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.     

Morphological variation of the Oncocyclus irises (Iris: Iridaceae) in the southern Levant

Morphological traits of Iris section Oncocyclus (Siems.) Baker in the southern Levant (Israel, Jordan, The Palestinian Authority and Sinai/Egypt) were analysed in order to clarify taxonomic relationships among taxa and the validity of diagnostic characters. Floral and vegetative characters were measured in 42 populations belonging to nine species during the peak of the flowering season in 1998–2000. Pearson's Coefficient of Racial Likelihood (CRL) was used to calculate morphological distances between populations. Twelve of the measured populations, distributed along the north-south aridity gradient in Israel, were further explored for morphological changes along the gradient. Cluster analysis revealed two major clusters: the first includes most of the dark-coloured Iris populations, with populations of I. petrana Dinsmore and I. mariae W. Barbey forming a subcluster; the second consists of all the light-coloured populations but also some dark-coloured populations. Pearson's CRL and geographical distance were significantly correlated among the dark-coloured populations. Along the geographical gradient, flower, stem and leaf size traits decrease towards the south, probably as an adaptation to aridity. This suggests that natural selection promoted the differences between populations. Almost no discrete phenotypic groups exist within the Oncocyclus species of the southern Levant except for variation in the floral colours. Most of the suggested diagnostic characters proved unreliable in that they varied continuously across populations. The taxonomical difficulties encountered in this study reflect the special evolutionary state of the Oncocyclus irises as a group in the course of speciation.  © 2002 The Linnean Society of London. Botanical Journal of the Linnean Society , 2002, 139 , 369–382.     

Pollination efficiency and foraging behaviour of honey bees and non‐Apis bees to sweet cherry

1. Crop pollination generally increases with pollinator diversity and wild pollinator visitation. To optimize crop pollination, it is necessary to investigate the pollination contribution of different pollinator species. In the present study, we examined this contribution of honey bees and non‐Apis bees (bumble bees, mason bees and other solitary bees) in sweet cherry.
2. We assessed the pollination efficiency (fruit set of flowers receiving only one visit) and foraging behaviour (flower visitation rate, probability of tree change, probability of row change and contact with the stigma) of honey bees and different types of non‐Apis bees.
3. Single visit pollination efficiency on sweet cherry was higher for both mason bees and solitary bees compared with bumble bees and honey bees. The different measures of foraging behaviour were variable among non‐Apis bees and honey bees. Adding to their high single visit efficiency, mason bees also visited significantly more flower per minute, and they had a high probability of tree change and a high probability to contact the stigma.
4. The results of the present study highlight the higher pollination performance of solitary bees and especially mason bees compared with bumble bees and honey bees. Management to support species with high pollination efficiency and effective foraging behaviour will promote crop pollination.

     

Phenological regulation of opportunities for within-inflorescence geitonogamy in the clonal species,iris versicolor (iridaceae)

Opportunities for selfing through geitonogamy are possible if more than one flower within the same clone, inflorescence, or floral unit is open at the same time. In a total of 200 inflorescences in two natural populations of Iris versicolor, flowers were observed and classified daily on the basis of anther dehiscence and stigma receptivity. Analysis of the flowering phenology demonstrated that (1) protandry limits opportunities for autogamy, (2) flowers mature sequentially within a floral unit (defined as a cluster of flowers borne on a single branch within an inflorescence), preventing the opportunity for geitonogamous fertilization between flowers of the same floral unit, and (3) 77% of all flowers had no opportunity to be pollinated by another flower within the same inflorescence. Both the number and the proportion of flowers with opportunities for geitonogamy increased with the number of flowers open in each population, and opportunities for geitonogamy also increased with the number of floral units within inflorescences. These morphological and phenological controls suggest that when selfing occurs in this species, it is most likely to occur between flowers on different inflorescences within the same clone. Since the organization of whole inflorescences in space is determined primarily by rhizome placement, clonal architecture may play an important role in mating system regulation in this species.     

