Can floral field margins improve pollination and seed production in coriander Coriandrum sativum L. (Apiaceae)?

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Extranuptial nectaries in flowers: ants increase the reproductive success of the ant‐plant Miconia tococa (Melastomataceae)

• Although the production of extranuptial nectar is a common strategy of indirect defence against herbivores among tropical plants, the presence of extranuptial nectaries in reproductive structures is rare, especially in ant‐plants. This is because the presence of ants in reproductive organs can generate conflicts between the partners, as ants can inhibit the activity of pollinators or even castrate their host plants. Here we evaluate the hypothesis that the ant‐plant Miconia tococa produces nectar in its petals which attracts ants and affects fruit set.
• Floral buds were analysed using anatomical and histochemical techniques. The frequency and behaviour of floral visitors were recorded in field observations. Finally, an ant exclusion experiment was conducted to evaluate the effect of ant presence on fruit production.
• The petals of M. tococa have a secretory epidermis that produces sugary compounds. Nectar production occurred during the floral bud stage and attracted 17 species of non‐obligate ants (i.e. have a facultative association with ant‐plants). Ants foraged only on floral buds, and thus did not affect the activity of pollinators in the neighbouring open flowers. The presence of ants in the inflorescences increased fruit production by 15%.
• To our knowledge, the production of extranuptial nectar in the reproductive structures of a myrmecophyte is very rare, with few records in the literature. Although studies show conflicts between the partners in the ant–plant interaction, ants that forage on M. tococa floral buds protect the plant against floral herbivores without affecting bee pollination.

     

Influence of reproductive traits on pollination success in two <Emphasis Type="Italic">Daphne</Emphasis> species (Thymelaeaceae)

Taxonomically related species can differ in a number of reproductive traits, which may translate into a differential mating system and pollination success. Here we compare two hermaphroditic insect-pollinated Daphne species (D. rodriguezii and D. gnidium) which differ in distribution (island endemic vs. mediterranean) and floral traits (long- vs. short-tube corolla). We investigated their mating system and pollen limitation by means of hand-pollination experiments and quantified the diversity and abundance of flower visitors by direct observations. Plant size and five reproductive traits (flower production, proportion of viable anthers, pollen production, flower tube length and tepal area) were studied to assess how they contribute to reproductive success, measured as proportion of pollen grains germinated per stigma and fruit set. Selfing was very low and pollen limitation existed in both species, though was higher in D. rodriguezii probably due to the scarcity of flower visitors. The low fruit set in both species suggests that most of the pollen grains found on stigmas are self-pollen. Pollinators appeared to favour some floral traits (specifically, flower tube length or tepal area) in both species, although flower crop in D. rodriguezii was the only reproductive trait influencing fruit set. In both species, the highest variability in reproductive traits and pollination success was within individuals. Our findings suggest that despite both species showed similar mating system, dependency on outcrossing pollen and selection of floral traits, pollen limitation was higher in D. rodriguezii, probably as a higher proportion of self-pollen arrives to its stigmas.     

Reproductive biology of Ferocactus recurvus (Mill.) Borg subsp. recurvus (Cactaceae) in the Tehuacán‐Cuicatlán Valley,Mexico

• Mexico has one of the highest diversities of barrel cacti species worldwide; however, all are threatened and require conservation policies. Information on their reproductive biology is crucial, but few studies are available. Ferocactus recurvus subsp. recurvus is a barrel cactus endemic to the Tehuacán‐Cuicatlán Valley. Our research aimed to characterise its floral and pollination biology. We hypothesised bee pollination, as suggested by its floral morphology and behaviour, and self‐incompatibility, like most barrel cacti studied.
• Three study sites were selected in the semiarid Zapotitlán Valley, Mexico. We examined 190 flowers from 180 plants to determine: morphometry and behaviour of flowers, flower visitors and probable pollinators, and breeding system.
• Flowers showed diurnal anthesis, lasting 2–5 days, the stigma being receptive on day 2 or 3 after the start of anthesis. Flowers produced scarce/no nectar and main visitors were bees (Apidae), followed by flies (Muscidae), ants (Formicidae), thrips (Thripidae) and hummingbirds (Throchilidae); however, only native bees and occasionally wasps contacted the stigma and anthers. Pollination experiments revealed that this species is self‐incompatible and xenogamous. In natural conditions, fruit set was 60% and cross‐pollination fruit set was 100%. Percentage seed germination resulting from cross‐pollination was higher than in the control treatment.
• Our results provide ecological information for conservation programmes to ensure a high probability of breeding and seed production in natural populations of F. recurvus.

