Floral divergence and temporal pollinator partitioning in two synchronopatric species of <Emphasis Type="Italic">Vigna</Emphasis> (Leguminosae-Papilionoideae)

Exclusivity of pollinators, temporal partitioning of shared pollinators and divergence in pollen placement on the shared pollinators’ bodies are mechanisms that prevent interspecific pollen flow and minimize competitive interactions in synchronopatric plant species. We investigated the floral biology, flower visitors, pollinator effectiveness and seasonal flower availability of two syntopic legume species of the genus Vigna, V. longifolia and V. luteola, in ‘restinga’ vegetation of an island in southern Brazil. Our goal was to identify the strategies that might mitigate negative consequences of their synchronous flowering. Vigna longifolia and V. luteola were self-compatible, but depended on pollinators to set seeds. Only medium to large bees were able to trigger the ‘brush type’ pollination mechanism. Vigna longifolia, with its asymmetrical corolla and hugging mechanism, showed a more restrictive pollination system, with precise sites of pollen deposition/removal on the bee’s body, compared to V. luteola, with its zygomorphic corolla and cymbiform keel. There was a daily temporal substitution in flower visitation by the main pollinators. Vigna longifolia and V. luteola had overlapping flowering phenology but the densities of their flowers fluctuated, resulting in a seasonal partitioning of flower visitation. The differences in corolla symmetry and mainly the temporal partitioning among pollinators throughout the day and the flowering season proved to be important factors in maintaining the synchronopatry of V. longifolia and V. luteola.     

Pollination-system diversity in <Emphasis Type="Italic">Epipactis</Emphasis> (Orchidaceae): new insights from studies of <Emphasis Type="Italic">E. flava</Emphasis> in Thailand

The rewarding orchid Epipactis flava was studied in NW Thailand. Its flowers were visited by a wide range of insects, most of which served as pollinators. The most frequent pollen bearers were (in decreasing order): the cricket Homoeoxipha lycoides, stingless bees of the Tetragonula testaceitarsis/hirashimai complex, hoverflies of subfam. Syrphinae, the wasp Polybioides gracilis and sweat bees of subfam. Halictinae. We found no evidence of a link between the rheophytic habit of E. flava and its pollinator fauna. Whereas most pollinators visited the flowers to feed on nectar, females of Episyrphus alternans (Syrphidae: Syrphinae) were observed to oviposit despite the absence of prey for their young. Hence, we suggest that dual pollination systems contribute to the opportunist strategy of E. flava, and we discuss, in a phylogenetic framework, how the strategy fits in with those previously reported for Epipactis sect. Arthrochilium. The elastic attachment of the epichile (a universal trait in sect. Arthrochilium) was found to promote outcrossing, and we hypothesize that loss of the elastic hinge has provided a key innovation facilitating recurrent evolution of obligate autogamy in sect. Epipactis (which is nested in sect. Arthrochilium).     

Facilitative interactions among co-flowering <Emphasis Type="Italic">Primula</Emphasis> species mediated by pollinator sharing

We investigated flowering phenology, pollinator visitation and visitor community composition in communities of self-incompatible sympatric Primula species in a high-elevation Himalayan ecosystem. Within the tight constraints imposed by short growing seasons in such ecosystems, interactions among co-occurring plants for pollinators may vary from competition to facilitation, depending on the specifics of the system. We found that pollinator community composition changed with elevation in this system: lepidopterans were the dominant visitors at lower elevations (2200–3000 masl), bees (other than bumblebees) dominated at mid-elevations (3000–3800 masl) and bumblebees dominated at higher elevations (3800–4600 masl). However, within an elevation zone, there were no significant differences in pollinators amongst co-occurring Primula species. At a focal study site where multiple Primula species co-occurred, our results showed that even while the overall flowering periods of these species broadly overlapped, the peak flowering periods of different Primula species were temporally segregated. Upon further inferring the nature of interaction amongst co-flowering Primula species, we found that plots with higher Primula diversity (≥?2 species) and density (80–100 individuals) experienced significantly higher pollinator visitation, compared with plots with single species and low flower densities (40–50 individuals). Our results suggest that in this community of sympatric, self-incompatible Primula species, a broadly aggregated, synchronous floral display of multiple species results in pollinator facilitation by attracting a greater number of pollinator visitors. Within this broadly synchronous display, the temporal segregation of peak flowering period of individual species may reduce competition for pollinators and limit heterospecific pollen transfer.     