Floral Lipid Chemistry of Byrsonima crassifolia (Malpigheaceae) and a Use of Floral Lipids by Centris Bees (Hymenoptera: Apidae)

Byrsonima rrassifolia (L.), a Neotropical malpighiaceous tree or treelet, has flowers that produce abundant lipids instead of nectar. Several species of Centris bees collect this oil. The floral oil shows variation between trees and can be separated into two types. One type is most common (11 of 14 samples) and consists of mono- and di-glycerides, some free fatty acids, a small amount of tri-glycerides and a trace of carbohydrate. The composition of the glycerides is predominantly (53–63%) esters of unsaturated 18C–22C fatty acids with the rest made up of 14C–18C saturated fatty acids. The second type of floral oil is similar to type one except that the samples contain large amounts of an unknown lipid more polar than the standards. Oil collected from the scopae of two representative Centris species, C. adanae and C. flavofasciata, was the same as the type one oil collected from B. rrassifolia floral elaiophores. The liquid provisions from the cells of several groundnesting Centris species was also found to be identical to B. crassifolia type one floral elaiophore oil with the exception of several minor ninhydrin positive compounds which may have been derived from the pollen which makes up part of the larval food provisions.     

Pollination and Breeding System of Vellozia squamata (Liliales: Velloziaceae): A Species of the Brazilian Cerrados

The pollination biology and breeding system of Vellozia squamata (Velloziaceae), a species of cerrado vegetation in Central Brazil, were studied. V. squamata is unusual in being pollinated by a few, generalist bee species despite having very large flowers, and having a distinctive pulsed flowering phenology. The species is self-incompatible but with a late-acting, post-fertilization rejection mechanism.     

Pollinators shift to nectar robbers when florivory occurs,with effects on reproductive success in Iris bulleyana (Iridaceae)

• Studies have indicated that florivory and nectar robbing may reduce reproductive success of host plants. However, whether and how these effects might interact when plants are simultaneously attacked by both florivores and nectar robbers still needs further investigation.
• We used Iris bulleyana to detect the interactions among florivory, nectar robbing and pollination, and moreover, their effects on plant reproductive success. Field investigations and hand‐pollination treatments were conducted on two experimental plots from a natural population, in which Experimental plot was protected from florivores and Control plot was not manipulated.
• The flower calyx was bitten by sawflies to consume the nectary, and three bumblebee species were pollinators. In addition, the short‐tongued pollinator, Bombus friseanus, was the only robber when there was a hole made by a sawfly. The bumblebee had significantly shortened flower handling time when robbing, as compared to legitimate visits. Pollinator visitation and seed production decreased significantly in damaged flowers. However, seed production per flower after supplementary hand‐pollination did not differ significantly between damaged and undamaged flowers. Compared to the Experimental plot, bumblebees visited fewer flowers per plant in a foraging bout in the Control plot.
• The flowers damaged by florivory allowed B. friseanus to shift to a nectar robber. Florivory and nectar robbing collectively decreased plant reproductive success by consuming nectar resources, which may reduce attractiveness to pollinators of the damaged flowers. However, the changes in pollinator behaviour might be beneficial to the plant by reducing the risk of geitonogamous mating.

     

Pollination by flies, bees, and beetles of Nuphar ozarkana and N. advena (Nymphaeaceae)

Nuphar comprises 13 species of aquatic perennials distributed in the temperate Northern Hemisphere. The European species N. lutea and N. pumila in Norway, the Netherlands, and Germany are pollinated by bees and flies, including apparent Nuphar specialists. This contrasts with reports of predominant beetle pollination in American N. advena and N. polysepala. We studied pollination in N. ozarkana in Missouri and N. advena in Texas to assess whether (1) there is evidence of pollinator shifts associated with floral-morphological differences between Old World and New World species as hypothesized by Padgett, Les, and Crow (American Journal of Botany 86: 1316-1324. 1999) and (2) whether beetle pollination characterizes American species of Nuphar. Ninety-seven and 67% of flower visits in the two species were by sweat bees, especially Lasioglossum (Evylaeus) nelumbonis. Syrphid fly species visiting both species were Paragus sp., Chalcosyrphus metallicus, and Toxomerus geminatus. The long-horned leaf beetle Donacia piscatrix was common on leaves and stems of N. ozarkana but rarely visited flowers. Fifteen percent of visits to N. advena flowers were by D. piscatrix and D. texana. The beetles' role as pollinators was investigated experimentally by placing floating mesh cages that excluded flies and bees over N. advena buds about to open and adding beetles. Beetles visited 40% of the flowers in cages, and flowers that received visits had 69% seed set, likely due to beetle-mediated geitonogamy of 1st-d flowers. Experimentally outcrossed 1st-d flowers had 62% seed set, and open-pollinated flowers 76%; 2nd-d selfed or outcrossed flowers had low seed sets (9 and 12%, respectively). Flowers are strongly protogynous and do not self spontaneously. Flowers shielded from pollinators set no seeds. A comparison of pollinator spectra in the two Old World and three New World Nuphar species studied so far suggests that the relative contribution of flies, bees, and beetles to pollen transfer in any one population depends more on these insects' relative abundances (and in the case of Donacia, presence) and alternative food sources than on stamen length differences between Old World and New World pond-lilies.     

