Armament imbalances: match and mismatch in plant-pollinator traits of highly specialized long-spurred orchids

Background

Some species of long-spurred orchids achieve pollination by a close association with long-tongued hawkmoths. Among them, several Habenaria species present specialized mechanisms, where pollination success depends on the attachment of pollinaria onto the heads of hawkmoths with very long proboscises. However, in the Neotropical region such moths are less abundant than their shorter-tongued relatives and are also prone to population fluctuations. Both factors may give rise to differences in pollinator-mediated selection on floral traits through time and space.

Methodology/Principal Findings

We characterized hawkmoth assemblages and estimated phenotypic selection gradients on orchid spur lengths in populations of three South American Habenaria species. We examined the match between hawkmoth proboscis and flower spur lengths to determine whether pollinators may act as selective agents on flower morphology. We found significant directional selection on spur length only in Habenaria gourlieana, where most pollinators had proboscises longer than the mean of orchid spur length.

Conclusions/Significance

Phenotypic selection is dependent on the mutual match between pollinator and flower morphologies. However, our findings indicate that pollinator-mediated selection may vary through time and space according to local variations in pollinator assemblages.     

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.     

Possible tobacco progenitors share long-tongued hawkmoths as pollen vectors

The putative ancestors of the allopolyploid hybrid Nicotiana tabacum have distinct flower features, apparently suited either for hawkmoth or bat pollination. This suggests that progenitors were reproductively isolated by mechanical and ethological barriers. However, the present data show that in natural populations pollen vectors could be shared by two of the possible progenitors. Pollen vectors of one of the possible male progenitors (N. otophora) were short- and long-tongued hawkmoths and a nectar-feeding bat, while those of the female ancestor (N. sylvestris) were only long-tongued hawkmoths. The latter are then the most likely vectors responsible for the presumed spontaneous hybridization. These data also suggest that interspecific pollen transfer occurred more likely in one direction.     

Nectar, nectaries, flower visitors, and breeding system in five terrestrial Orchidaceae from central Argentina

Floral nectar sugar composition, nectary anatomy, and visitors are studied in five Argentine Orchidaceae, from 18 populations. Hand-pollinations were performed to evaluate their breeding system. We found two different types of perigonal nectaries located either in the spur (Habenaria gouriieana, H. hieronymi, Habenariinae), or in the basal lateral parts of the labellum (Beadlea dutraei, Pelexia bonariensis, Stenorrhynchos orchioides, Spiranthinae). The spur ofHabenaria is a nonvascularised and nonstructural nectary. The inner epidermis bears one-celled long papillae. In bud stage, the papillae are filled with starch grains, but when the flower opens and nectar secretion starts, they show no starch grains. This fact may indicate that starch is a source for some of the secreted nectar. In the remainder genera, the lateral basal parts of the labellum are secretory. The two glands are located in the adaxial basal lateral faces of the labellum. These nectaries are structural and nonvascularised.Stenorrhynchos produces abundant, concentrated nectar (40–50%).Habenaria gourlieana accumulates copious nectar in a lower concentration (<20%), whereas the other species produce small quantities of concentrated nectar (ca. 50%). Three of the studied species have sucrose predominant nectar (Beadlea dutrael, Habenaria gourlieana, andPelexia bonariensis) whileH. hieronymi, Stenorrhynchos orchioides have hexose predominant ones. Nectar removal and/or pollination induce flower senescence.H. gouriieana is visited by sphingids,S. orchioides by hummingbirds, andB. dutrael by bees. For the two other species we did not record flower visitors.Pelexia bonariensis, B. dutrael, andS. orchiodes are self-compatible species but a pollinator is needed.     

Pollinator-Mediated Selection on Flower Color,Flower Scent and Flower Morphology of Hemerocallis: Evidence from Genotyping Individual Pollen Grains On the Stigma