     

The generalist Inga subnuda subsp. luschnathiana (Fabaceae): negative effect of floral visitors on reproductive success?

Inga species are characterised by generalist or mixed pollination system. However, this feature does not enhance reproductive rates in species with very low fruit set under natural conditions. Some ecological and genetic factors are associated with this feature, and to test the effect of massive visits on pollination success in Inga subnuda subsp. luschnathiana, we studied the efficacy of polyads deposited on stigmas of flowers isolated from visitors and polyads exposed to visitors. The proportion of polyads fixed in stigmas decreased after exposure to visitors (24 h) in comparison to stigmas isolated from visitors (hummingbirds, bees, wasps, hawkmoths and bats), and fruit set was very low. Furthermore, nectar production, sugar composition and other floral biology traits were evaluated. Increased nectar production, sugar availability and sucrose dominance during the night indicates adaptation to nocturnal visitors and supports their role as main pollinators; although the brush‐flower morphology, time of anthesis, nectar dynamics and chemical composition also allow daytime visitors. Thus the species is an important resource for a diverse group of floral visitors. We conclude that excess visits (diurnal and nocturnal) are responsible for the decrease in fixed polyads in stigmas of I. subnuda subsp. luschnathiana flowers, thus contributing, with others factors, to its low fruit set. Therefore, the generalist pollination system does not result in reproductive advantages because the low fruit set in natural conditions could be the result of a negative effect of visitors/pollinators.     

Refuse attracts? Effect of refuse dumps of leaf‐cutting ants on floral traits

The nutrient‐rich organic waste generated by ants may affect plant reproductive success directly by enhancing fruit production but also indirectly, by affecting floral traits related with pollinator attraction. Understanding how these soil‐nutrient hot spots influence floral phenotype is relevant to plant–pollination interactions. We experimentally evaluated whether the addition of organic waste from refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae: Attini) alters floral traits associated with pollinator attraction in Eschscholzia californica (Ranunculales: Papaveraceae), an entomophilous herb. We analysed flower shape and size using geometric morphometric techniques in plants with and without the addition of refuse‐dumps soil, under greenhouse conditions. We also measured the duration of flowering season, days with new flowers, flower production and floral display size. Plants growing in refuse‐dumps soil showed higher flower shape diversity than those in control soil. Moreover, plants in refuse‐dumps soil showed bigger flower and floral display size, longer flowering season, higher number of flowering days and flower production. As all these variables may potentially increase pollinator visits, plants in refuse‐dumps soil might increase their fitness through enhanced attraction. Our work describes how organic waste from ant nests may enhance floral traits involved in floral attraction, illustrating a novel way of how ants may indirectly benefit plants.     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

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.     

Day–night fluctuations in floral scent and their effects on reproductive success in <Emphasis Type="Italic">Lilium auratum</Emphasis>

We examined the contribution of diurnal and nocturnal pollination to male and female reproductive success in Lilium auratum. Plants were bagged for either 12 h during the day or at night to allow either only nocturnal or only diurnal visitors to forage throughout the flowering period. We found that there was no significant difference in the seed:ovule ratio among diurnally pollinated, nocturnally pollinated, or control flowers. However, in terms of male reproductive success, it was more advantageous for the plants to be pollinated both diurnally and nocturnally: the numbers of pollen grains remaining in diurnally pollinated or nocturnally pollinated flowers were significantly greater than those in control flowers. The total amount of floral volatiles of L. auratum was significantly higher at night than during the day. The constituents of floral scent of all time series examined were mostly monoterpenoids, many of which serve as attractants for nocturnal hawkmoths. Such nocturnally biased floral scent emission of L. auratum might achieve male reproductive success by attracting nocturnal visitors, which may suggest that the relative contribution of floral scent in this species is biased towards male reproductive success.     

