

As exploiters of plant-pollinator mutualisms, nectar-robbers remove rewards (nectar) without providing pollination services. Though one might expect nectar-robbing to be costly to plants, it may instead benefit plants by indirectly increasing pollen dispersal. I investigated the direct effects of nectar-robbing bees (Xylocopa californica) on floral rewards and behaviors of pollinators visiting desert willow (Chilopsis linearis) and indirect effects of robbing on the reproductive success of the plant. Nectar-robbers reduced nectar; while unrobbed and robbed flowers were equally likely to contain nectar, nectar volumes were smaller in robbed flowers with nectar. Apis mellifera (honeybees), ineffective pollinators in terms of pollen deposition, avoided robbed flowers. In contrast, Bombus sonorus (bumblebees), effective pollinators, did not avoid robbed flowers. While bumblebees tended to spend less time in robbed flowers, the time that they spent in flowers was not correlated with pollen deposition. Using powder mimicking pollen, I found that on some days, powder was dispersed farther or to more flowers from robbed flowers, indicating that robbing may sometimes benefit plants by increasing male reproductive success. Powder movement suggested that the effect of robbing on male reproductive success ranged from costly to beneficial. The outcome for flowers that were marked early each morning was a function of prevalence of robbing and abundances of effective pollinators, but not a function of spatial variability among trees in prevalence of robbing or the abundance of ineffective honeybees. Unlike powder dispersal, female reproductive success, measured by fruit set and the number of pollen tubes growing in styles, was not affected by robbing. Thus, robbers did not reduce plants female reproductive success either directly by damaging flowers or indirectly by reducing pollen deposition by pollinators. Overall, this study indicates that nectar-robbers were not often costly to plants, and sometimes even benefited plants.     

Nectar consumption of warblers after long-distance flights during spring migration

The consumption of nectar by European passerines has been reported only occasionally. In this study we investigated the occurrence and significance of nectar consumption of small passerine birds on spring migration after crossing the Mediterranean Sea. On Ventotene Island in the Tyrrhenian Sea, four migrating species of Sylvia warblers [Garden Warbler S. borin , Subalpine Warbler S. cantillans , Whitethroat S. communis , Blackcap S. atricapilld ) regularly foraged on the two most common flowering species at that time of year, Brassica fruticulosa (Cruciferae) and giant fennel Ferula communis (Umbelliferae), while other species visited flowers only occasionally or not at all. Feeding behaviour, pollen traces on the head, and the examination of pollen and sugar remains in droppings indicated that nectar was the main target of the Sylvia warblers, rather than pollen or insects on the flowers. This was confirmed by food choice experiments indicating a clear preference by Garden Warblers and Whitethroats for nectar from artificial flowers over mealworms. Although conclusive experiments are not available, we hypothesize that nectar might be a diet easy to obtain and to absorb for birds after a long-distance flight in which they have incurred a depletion of energy stores and a reduction of the digestive tract.     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

Nectar Sugar Production across Floral Phases in the Gynodioecious Protandrous Plant Geranium sylvaticum

Many zoophilous plants attract their pollinators by offering nectar as a reward. In gynodioecious plants (i.e. populations are composed of female and hermaphrodite individuals) nectar production has been repeatedly reported to be larger in hermaphrodite compared to female flowers even though nectar production across the different floral phases in dichogamous plants (i.e. plants with time separation of pollen dispersal and stigma receptivity) has rarely been examined. In this study, sugar production in nectar standing crop and secretion rate were investigated in Geranium sylvaticum, a gynodioecious plant species with protandry (i.e. with hermaphrodite flowers releasing their pollen before the stigma is receptive). We found that flowers from hermaphrodites produced more nectar than female flowers in terms of total nectar sugar content. In addition, differences in nectar production among floral phases were found in hermaphrodite flowers but not in female flowers. In hermaphrodite flowers, maximum sugar content coincided with pollen presentation and declined slightly towards the female phase, indicating nectar reabsorption, whereas in female flowers sugar content did not differ between the floral phases. These differences in floral reward are discussed in relation to visitation patterns by pollinators and seed production in this species.     

