Native flower strips increase visitation by non-bee insects to avocado flowers and promote yield

Pollination is an essential ecosystem service for pollinator-dependent crops and plant communities. Apis mellifera L. is by far the most commonly used species to obtain this service in agriculture. However, there is growing evidence of the importance for crop yields of the service provided by wild bees and non-bee insect pollinators. Establishing flower resources in agricultural landscapes is a management practice that can increase insect pollinator populations and improve crop yields. We established perennial native flower strips (NFS) in four avocado orchards in central Chile during autumn 2017. We monitored flower visitors and counted newly formed fruits in avocados near and far from NFS in spring 2019, to assess flower visitor groups, flower visitation rates and fruit numbers. Only A. mellifera visited avocado flowers within bees, whereas both the managed A. mellifera and wild bees were the main visitors to the NFS. NFS increased visitation rates to adjacent avocado of flies, and with a trend for the sum of all non-managed flower visitors (i.e. excluding A. mellifera). However, there were no differences in the rates of A. mellifera visitation to avocados near and far from NFS. Avocado fruit numbers were higher among avocados near NFS than among those farther away. This difference could be due to better pollination by the increased visits to avocado flowers by flies or other wild insects. Therefore, NFS could contribute to crop fruit number, as well as conservation of native flora, wild bees and non-bee pollinators on fruit farms in the “Central Chile” biodiversity hotspot.     

Moths complement bumblebee pollination of red clover: a case for day-and-night insect surveillance

Recent decades have seen a surge in awareness about insect pollinator declines. Social bees receive the most attention, but most flower-visiting species are lesser known, non-bee insects. Nocturnal flower visitors, e.g. moths, are especially difficult to observe and largely ignored in pollination studies. Clearly, achieving balanced monitoring of all pollinator taxa represents a major scientific challenge. Here, we use time-lapse cameras for season-wide, day-and-night pollinator surveillance of Trifolium pratense (L.; red clover) in an alpine grassland. We reveal the first evidence to suggest that moths, mainly Noctua pronuba (L.; large yellow underwing), pollinate this important wildflower and forage crop, providing 34% of visits (bumblebees: 61%). This is a remarkable finding; moths have received no recognition throughout a century of T. pratense pollinator research. We conclude that despite a non-negligible frequency and duration of nocturnal flower visits, nocturnal pollinators of T. pratense have been systematically overlooked. We further show how the relationship between visitation and seed set may only become clear after accounting for moth visits. As such, population trends in moths, as well as bees, could profoundly affect T. pratense seed yield. Ultimately, camera surveillance gives fair representation to non-bee pollinators and lays a foundation for automated monitoring of species interactions in future.     

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.     

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.     

Reproductive biology of Echinopsis terscheckii (Cactaceae): the role of nocturnal and diurnal pollinators

The aim of this study was to analyse the reproductive biology of Echinopsis terscheckii, a species endemic to northwest Argentina that has nocturnal flowers. We expected that this species had a generalised pollination system, with moths and diurnal visitors as the primary pollinators. To test this, we studied the floral biology, breeding system and floral visitors of this species and the effectiveness of nocturnal and diurnal visitors. Floral biology was defined based on floral morphology, floral cycle and nectar production of the flowers. The breeding system and relative contributions of diurnal and nocturnal visitors to fruit and seed set were analysed through field experiments. E.?terscheckii flowers opened at sunset and closed the following day. The peak of nectar production occurred at midnight. Flowers were determined to be self-incompatible. Moths, bees and birds were identified as floral visitors. Moths were the most frequent visitors at night, whereas bees were the most frequent visitors during the day. Fruit production by diurnal pollinators was less than that by nocturnal pollinators; among all floral visitors, moths were the most effective pollinators. We have demonstrated for the first time that moths are the primary pollinators of columnar cacti of the genus Echinopsis. Our results suggest that moths might be important pollinators of columnar cactus species with nocturnal flowers in the extra-tropical deserts of South America.     