Reproductive biology of Pachira aquatica Aubl. (Malvaceae: Bombacoideae): a tropical tree pollinated by bats,sphingid moths and honey bees

We investigated the reproductive biology, including the floral biology, pollination biology, breeding system and reproductive success, of Pachira aquatica, a native and dominant tropical tree of fresh water wetlands, throughout the coastal plain of the Gulf of Mexico. The flowers present nocturnal anthesis, copious nectar production and sugar concentration (range 18–23%) suitable for nocturnal visitors such as bats and sphingid moths. The main nocturnal visitors were bats and sphingid moths while bees were the main diurnal visitors. There were no differences in legitimate visitation rates among bats, moths and honey bees. Bats and honey bees fed mainly on pollen while moths fed on nectar, suggesting resource partitioning. Eight species of bats carried pollen but Leptonycteris yerbabuenae is probably the most effective pollinator due to its higher pollen loads. The sphingid moths Manduca rustica, Cocytius duponchel and Eumorpha satellitia were recorded visiting flowers. Hand pollination experiments indicated a predominant outcrossing breeding system. Open pollination experiments resulted in a null fruit set, indicating pollen limitation; however, mean reproductive success, according to a seasonal census, was 17 ± 3%; these contrasting results could be explained by the seasonal availability of pollinators. We conclude that P. aquatica is an outcrossing species with a pollination system originally specialized for bats and sphingid moths, which could be driven to a multimodal pollination system due to the introduction of honey bees to tropical America.     

Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado

• Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self‐compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self‐compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics.
• We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors.
• We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self‐incompatible and quite generalist regarding floral visitors.
• Enantiostyly in self‐incompatible plants seems to confer a reproductive advantage by reducing self‐interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

     

Variation in pollinator assemblages in a fragmented landscape and its effects on reproductive stages of a self-incompatible treelet, <Emphasis Type="Italic">Psychotria suterella</Emphasis> (Rubiaceae)

Few studies of plant–pollinator interactions in fragmented landscapes evaluate the consequences of floral visitor variation on multiple stages of plant reproduction. Given that fragmentation potentially has positive or negative effects on different organisms, and that self-incompatible plant species depend on pollinators for sexual reproduction, differences in floral visitor assemblages may affect certain plant reproductive stages. We evaluated how pollinator assemblage, availability of floral resources, pollination, reproductive output, and seed and seedling performance of Psychotria suterella Muell. Arg. varied among three fragmentation categories: non-fragmented habitats, fragments connected by corridors, and isolated fragments. Richness and frequency of floral visitors were greater in fragments than in non-fragmented sites, resulting mainly from the addition of species typically found in disturbed areas. Although 24 species visited Psychotria suterella flowers, bumblebees were considered the most important pollinators, because they showed the highest frequency of visits and were present in eight out of ten sites. Additionally, the number of pollen tubes per flower per visit was lower in areas without bumblebees. The increased visitation in fragments seemed to enhance pollination slightly. However, fruit and seed output, germination, and seed and seedling mass were similar in non-fragmented sites, connected sites, and isolated fragments. Our results suggested that, even for a self-incompatible species, responses to habitat fragmentation at different stages of plant reproduction might be decoupled from the responses observed in floral visitors, if fruit set is not pollen limited. If all reproductive stages were considered, variation on the small scale was more important than the variation explained by fragmentation category. In spite of its self-incompatible breeding system, this plant–pollinator system showed resilience to habitat fragmentation, mainly as a result of high availability of potential mates to P. suterella individuals, absence of pollen limitation, and the presence of bumblebees (Bombus spp.) throughout this highly connected landscape.     