Pollination ofPlectranthus vestitus (Lamiaceae) by trap-lining hovering bees in Madagascar

In a primary forest on the Central Plateau of MadagascarPlectranthus vestitus (Lamiaceae) was principally pollinated by the beePachymelus limbatus (Hymenoptera, Anthophoridae). A species ofStylogaster (Diptera, Conopidae) acted as a co-pollinator. Flower — pollinator interactions are described. The bee performed pollination while hovering, a flower-visit lasting onlyc. 0.3 seconds. Floral features such as shape and size of the corolla tube, and the lack of a landing place suggest specialization to hovering anthophorid beepollinators. Floral biology and pollination in the genusPlectranthus are discussed.     

Reproductive biology and pollinators of the invasive shrub Autumn olive (Elaeagnus umbellata Thunberg)

This study examined the reproductive biology of the invasive nitrogen-fixing shrub Elaeagnus umbellata. Hand-pollination experiments and pollinator-exclusion experiments were performed in four Illinois, U.S.A. populations to determine the breeding system of E. umbellata, and floral visitors were collected to determine pollinators in the invasive range. Although self-compatibility is a trait shown to confer invasiveness, our experiments revealed that E. umbellata is a mostly outcrossing species with a self-incompatible breeding system. Variation does exist in that a small percentage of individuals allow self-fertilization through autogamy. There is also variability among plants in the separation of male and female floral parts that may further affect selfing potential. The majority of floral visitors to E. umbellata were generalist pollinators, including bees, flies, and moths. Many of the larger insect visitors are pollinators of E. umbellata based on analysis of pollen on insect specimens, but smaller insects do not pollinate as frequently. Its ability to attract generalist pollinators means that E. umbellata will produce fruit wherever pollinators and mates occur; however, the low fruit set on open-pollinated branches contrasts with the idea of a prolifically fruiting plant. E. umbellata seems to serve as a reliable food source for many ecologically and economically significant insects, including native bumble bees (Bombus), the exotic honey bee (Apis mellifera), and armyworm (Mythimna unipuncta), a crop pest.     

The evolution of floral sonication,a pollen foraging behavior used by bees (Anthophila)

Over 22,000 species of biotically pollinated flowering plants, including some major agricultural crops, depend primarily on bees capable of floral sonication for pollination services. The ability to sonicate (“buzz”) flowers is widespread in bees but not ubiquitous. Despite the prevalence of this pollinator behavior and its importance to natural and agricultural systems, the evolutionary history of floral sonication in bees has not been previously studied. Here, we reconstruct the evolutionary history of floral sonication in bees by generating a time‐calibrated phylogeny and reconstructing ancestral states for this pollen extraction behavior. We also test the hypothesis that the ability to sonicate flowers and thereby efficiently access pollen from a diverse assemblage of plant species, led to increased diversification among sonicating bee taxa. We find that floral sonication evolved on average 45 times within bees, possibly first during the Early Cretaceous (100–145 million years ago) in the common ancestor of bees. We find that sonicating lineages are significantly more species rich than nonsonicating sister lineages when comparing sister clades, but a probabilistic structured rate permutation on phylogenies approach failed to support the hypothesis that floral sonication is a key driver of bee diversification. This study provides the evolutionary framework needed to further study how floral sonication by bees may have facilitated the spread and common evolution of angiosperm species with poricidal floral morphology.     

Pollen specialisation is associated with later phenology in Osmia bees (Hymenoptera: Megachilidae)

1. Species exhibit a range of specialisation in diet and other niche axes, with specialists typically thought to be more efficient in resource use but more vulnerable to extinction than generalists. Among herbivorous insects, dietary specialists seem more likely to lack acceptable host plants during the insect's feeding stage, owing to fluctuations in host-plant abundance or phenology. Like other herbivores, bee species vary in host breadth from pollen specialisation (oligolecty) to generalisation (polylecty).
2. Several studies have shown greater interannual variation in flowering phenology for earlier-flowering plants than later-flowering plants, suggesting that early-season bees may experience substantial year-to-year variation in the floral taxa available to them.
3. It was therefore reasoned that, among bees, early phenology could be a more viable strategy for generalists, which can use resources from multiple floral taxa, than for specialists. Consequently, it was expected that the median dates of collection of adult specimens to be earlier for generalist species than for specialists. To test this, phenology data and pollen diet information on 67 North American species of the bee genus Osmia was obtained.
4. Controlling for latitude and phylogeny, it was found that dietary generalisation is associated with significantly earlier phenology, with generalists active, on average, 11–14 days earlier than specialists.
5. This result is consistent with the generalist strategy being more viable than the specialist strategy for species active in early spring, suggesting that dietary specialisation may constrain the evolution of bee phenology—or vice versa.