Attributes of Individual Flowers of Brassica napus L. are Affected by Defoliation but not by Intraspecific Competition

To investigate the effect of intraspecific competition on floweringin Brassica napus‘Westar’, a series of 30 pots wasestablished that spanned a range of one–96 plants perpot. In each pot, the following attributes of individual flowerswere quantified: petal length; petal width; stamen length; pistillength; pollen production; diameter of pollen grains; and nectarproduction. Certain plants contained a transgenic marker thatenabled the vigour of their pollen to be quantified by screeningthe progeny from post-pollination trials against conventionalmale competitors. Plant size was strongly affected by plantdensity; plants in the lowest density pots comprised ten-timesmore dry biomass than individuals in the highest density pots.However, none of the attributes of individual flowers variedwith density. In contrast, the number of flowers per plant declinedrapidly with density. In the face of resource scarcity, theplants apparently conserved flower size rather than flower number.There was no difference in the response to density between transgenicand conventional plants, but transgenic plants obtained morefertilizations than expected in post-pollination competitionagainst conventional competitors. A separate experiment demonstratedthat B. napus is, nevertheless, capable of plasticity in certainflower attributes (petal size, stamen length) in response todefoliation. Explanations for the stability of flower size relativeto flower number are discussed. Copyright 2001 Annals of BotanyCompany Brassica napus, defoliation, density effects, flowers, flower size, intraspecific competition, nectar production, oilseed rape, pollination, transgenic plants     

Nectar robbing improves male reproductive success of the endangered Aconitum napellus ssp. lusitanicum

Nectar robbery is usually thought to impact negatively on the reproductive success of plants, but also neutral or even positive effects have been reported. Very few studies have investigated the effects of nectar robbing on the behaviour of legitimate pollinators so far. Such behavioural changes may lead to the reduction of geitonogamy or to increased pollen movement. We simulated nectar robbing in experimental sites as well as in natural populations of Aconitum napellus ssp. lusitanicum, a rare plant pollinated by long-tongued bumblebees. In an experimental setup, we removed the nectaries of 40 % of the flowers, which is similar to rates of robbing observed in wild populations. Patches of plants with experimentally robbed flowers were compared with control patches containing plants with untreated flowers. We observed pollinator behaviour, mimicked male reproductive success (pollen dispersal) using fluorescent dye, and measured female reproductive success (seed set). The main legitimate visitors were bumblebees while honeybees were often observed robbing nectar. They did so by “base working”, i.e. sliding between tepals. Bumblebees tended to visit fewer flowers per plant and spent less time per single flower when these had been experimentally robbed. This change in behaviour consequently increased the proportion of flowers visited by bumblebees in patches with robbed flowers. Fluorescent dye mimicking pollen flow was dispersed larger distances after pollinators had visited patches with robbed flowers compared to control patches. Average seed set per plant was not affected by nectar robbing. Our results demonstrated that A. napellus does not suffer from nectar robbery but may rather benefit via improved pollen dispersal and thus, male reproductive success. Knowledge on such combined effects of behavioural changes of pollinators due to nectar robbery is important to understand the evolutionary significance of exploiters of such mutualistic relationships between plants and their pollinators.     

Ultrastructure and Function of Floral Nectaries of Chamelaucium uncinatum (Myrtaceae)

Nectar is secreted for up to 11d after anthesis inChamelauciumuncinatum . The volume and sucrose concentration secreted variesbetween flowers, plants and days. The period of nectar secretioncoincides with the period of pollen presentation and stigmaticreceptivity suggesting nectar is part of an efficient reproductivestrategy inC. uncinatum . The nectary ofC. uncinatum consistsof the entire upper surface of the ovary and hypanthium. Theepidermis of the nectary is covered by a thickened cuticle whichis only broken at the sites of the numerous modified stomatawhich are scattered across its surface. It is suggested thatnectar is secreted onto the surface of the ovary via these modifiedstomata. The presence of extensive and well developed endoplasmicreticulum, mitochondria and Golgi bodies in the nectar secretingcells indicates that a granulocrine mechanism of secretion isoccurring inC. uncinatum . Chamelaucium uncinatum ; Geraldton Waxflower; floral nectaries; nectar production; modified stomata     

Pollen Presentation on Petal Combs in the Geof lorous Heath Acrotriche serrulata (Epacridaceae)

