The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees,night-sheltering male bees and female solitary bees as pollinators

Background and Aims

The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.

Methods

Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.

Key Results

The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.

Conclusions

Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.     

A new species of Annona (Annonaceae) from the northeastern Guayana Shield

Annona prevostiae, a new species from the northeastern margin of the Guayana Shield, is described and illustrated.     

Non-Euglossine bees also function as pollinators of Sinningia species (Gesneriaceae) in southeastern brazil

Pollination by male and female Euglossini bees, euglossophily, was suggested for a number of neotropical Gesneriaceae species. Information on bee species other than Euglossini as pollinators of neotropical members of this family is limited, and in the tribe Sinningieae data about bee pollination are still lacking. Here, we report on floral biology and bee pollination of four Sinningia species: S. schiffneri, S. eumorpha, S. villosa, and Sinningia "canastrensis". The flower features, such as corolla size, shape, and colour, are very different among the four species, but all conform to the melittophilous syndrome. The average nectar volume and sugar amount is low in S. schiffneri, S. eumorpha, and Sinningia "canastrensis", when compared to that of S. villosa, but low nectar amounts is a general feature of Sinningia species. The main pollinators of the four species are: small Tapinotaspidini (Trigonopedia ferruginea) of S. schiffneri, large Bombini (Bombus morio) and large Centridini (Epicharis morio) of S. eumorpha, large Euglossini (Eulaema cingulata and Eufriesea surinamensis) of S. villosa, and large Euglossini (Eufriesea violascens) and Megachilini (Megachile sp.) of Sinningia "canastrensis". Out of the four species, only S. villosa is exclusively Euglossini-pollinated. The marked differences in flower features and nectar production of these Sinningia species may reflect their pollination by distinct groups of bees. These results strengthen the idea of multiple origins for the pollination systems involving bees within this genus, which is highly supported by molecular phylogenetic analyses.     

Small sweat bees (Hymenoptera: Halictidae) as potential major pollinators of melon (Cucumis melo) in the Mediterranean

In the current scenario of a general decline of the honeybee worldwide, studies on the potential of alternative bee species in pollinating cultivated plants are important. Although melon, Cucumis melo, is a crop with great commercial importance, there is very little information on its pollinating fauna in Europe, and none from the southern Mediterranean area. In a locality in central Spain, using both pan‐traps and net collections, we found that melon flowers are visited by 31 species of bees spanning four families, though only four were both dominant and constant. These four species belonged to the family Halictidae (sweat bees) and mostly (three species) to the genus Lasioglossum. Five other species could be defined as accessory: honeybee, Apis mellifera, and four other halictids. Individuals of the dominant species were smaller, on average, than those from all the other species. Observations on the frequency of pollen and nectar foraging and on flower visit duration further suggested L. malachurum as the potential key pollinator. Females of this species started to forage on melon early in the flowering season and exhibited two activity peaks in summer, thus covering the whole season. Although in other sites across continents melon seems to be more heavily pollinated by honeybees, this seems to be not the case in the Mediterranean, where sweat bees seem to be the major pollinators of this crop.     

Reproductive ecology of the basal angiosperm Trithuria submersa (Hydatellaceae)

Background and Aims

Trithuria, the sole genus in the family Hydatellaceae, is an important group for understanding early angiosperm evolution because of its sister relationship to the ancient lineage, Nymphaeales (water lilies). Although also aquatic, Trithuria differs from water lilies in that all species are extremely small, and most have an annual life form and grow in seasonal wetlands. Very little is known about their reproductive ecology. This paper reports on reproductive timing, mode of pollination and characteristics of the breeding system of Trithuria submersa in Western Australia.

Methods

Mass collections of open-pollinated plants from different ecological settings were used to characterize the reproductive developmental sequence and natural pollen reception. Hand-pollination, caging and emasculation experiments were used to measure outcross + geitonogamous pollen reception versus autonomous self-pollination in two populations over two field seasons.

Key Results

Natural outcross or geitonogamous pollination was by wind, not by water or insects, but pollen reception was extremely low. Pollen production was very low and pollen release was non-synchronous within populations. The pollen to ovule (P/O) ratio was 23·9, compared with 1569·1 in dioecious Trithuria austinensis. Stigmas became receptive before male phase and remained so until anthers dehisced and autonomous self-pollination occurred. Natural pollen loads are composed primarily of self pollen. Self- and open-pollinated plants had equivalent seed set (both >70 %). Self-pollinated plants produced seed within 17 d.