     

Perfume-collecting male euglossine bees as pollinators of a basal angiosperm: the case of Unonopsis stipitata (Annonaceae)

Pollination of Unonopsis stipitata (Annonaceae) by males of two perfume-collecting bees, Euglossa imperialis and Eulaema bombiformis (Euglossini) is described. This is the first detailed account of this pollination mode in a member of a basal angiosperm family. Pollinator behaviour, identification of the odour bouquet and electrophysiological reaction of one of the two pollinators to the odour bouquet were determined. The collected odour is produced by 'osmophores' located adaxially on the petals. Starch and polysaccharides accumulated in petals are metabolized during odour emission. Mainly monoterpenes were detected in the scent samples, among them trans-carvone oxide. This molecule is thought by several authors to be the key attractant for male Eulaema bees and may be pivotal for convergent evolution of the perfume-collecting syndrome among dicotyledonous and monocotyledonous plants. It is speculated that Unonopsis , which on the basis of molecular age dating is considered a relatively recent genus of the Annonaceae (being 15–30 million years old), has diversified in relation to male euglossine bee pollinators.     

Pollination biology and breeding system of Zeyheria montana (Bignoniaceae)

The reproductive biology of Zeyheria montana was studied through field observations of flower visitors and floral events, controlled manual pollinations, and observations of pollen tube growth and ovule penetration by fluorescence microscopy. Analysis of secretory areas of the nectar chamber was made by flower dissections and histology of serial sections. The flower lasted 6–8 days, but pollen exposure and stigma receptivity occurred only up to the end of the first and second days, respectively. Pollination was effected by several species of hummingbirds, especially Colibri serrirostris. The flowers present a rudimentary, non-functional disc, and secretion of nectar is performed by corolla-borne glandular trichomes. Only hand cross-pollinated and natural-pollinated flowers set fruits. Artificially self-pollinated and non-pollinated flowers dried off after anthesis without presenting any swelling of the ovary. Almost all the ovules in selfed and crossed pistils were penetrated 96 h after pollination. However, a delay in ovule penetration in self-pollinated pistils was verified, which indicates the occurrence of late-acting self-incompatibility.We are grateful to Fundo de Apoio ao Ensino e à Pesquisa (UNICAMP) for financial support, and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the fellowship granted to the first author (this work is part of a Ph.D. thesis presented to UNICAMP, Campinas, SP, Brazil).     

Pollination ecology and pollination system of Impatiens reptans (Balsaminaceae) endemic to China

BACKGROUND AND AIMS: China is one of the centres of geographical distribution of Impatiens L. Studies of the pollination ecology of this genus in China have, until now, been unreported. Impatiens reptans, a species endemic to China, was studied. The aims were to examine the pollination ecology and pollination system of this species, to compare its pollination ecology with other Impatiens species growing in Sumatra and Japan, and to discuss possible reasons for its limited distribution. METHODS: The pollination ecology of I. reptans was studied by carrying out continuous observations within three naturally growing populations. Its pollination system was studied using different pollination methods, marking and counting pollen grains, assessing pollen viability and observing pollinator behaviour. KEY RESULTS: The flowering phase of the protandrous I. reptans lasted for 89 d. The life span of an individual flower was 3.6 d. Primary pollinators were honey-bees and bumble-bees. Secondary pollinators were diurnal hawk moths and butterflies. Bombus briviceps and Bombus sp. were nectar gatherers. The mean nectar sugar concentration was 29.5 %, and the mean value of sucrose/glucose + fructose was 0.82. The proportion of seed set ranged from 0.857 to 0.873. Distances that seeds were ejected ranged from 0.58 to 1.17 m. Percentage seed germination under controlled conditions was 23.1. Pollen viability was highest on the day of anthesis and thereafter decreased. Ratios of pollen : ovules ranged from 958.8 to 970.6. CONCLUSIONS: Impatiens reptans reproduces by means of cross-pollination. Its dependence on a specialized habitat, a narrow environmental niche, a low percentage of seed germination, and habitat loss could be reasons for its limited distribution and endemism.     