To trace the fate of individual pollen grains through pollination processes, we determined genotypes of single pollen grains deposited on Hemerocallis stigmas in an experimental mixed-species array. Hemerocallis fulva, pollinated by butterflies, has diurnal, reddish and unscented flowers, and H. citrina, pollinated by hawkmoths, has nocturnal, yellowish and sweet scent flowers. We observed pollinator visits to an experimental array of 24 H. fulva and 12 F2 hybrids between the two species (H. fulva and H. citrina) and collected stigmas after every trip bout of swallowtail butterflies or hawkmoths. We then measured selection by swallowtail butterflies or hawkmoths through male and female components of pollination success as determined by single pollen genotyping. As expected, swallowtail butterflies imposed selection on reddish color and weak scent: the number of outcross pollen grains acquired is a quadratic function of flower color with the maximum at reddish color, and the combined pollination success was maximal at weak scent (almost unrecognizable for human). This explains why H. fulva, with reddish flowers and no recognizable scent, is mainly pollinated by swallowtail butterflies. However, we found no evidence of hawkmoths-mediated selection on flower color or scent. Our findings do not support a hypothesis that yellow flower color and strong scent intensity, the distinctive floral characteristics of H. citrina, having evolved in adaptations to hawkmoths. We suggest that the key trait that triggers the evolution of nocturnal flowers is flowering time rather than flower color and scent.     

Flower morphology evolution in Jaborosa (Solanaceae): shape and size variation associated with contrasting pollination modes

• Flower morphology is considered an important factor in species diversification because it may influence the efficiency of pollination in different ways (e.g. attraction and mechanical fit with different groups of pollinators). In the present study, we quantified the variation in flower morphology (i.e. shape and size) of the diverse South American genus Jaborosa Juss. (Solanaceae) in relation to contrasting pollination modes: rewarding pollination either by moths or by generalist small insects versus brood-site deceptive pollination by saprophilous flies.
• We examined variations of flower morphology in frontal (pollinator attraction) and sagittal (functional fit with pollinators) views in 12 Jaborosa species using geometric morphometric methods and comparative approaches to infer whether flower shape evolution, not attributable to flower allometry or phylogenetic relationship, is associated with shifts in pollination modes.
• We found remarkable variation in flower morphology among both Jaborosa species and pollination modes, largely in sagittal view. Evolutionary trends in shape of fly-pollinated flowers were mainly attributable to changes in developmental trajectories. Variation in flower architecture facilitated differential pollen placement – on the proboscis of moths, and either on the back or ventral region of saprophilous flies – promoting diversification of the genus.
• Diversification of shape, independent of size, in most of the studied Jaborosa species would indicate adaptation to contrasting pollination modes.

     

Interactions between hawkmoths and flowering plants in East Africa: polyphagy and evolutionary specialization in an ecological context

Hawkmoths (Lepidoptera, Sphingidae) are considered important pollinators in tropical regions, but the frequency and degree of reciprocal specialization of interactions between hawkmoths and flowers remain poorly understood. Detailed observations at two sites in Kenya over a two‐year period indicate that adult hawkmoths are routinely polyphagous and opportunistic, regardless of their proboscis length. About 700 individuals of 13 hawkmoth species were observed visiting a wide range of plant species at the study sites, including 25 taxa that appear to be specifically adapted for pollination by hawkmoths. We estimate that 277 plant species in Kenya (c. 4.61% of the total angiosperm flora) are adapted for pollination by hawkmoths. Floral tube lengths of these plants have a bimodal distribution, reflecting the existence of two hawkmoth guilds differing in tongue length. Hawkmoths exhibited strongly crepuscular foraging patterns with activity confined to a 20‐min period at dusk and, in some cases, a similar period just before dawn. Corolla tube length appears to act as a mechanical filter as the longest‐tubed plants were visited by the fewest hawkmoth species and these were exclusively from the long‐tongued guild. Tube length showed a strong positive relationship with nectar volume, even after phylogenetic correction, which implies that plants with long corolla tubes are under selection to offer relatively large amounts of nectar to entice visits by polyphagous long‐tongued hawkmoths. Our study shows that diffusely co‐evolved pollination systems involving long‐tongued hawkmoths are clearly asymmetrical, with plants exhibiting a high degree of floral specialization, while hawkmoths exhibit polyphagous behaviour. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 199–213.     

