Pollination by hummingbirds and bees in eight syntopic species and a putative hybrid of Ericaceae in Southeastern Brazil

We report on flowering phenology, floral morphology, pollinators, and nectar for eight species and a putative natural hybrid belonging to Agarista, Gaultheria and Gaylussacia that occur syntopically in a montane area. The campanulate to tubular flowers of eight out of nine Ericaceae taxa are primarily pollinated by either hummingbirds or bees. Flowering overlaps in all species but slight differences of floral shape, colour, and nectar characterize pollination by each pollinator group. Differences in floral traits are not large enough to exclude secondary pollinators. Thus, either the main pollinators of a species belonging to its syndrome, or secondary pollinators of a species belonging to different syndromes, may allow for inter-specific crosses.     

Herbivory-induced extrafloral nectar increases native and invasive ant worker survival

Ascertaining the costs and benefits of mutualistic interactions is important for predicting their stability and effect on community dynamics. Despite widespread designation of the interaction between ants and extrafloral nectaries (EFNs) as a mutualism and over 100 years of studies on ant benefits to plants, the benefits to ants have never been experimentally quantified. The success of invasive ants is thought to be linked to the availability of carbohydrate-rich resources, though reports of invasive ant visits to EFNs are mixed. In two laboratory experiments, we compared worker survival of one native (Iridomyrmex chasei) and two invasive ant species (Linepithema humile and Pheidole megacephala) exposed to herbivorized or non-herbivorized EFN-bearing plants (Acacia saligna) or positive and negative controls. We found that non-herbivorized plants did not produce any measurable extrafloral nectar, and ants with access to non-herbivorized plants had the same survival as ants with access to an artificial plant and water (unfed ants). Ants given herbivorized plants had 7–11 times greater worker survival relative to unfed ants, but there were no differences in survival between native and invasive ants exposed to herbivorized plants. Our results reveal that ants cannot induce A. saligna extrafloral nectar production, but workers of both native and invasive ant species can benefit from extrafloral nectar as much as they benefit from sucrose.     

伤害和挥发物对棉花花外蜜的诱导效应

植物花外蜜的分泌是一种植物间接防御反应。为了明确植食性昆虫、机械伤和机械伤诱导的挥发性气体在植物花外蜜诱导分泌中的作用,分析了咀嚼式口器昆虫棉铃虫Helicoverpa armigera(Hübner)、刺吸式口器昆虫棉蚜Aphis gossypii Glover取食、剪刀机械伤、剪刀机械伤+棉铃虫反吐物、针刺机械伤以及机械伤诱导挥发物、顺式-茉莉酮对棉花Gossypium hirsutum L.叶片花外蜜分泌量的影响。结果表明,棉铃虫取食、剪刀机械伤、剪刀机械伤+棉铃虫反吐物处理均显著增加了被处理叶片花外蜜的分泌量。棉花花外蜜的诱导效应在处理叶片上表现明显,并且在较幼嫩的第3片真叶上也有系统性增长。顺式-茉莉酮和机械伤挥发物处理1 d对棉花较幼嫩的第4、5片真叶花外蜜有诱导效应。棉花叶片花外蜜的诱导主要与植物组织损伤有关;不同口器类型的昆虫对棉花叶片花外蜜的诱导量不同,咀嚼式口器的棉铃虫对棉花花外蜜的诱导强度显著高于刺吸式口器的棉蚜;顺式-茉莉酮和机械伤诱导的挥发物能作为棉花植株间交流的信息物质诱导棉花幼嫩叶片花外蜜的分泌。     

The flower biology ofCephalanthera longifolia (Orchidaceae)—pollen imitation and facultative floral mimicry

Solitary bees (Halictus sp.) were found to be the effective pollinators ofCephalanthera longifolia. In the same foraging flight the bees also visit flowers ofCistus salviifolius which has a similar colour pattern.Cephalanthera offers no reward to its pollinators, but orange papillae on its labellum successfully imitate pollen ofCistus. AsCephalanthera also attracts pollinators in the absence ofCistus, this is regarded as facultative floral mimicry.     