Pollinator-mediated impacts of alien invasive plants on the pollination of native plants: the role of spatial scale and distinct behaviour among pollinator guilds

Alien invasive plant species can affect pollination, reproductive success and population dynamics of co-flowering native species via shared pollinators. Consequences may range from reproductive competition to facilitation, but the ecological drivers determining the type and magnitude of such indirect interactions remain poorly understood. Here, we examine the role of the spatial scale of invader presence and spatially contingent behavioural responses of different pollinator groups as potential key drivers, using the invasive Oxalis pes-caprae and the self-incompatible native annual Diplotaxis erucoides as a model system. Three treatments were assigned to native focal plants: (1) invader present at the landscape scale (hectares) but experimentally removed at the floral neighbourhood scale (pa); (2) invader present at both scales (pp); (3) invader absent at both scales (aa). Interestingly, we found pronounced spatially contingent differences in the responses of pollinators: honeybees and bumblebees were strongly attracted into invaded sites at the landscape scale, translating into native plant visitation facilitation through honeybees, while bumblebees almost exclusively visited Oxalis. Non-corbiculate wild bees, in contrast, showed less pronounced responses in foraging behavior, primarily at the floral neighborhood scale. Average heterospecific (Oxalis) pollen deposition onto stigmas of Diplotaxis was low (<1 %), but higher in the pp than in the pa treatment. Hand-pollination of Diplotaxis with Oxalis and conspecific pollen, however, reduced seed set by more than half when compared to hand-pollination with only conspecific pollen. Seed set of Diplotaxis, finally, was increased by 14 % (reproductive facilitation) in the pp treatment, while it was reduced by 27 % (reproductive competition) in the pa treatment compared to uninvaded populations. Our study highlights the crucial role of spatial scale and pollinator guild driving indirect effects of invasive on co-flowering native plant species.     

Pollen load diversity and foraging niche overlap in a pollinator community of the rare <Emphasis Type="Italic">Dictamnus albus</Emphasis> L.

Bees collect pollen as an important resource for offspring development while acting as pollen vectors for the plants visited. Foraging preferences of pollinators together with plant species availability shape the web of interactions at the local scale. In this study, we focused on the bee pollinator community of a population of the rare protected perennial herb Dictamnus albus, with the aim to characterise the pollen preferences and the foraging niche overlap among species through time. Bees were sampled during four consecutive years in a natural population of D. albus, throughout the blooming period of the plant species. We performed an analysis of insect pollen loads to investigate the interactions with the study species and the co-flowering plants in the area, and to evaluate the degree of foraging overlap among pollinators. Over the study years, all bee species showed a high fidelity to D. albus (60–80%), even if some taxa preferentially collected pollen from other flowering species. The foraging niche overlap in the pollinator community decreased together with an increased diversity of co-flowering plant species. The results obtained indicate that bees preferentially forage on D. albus in the studied area, but that co-flowering species contribute to complement their diet and likely reduce competition for foraging resources. It appears therefore important to maintain a high diversity of co-flowering plants to preserve the diversity in the studied pollinator community of D. albus.     

Flowers as sleeping places for male bees: somehow the males know which flowers their females prefer

Males of solitary bees usually spend the night out of the nests. In the middle or late afternoon, they stop the patrolling behavior and move on to their sleeping places. Usually, they hang with the mandibles on small branches of the vegetation or stay inside flowers until the next day. We report the sleeping places of males of four Tapinotaspidini species on flowers of six plant species of four families. Flowers of three Iridaceae species were the most sought by males, especially flowers of Sisyrinchium scariosum which show high synchrony between anthesis and activity period of Lanthanomelissa discrepans males. Moreover, S. scariosum flowers are the most visited by females of L. discrepans which are the main pollinators; however, the role of the males as pollinators is unclear. Similar situation is evident for the interaction between males of Arhysoceble picta and Cuphea glutinosa (Lythraceae), where the males take nectar and may act as pollinators, like their females. We believe the plants are indirectly benefited by these interactions through the maintenance of the male populations of the pollinator bee species.     