Contribution of diurnal and nocturnal insects to the pollination of Jatropha curcas (Euphorbiaceae) in southwestern China

Jatropha curcas L. (Euphorbiaceae) is being increasingly planted worldwide, but questions remain regarding its pollination biology. This study examined the contribution of diurnal and nocturnal insects to the pollination of monoecious J. curcas, through its floral biology, pollination ecology, and foraging behavior of potential pollinators. Nectar production of both male and female flowers peaked in the morning, declined in the afternoon, and rapidly bottomed during the night in all of their anthesis days. The diurnal visitors to the flowers of J. curcas are bees and flies, and the nocturnal visitors are moths. Flowers received significantly more visits by diurnal insects than by nocturnal insects. Through bagging flowers during night or day or both or exclusion, we compared fruit and seed production caused by diurnal and nocturnal pollinators. Both nocturnal and diurnal visitors were successful pollinators. However, flowers exposed only to nocturnal visitors produced less fruits than those exposed only to diurnal visitors. Thus, diurnal pollinators contribute more to seed production by J. curcas at the study site.     

Generalized pollination system: Are floral traits adapted to different pollinators?

Flowers that are pollinated both during the day and at night could exhibit two different groups of pollinators and produce two different sets of attractants and rewards. We explored the patterns of emission of flower scents and production of nectar in the cactus Echinopsis chiloensis ssp. chiloensis, in relation to the patterns of activity of its diurnal and nocturnal pollinators. We measured frequency of flower visitors, analyzed floral scents, measured nectar production and sugar concentration, and performed pollination exclusion experiments. Bees were the main visitors at daytime and hawkmoths at nighttime. Diurnal scents were dominated by several compounds that can attract a wide range of pollinators, whereas nocturnal scents were less diverse and were dominated by (E)-nerolidol, a compound eliciting antennal responses in hawkmoths. Nectar volume and sugar concentration at night were similar to those recorded in hawkmoth-pollinated flowers. Daytime nectar volume was higher than those commonly found in bee-pollinated flowers, but similar to those found in flowers pollinated by several pollinators. Daytime sugar concentration was similar to those recorded in bee-pollinated flowers. Flowers of E. chiloensis ssp. chiloensis seem morphologically adapted to hawkmoth pollination, but diurnal and nocturnal pollinators contribute to similar extents to reproductive success. Additionally, diurnal and nocturnal pollinators showed a synergic effect on the product of fruit set and seed set. The results are discussed in terms of the linkage between floral traits and perception abilities and requirements of pollinators.     

Flower‐visiting bat species contribute unequally toward agricultural pollination ecosystem services in southern Thailand

The large majority of angiosperm species depend on animals for pollination, including many agricultural crops, and plant‐pollinator interactions have been extensively studied. However, not all floral visitors actually transfer pollen, and efforts to distinguish true pollinators from mere visitors are particularly scarce among the bat pollination literature. To determine whether Old World bat species are equally effective pollinators in mixed‐agricultural areas of southern Thailand, we examined six night‐blooming plant taxa and quantified pollinator importance (PI) of seven common nectarivorous bat species. PI was calculated as the product of nightly bat visitation rate (obtained from mist‐netting data) and pollen transfer efficiency (estimated from bat pollen loads). We found that PI varied by both bat species and plant species. In general, the nectar‐specialist bat species were more important pollinators, yet their order of importance differed across our focal plant species. In addition, PI was dictated more by pollen transfer effectiveness than visitation rate. Our findings highlight the importance of Old World bat pollinators within southern Thailand's mixed‐agricultural landscape and illustrate how seemingly similar floral visitors can have very different contributions toward plant pollination success.     

Native bees buffer the negative impact of climate warming on honey bee pollination of watermelon crops

If climate change affects pollinator‐dependent crop production, this will have important implications for global food security because insect pollinators contribute to production for 75% of the leading global food crops. We investigate whether climate warming could result in indirect impacts upon crop pollination services via an overlooked mechanism, namely temperature‐induced shifts in the diurnal activity patterns of pollinators. Using a large data set on bee pollination of watermelon crops, we predict how pollination services might change under various climate change scenarios. Our results show that under the most extreme IPCC scenario (A1F1), pollination services by managed honey bees are expected to decline by 14.5%, whereas pollination services provided by most native, wild taxa are predicted to increase, resulting in an estimated aggregate change in pollination services of +4.5% by 2099. We demonstrate the importance of native biodiversity in buffering the impacts of climate change, because crop pollination services would decline more steeply without the native, wild pollinators. More generally, our study provides an important example of how biodiversity can stabilize ecosystem services against environmental change.     