The urn-shaped flowers of Acrotriche serrulata R.Br. are unusualin two features: the mode of pollen presentation, and the corollatube which is filled with nectar at maturity, submerging thestyle and stigma. At the end of the corolla tube, the petallobes bear conspicuous subterminal hairs that usually containclumps of pollen embedded in viscous pollencoat materials. Developmentalstudies show that at anthesis, pollen from the dehisced anthersadheres to the petal hairs. When the corolla lobes reflex, thepollen is presented on the newly exposed hairs. The mature corollatube either abscises or may be removed by nectar-seeking non-flyingmarsupials, putative agents of pollination. Acrotriche serrulata, Epacridaceae, pollen presentation, pollination, geoflory, floral structure, nectar, stigma     

大和切叶蜂对其蜜源植物披针叶黄华的盗蜜行为

【目的】调查和观测内蒙古毛乌素沙地大和切叶蜂Megachile (Xanthosaurus) japonica Alfken对其蜜源植物披针叶黄华Thermopsis lupinoides (L.)的盗蜜行为。【方法】在披针叶黄华花期内, 设置样方观测披针叶黄华的主要访花昆虫。采用目测, 拍照等方法对大和切叶蜂盗蜜行为进行观测, 记录和统计花被盗蜜后留下的盗蜜孔的数量和在花上的位置。【结果】大和切叶蜂在披针叶黄华传粉蜂中数量上占有绝对的优势。作为初级盗蜜者时, 用上颚在花基部切割出一个纵向裂口, 将口器伸入孔内吸取花蜜。作为次级盗蜜者时, 利用已有的孔洞来吸蜜。在盗蜜时没有表现出寻找已经存在的盗蜜孔来吸蜜的现象, 同时其个体在盗蜜时表现出“偏好”花基部一侧的行为。在13个样地, 已开放花朵被盗蜜率最低为95.4%, 最高达到100%, 而未开放花朵的被盗蜜率最高则达到64.7%。【结论】在毛乌素沙地大和切叶蜂既是披针叶黄华的主要传粉者, 也是其初级盗蜜者和次级盗蜜者。     

Nectar Production and Floral Characteristics of Tropaeolum majus L. Grown in Ambient and Elevated Carbon Dioxide

Tropaeolum majus (nasturtiums) were grown from seed in growthcabinets, under 380 and 750 ppmv CO₂. Elevated CO₂significantlyincreased nectar secretion rate, both in flowers milked of nectardaily and in once sampled, 3-d-old flowers. Elevated CO₂didnot affect time to flowering, total number of flowers produced,pollen to ovule ratio, or the total or individual concentrationsof nectar amino acids. The dry weight and longevity of individualflowers was also unchanged. Nectar sugar content was unchangedby elevated CO₂in a subset of flowers used to assess the 3-d-oldnectar volume. This subset did not show the same increase innectar volume under elevated CO₂as the full set, resulting inthe concentration of sugars remaining unchanged. Overall, thequantity rather than the quality of the nectar changed underelevated CO₂while flower characteristics remained constant,implying that the identity of pollinators may remain the samewhile foraging behaviour (e.g. number of visits per plant, distancetravelled) may change in the future. Copyright 1999 Annals ofBotany Company Tropaeolum majus, nasturtiums, elevated CO₂, nectar, phenology, floral characteristics, amino acids, pollinator-plant interactions.     

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.     

Nectar replenishment and pollen receipt interact in their effects on seed production of Penstemon roseus

Resource supply and pollen delivery are often thought to equally limit seed production in animal-pollinated plants. At equilibrium, plants should show no response to experimental pollen supplementation because resources limit seed set above the current level of pollen attraction, while experimental reduction in pollen deposition below the equilibrium level would reduce seed set. The predicted equilibrium may be disrupted, however, if plants expend additional energy to replenish removed nectar. We investigated the combined effects of nectar removal and pollen delivery on female reproductive success of Penstemon roseus (Plantaginaceae), a hummingbird-pollinated plant that replenishes removed nectar. We first documented that the frequency of experimental nectar removal was correlated with total nectar secretion; and increased frequency of nectar removal resulted in increased female reproductive costs to the plant. Trade-offs between investing resources in nectar and investing resources in seeds were then investigated in two contrasting natural populations by removing nectar from flowers at increasing frequencies while simultaneously hand-pollinating flowers with increasing amounts of pollen. Seed set was lowest at low levels of pollen deposition, highest at medium-sized pollen loads, and intermediate when pollen loads were highest. At both sites, the frequency of nectar removal and pollen deposition had an interactive effect on seed production, in that intermediate levels of nectar removal result in the absolute highest seed set, but only at intermediate pollen loads. At high pollen loads, seed set was higher following little to no nectar removal, and at low pollen loads, all rates of nectar removal affected fecundity equally. Seed mass responded to nectar removal and pollination differently than did seed set. High levels of nectar removal and pollen delivery both lowered seed mass, with little interaction between main effects. Our findings are among the first to demonstrate that nectar replenishment costs and pollination intensity jointly affect seed production. This conflict between nectar replenishment costs and pollen-limiting factors results in trade-offs between pollinator attraction and seed provisioning. Thus, resource allocation towards nectar production should more often be considered in future studies of pollen limitation.     