Flower scent bouquet variation and bee pollinator visits in <Emphasis Type="Italic">Stevia rebaudiana</Emphasis> Bertoni (Asteraceae), a source of natural sweeteners

Pollinators provide a key service to both natural and agricultural ecosystems. Little is reported on the pollination chemoecology of Stevia rebaudiana (Asteraceae), a hermaphroditic species producing self-incompatible florets in small corymbs. We investigated the chemistry of volatiles potentially involved in its pollination system. The VOCs emitted by the corymbs of 27 F1 open-pollinated genotypes were collected by solid-phase micro-extraction and analyzed by gas chromatography–mass spectrometry (GC–MS), as well as morphometric data of the genotypes were recorded. Finally, we quantified the abundance of pollinators for each genotype. S. rebaudiana flowers were mainly visited by bees (Apidae and Halictidae), followed by hoverflies (Diptera: Syrphidae). GC–MS indicated that S. rebaudiana was characterized by a complex scent profile with large variability among F1 plants. Discriminant analysis showed that limonene, δ-elemene and bicyclogermacrene were the compounds explaining most of the scent bouquet difference between high attractive (>40 pollinators/plant) from low attractive pollinator power (<40 pollinators/plant). Limonene was the most representative VOC among plants that are more attractive to pollinators, while high emissions of δ-elemene and bicyclogermacrene were linked to plants that are less attractive to pollinators. S. rebaudiana morphometric data highlighted that, besides floral VOCs, corymb abundance and size, as well as plant height, may route pollinator visits. Overall, this study adds knowledge on floral phenology and pollinator ecological traits of S. rebaudiana, allowing a deeper understanding of its chemical ecology and pollination.     

Management affects the pollinator abundance but not the reproductive success of butterfly orchids

Management of semi-natural grasslands should be based on the requirements of plants as well as their pollinators since conditions beneficial for plants are not necessarily beneficial for their pollinators and vice versa. The factors affecting the reproductive success (fruit set) of Platanthera bifolia and Platanthera chlorantha and their pollinators in agricultural landscape and woodlands were studied. In the years 2014–2016, we observed and caught moths during the flowering period of Platanthera species (late June–mid July) in four pure P. bifolia, five pure P. chlorantha and nine mixed populations under management or no management in Estonia. We determined pollinator species richness, pollinator abundance, fruit set and visibility of Platanthera plants in each population. We found that pollinator assemblages of P. bifolia and P. chlorantha did not differ between managed and unmanaged sites. Pollinator abundance had an effect on the fruit set of P. bifolia but not on that of P. chlorantha. Presence or absence of management, visibility of plants and rarefied pollinator species richness did not affect the fruit set of either plant species. Pollinator abundance was significantly higher in unmanaged populations of both plant species but rarefied pollinator species richness was higher only in unmanaged populations of P. chlorantha. Based on our findings the recommendations for management of semi-natural grasslands are to promote larger landscape diversity for the benefit of moth abundance by leaving unmanaged patches in different parts of a grassland in different years and rotational and post-fruiting management for higher reproductive success of orchids.     

Nectar trichome structure of aquatic bladderworts from the section <Emphasis Type="Italic">Utricularia</Emphasis> (Lentibulariaceae) with observation of flower visitors and pollinators

In Utricularia, the flower spur is a nectary and in this organ, nectar is produced and stored. This study aimed to examine the structure of the nectary trichomes in four Utricularia species (Utricularia vulgaris L., U. australis R.Br., U. bremii Heer and U. foliosa L.) from the generic section Utricularia. We have investigated whether species with different spur morphology had similar spur anatomy and nectary trichome structure. In Utricularia flowers, nectar is produced by spur capitate trichomes (sessile or stalked). Our results showed that regardless of the various spur morphology, trichomes have similar architecture and ultrastructure. Head cells of these trichomes are transfer cells with an eccrine nectar secretion. Examined species differed in the micromorphology of papillae in spurs. The fly Eristalis tenax was found to be a pollinator of U. vulgaris. Small Halictidae bees seem to be pollinators of U. foliosa.     