Diurnal and nocturnal pollination of Silene alba (Caryophyllaceae)

Flowers that are open for >12 h may be visited by both diurnal and nocturnal pollinators. I compared the effectiveness (measured as seed production and pollen movement distance) of diurnal and nocturnal pollinators of Silene alba, a species whose flowers open in evening but close by midmorning the following day. By bagging flowers either during evening hours or during daylight hours or both day and night, I compared seed production caused by diurnal and nocturnal pollinators. Flowers exposed only to nocturnal visitors (mostly sphingid and noctuid moths) produced significantly more seeds than flowers exposed only to diurnal visitors (bees, flies, and wasps). Fluorescent dye applied to anthers moved significantly further and to more stigmas at night than during the day. In both measures of pollination effectiveness, nocturnal-visiting moths are better pollinators of S. alba than are the diurnal-visiting bees, flies, and wasps. These data support the hypothesis that floral phenology is an adaptation to expose flowers to the most effective pollinators.     

Of feathers and fur: Differential pollinator roles of birds and small mammals in the grassland succulent Aloe peglerae

The pollination syndrome of the genus Aloe, with gaudy inflorescences and orange–red flowers, suggests bird pollination. However, the diversity of flowering phenologies and structures suggests that generalizations within the group are currently uninformative because few studies have addressed the role of multiple pollinator guilds, especially mammals. Aloe peglerae, endemic to montane grassland of the Magaliesberg Mountains, South Africa, is primarily bird pollinated, although small mammals are recorded nocturnal visitors. We compared the independent contributions of diurnal and nocturnal visitors, i.e. birds and small mammals respectively, to reproductive success by assigning 12 flowering A. peglerae plants to each of four pollinator selective exclusion treatments: (i) no‐visitors; (ii) nocturnal‐visitors; (iii) diurnal‐visitors; and (iv) all‐visitors (control). Birds alone contributed more to fruit set (1.2–2.6 times), average seed/fruit (1.0–1.8 times) and total seed production (1.1–2.8 times) than both the all visitors (birds and small mammals combined) and the small mammal visitors respectively. The exclusion of all visitors resulted in no fruit set, suggesting that A. peglerae requires pollinators to set seed. Germination trials over six weeks and viability testing with tetrazolium staining identified no significant differences among the three treatments that produced fruit and seed. Germination success was 90–97%, indicating that seed quality of small‐mammal and bird‐pollinated plants are similar. Aloe peglerae inflorescences were visited continuously throughout the day and night, with the Cape rock‐thrush (Monticola rupestris) and dark‐capped bulbul (Pycnonotus tricolor) being the most abundant pollinators, accounting for 68.2% and 13.7% of all visits respectively in the control treatment. Small mammals, i.e. Namaqua rock mouse (Micaelamys namaquensis) and eastern rock sengi (Elephantulus myurus), while not increasing reproductive success in the absence of birds, are suitable alternative pollinators. The presence of diurnal bird and nocturnal small mammal visitors provides a diversity of pollinators in this resilient pollination syndrome.     

Mass flowering crops enhance pollinator densities at a landscape scale

To counteract the decline of pollinators in Europe, conservation strategies traditionally focus on enhancing the local availability of semi‐natural habitats, as supported by the European Union's Common Agriculture Policy. In contrast, we show that densities of bumblebees, an important pollinator group in agroecosystems, were not determined by the proportion of semi‐natural habitats in agricultural landscapes. Instead, bumblebee densities were positively related to the availability of highly rewarding mass flowering crops (i.e. oilseed rape) in the landscape. In addition, mass flowering crops were only effective determinants of bumblebee densities when grown extensively at the landscape scale, but not at smaller local scales. Therefore, future conservation measures should consider the importance of mass flowering crops and the need for management schemes at landscape level to sustain vital pollination services in agroecosystems.     