Pollination of Richea dracophylla (Ericaceae) and the role of the nocturnal ant Notoncus hickmani (Formicinae)

Our observation of large numbers of ants Notoncus hickmani (Formicinae) on the flowers of a Tasmanian endemic shrub Richea dracophyllum (Ericacea) during the night in winter prompted us to carry out a more extensive study of pollination of R. dracophyllum at two sites. Terminal panicles bear inflorescences of 300+ flowers whose petals are fused into calyptrae covering the reproductive organs, and the calyptrae must be removed before pollination can occur. To exclude birds, we placed mesh bags over some inflorescences. Inflorescences with exclusion bags had a high proportion of retained calyptra and the numbers of viable seeds were greatly reduced. We found R. dracophyllum to have a very long flowering period (more than 6 months) and a range of floral visitors, including honey-eaters (Meliphagidae) and flying insects. At one site we observed large numbers (up to 100) of N. hickmani on inflorescences between dusk and midnight during winter at air temperatures down to 2.5°C. They fed on nectar from flowers that had lost their calyptra where they were exposed to pollen from the anthers and scanning electron microscopy showed high numbers of pollen grains being carried on the ants. The feeding behaviour of these ants would facilitate pollination between adjacent flowers (geitonogamy). Smaller numbers of dolichoderine ants (Iridomyrmex bicknelli?) were observed during the daytime in spring but mainly fed under partially detached calyptrae where they had little exposure to pollen. We suggest that R. dracophylla is pollinated by a range of species and functional groups—birds, flying insects and ants, and that the pollinating species will vary during the year and between sites, although insects depend on birds to remove the calyptrae. Ant pollination is rare, nocturnal ant pollination is extremely rare, and pollination by nocturnal cryophilic ants has not been documented previously.     

An example of flexistyly in a wild cardamom species (Amomum maximum (Zingiberaceae))

We studied the floral and pollination biology of Amomum maximum. Our observations suggest that flexistyly is present in this species, and stigmatic behavior is similar as reported for Alpinia species. Two phenotypes occur in a ratio of 1:1 in natural populations. No fruit set occurred in the unpollinated bagged plants. Significant differences in seed set between selfing and crossing manipulations demonstrate that inbreeding depression or partial self-incompatibility is apparent in this species. Despite the resemblance in floral behavior, there are still some differences between the previous studies and ours: with special floral design, the body size of legitimate visitors ranged wider in Amomum maximum than in Alpinia species; the presence of anther appendage is suggested to prevent competing and delayed selfing. This is the first report of flexistyly in a wild cardamom species, which is the second largest group in ginger family (Zingiberaceae).     

Pollination biology in an endangered rocky mountain toadflax (Linaria cavanillesii)

Knowledge about the reproductive system of species inhabiting rocky habitats is scarce. The reproductive biology (floral biology, experimental pollination, insect visits, inbreeding depression, and seed predation) of the rupicolous Linaria cavanillesii was analyzed under field and experimental conditions. Self-compatibility was revealed by the high fruit set, seed set, and seed mass in the pollination experiment. Furthermore, results disclose that this species does not need insect visitors for seed production since fruit set after autonomous self-pollination was similar to that by hand cross-pollination. Self-offsprings were not affected by a strong degree of inbreeding depression in early life-cycle stages. Seed predation by Cucurlionidae is the main limiting factor of fruit and seed production in this species. Contrary to other perennial species of Linaria previously studied, L. cavanillesii represents the first clear report of self-fertility.     

Ant pollination of <Emphasis Type="Italic">Syzygium occidentale</Emphasis>, an endemic tree species of tropical rain forests of the Western Ghats,India

Although mutualism between ants and flowering plants is wide spread, ant pollination has not evolved as a major pollination syndrome. So far ant pollination has been reported largely in herbaceous species, growing in warm and dry habitats. While studying pollination ecology of Syzygium species (Myrtaceae), growing in tropical forests of the Western Ghats, India, we observed one of the ant species, Technomyrmex albipes, to be the dominant floral visitor in S. occidentale (Bourd.) Chithra among a range of other insect (species of Xylocopa and Trigona, and Apis cerana) and bird visitors. We studied the role of ant species in pollination when compared to other floral visitors. The fruit set in flowers exclusively visited by T. albipes was significantly higher than those visited by any other visitor. The day and night exclusive pollination experiments allowing only T. albipes indicated diel pollination by T. albipes, which was the only active flower visitor during the night. The breeding system of the species was studied through controlled pollinations. The species is partially self-compatible and exhibits considerable autogamy.     