Specific features of pollination in the Manchurian birthwort, <Emphasis Type="Italic">Aristolochia manshuriensis</Emphasis>

An analysis of pollination system in Aristolochia manshuriensis has shown that flower structure in this species is strictly adapted to cross-pollination, but the possibility of an autogamous or geitonogamous type of self-pollination with the involvement of insects is not excluded. The flowers of A. manshuriensis are most frequently visited by flies of the family Anthomyiidae, which markedly contribute to their pollination. Males account for 65% of pollinator insects collected from the flowers.     

Investigating the floral and reproductive biology of the endangered microendemic cactus <Emphasis Type="Italic">Uebelmannia buiningii</Emphasis> Donald (Minas Gerais,Brazil)

The Cactaceae are known to be amongst the most endangered plant families of the world due to reduction of their habitats and activities of collectors. As the species of the family are dependent on animals to perform cross pollination, and hence seed production, their population performance may be further negatively affected by interrupted biotic interactions. For efficient conservation of rare species, knowledge on reproductive biology and pollinators is of prime importance. In our study we focused on Uebelmannia buiningii Donald, a microendemic cactus from the Serra Negra State Park, Minas Gerais state, Brazil. During four field expeditions to three localities of the species between September 2012 and September 2013, we measured flowers, detected nectar-guides and osmophores and performed pollen viability tests. We studied the reproductive system of the species using manual self- and cross-pollination tests and observed pollinators. Our results revealed that the flowering period takes place during the dry season, between April and October, and that the diurnal flowers open between 7:00 a.m. and 5 p.m. The flowers are shortly tubular with yellow perianth-segments. We found neither nectar nor nectar-guides, and osmophores appeared as glands within the flower tube. Whereas pollen viability was 90.25%, manual cross-pollination tests have shown cross-pollination with gametophytic incompatibility. We observed two bee species visiting the flowers and acting as effective pollinators: Dialictus opacus and Plebeia sp. The combination of low reproductive activity with gametophytic incompatibility, together with the reduced number of individuals in a population and low number of populations, makes the endemic cactus U. buiningii a critically endangered species.     

Delayed selfing ensures reproductive assurance in <Emphasis Type="Italic">Utricularia praeterita</Emphasis> and <Emphasis Type="Italic">Utricularia babui</Emphasis> in Western Ghats

Numerous bladderwort (Utricularia) species are distributed worldwide, but their reproductive biology is rarely investigated. Bladderworts are known to depend on tiny organisms to meet a significant proportion of their energy requirement by trapping them in bladders. However, information on the extent of their reliance on insects for pollination success is limited. We examined the reproductive strategy of two Utricularia species viz. Utricularia praeterita and U. babui, endemic to Western Ghats, India. The main aspects of the investigation involved floral biology, breeding system, pollination mechanism, and reproductive success. Flowers of both the species are structured for outbreeding through entomophilous floral suites, herkogamy, protandrous dichogamy and sensitive lobes of the stigma. With nearly 65% natural fruit-set, both the species appeared to be sufficiently open-pollinated. However, pollinators failed to show in plants of U. praeterita while in U. babui there was an apparent mismatch between the extent of fruit-set and pollinator visits. The study demonstrated that in the absence/insufficient visits of pollinators, the two species resort to autonomous selfing. In U. babui, denser patches of plants appeared to be crucial for attracting the pollinators. Both species are self-compatible, and reproductive success is predominantly achieved by delayed autonomous selfing. The sensitive stigma in the species fails to prevent selfing due to diminished herkogamy during the late anthetic stages. It is inferred that in the pollinator-limited environment, delayed selfing contributes to absolute natural fecundity in U. praeterita, while it produces a mixed progeny in U. babui.     