Culturally valuable minority crops provide a succession of floral resources for flower visitors in traditional orchard gardens

Agricultural intensification typically has detrimental effects on pollinator communities, but diverse cropping systems that contain sequentially-flowering crops have the potential to benefit pollinators through the provision of additional floral resources. In this study we investigate the importance of cultivated flora for flower visitors in ten agricultural gardens in South Sinai, Egypt. Insect-flower interactions in gardens and unmanaged plots were surveyed across a four-month period in two environmentally distinct years (pre-flood and post-flood). Despite containing an equal abundance and diversity of wild plants as unmanaged habitat, gardens supported a higher abundance and diversity of flower visitors due to the additional presence of cultivated flora. Visitation networks exhibited dramatic intra-annual changes in composition, with cultivated plants becoming increasingly important in later months. Trends were highly conserved across 2 years despite highly contrasting rainfall. Several key crop species were strongly involved in shaping the structure of the networks, the majority of which were herbs with strong cultural significance (fennel, rosemary, mint) and grown incidentally alongside the primary orchard crops. Minority crops are frequently overlooked in agricultural systems due to their low economic value, but we show that they can have a dramatic influence upon the structure of visitation networks, increasing both pollinator abundance and diversity, and emphasising the link between cultural practices and biodiversity conservation.     

Generalized and complementary pollination system in the Andean cactus Echinopsis schickendantzii

The considerable floral diversity present in the cactus family has often been associated with the specificity of its pollinators. However, many cactus pollination systems are generalized as their flowers are pollinated by a wide spectrum of animals. For example, cactus species with white flowers, nocturnal anthesis and extended floral cycles would present generalized pollination systems in which both nocturnal and diurnal visitors could be effective pollinators. In this article, we tested this hypothesis by studying the pollination biology of Echinopsis schickendantzii, an Andean cactus with sphingophilous flowers. In addition, we evaluated whether the cactus’s pollination system is complementary or redundant regarding the relative contributions of nocturnal and diurnal pollinators. Specifically, we studied the floral cycle, the reproductive system and the pollination effectiveness of floral visitors. The flowers of E. schickendantzii are self-incompatible; they opened at crepuscule and have an extended floral cycle. Moths were frequent visitors at night, whereas bees were frequent visitors during the day; both were effective pollinators of the cactus. Our results indicated that the flowers of this species present phenotypic, functional and ecological generalization, and their fruit set is determined by the contributions of both pollinator functional groups, i.e., they have complementary pollination systems. These results support the hypothesis that cacti in the extra-tropical deserts of South America have generalized pollination systems.     

Pollination crisis in the butterfly-pollinated wild carnation Dianthus carthusianorum?

Knowledge of pollination services provided by flower visitors is a prerequisite for understanding (co)evolutionary processes between plants and their pollinators, for evaluating the degree of specialization in the pollination system, and for assessing threats from a potential pollination crisis. This study examined pollination efficiency and visitation frequency of pollinators--key traits of pollinator-mediated fecundity--in a natural population of the wild carnation Dianthus carthusianorum. The five lepidopteran pollinator species observed differed in pollination efficiency and visitation frequency. Pollinator importance, the product of pollination efficiency and visitation frequency, was determined by the pollinator's visitation frequency. Pollination of D. carthusianorum depended essentially on only two of the five recorded pollinator species. Seed set was pollen-limited and followed a saturating dose-response function with a threshold of c. 50 deposited pollen grains for fruit development. Our results confirm that D. carthusianorum is specialized to lepidopteran pollinators, but is not particularly adapted to the two main pollinator species identified. The local persistence of D. carthusianorum is likely to be at risk as its reproduction depends essentially on only two of the locally abundant, but generally vulnerable, butterfly species.     