Moths pollinate four crops of Cucurbitaceae in Asia

Pollination ecology of many crops is not fully known, especially in tropical and subtropical regions. Non-bee pollinators may contribute substantially to crop yield, even if they do not receive much attention. Although moth pollination has fascinated ecologists and evolutionary biologists since Darwin, crop pollination by moths has not been well investigated and experimentally examined. Hence, we explored the pollination ecology of four cucurbit crops with crepuscular or nocturnal flowers. Lagenaria siceraria (Molina) Standl., Luffa acutangula (L.) Roxb., Trichosanthes anguina L., and Trichosanthes kirilowii Maxim. all display floral traits suggestive of moth pollination, such as opening around dusk or in evenings and secreting ample dilute nectar. We demonstrated that these crops’ flowers attracted a wide range of moth species, especially hawkmoths. The assemblage of flower-visiting moths varied according to location and season. Pollination treatments and pollen load analysis confirmed the pollination of the four crops by moths, especially hawkmoths. Our results provide evidence for the value as wild pollinators for the four crops, for which domesticated bees cannot provide reliable pollination services in practice. This study lends support to the proposal to pay more attention to the value of non-bee pollinators and to leave some areas unexploited in rural landscapes for the conservation of wild pollinators, including moths.     

UV bullseye contrast of Hemerocallis flowers attracts hawkmoths but not swallowtail butterflies

The color and patterns of animal‐pollinated flowers are known to have effects on pollinator attraction. In this study, the relative importance of flower color and color contrast patterns on pollinator attraction was examined in two pollinator groups, swallowtail butterflies and hawkmoths using two Hemerocallis species; butterfly‐pollinated H. fulva and hawkmoth‐pollinated H. citrina, having reddish and yellowish flowers in human vision, respectively. Flowers of both species have UV bullseye patterns, composed of UV‐absorbing centers and UV‐reflecting peripheries, known to function as a typical nectar guide, but UV reflectance was significantly more intense in the peripheries of H. citrina flowers than in those of H. fulva flowers. Comparison based on the visual systems of butterflies and hawkmoths showed that the color contrast of the bullseye pattern in H. citrina was more intense than that in H. fulva. To evaluate the relative importance of flower color and the color contrast of bullseye pattern on pollinator attraction, we performed a series of observations using experimental arrays consisting of Hemerocallis species and their hybrids. As a result, swallowtail butterflies and crepuscular/nocturnal hawkmoths showed contrasting preferences for flower color and patterns: butterflies preferred H. fulva‐like colored flower whereas the preference of hawkmoths was affected by the color contrast of the bullseye pattern rather than flower color. Both crepuscular and nocturnal hawkmoths consistently preferred flowers with stronger contrast of the UV bullseye pattern, whereas the preference of hawkmoths for flower color was incoherent. Our finding suggests that hawkmoths can use UV‐absorbing/reflecting bullseye patterns for foraging under light‐limited environments and that the intensified bullseye contrast of H. citrina evolved as an adaptation to hawkmoths. Our results also showed the difference of visual systems between pollinators, which may have promoted floral divergence.     

Context-dependent reproductive isolation mediated by floral scent and color

Reproductive isolation due to pollinator behavior is considered a key mode of speciation in flowering plants. Although floral scent is thought to mediate pollinator behavior, little is known about its effects on pollinator attraction and floral visitation in the wild. We used field experiments with wild hawkmoths and laboratory experiments with naïve hawkmoths to investigate attraction to and probing of flowers in response to indole, a volatile emitted by Ipomopsis tenuituba but not its close relative I. aggregata, both alone and in combination with floral color differences. We demonstrated that indole attracts wild hawkmoths to flowers, but has little effect on the rate at which those attracted moths probe flowers. In contrast, white flower color did not influence hawkmoth attraction in the field, but caused more attracted moths to probe flowers. Thus, the moths require both scent and high visual contrast, in that order, to feed at flowers at dusk. Their preference for indole-scented flowers is innate, but species-specific preference is mitigated by previous experience and plant spatial patterning. This context-dependent behavior helps explain why these Ipomopsis species show geographical variation in the extent of hybridization and may potentially explain formation of hybrid bridges in other systems of hawkmoth-pollinated plants.     