Pollination ofDianthus gratianopolitanus (Caryophyllaceae)

Pollination ofDianthus gratianopolitanus was studied in a population of the Swiss Jura mountains. Pollinators of this plant species are reported here for the first time. The flowers were not only visited by butterflies as postulated in the literature, but also by diurnal hawkmoths (Macroglossum stellatarum) and by diurnal and nocturnal noctuid moths. — Nectar is sucrose-dominant, the sugar concentration is moderate but the amino acid concentration is high. Nectar characteristics correspond well with the syndrome ofLepidoptera-pollinated flowers. — Field observations and flower characters (colour, range of the calyx length) suggest thatDianthus gratianopolitanus is an intermediate species in the transition of butterfly to moth pollination. — Lack of reproductive success inDianthus gratianopolitanus can not be attributed to lack of suitable pollinators.     

A generalised mimicry system involving angiosperm flower colour, pollen and bumblebees’ innate colour preferences

Flower colour is a major advertisement signal of zoophilous plants for pollinators. Bees, the main pollinators, exhibit innate colour preferences, which have often been attributed to only one single floral colour, though most flowers display a pattern of two or several colours. The existing studies of floral colour patterns are mostly qualitative studies. Using a model of bee colour vision we quantitatively investigate two questions: whether or not component colours of floral colour patterns may mimic pollen signals, and whether or not bumblebees exhibit innate preferences for distinct parameters of naturally existing floral colour patterns. We analysed the spectral reflectances of 162 plant species with multicoloured flowers and inflorescences, distiniguishing between inner and outer colours of floral colour patterns irrespective of the particular structures so coloured.We found that:– The inner colour of radially symmetrical flowers and inflorescences and of zygomorphic flowers appears less diverse to bees than the peripheral colour.– The inner colour of most radial flowers and inflorescences as well as the inner colour of a large number of non-related zygomorphic flowers appears to bees to be very similar to that of pollen.– Bumblebees (Bombus terrestris) exhibit innate preferences for two-coloured over single-coloured dummy flowers in a spontaneous choice test.– Bumblebees exhibit innate preferences for dummy flowers with a large over those with a small centre area.– Bumblebees exhibit innate preferences for dummy flowers with a centre colour similar to that of pollen over those with another centre colour.Our findings support the hypotheses that the inner component of floral colour patterns could be interpreted as a generalised and little recognised form of mimicry of the colour of visually displayed pollen, that bumblebees exhibit innate preferences regarding colour and size parameters of floral colour patterns, and that these correspond to visually displayed pollen. These findings together suggest a prominent role of floral colour patterns in advertisement to and guidance of naive flower visitors.     

Contrasting effects of invasive plants in plant–pollinator networks

The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity. However, there is little information available about the effect of introduced species on the structure of such networks. We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant–pollinator networks. To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders. Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators. Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively. Invasive plants acted as pollination super generalists. The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant–pollinator networks. Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant–pollinator network. These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant–pollinator network.     

Oriole pollination ofErythrina breviflora (Leguminosae): Evidence for a polytypic view of ornithophily

Erythrina breviflora is visited by large numbers of passerine birds of which orioles (Icterus: Icteridae) are the primary pollinators. The flowers produce large quantities of nectar but they are rarely visited by hummingbirds. Inflorescence and floral morphology, and low levels of sucrose in the nectar probably explain the rarity of foraging hummingbirds. A comparison of Old WorldErythrina and their pollinators with New World species pollinated by orioles and hummingbirds suggests that parallel evolution has occurred. When the comparison is expanded to include other species pollinated by orioles, it is clear that various New WorldIcteridae, Thraupidae, etc. are ecological equivalents of Old WorldOriolidae, Pycnonotidae, Sturnidae, etc. and that flowers pollinated by these birds have similar characteristics.     