Conclusions

Autonomous self-pollination and self-fertilization are predominant in T. submersa. The low P/O ratio is not an artefact of small plant size and is inconsistent with long-term pollination by wind. It indicates that T. submersa has evolved a primarily autogamous breeding system. Selfing, along with the effect of small plant size on the speed of reproduction, has enabled T. submersa to colonize marginal ephemeral wetlands in the face of unpredictable pollination.     

Morning floral heat as a reward to the pollinators of the Oncocyclus irises

Relationships between flowering plants and their pollinators are usually affected by the amount of reward, mainly pollen or nectar, offered to pollinators by flowers, with these amounts usually positively correlated with floral display. The large Oncocyclus iris flowers, despite being the largest flowers in the East Mediterranean flora, are nectarless and have hidden pollen. No pollinators visit the flowers during daytime, and these flowers are pollinated only by night-sheltering solitary male bees. These iris flowers are partially or fully dark-colored, suggesting that they gather heat by absorbing solar radiation. Here we test the hypothesis that the dark-colored flowers of the Oncocyclus irises offer heat reward to their male solitary bee pollinators. Floral temperature was higher by 2.5°C than ambient air after sunrise. Solitary male bees emerged earlier after sheltering in Oncocyclus flowers than from other experimental shelter types. Pollination tunnels facing east towards the rising sun hosted more male bees than other aspects. We suggest that floral heat reward can explain the evolution of dark floral colors in Oncocyclus irises, mediated by the pollinators’ behavior.     

Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome

Background and Aims

If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.

Methods

Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.

Key Results

Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.

Conclusions

This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.     

Pollination ecology of Anthyllis vulneraria subsp. vulgaris (Fabaceae): nectar robbers as pollinators

This paper examines the hypothesis that nectar robbing can affect plant reproductive success either positively or negatively. To this end, I investigated various aspects of the pollination ecology of a population of the herb Anthyllis vulneraria subsp. vulgaris in northwest Spain over 5 yr. By observing floral visitors, I found that the most important pollinator species was the long-tongued bee Anthophora acervorum, which accounted for ～45% of recorded insect visits. However, just over 45% of visits were by the nectar-robbing bumble bees Bombus terrestris and B. jonellus. Although the incidence of robbing differed considerably over 5 yr of study, the frequency in every season was very high (66.4-76.5% of robbing) except for 1997 (0% robbing). Despite this high frequency of robbing, robbed flowers had a higher probability of setting fruit than nonrobbed flowers in all years of the study (mean: 82.0 vs. 51.0%; excluding 1997). This increased fruit set in robbed flowers is directly related to bumble bee behavior because the robbers' bodies came into contact with both the anthers and stigmas while robbing. Thus, the robbers effect pollination. These results suggest that the effect of nectar robbers on plant reproductive success is dependent both on the robbers' behavior and on flower/inflorescence structure. The importance of nectar-robbing bumble bees on the reproductive success of A. vulneraria and its yearly high frequency suggest that the relationship between robbers and this plant is part of a successful long-term mutualism.     

Studies in Annonaceae. XXVIII. Macromorphological variation of recent invaders in northern Central America: the case of Malmea (Annonaceae)

Cluster analysis is used to reveal patterns of macromorphological variation in a species complex of Malmea (Annonaceae) distributed in eastern Mexico, Guatemala, Belize, and Honduras. Of 53 characters, 24 are important for the clustering of 238 herbarium specimens into 12 clusters. No cluster is exclusively specified by any character or combination of characters. Neither can any geographical pattern be detected, except for the clustering of specimens from Los Tuxtlas Tropical Biological Station, Veracruz. A new subspecies from this area, M. depressa subsp. abscondita, is described. Malmea gaumeri and M. leiophylla are brought into synonymy with M. depressa. Distribution patterns of Mexican and Central American taxa of Malmea are largely concordant with those of South American taxa, which spread into Cental America after the Pliocene closure of the Isthmus of Panama. A phytogeographical novelty is presented, however, as the distribution of M. depressa subsp. abscondita within the Sierra de los Tuxtlas shows an hitherto unknown pattern: the new subspecies is distributed only on the northeastern slopes of this mountain range, and is geographically separated from M. depressa subsp. depressa, which occurs only on the southwestern slopes.     

The attractiveness of flowering herbaceous plants to bees (Hymenoptera: Apoidea) and hoverflies (Diptera: Syrphidae) in agro‐ecosystems of Central Spain

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Choice of flowers by foraging honey bees (Apis mellifera): possible morphological cues

Abstract.