Feedback effects between plant and flower-visiting insect communities along a primary succession gradient

Primary successions of glacier forelands are unique model systems to investigate community dynamics and assembly processes. However, successional changes of plant and insect communities have been mainly analysed separately. Therefore, changes in plant–insect interactions along successional gradients on glacier forelands remain unknown, despite their relevance to ecosystem functioning. This study assessed how successional changes of the vegetation influenced the composition of the flower-visiting insect assemblages of two plant species, Leucanthemopsis alpina (L.) Heyw. and Saxifraga bryoides L., selected as the only two insect-pollinated species occurring along the whole succession. In addition, we investigated the links between reproductive output of these plants and pollinator abundance through experimental exclusion of pollinators. Plant community structure changed along the succession, affecting the distribution and the abundance of insects via idiosyncratic responses of different insect functional groups. L. alpina interacted with ubiquitously distributed pollinators, while S. bryoides pollinators were positively associated with insect-pollinated plant species density and S. bryoides abundance. With succession proceeding, insect assemblages became more functionally diverse, with the abundance of parasitoids, predators and opportunists positively related to an increase in plant cover and diversity. The reproductive output of both plant species varied among successional stages. Contrary to our expectation, the obligate insect-pollinated L. alpina showed a reproductive output rather independent from pollinator abundance, while the reproductive output of the self-fertile S. bryoides seemed linked to pollinator abundance. Observing ecological interactions and using functional traits, we provided a mechanistic understanding of community assembly processes along a successional gradient. Plant community diversity and cover likely influenced insect community assembly through bottom-up effects. In turn, pollinators regulate plant reproductive output through top-down control. We emphasise that dynamics of alpine plant and insect communities may be structured by biotic interactions and feedback processes, rather than only be influenced by harsh abiotic conditions and stochastic events.     

Interpopulation variation in pollinators and floral scent of the lady’s-slipper orchid <Emphasis Type="Italic">Cypripedium calceolus</Emphasis> L.

Floral scent is a key mediator in many plant–pollinator interactions. It is known to vary not only among plant species, but also within species among populations. However, there is a big gap in our knowledge of whether such variability is the result of divergent selective pressures exerted by a variable pollinator climate or alternative scenarios (e.g., genetic drift). Cypripedium calceolus is a Eurasian deceptive lady’s-slipper orchid pollinated by bees. It is found from near sea level to altitudes of 2500 m. We asked whether pollinator climate and floral scents vary in a concerted manner among different altitudes. Floral scents of four populations in the Limestone Alps were collected by dynamic headspace and analyzed by gas chromatography coupled to mass spectrometry (GC/MS). Flower visitors and pollinators (the subset of visitors with pollen loads) were collected and identified. Preliminary coupled gas chromatographic and electroantennographic measurements with floral scents and pollinators revealed biologically active components. More than 70 compounds were detected in the scent samples, mainly aliphatics, terpenoids, and aromatics. Although several compounds were found in all samples, and all samples were dominated by linalool and octyl acetate, scents differed among populations. Similarly, there were strong differences in flower visitor spectra among populations with most abundant flower visitors being bees and syrphid flies at low and high altitudes, respectively. Pollinator climate differed also among populations; however, independent of altitude, most pollinators were bees of Lasioglossum, Andrena, and Nomada. Only few syrphids acted as pollinators and this is the first record of flies as pollinators in C. calceolus. The electrophysiological tests showed that bees and syrphid flies sensed many of the compounds released by the flowers, among them linalool and octyl acetate. Overall, we found that both floral scent and visitor/pollinator climate differ among populations. We discuss whether interpopulation variation in scent is a result of pollinator-mediated selection.     