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

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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.     

Centris pallida (Hymenoptera: Apidae) male body size decreases across five decades

Historical data suggest that many bee species have declined in body size. Larger-bodied bees with narrow phenological and dietary breadth are most prone to declines in body size over time. This may be especially true in solitary, desert-adapted species that are vulnerable to climate change such as Centris pallida (Hymenoptera: Apidae). In addition, body size changes in species with size-linked behaviours could threaten the prevalence of certain behavioural phenotypes long-term. C. pallida solitary bees are found in the Sonoran Desert. Males use alternative reproductive tactics (ARTs) and are dimorphic in both morphology and behaviour. C. pallida male body size has been studied since the 1970s in the same population. The authors collected body size data in 2022 and combined it with published records from 1974–2022. The authors find a persistent decline in the mean head width of patrolling males, and shifts towards smaller body sizes in the populations of males found foraging and hovering. Both morphs declined in average body size, and the proportion of large-morph males in the population decreased by 8%. Mating males did not decline in mean body size over the last five decades. The authors discuss hypotheses related to the decline in C. pallida male head width. Finally, the authors advocate for C. pallida as an excellent study system for understanding the stability of ARTs with size-linked behavioural phenotypes.     

草豆蔻传粉生物学的研究

比较研究了花柱卷曲性植物草豆蔻 Alpinia hainanensis两种表型(上举型和下垂型)的传粉生物学特性。结果表明:草豆蔻的花柱卷曲运动节律与其他已报道的山姜属Alpinia植物基本一致,而下垂型花的花柱卷曲运动明显滞后于上举型花约2 h。草豆蔻的花柱卷曲运动中存在一些不同步的现象,甚至在同一花序上的两朵花的花柱运动节律亦会表现出不一致的现象,但上举型花的花药开裂时间却严格同步,都发生在同类型个体的柱头全部位于花药上方之后进行。草豆蔻上举型花的花蜜分泌量、单花花粉量、花粉/胚珠比率(P/O)均明显比下垂型花多,而两种表型的胚珠数、花蜜糖含量、氨基酸含量无显著差异。在草豆蔻的单花期内不同时间段进行人工授粉,上举型花在其柱头位于花药下方时进行人工授粉,其结实率明显比柱头位于花药上方时人工授粉的处理高,下垂型花则没有显著差异。在自然居群中,草豆蔻的主要传粉者是无垫蜂Ameglla sp.和两种木蜂Xylocopa sp.,但存在传粉者不足而影响结实率的现象。完全套袋、去雄和去雌(去柱头)处理的均不结实,表明草豆蔻中不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象。而人工自交和异交均具有较高的结实率,表明草豆蔻为自交亲和植物。草豆蔻的繁育系统是具花柱卷曲性运动的异花授粉的交配系统。     

Pollination in a cryptically dioecious plant Decaspermum parviflorum (Lam.) A. J. Scott (Myrtaceae) by pollen-collecting bees in Sulawesi, Indonesia

Decaspermum parviflorum is a common shrub or treelet in clearings in eastern Sulawesi, Indonesia. It is widely distributed in SE Asia. It produces numerous white flowers and small blue-black fruits. Individual shrubs were of two types in 1:1 ratio, one bearing staminate flowers, the other bearing flowers with both stamens and carpels. The 'hermaphrodite' flowers were shown to have sterile pollen so the species is functionally dioecious. The flowers and floral sprays of the two sexes differed in several measurements: the male flowers had more anthers and were larger and more numerous, making the corporate visual image of male sprays more than twice as large as that for sprays of female flowers. Flowers, individuals of which last only one day, opened only every second day, when all bushes flower synchronously. When the anthers burst in the morning, pollen and sterile pollen was collected vigorously by a variety of bees, mostly Apis dorsata and Nomia spp. Most of the pollen was collected within 1 h of anthesis. Anthesis in males takes place about 20 min earlier than in females; insect visitation follows this pattern. The minute quantity of nectar was collected by only few insects and mostly after the pollen had gone, if at all. Fruits were eaten by birds. The species displays many features typical of dioecious tropical plants with the unusual feature of pollen being the main food reward for pollinators. It was pollinated effectively in Sulawesi.