• 1 Honey bees foraging for nectar on lavender (Lavandula stoechas) chose inflorescences with more of their flowers open. The number of open flowers predicted whether an inflorescence was visited by bees, inspected but rejected, or ignored. Inflorescences chosen arbitrarily by observers had numbers of open flowers intermediate between those of visited and ignored inflorescences.
• 2 Differences in morphological characters between types of inflorescence correlated with nectar volume and sugar weight per flower so that visited inflorescences had a disproportionately greater volume of nectar and weight of sugar per flower and greater variance in nectar volume.
• 3 Although there were significant associations between nectar content and the morphological characters of inflorescences, discriminant function analysis revealed discrimination on the basis of morphology rather than nectar content.
• 4 Visited inflorescences tended to have smaller than average flowers but bees tended to probe the largest flowers on visited inflorescences.
• 5 Choice of flowers within inflorescences is explicable in terms of the relationship between flower size and nectar content.

     

Pollination biology and reproductive success in four Brazilian species of Gomesa (Orchidaceae: Oncidiinae): Specific pollinators,but high pollen loss and low fruit set

Four species of Gomesa (Gomesa flexuosa, Gomesa ranifera, Gomesa cornigera, and Gomesa riograndensis), a horticulturally important, albeit scarcely studied orchid genus, were investigated. Pollination biology was studied through fieldwork in Rio Grande do Sul in Southern Brazil during two flowering seasons. Breeding systems were studied through controlled pollinations in cultivated plants. All studied species secrete floral oils through elaiophores and are exclusively pollinated by native, oil-collecting bees of the Centridini or Tapinotaspidini tribes. Pollinarium reconfiguration after withdrawal, a feature likely promoting cross-pollination, was recorded in G. cornigera and G. riograndensis. All studied species are pollinator-dependent and either predominantly or fully self-incompatible, that is, unable to set fruit following self-pollination. The used indexes of pollination efficiency consistently scored values below 1, indicating that in all studied species less than one flower is pollinated per pollinarium removed. Accordingly, pollen loss was high, ranging from 40% to 75%. Furthermore, inflorescences in natural populations displayed low fruit sets (below 13%). Gomesa flexuosa presented the highest values of pollinarium removal (36.24%) and deposition (15.34%). Significantly, this is the only studied species bearing a tabula infraestigmatica, a column swelling that bees hold to stabilize themselves while gathering the floral oils. Since pollinators were observed in all studied species spending considerable time (up to 3 min 46 s) and visiting several (up to 19) flowers, we propose that the observed low fruit set is the result of a combination of self-incompatibility, pollinator-mediated self-pollination, and pollen loss.     

Why pollinators visit only a fraction of the open flowers on a plant: The plant's point of view

We analyze the so-called ‘exponential decay’ model of pollen transfer for the case in which pollinators visit a fraction of the open flowers on a plant. The analysis is limited to the simplest case of a self-incompatible species, in which self pollen does not interfere with seed set. Plants can manipulate the number of flowers visited per pollinator approach by adjusting attractiveness of flowers so as to maximize pollen export (male fitness). In the model the length of the visitation sequence that is optimal for the plant f? always decreases with pollen deposition rate k₁, pollen uptake rate k₂, and the number of pollinator approaches to the plant X, but increases with the total number of flowers F. We assume that the average number of pollinator approaches X increases with the total number of flowers F, but slower than proportionality. It then follows that the optimal fraction of flowers a pollinator visits after arriving on the plant (f?/F) decreases with the total number of flowers F. Furthermore, the male fitness gain per flower generally decreases with number of flowers, except for the condition of ‘inefficient’ pollen transfer with very small values of k₁, k₂, and X, or when the plant has only a few flowers. The latter conditions should favour high nectar production and mass blooming. The known range of parameters suggests that male fitness increases with the total number of flowers slower than proportionality.     