Co-dependency between a specialist <Emphasis Type="Italic">Andrena</Emphasis> bee and its death camas host, <Emphasis Type="Italic">Toxicoscordion paniculatum</Emphasis>

Among associations of plants and their pollinating bees, mutually specialized pairings are rare. Typically, either pollen specialist (oligolectic) bees are joined by polylectic bees in a flowering species’ pollinator guild, or specialized flowers are pollinated by one or more polylectic bees. The bee Andrena astragali is a narrow oligolege, collecting pollen solely from two nearly identical species of death camas (Toxicoscordion, formerly Zigadenus). Neurotoxic alkaloids of these plants are implicated in sheep and honey bee poisoning. In this study, T. paniculatum, T. venenosum and co-flowering forbs were sampled for bees at 15 sites along a 900-km-long east–west transect across the northern Great Basin plus an altitudinal gradient in northern Utah’s Bear River Range. Only A. astragali bees were regularly seen visiting flowering panicles of these Toxicoscordion. In turn, this bee was never among the 170 bee species caught at 17 species of other prevalent co-occurring wildflowers in the same five state region (38,000 plants surveyed). Our field pollination experiments show that T. paniculatum is primarily an outcrosser dependent on pollinator visitation for most capsule and seed set. Thus, both A. astragali and two sister species of Toxicoscordion are narrowly specialized and co-dependent on each other for reproduction, illustrating a rare case of obligate mutual specialization in bee–plant interactions.     

Comparative reproductive biology reveals two distinct pollination strategies in Neotropical twig-epiphyte orchids

Members of Oncidiinae are widely known for their interactions with oil-collecting bees that explore lipophilic secretions on flowers. They may also be pollinated through food deception and the offering of nectar. Although data on breeding systems are available for many Oncidiinae orchids, little is known about the reproductive strategies in Rodriguezia, a neotropical genus of ca. 55 species. In this paper, we explore the reproductive biology of two species of Rodriguezia with distinctive morphologies: R. decora and R. lanceolata. Floral features, spectral reflectance, pollinators and pollination mechanisms, and breeding systems were studied. Both species are scentless and produce nectar as a reward. Floral nectar is secreted by a gland at the base of the labellum and stored into the sepaline spur. Rodriguezia decora reflects mainly in the blue and red regions of the light spectrum, while R. lanceolata reflects in the red region. Rodriguezia decora is exclusively visited and pollinated by butterflies, while Trochilidae hummingbirds are the pollinators of R. lanceolata. Pollinaria attach to the upper third of the proboscis of butterflies (R. decora), and to the bill of hummingbirds (R. lanceolata), during the collection of nectar from the spur. Both Rodriguezia species are self-sterile. Flower features and floral reflectance support the occurrence of psychophily in R. decora and ornithophily in R. lanceolata.     

Local extinction of a rare plant pollinator in Southern Utah (USA) associated with invasion by Africanized honey bees

Twenty-five years ago, Arctomecon humilis, a pollinator-dependent, endangered poppy globally restricted to the extreme northeastern Mojave Desert in southwestern Utah, was pollinated by native bee species and the European honey bee. Follow-up studies beginning in 2012 failed to find the two most important native bee pollinator species, one of which, Perdita meconis, is a strict poppy specialist. We had four objectives: (1) confirm the status of formerly important native bee pollinators; (2) determine the role of the Africanized honey bee which reportedly invaded southern Utah in 2008; (3) examine the effect of the ostensible change in pollinator fauna on fruit set in four populations; (4) describe the pollination proficiency of species that presently visit poppy flowers. For the fourth consecutive survey, P. meconis was absent; its local extinction in Utah now seems certain. Another previously important native pollinator, Eucera quadricincta, was very rare. Also uncommon was the European honey bee, having been largely replaced by Africanized honey bees which have become, in most populations, the prevalent pollinator. Africanized bees forage early in the day and quickly strip flowers of their copious pollen leaving little for native bees. We argue that the invasion of southern Utah by Africanized bees is the most likely cause of the severe disruption of the A. humilis pollination system. The ascension of the Africanized bee is also associated with reduced fruit set in all poppy populations, especially those where plants are sparse. Arctomecon humilis now appears to depend mostly on an invasive species for pollination.     