Bird pollination in South African species ofSatyrium (Orchidaceae)

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

Pollination by deceit and floral mimesis inThelymitra antennifera (Orchidaceae)

Thelymitra antennifera is found to be a general mimic of the search image of other co-blooming yellow or cream-flowered species including members of the following genera:Hibbertia, Goodenia and, to a lesser extent,Helichrysum, Pimelia andStackhousia. The most common pollinators areLasioglossum (Chilalictus) spec. and the introducedSyrphus damaster (hover fly) as well as a solitary wasp,Eurys spec.—T. antennifera acts by deceit, involving opticals (yellow colour and pollen imitation) as well as olfactory (sweet odour) signals. Capsule production is low and density dependent, a situation demonstrated in other species which have evolved pollination syndromes involving deceitful mimicry.     

Reward partitioning in Capparis spp. along ecological gradient

Summary The genus Capparis is represented in Israel by three taxons of the section spinosa. The pollinators of the three taxa are hawkmoths, crepuscular bees of the genus Proxylocopa and various daily bees. However, the efficiency of the different pollinators is very varied with respect to the individual taxons. Hawkmoths were frequently observed as pollinators of C. spinosa var. aravensis and C. ovata, but rarely on C. spinosa var. aegyptia. All the other bees, on the other hand, were found abundantly on all three taxons, all over the country. The three taxa all provide different rewards for their particular pollinators. Bees visiting C. spinosa var. aegyptia are provided with a lot of pollen but a relatively small amount of nectar, with a low sugar content. C. spinosa var. aravensis and C. ovata (the flowers visited more by hawkmoths), have less pollen but a higher amount of nectar which has a higher concentration of sugar. In drier habitats the emphasis of the reward provided by the flowers is nectar, whilst in the Mediterranean habitats it is the pollen which is the reward, without any connection to the systematic origin of the plant.     

On flower visitors and true pollinators: The case of protandrous Heracleum sphondylium L. (Apiaceae)

Hogweed (Heracleum sphondylium L.), a common European umbellifer, is very variable in terms of flower and inflorescence morphology. Its flowers are visited by numerous insects, yet little is known about the importance of the particular insect taxa. I observed umbels of two colour morphs (subspecies) of Heracleum sphondylium growing in NE Poland, which were visited by more than 108 insect species during two study seasons. Analysis of the insects' importance suggests that the most efficient pollinators are the medium-sized flies Eriozona syrphoides, and Lucilia spp. (Diptera). Bumblebees Bombus terrestris (Hymenoptera), beetles of genus Stenurella spp. and Dasytes spp. (Coleoptera) and flies Eristalis spp., Meliscaeva cinctella, Phaonia angelicae and Thricops nigrifrons also contribute to pollination of the studied plants, but their efficiency shows considerable seasonal variation. Although the dense umbels of the white flowered H. sphondylium subsp. sphondylium are generally more attractive for insect visitors than the loose yellowish inflorescences of H. sphondylium subsp. sibiricum, these taxa do not seem to attract different sets of the pollinators. For both subspecies, flowers in the staminate phase were visited significantly more often than those in the pistillate phase. Some flower visitors visited the staminate phase only, which suggests they may be parasites rather than pollinators.     