Crane flies and microlepidoptera also function as pollinators in Epidendrum (Orchidaceae: Laeliinae): the reproductive biology of E. avicula

Crane flies and microlepidoptera have been recorded as pollinators in unrelated orchid groups, but these insects have never been recorded in Epidendroideae, the most species‐rich orchid subfamily, which includes one of the most diverse genera among Orchidaceae, Epidendrum. Based on data on phenology, floral morpho‐anatomy, pollinators, pollination mechanisms and breeding system, the reproductive biology of E. avicula was studied in south‐eastern Brazil. Epidendrum avicula possess osmophores that produce a citric fragrance at night. The flowers attract Tipulidae flies and several families of microlepidoptera that drink the nectar produced in a tube formed by the adnation of the labellum and column. As is common in Epidendrum, after removing the pollinarium, both crane flies and micro‐moths get trapped by the proboscis, which frightens the insects and inhibits any possible intent to immediately visit another flower. The behavior of the pollinators on flowers, plus the retention of the anther cap by the pollinarium, results in a reduction in the occurrence of geitonogamy. Because E. avicula is self‐incompatible, the consequence of pollinator behavior and the floral mechanisms tend to reduce the pollen loss. As far as we know, this is the first study to report the reproductive biology of a species of Epidendroideae pollinated by crane flies and microlepidoptera. Based on more recent concepts of plant–pollinator interactions, although E. avicula is pollinated by several species belonging to two distinct orders, suggesting an unspecialized pollination system is involved, nectar‐seeking microlepidoptera and Tipulidae flies can be recognized as a single functional group.     

Pollination of Kadsura longipedunculata (Schisandraceae), a monoecious basal angiosperm, by female, pollen-eating Megommata sp. (Cecidomyiidae: Diptera) in China

The plants of Kadsura longipedunculata (Schisandraceae) are monoecious and possess either red or yellow male flowers (the androecium), with yellow tepals, and yellow female flowers. All flower types simultaneously produce heat and floral odours (dominated by methyl butyrate) throughout a 4–5-h nocturnal period. The flowers are pollinated only by female, pollen-eating Megommata sp. (Cecidomyiidae). Pollen is the only reward, and female flowers use the same attractants as male flowers but offer no food (pollination by deceit). Open pollinated flowers in nature varied in fruit set from 8 to 92%. Megommata (subfamily Cecidomyiinae, supertribe Cecidomyiidi), consists of six described species, which feed on Coccoidea (scale insects) and are distributed worldwide.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 523–536.     

Flower specialisation: the occluded corolla of snapdragons (Antirrhinum) exhibits two pollinator niches of large long‐tongued bees

• Flower specialisation of angiosperms includes the occluded corollas of snapdragons (Antirrhinum and some relatives), which have been postulated to be one of the most efficient structures to physical limit access to pollinators. The Iberian Peninsula harbours the highest number of species (18 Iberian of the 20 species of Antirrhinum) that potentially share similar pollinator fauna.
• Crossing experiments with 18 Iberian species from this study and literature revealed a general pattern of self‐incompatibility (SI) – failure in this SI system has been also observed in a few plants – which indicates the need for pollinator agents in Antirrhinum pollination.
• Field surveys in natural conditions (304 h) found flower visitation (>85%) almost exclusively by 11 species of bee (Anthophora fulvitarsis, Anthophora plumipes, Anthidium sticticum, Apis mellifera, Bombus hortorum, Bombus pascuorum, Bombus ruderatus, Bombus terrestris, Chalicodoma lefebvrei, Chalicodoma pyrenaica and Xylocopa violacea). This result covering the majority of Antirrhinum species suggests that large bees of the two long‐tongued bee families (Megachilidae, Apidae) are the major pollinators of Antirrhinum.
• A bipartite modularity analysis revealed two pollinator systems of long‐tongued bees: (i) the long‐studied system of bumblebees (Bombus spp.) associated with nine primarily northern species of Antirrhinum; and (ii) a newly proposed pollinator system involving other large bees associated with seven species primarily distributed in southern Mediterranean areas.

     

Detection of aerosolized bacterial spores (Bacillus atrophaeus) using free-flying honey bees (Hymenoptera: Apidae) as collectors

An aerosol cloud of Bacillus atrophaeus (previously B. subtilis variety niger) spores, an anthrax surrogate, was created in a large 0.4 ha (1 ac), bee-containing, open-mesh tent. Bees from a B. atrophaeus uncontaminated hive flying through the cloud adsorbed the spores in statistically significant quantities. After removal of the B. atrophaeus contaminated hive and introduction of another B. atrophaeus uncontaminated hive, the bees again were monitored for the next few days for B. atrophaeus spores. B. atrophaeus spores accumulated on the bees bodies following their exposure to the residual B. atrophaeus contamination in the tent. The spore loads on the bees quickly returned to background levels after the hives were removed from the contaminated tent area. It may therefore be practical to use honey bee colonies to monitor foraging areas for disease-causing spores.