Native or Exotic? Double or Single? Evaluating Plants for Pollinator-friendly Gardens

In a series of dawn-to-dusk studies, we examined the natureand accessibility of nectar rewards for pollinating insectsby monitoring insect visits and the secretion rate and standingcrop of nectar in the British native plant species Salvia pratensis,Stachys palustris, S. officinalis, Lythrum salicaria, Linariavulgaris, the non-native Calendula officinalis, Petunia x hybrida,Salvia splendens, and the possibly introduced Saponaria officinalis.We also compared single with double variants ofLotus corniculatus, Saponaria officinalis, Petunia x hybrida andCalendula officinalis. All the British species studied are nectar-rich and are recommendedfor pollinator-friendly gardens. They showed maximal secretionrates of about 10–90 µg sugar per flower h^-1, andmost had mean standing crops of about 5–60 µg sugarper flower. In all British species studied, the corolla wasdeep enough for the relatively long-tongued bumblebee Bombuspascuorum, but the shallower flowers of Lythrum salicaria werealso much visited by shorter-tongued bees and hoverflies, aswell as by butterflies. The exotic Salvia splendens, presumablycoevolved with hummingbirds in the Neotropics, has such deepflowers that British bees cannot reach the nectar except bycrawling down the corolla. With a secretion rate approaching300 µg sugar per flower h^-1and little depletion by insects,S. splendens accumulated high standing crops of nectar. S. splendens,and single and double flowers of the two probably moth-pollinatedspecies Petunia x hybrida and Saponaria officinalis, receivedfew daytime visits despite abundant nectar but Calendula waswell visited by hoverflies and bees. We compared single anddouble variants of Lotus corniculatus,Petunia x hybrida andCalendula officinalis, and also Saponaria officinalis, the lastbeing probably introduced in Britain (Stace, 1997 New floraof the British Isles. 2nd edn. Cambridge: Cambridge UniversityPress). In Petunia, Saponaria and Lotus, double flowers secretedlittle or no nectar. In Calendula, where doubling involved achange in the proportion of disc and ray florets rather thanmodification of individual flower structure, double and singlecapitula had similar standing crops of nectar. Except inCalendula, exotic or double flowers were little exploited by insect visitors.In the exotics, this was probably due to the absence or scarcityof coevolved pollinators, coupled, in double flowers, with theabsence of nectar. Copyright 2001 Annals of Botany Company Salvia pratensis, Salvia splendens, Stachys palustris, Stachys officinalis,Lythrum salicaria , Linaria vulgaris, Lotus corniculatus, Saponaria officinalis,Petunia x hybrida, Calendula officinalis, wild flowers, double flowers, gardens, nectar, secretion rate, standing crop, pollinators, bumblebees, Bombus, honeybees, Apis, hoverflies, butterflies,Anthidium manicatum     

Floral color change and the attraction of insect pollinators in lungwort (Pulmonaria collina)

We studied the effect of floral color change on long- and short-distance attraction of insect pollinators to the herb lungwort, Pulmonaria collina. Lungwort flowers change color with age from red to blue. Young red flowers had a significantly greater pollen and nectar reward and were significantly more often unpollinated than old blue ones. Red and blue flowers both influenced long-distance attractiveness of plants, defined as the number of insect approaches towards an individual plant. After reaching a plant, flower visitors preferred to visit young red flowers. Therefore, short-distance attractiveness, defined as the number of flowers visited successively on an individual plant, was influenced mainly by the number of young red flowers. The co-occurrence of the change in reproductive ability, in amount of reward, and in flower color enabled lungwort plants to direct pollinators to reproductive, highly rewarding red flowers. The data suggest that by maintaining changed flowers lungwort plants can increase their long-distance attraction and simultaneously enhance the probability of flower visits to pre-changed flowers. Thus, we propose floral color change as a mechanism that can increase the efficiency of pollen transfer to enhance plant fitness. Received: 2 November 1998 / Accepted: 14 July 1999     