植物园迁地保护环境下纹瓣兰的繁殖生物学研究

深入了解濒危物种的繁殖特性,能为物种制定有效保护策略提供科学依据。于2022和2023年对中国科学院西双版纳热带植物园迁地保存的纹瓣兰(Cymbidium aloifolium)的繁殖生物学特征进行了观察和研究,主要包括开花物候、繁育系统、花部形态特征、昆虫传粉特征以及花的挥发性成分等。结果表明,纹瓣兰的盛花期在4月中旬,单花花期7 d;群体花期约为36 d。花粉活力在开花第1天时最高,约为47.15%,之后呈下降趋势。柱头在整个单花期一直具可授性,开花第2天可授性最强。人工授粉实验表明纹瓣兰自交亲和,不存在无融合生殖和自动自花授粉,繁育系统属于兼性异交繁育并依赖传粉者,自然结实率较低为6.9%。唇瓣的先端和基部的表皮细胞分别为锥形和指状突起,基部的指状突起细胞外附着有油脂颗粒,可能是传粉者的报酬。药帽黄色,其表皮细胞为锥形细胞,能够反射光线,可能起到吸引昆虫的作用,中华蜜蜂(Apis cerana)是纹瓣兰唯一的有效传粉昆虫。纹瓣兰的挥发性气味中(E)-乙酸-2-癸烯-1-醇酯含量最高。因此,纹瓣兰的花部结构与传粉昆虫具有显著的适应性。     

Relationships between the ant Brachymyrmex obscurior (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae)

Summary. In central Mexico, the ant Brachymyrmex obscurior Forel feeds on nectar produced by extrafloral nectaries of Acacia pennatula (Schlecht. & Cham.) Benth. However, no studies have determined whether the ants visitation is related to plant nectar availability and whether ants protect A. pennatula from herbivory. The objectives of this 2-yr study (2000–2001) were to assess whether seasonal changes in ant visitation coincide with extrafloral nectar productivity in A. pennatula and to determine whether ants protect the plant. At the end of the dry season (April–June) B. obscurior was the only ant species on A. pennatula and extrafloral nectar production is limited to this period. Exclusion experiments, performed at the end of the dry season showed that A. pennatula did not receive a protective benefit when visited by ants. Branches with ants and branches where ants are excluded had similar numbers of the nonmyrmecophile leafhopper Sibovia sp. which was the only herbivore observed under natural conditions.Received 24 March 2004; revised 4 September 2004; accepted 8 September 2004.     

Catching ants with honey: an experimental test of distraction and satiation as alternative modes of escape from flower-damaging ants

According to the distraction hypothesis, extrafloral nectaries (EFN) evolved under selection to entice ants away from floral nectaries, reducing ant-mediated damage to flowers and/or interference with pollinators. Predator-satiation, through production of nectar in either surplus flowers or EFN, provides an alternative mechanism for reducing the impact of ants as flower visitors. I tested these two hypotheses by experimentally adding EFN to flowering plants of the alpine wildflower, Polemonium viscosum, and by surveying the relationship between ant visitation and nectary number in nature. Plants of P. viscosum lack EFN and experience flower damage by ants of Formica neorufibarbus gelida. Ant behavior was compared on plants with five flowers and three experimental EFN and on controls with equal floral display, but no EFN. Addition of EFN increased flower visitation by ants. The effect of EFN on flower visitation did not depend on proximity of EFN to flowers or attractiveness of EFN to ants. Findings suggest that ants perceived patch quality on a whole plant basis, rather than responding to EFN and flowers as distinct nectar patches. Ant visitation did not keep pace with nectary number in nature. The relationship between ant visitation and nectary number per plant was weak and shallow as predicted under satiation. Ant foraging choices on experimental inflorescences showed that ants bypass flowers avoided by earlier ants, enhancing probability of escape via satiation. Results do not support the idea that EFN evolve to reduce flower visitation by ants, but show instead that nectar in surplus flowers can satiate ants and reduce their negative impacts on flower function and integrity.     

Chiropterophily and ornithophily inFreycinetia (Pandanaceae) in Samoa

In Samoa, field observations, experiments, and SEM micrographs of hair and feather samples indicate the dioecious lianaFreycinetia reineckei to be both chiropterophilous and ornithophilous with the principal pollinators being an endemic flying fox,Pteropus samoensis, and an endemic starling,Aplonis atrifuscus. Nutritional analysis of staminate and pistillate inflorescenses indicate qualitative and quantitative differences between the rewards offered by the two sexes. No nectar is presented, the principal pollinator rewards being fleshy bracts and pollen. As a result, staminate and bisexual inflorescenses sustain higher levels of pollinator damage, a factor which may tend to maintain dioecism within the population.     