Experimental study of pollination by ants in Mediterranean high mountain and arid habitats

In this paper, we report the results of an experimental study on ant pollination of three plant species inhabiting the Mediterranean high mountains (Alyssum purpureum, Arenaria tetraquetra and Sedum anglicum) and four species inhabiting the aridlands (Lepidium subulatum, Gypsophyla struthium, Frankenia thymifolia and Retama sphaerocarpa) of South-eastern Spain. We determined several plant and ant traits, as well as the composition and abundance of the pollinator assemblage. Insects belonging to 29 families and five orders visited the flowers of the plant species studied. In all but two, L. subulatum and G. struthium, the ants comprised 70–100% of the flower visitors. The results clearly show that five out of seven of these plant species were pollinated by ants. The role of the ants as pollinators seems to depend heavily on the relative abundance of the ants with respect to the other species of the pollinator assemblage, ant pollination becoming evident when ants outnumber other floral visitors. The ant-pollination systems analysed in this study may be the result of prevailing ecological conditions more than an evolutionary result of a specialized interaction.     

Pollination ecology of Yucca elata

The pollination biology of a population of 250 Yucca elata (Liliaceae) plants was studied in southern New Mexico. Yucca elata and the prodoxid yucca moth Tegeticula yuccasella have a mutualistic association that is essential for the successful sexual reproduction of both species. However, a wide range of other invertebrate species visit flowers during the day and at night. Our aim was to quantify the role of yucca moths and other invertebrate visitors in pollination and fruit set, using manipulative field experiments. Inflorescences were bagged during the day or night (N=12 inflorescences) to restrict flower visitors to either nocturnal or diurnal groups. Yucca moths were active exclusively nocturnally during the flowering period and thus did not visit inflorescences that were unbagged during the day. None of the 4022 flowers exposed only to diurnal visitors set fruit, whereas 4.6% of the 4974 flowers exposed only to nocturnal visitors (including yucca moths) produced mature fruit. The proportion of flowers producing fruit in the latter treatment was not significantly different from unbagged control inflorescences. In a series of experimental manipulations we also determined that: (1) flowers opened at dusk and were open for two days on average, but were only receptive to pollen on the first night of opening; (2) pollen must be pushed down the stigmatic tube to affect pollination; and (3) most plants require out-cross pollination to produce fruit. The combination of these results strongly suggests that yucca moths are the only species affecting pollination in Y. elata, and that if another species was to affect pollination, it would be a rare event.     

The effects of nutrient addition on floral characters and pollination in two subalpine plants, <Emphasis Type="Italic">Ipomopsis aggregata</Emphasis> and <Emphasis Type="Italic">Linum lewisii</Emphasis>

The availability of soil and pollination resources are main determinants of fitness in many flowering plants, but the degree to which each is limiting and how they interact to affect plant fitness is unknown for many species. We performed resource (water and nutrients) and pollination (open and supplemental) treatments on two species of flowering plants, Ipomopsis aggregata and Linum lewisii, that differed in life-history, and we measured how resource addition affected floral characters, pollination, and reproduction (both male and female function). We separated the direct effects of resources versus indirect effects on female function via changes in pollination using a factorial experiment and path analysis. Resource addition affected I. aggregata and L. lewisii differently. Ipomopsis aggregata, a monocarp, responded to fertilization in the year of treatment application, increasing flower production, bloom duration, corolla width, nectar production, aboveground biomass, and pollen receipt relative to control plants. Fertilization also increased total seed production per plant, and hand-pollination increased seeds per fruit in I. aggregata, indicating some degree of pollen limitation of seed production. In contrast, fertilization had no effect on growth or reproductive output in the year of treatment on L. lewisii, a perennial, except that fertilization lengthened bloom duration. However, delayed effects of fertilization were seen in the year following treatment, with fertilized plants having greater aboveground biomass, seeds per fruit, and seeds per plant than control plants. In both species, there were no effects of resource addition on male function, and the direct effects of fertilization on female function were relatively stronger than the indirect effects via changes in pollination. Although we studied only two plant species, our results suggest that life-history traits may play an important role in determining the reproductive responses of plants to soil nutrient and pollen additions.     

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.