Hand pollination of global crops – A systematic review

Global pollinator declines and land-use change can lead to pollination limitation with implications for agricultural productivity. Hand pollination is used in agricultural production as a technique to manually pollinate crops. But the prevalence of hand pollination, as well as benefits and costs, remain unknown. We systematically reviewed the literature for examples, methods, drivers, and economic motivations of hand pollination. Furthermore, we discuss the risks, constraints, and opportunities of hand pollination. We found evidence for 20 hand-pollinated crops, including minor but also economically important crops (e.g. apple, oil palm, cacao). The lack of pollinators was the most important reason for the application of hand pollination (50% of crops), while insufficient proportion or proximity of pollinizers (8% of crops) and skewed sex ratio or dichogamy (8% of crops) were second most important. The main economic motivations for practicing or recommending hand pollination were to increase fruit set, and/or fruit quality (78% of crops). Hand pollination is practiced in large- and small-scale farming, home gardens, and greenhouses. Opportunities of hand pollination are the control of pollen origin and quantity, pollination timing and frequency as well as independence from environmental fluctuations. Farmers can increase yields, improve fruit quality, avoid fruit abortion, increase employment, and secure subsistence food. The main constraints of hand pollination are high labor inputs, high material costs, and required skills. Major risks of hand pollination include management ignoring pollinator conservation, high food prices, over-pollination, labor accidents, and unfair labor. We conclude that in the face of global change, hand pollination allows improved control of pollination and is likely to increase in importance. The benefits of hand pollination need to outweigh the costs and fair labor is essential. Altogether, hand pollination can be a valuable tool for crop systems where pollinators are absent or are not reliable for sustaining high-quality crop production.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

THE POLLINATION BIOLOGY OF TILIA

An anthecological study of Tilia in the Great Plains and New England was undertaken for both native and introduced species. The floral bracts are postulated as being at least as important in pollinator attraction as they are in fruit dispersal. A characteristic sweet odor is always present, but becomes stronger at dusk. Flowers are protandrous with anthers opening for the first time late in the afternoon and releasing pollen abundantly for about 24 hr. Initial nectar production coincides with stigma receptivity which begins late in the afternoon, but on the second day a flower is open. Sixty-six species of insects in 29 families were collected. Bees and flies are the most common diurnal visitors and moths are the primary nocturnal visitors. Samples of pollen taken from the insects indicate a relatively high constancy. Experimental tests show that Tilia is not apomictic or self compatible. Anemophily plays a secondary role to entomophily in pollination. Nocturnal pollinators are slighly less effective than diurnal pollinators in effecting fruit set. Although nocturnal pollinators are favored by the syndrome of floral characteristics, it is concluded that Tilia is a generalist in regard to pollination. It is proposed that the lack of clear morphological differences between Tilia species leads to pollinator inconstancy and thus hybridization.     

Pollination and fruit‐set of Grevillea robusta in western Kenya

The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.     

Pollination of Campomanesia phaea (Myrtaceae) by night‐active bees: a new nocturnal pollination system mediated by floral scent

• Bees are the most important diurnal pollinators of angiosperms. In several groups of bees a nocturnal/crepuscular habit developed, yet little is known about their role in pollination and whether some plants are adapted specifically to these bees. We used a multidisciplinary approach to investigate the reproductive biology and to understand the role of nocturnal/crepuscular bees in pollination of Campomanesia phaea (Myrtaceae), popularly named cambuci.
• We studied the floral biology and breeding system of C. phaea. We collected the floral visitors and tested the pollinators' effectiveness. We also determined the floral scents released at night and during daytime, and studied behavioural responses of crepuscular/nocturnal bees towards these scents.
• The flowers of cambuci were self‐incompatible and had pollen as the only resource for flower visitors. Anthesis lasted around 14 h, beginning at 04:30 h at night. The flowers released 14 volatile compounds, mainly aliphatic and aromatic compounds. We collected 52 species of floral visitors, mainly bees. Nocturnal and crepuscular bees (four species) were among the most frequent species and the only effective pollinators. In field bioassays performed at night, nocturnal/crepuscular bees were attracted by a synthetic scent blend consisting of the six most abundant compounds.
• This study describes the first scent‐mediated pollination system between a plant and its nocturnal bee pollinators. Further, C. phaea has several floral traits that do not allow classification into other nocturnal pollination syndromes (e.g. pollinator attraction already before sunrise, with pollen as the only reward), instead it is a plant specifically adapted to nocturnal bees.