Pollination biology ofOpuntia polyacantha andOpuntia phaeacantha (Cactaceae) in southern Colorado

The phenology, compatibility system, and pollinator fauna ofOpuntia polyacantha Haw. andO. phaeacantha Engelm. in southern Colorado were studied and compared. The total blooming periods overlap, but the peak of blooming differs between the species withO. polyacantha blooming first and for longer. Neither species is apomictic,O. polyacantha is largely self-incompatible andO. phaeacantha is self-compatible but may not automatically fully self-pollinate. Examination of flowering structures showed that the former has larger flowers with more perianth parts and heavier (but fewer) anthers. Further the flowers of the allogamous species are visited by greater diversity (but similar abundance) of pollinators. Medium to large bees of the generaDiadasia, Lithurge, Melissodes, Bombus, Agapostemon andMegachile were found to be effective pollinators of the Opuntias studied. The two species were compared as to their floral attributes, breeding systems, and pollinators, and the possible role of competition for pollinators is discussed with respect to its role in their evolutionary paths.     

Pollination of New CaledonianWinteraceae: Opportunistic shifts or parallel radiation with their pollinators?

Flower visitors on 12 species of New CaledonianWinteraceae were studied. The visitors were two species of ancestral moths (Sabatinca; Micropterigidae), three species of weevils (Palontus; Curculionidae), and a species of thrips. Behavior observations and pollen records suggest that the beetles and occasionally the moths serve as pollinators ofZygogynum and Exospermum, andBelliolum is pollinated primarily by thrips. The floral volatiles are simple in composition, usually dominated by short esters. Preliminary experiments showed that ethyl acetate elicited alighting, and a distinctive huddling behavior was elicited by artificially mixed fragrance. The host associations of otherSabatinca andPalontus spp. do not support the hypothesis that theWinteraceae have radiated in association with their pollinators. The available evidence supports the notion of opportunistic isolated host colonizations at some point in the radiation of the pollinator groups.     

Nectary structure and nectar presentation in Aloe castanea and A. greatheadii var. davyana (Asphodelaceae)

This paper deals with the nectary structure and nectar presentation of two species belonging to different sections of the genus Aloe: A. castanea (Anguialoe) and A. greatheadii var. davyana (Pictae). The development of the nectary was studied by means of bright field and fluorescence light microscopy and scanning electron microscopy (SEM) in three flower stages (young, intermediate, old). Both species have septal nectaries. In A. castanea, a subsidiary tissue, not present in A. greatheadii var. davyana, was found beneath the nectary epithelium. This tissue accumulated starch that was hydrolyzed during secretion. Starch was slightly accumulated around the nectary in A. greatheadii var. davyana. The distribution of chlorophyll in the ovary was also different in the two species. These anatomical differences are not, however, correlated with greater nectar production in A. castanea. In this species, the nectary seems to degenerate after secretion, while in A. greatheadii var. davyana no sign of degeneration was observed. Differences in nectar presentation among the two species may account for different pollinators visiting their flowers.     

Floral colour change in Pedicularis monbeigiana (Orobanchaceae)

We examined the effects of the retention of colour-changed flowers on long- and short-distance attractiveness of bumblebees and the likelihood of successive flower visits by bumblebees in Pedicularis monbeigiana. The lower lip changed colour with age from white to purple. Hand geitonogamous pollination significantly reduced seed production. No pollen limitation occurred in this species. Purple-phase flowers contributed minimally to pollinator attractiveness at long distance. The combination of less reproductive flowers with a lower amount of reward and floral colour change enabled plants to direct pollinators to reproductive, highly rewarding white flowers at close range. A high percentage of purple-phase flowers in an inflorescence was associated with a marked reduction in the frequency of successive flower visits to individual plants. We suggest floral colour change in P. monbeigiana may serve as a mechanism for enhancing inter-individual pollen transfer and reducing intra-individual pollen transfer.