Floral contrivances and specialised pollination mechanism strongly influence mixed mating in Wrightia tomentosa (Apocynaceae)

• Reproductive success of a plant species is largely influenced by the outcome of mating pattern in a population. It is believed that a significantly larger proportion of animal‐pollinated plants have evolved a mixed‐mating strategy, the extent of which may vary among species. It is thus pertinent to investigate the key contributors to mating success, especially to identify the reproductive constraints in depauperate populations of threatened plant species.
• We examined the contribution of floral architecture, pollination mechanism and breeding system on the extent of outcrossing rate in a near‐threatened tree species, Wrightia tomentosa. The breeding system was ascertained from controlled pollination experiments. In order to determine outcrossing rate, 60 open‐pollinated progeny were analysed using an AFLP markers.
• Although the trees are self‐compatible, herkogamy and compartmentalisation of pollen and nectar in different chambers of the floral tube effectively prevent spontaneous autogamy. Pollination is achieved through specialised interaction with moths. Differential foraging behaviour of settling moths and hawkmoths leads to different proportions of geitonogamous and xenogamous pollen on the stigma. However, most open‐pollinated progeny were the result of xenogamy (outcrossing rate, tm = 0.68).
• The study shows that floral contrivances and pollination system have a strong influence on mating pattern. The differential foraging behaviour of the pollinators causes deposition of a mixture of self‐ and cross‐pollen to produce a mixed brood. Inbreeding depression and geitonogamy appear to play a significant role in sustaining mixed mating in this species.

     

Nectar usage in a southern Arizona hawkmoth community

Abstract.  1. Hawkmoths (Sphingidae) are important plant associates at two life-history stages: larvae are herbivorous, whereas adults are nectar feeders and often pollinators. The diversity and identities of plants used for nectar is poorly known, however.
2. This study takes a community-level approach to hawkmoth nectar plant usage in a semi-arid grassland habitat in southern Arizona, U.S.A.
3. Pollen carried on the proboscis was identified from over 700 individuals of 14 hawkmoth species attracted to lights over a 2-year period.
4. Two plant species dominated pollen loads, suggesting that hawkmoths use these species extensively as nectar sources: Datura wrightii (Solanaceae), a classic hawkmoth-pollinated plant, and Agave palmeri (Agavaceae), which is known to be used extensively by bats. Field surveys indicate that both species are relatively rare in the flowering community. Little or no pollen was present on the moths from the most common plant species in flower during the study.
5. The dominance of Agave in pollen loads suggests that this typically bat-pollinated species may be subsidising pollinator populations of the hawkmoth-pollinated flora.
6. Three groups of hawkmoths within this community are identified based on larval diets (reported in the literature) and adult diets (documented here): those that, at a given site, heavily exploit the same plant species at both life-history stages ( Manduca sexta and M. quinquemaculata ); those that have broad local associations at both life-history stages ( Hyles lineata ); and those that exhibit narrow but non-overlapping local associations at the two life-history stages (all other hawkmoths at this site).     

Relative importance of diurnal and nocturnal pollinators for reproduction in the early spring flowering shrub Stachyurus praecox (Stachyuraceae)

Generalized pollination systems may be favored in early spring flowering plants, as during this period pollinator activity is unpredictable. Many previous studies have concentrated on the importance of diurnal visitors in pollination, and consequently, information on the contribution of nocturnal visitors to pollination in early spring is limited. This study was conducted to evaluate the relative importance of diurnal and nocturnal pollinators in the early spring flowering dioecious shrub Stachyurus praecox (Stachyuraceae), in two temperate forests in central Japan. Visitors to the female and male flowers were observed during day and night, and their relative contributions to seed set were compared. The pollinator observations revealed that the diurnal and nocturnal insects visited both male and female flowers, and that the main flower visitors were diurnal small bees and flies as well as nocturnal settling moths. The diurnal and nocturnal flower visitors also acted as pollinators, as the pollen grains of S. praecox were attached to the insects collected from the female flowers. Pollination experiments demonstrated that the contributions of diurnal pollinators to the seed set were higher than those of the nocturnal pollinators. The results of this study indicate that S. praecox has a generalized pollination system, comprising both diurnal insects and nocturnal settling moths. Although the roles of diurnal insects are more important in the pollination of S. praecox, nocturnal settling moths may have a complementary role in early spring.     