The influence of nectar production on hawkmoth behavior,self pollination,and seed production in Mirabilis multiflora (Nyctaginaceae)

Nectar is the most common floral reward that plants produce to attract pollinators. To determine the effect of nectar production on hawkmoth behavior, pollen movement, and reproductive success in Mirabilis multiflora, I manipulated nectar volumes and observed the subsequent foraging behavior of the hawkmoth Hyles lineata and the resulting pollen movement patterns. Individual hawkmoths visited significantly more flowers on plants with more nectar. The increase in flower visits significantly increased pollen deposition on stigmas and pollen removal from anthers when nectar volume was raised to twice the highest level found in nature. As hawkmoths visited flowers consecutively on a plant, the proportion of self pollen deposited on stigmas increased significantly and rapidly. Based on simulated hawkmoth visits, seed set was significantly reduced for flowers later in a visit sequence. A simple model combining these results predicts that the form of selection on nectar production varies depending on pollinator abundance. Using a multiple regression analysis a nearly significant (P < 0.08) effect of stabilizing selection was detected during a single season as predicted by the model for the prevailing hawkmoth abundance. Although increased nectar production may indirectly affect plant fitness by reducing resources available for other plant functions, the direct effect of high nectar production on pollinator behavior and self pollination may generally limit floral nectar production.     

Causes and consequences of floral damage in Aconitum lycoctonum at high and low elevations in Switzerland

The outcomes of interactions among plants and the insects that use their flowers are likely to vary among the physical environments and the communities in which they grow. In this study we quantified floral damage of Aconitum lycoctonum in high (>2000 m) and low (c. 500 m) elevation populations in Switzerland. At high elevation, floral damage was frequent and was caused by nectar-robbing short-tongued bumblebees. Nectar robbers make a hole in the flower when they collect nectar. A nectar robber exclusion experiment showed that nectar robbery by short-tongued bumblebees had no effect on the female reproductive success of plants; robbing bees rarely damaged the nectaries, and damage to the petals probably does not decrease flower longevity. In addition, nectar robbers tended to collect pollen during about 10% of their visits. Thus, these bees may act as low-efficiency pollinators and may, at times, be mutualistic associates. At low elevation, the holes in the flowers were caused by beetles (Meligethes viridescens) and not by short-tongued bumblebees. The beetles eat pollen and might also consume nectar. Since the beetles gain access to pollen and nectar by entering the flower through its opening, and later chew holes while foraging on floral tissue, the beetles are pollen eaters rather than nectar robbers. Overall, our results show that not all floral damage is caused by nectar robbers and that there can be strong altitudinal variation in the causes and consequences of floral damage.     

海切叶蜂的筑巢和访花行为

【目的】研究毛乌素沙地重要野生传粉昆虫海切叶蜂Megachile maritima的筑巢和访花行为对保护其栖息环境具有重要意义。【方法】采用目测及拍照等方法对海切叶蜂的整个筑巢过程进行了连续观测;以2 m×2 m 样方的方式观测海切叶蜂的访花频率、单花停留时间及日活动规律等访花行为,其中日活动规律每天连续观测,共观测7天。【结果】海切叶蜂在沙土中筑巢,每巢只有一个巢室,其筑巢过程为：寻找合适的筑巢地点,挖巢,构建巢室,采集蜂粮,产卵,封住巢室,筑完一个巢。它连续筑完一个巢大约需要9 h。海切叶蜂构建一个巢室需要切取26～29片叶子,为每巢室采集蜂粮11～12次,每巢室内产卵1粒;在塔落岩黄芪和细叶益母草上的平均访花频率分别为（13.23±6.49）朵/min和（16.72±4.84）朵/min,平均单花停留时间分别为（3.08±2.48）s和（2.49±1.31）s。晴天,海切叶蜂在12:00～14:00期间活动较活跃。【结论】海切叶蜂不同个体之间的筑巢行为相似。该蜂在塔落岩黄芪和细叶益母草上的访花过程、访花频率及单花停留时间具有显著的差异。