Increase in ant density promotes dual effects on bee behaviour and plant reproductive performance

Floral rewards do not only attract pollinators, but also herbivores and their predators. Ants are attracted by extrafloral nectaries (EFNs), situated near flowers, and may interfere with the efficiency and behaviour of pollinators. We tested the hypothesis that the impacts of ant–pollinator interactions in plant–pollinator systems are dependent on (1) the seasonal activity of EFNs, which increase ant abundance closer to flowers; (2) consequently, an ant effect, where ants decrease the temporal niche overlap of bees due to predator avoidance; and (3) ant density, where higher densities may negatively affect plant–pollinator interactions and plant performance. We studied two ant–plant–pollinator systems based on Banisteriopsis campestris and Banisteriopsis malifolia plant species. The periods of high ant abundance coincided with plant species blooming. The presence of ants around flowers reduced the visitation rates of the smaller bees and the temporal niche overlap between bee species was not higher than randomly expected when ants had free access. Additionally, we observed variable ant effects on fruit set and duration of bee visits to both Malpighiaceae species when ant density was experimentally kept constant on branches, especially on B. campestris. Our goal was to show the dual role of ant density effects, especially because the different outcomes are not commonly observed in the same plant species. We believe that reduced temporal niche overlap between floral visitors due to ant presence provides an opportunity for smaller bees to improve compatible pollination behaviour. Additionally, we concluded that ant density had variable effects on floral visitor behaviours and plant reproductive performance.     

Intersexual mimicry and flowering phenology facilitate pollination in a dioecious habitat specialist species, <Emphasis Type="Italic">Myristica fatua</Emphasis> (Myristicaceae)

Myristica fatua is a dioecious specialist species restricted to the endangered, freshwater Myristica swamp forests in the Western Ghats, India. Earlier studies have alluded to pollination by deception in members of the Myristica genus, and thus we examined the pollination ecology comprising floral biology, flower production, flower visitors, and reproductive success in M. fatua and inferred the potential strategies that could permit such deception in this habitat specialist tree. Male flowers provide pollen rewards for an extended period of time while female flowers are rewardless and both sexes are visited by generalist insects, mainly by honeybees and stingless bees. Bee visits were significantly more frequent and longer on male than on female flowers as bees collected pollen from male flowers. We found that flower production patterns create a preponderance of males compared to females in the swamp populations. Using a model of honeybee color vision, we found the distance between the color loci of male and female flowers and based on minimum visual angle subtended by these flowers, we suggest that the two floral sexes cannot be discriminated by bees. Bees are likely deceived by the perceptual similarity of rewardless female flowers to pollen-offering male flowers and pollination is the consequence of foraging errors made by pollinators that encounter largely male–rarely female flower mosaics as they forage among clump-distributed M. fatua trees in the swamp habitat.     

Effects of alien invasion by <Emphasis Type="Italic">Bombus terrestris</Emphasis> L. (Apidae) on the visitation patterns of native bumblebees in coastal plants in northern Japan

When alien pollinator species enter a native community of pollinators in which resource partitioning has been established, the pollination network between plants and pollinators may be modified through the interactions between the pollinators over the use of floral resources. We observed the floral-use patterns of native (Bombus hypocrita and B. deuteronymus) and alien (B. terrestris) bumblebee species in a coastal grassland in northern Japan. We analyzed the factors determining resource partitioning patterns. B. hypocrita tended to visit flowers with shallow or wide open corollas, such as Rosa rugosa, whereas B. deuteronymus visited flowers with complex or deeper corollas, such as Lathyrus japonicus. Given the wider floral preference of B. terrestris, floral use by the alien bumblebees consistently overlapped with that of native bumblebees. The visitation of B. terrestris to R. rugosa flowers was positively correlated with that of B. hypocrita. These bumblebee species frequently used similar floral resources, in part because of the large overlap in the seasonality of their foraging activity. The visitation frequency of B. deuteronymus to L. japonicus flowers was independent of the visitation frequency of other bumblebee species. The major visitation periods of the bumblebees to L. japonicus flowers reciprocally differed between B. deuteronymus and B. terrestris, suggesting phenological resource partitioning between these species. Our study suggests that phenological niche partitioning is more common in specialized flowers (L. japonicus) than in generalized flowers (R. rugosa).