Pollination in Lilium sargentiae (Liliaceae) and the first confirmation of long‐tongued hawkmoths as a pollinator niche in Asia

The geographical distribution of pollinators is an important factor determining the biogeographical pattern of floral evolution and diversification. In Africa, a guild of plant species has converged in a floral syndrome for pollination by long‐tongued hawkmoths (predominantly Agrius convolvuli). It is hypothesized that such floral convergence could track the geographical distribution of long‐tongued hawkmoths, so it may not be confined to Africa. We investigated the pollination biology of Lilium sargentiae E. H. Wilson, which is endemic to China and exhibits traits suggestive of long‐tongued hawkmoth pollination. Lilium sargentiae was visited by A. convolvuli as well as small beetles (Nitidulidae) but pollination was mostly or totally effected by the moth. It was consistent with other hawkmoth‐pollinated plants in terms of floral tube length, nectar traits, tepal reflectance, and scent composition. We present the first experimental evidence for the hypothesis proposed above and for hawkmoth pollination in the widespread and ornamentally and economically important genus Lilium L. in a natural habitat. Our findings imply that long‐tongued hawkmoths (especially the extremely widespread A. convolvuli) as a distinctive pollinator niche may have underlain the evolution of long‐tubed (>8 cm) flowers across different continents.     

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

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

Pollen Transport in the Dark: Hawkmoths Prefer Non Crop Plants to Crop Plants in an Agricultural Landscape

In many previous studies hawkmoths have been found to be involved in nocturnal pollination of many plants. But their role in crop pollination is still unexplored. This study tried to evaluate their potential to carry pollens of crop. Hawkmoths were collected by light traps and slides were made from the pollen attached to their body. The hawkmoths and the pollens were identified. A pollen transport network was created using the data. The species level indices of each plant species in the network were estimated. Only a small fraction of pollens were made by the crops among the total types of pollens carried by the hawkmoths. Rest of the pollens belonged to that of the non crops. The species level indices suggest the crop pollens to be relatively less important in the network as reflected in the lower values of species degree and species strength. The higher species level specialisation values of crop pollens suggested they are restricted to be carried by few hawkmoth species only. It is clearly revealed that hawkmoths carry mostly non crop pollens, not crop pollens.     

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.     

茜草科二型花柱植物滇丁香传粉精确性研究

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

Pollination biology of four sympatric species of Habenaria (Orchidaceae: Orchidinae) from southern Brazil

The pollination process and breeding system of the sympatric Habenaria johannensis, H. macronectar, H. megapotamensis and H. montevidensis was documented for native populations from Rio Grande do Sul, Brazil. All species investigated offer a nectar reward (mean values of total sugars ranging from 18 to 26%) concealed in a spur. Habenaria montevidensis is pollinated by butterflies (Hesperiidae, especially of the genus Urbanus) that carry pollinaria on their eyes; the other three species are pollinated by Sphingidae. Habenaria johannensis is pollinated by the moths Manduca rustica and M. sexta that carry the pollinaria at the base of the proboscis. Habenaria macronectar is pollinated by the moths Eumorpha labrusca and M. cf. lucetius, and these bear pollinaria between the palpi. Habenaria megapotamensis is pollinated by moths of M. cf. lucetius that bear the pollinaria on the proboscis. All species studied are self‐compatible, but pollinator dependent. They also displayed high reproductive success (ranging from 69.48 to 97.40%) and male efficiency factors slightly higher than 1, suggesting that at least one flower was pollinated for each flower acting as pollen donor. At the study sites, the investigated Habenaria spp. are isolated (in terms of pollination) by a set of factors that includes differing floral morphologies, different pollinators and/or different pollinarium placement on the pollinator. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ??, ??–??.     

Reproduction and Pollination in Central European Populations of Silene and Saponaria Species

Flower morphology, flowering phenology, flower visitors, reproductive systems, and stigmatic receptivity of six species of Silene and Saponaria (Silene alba, S. dioica, S. vulgaris, S. nutans, S. noctiflora. Saponaria officinalis) were studied from April to October 1993 and from April to June 1994 in natural populations around Giessen in Hessen/Central Germany and, additionally, in individuals grown from seeds in the Botanical Garden of the University of Giessen. With the exception of Saponaria officinalis and S. noctiflora, all species were regularly visited and pollinated by crepuscular and nocturnal moths and hawkmoths, but only one species, S. alba, was exclusively pollinated by these night-active insects. The other species showed mixed pollination syndromes in which nocturnal and diurnal insects both promoted pollen transfer. Geitonogamy or even autogamy occurred in the gynodioecious and hermaphrodite species S. vulgaris, S. nutans, S. noctiflora, and Saponaria officinalis. Silene noctiflora, the only annual species, is pseudocleistogamous; the majority of its flowers did not open, and fruit set occurred after selfing in bud.