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.     

Opportunistic nectar-feeding birds are effective pollinators of bird-flowers from Canary Islands: experimental evidence from Isoplexis canariensis (Scrophulariaceae)

Insular floras, characterized by simple pollination networks, sometimes include novel mutualistic agents such as nonspecialist nectarivores. In this study we confirmed the effective pollination of Isoplexis canariensis by opportunistic nectar-feeding birds in Tenerife, Canary Islands, Spain. This plant is among the ornithophilous species of the Canarian flora that lack past and present specialist nectarivorous birds. Experimental hand pollinations revealed self-compatibility, but cross-pollinated flowers produced a greater percentage of viable seeds than self-pollinated ones. Flowers were visited by five species of birds (Phylloscopus canariensis, Parus caeruleus, Sylvia melanocephala, Serinus canarius, and Fringilla coelebs) and by the endemic lizard (Gallotia galloti, Lacertidae). Insect pollination was absent, and the few insect visitors acted as nectar thieves or secondary nectar robbers. Birds represented 93.1% of total visits, with the Canarian Chiffchaff, Ph. canariensis, being the most frequent visitor. Flowers visited by birds set more, larger, and heavier fruit than flowers from which birds were excluded. Bird visitation also enhanced seed viability. These results demonstrate the active role of these opportunistic birds as effective pollinators of this Canarian bird-flower species. Further, the results reveal the need to consider the effect of these birds on the evolution of ornithophilous floral traits in absence of specialist nectarivores.     

Reproductive ecology of Aloe plicatilis,a fynbos tree aloe endemic to the Cape Winelands,South Africa

While the pollination ecology of many Aloe species is well-documented, knowledge on aloe seed ecology, and hence aloe reproductive ecology in its entirety is limited. The aim of this study was to investigate the reproductive ecology of Aloe plicatilis, a Cape fynbos tree aloe endemic to the Cape Winelands, South Africa. Results from a pollinator exclusion experiment conducted at an A. plicatilis population on Paarl mountain suggests pollination primarily by insects, although bird visitation significantly increased seed set/fruit indicating possible co-pollination with insects. The species' long-tubed flowers and production of concentrated nectar, with observations of malachite sunbirds as the most common avian visitors to A. plicatilis flowers indicate the importance of long-billed specialist avian nectarivores as floral visitors. Analysis of the relationship between plant size and inflorescence production for five populations combined revealed a significant, positive linear relationship between plant size and the logarithm of the number of inflorescences/plant. Natural fruit and seed set determined for three populations (1325, 27,930 and 251,616 seeds/population) suggests low reproductive output compared to several other Aloe species. The smallest (31 individuals) and least dense (75 plants/ha) A. plicatilis population produced the lowest seed set/plant (128 seeds) and per population (1325 seeds), suggesting an Allee effect. Evaluation of seed dispersal potential showed that potential dispersal distances were approximately three times the canopy height; however, the occurrence of A. plicatilis on mountains isolated from more continuous mountain ranges on which the species also occurs suggests the possibility of long-distance dispersal by strong, gusty, summer winds. Soil seed bank samples collected from 13 populations yielded close to zero seedling emergence, indicating the absence of persistent seed banks. A. plicatilis seeds stored under ambient laboratory conditions for 3, 18 and 24 months were germinated in an environmental control chamber and a laboratory. High percentage germination was recorded for 18- and 24-month-seed (86 and 80%, respectively), while germination of 3-month-old seeds was three times lower, suggesting the need for after-ripening. Germination of fresh and one-year-old seed under ambient nursery conditions at the Karoo Desert National Botanical Garden in Worcester yielded emergence percentages of 67 and 44%, respectively, and were therefore less successful than germination under more controlled conditions. This is the first known study to investigate the reproductive ecology of a tree aloe species and that of a Cape fynbos aloe. The study highlights the importance of further studies on aloe seed ecology, particularly for rare and threatened species.     

High levels of fecundity in small and isolated populations of a self‐compatible Aloe pollinated by opportunistic birds and bees

• Plant species that are effective colonisers of transient habitats are expected to have a capacity for uniparental reproduction and show flexibility in pollination systems. Such traits may enable populations to be established from a small number of founding individuals without these populations succumbing to reductions in fecundity arising from pollinator limitation.
• We tested these predictions for Aloe thraskii (Xanthorrhoeaceae), a succulent treelet that colonises shifting coastal dunes and has both bird and bee pollinators. We performed hand‐pollination experiments, and selectively excluded bird visitors to determine differences in pollinator effectiveness. We measured pollinator visitation rates and fecundity in populations varying in their size, density and isolation distance.
• Controlled hand‐pollinations revealed that unlike most other Aloe species, A. thraskii is self‐compatible and thus capable of uniparental reproduction. The species does however depend on pollinators and is visited by various bird species as well as by bees. Fruit and seed set are not affected by selective exclusion of birds, thus indicating that bees are effective pollinators. Bird visitation rates increased with increasing plant height and population size, while bee visitation rates increased with increasing population size and density. We found that seed set per flower was lower in large populations than in small populations.
• These results suggest that establishment of populations of A. thraskii from a small number of individuals is unlikely to be limited by the fecundity of individual plants.

     

The value of insect pollination to yield of oilseed rape (Brassica rapa) in Bangladesh

Brassica rapa L., is a crop grown globally and studies have indicated that insect pollination can improve yields. However, the importance of insect pollination in this crop depends on cultivar reproductive biology, insect pollinator species and their abundances. In Bangladesh, the acreage of B. rapa for oilseed production has been expanding, but little is known about whether insects contribute to yield improvements. Using the commonly grown variety Tori-7, we found that plants with inflorescences left exposed to flower visiting insects (body width > 1.5 mm) had a 30.8% greater seed yield compared to those where insect flower visitors were excluded. Of 794 insect flower visiting individuals recorded from observation and trapping surveys conducted across four separate fields, Apis bees (four spp.) were the most abundant (66.1%), followed by flies Musca domestica (14.7%), Sepsis fulgens (6.3%) and hoverflies (3.0%; 3 spp; Syrphidae). Other flower visitors included ants, wasps, beetles, butterflies and moths. For the cultivar assessed, we calculated the economic value of oilseed rape was $US 87.5 million per annum in Bangladesh, of which the economic value of insect pollination was $US 26.92 million per annum. Thus, one in every four dollars earned by our Bangladeshi growers resulted from insect crop pollination. This demonstrates to growers the need to promote and protect insect pollination to optimize their economic returns.     

Reproductive ecology of <Emphasis Type="Italic">Agave colorata</Emphasis>: the importance of nectar-feeding bats and the germination consequences of self-pollination

Agave colorata is a paniculate agave distributed along the migratory route of the nectar-feeding bat Leptonycteris yerbabuenae. In this paper, we evaluate the importance of nectar-feeding bats in the reproduction of A. colorata in a population in Sonora, Mexico, and describe the germination consequences of self-pollination. We estimated abundance using five plots and set pollination treatments to evaluate the importance of bats. We recorded 14.8?±?6.8 plants/400 m², with a bimodal size distribution. Flowers are protandrous and visited mainly (>?20 visits/plant/30 min) by L. yerbabuenae. Pollination exclusion experiments showed that flowers excluded from diurnal visitors had maximum fruit set values (0.49?±?0.42), while the autonomous self-pollination treatment had the lowest value (0.03?±?0.06). Similarly, the greatest number of viable seeds per fruit was recorded in the diurnal exclusion treatment, while the greatest number of empty seeds was observed in the self-pollination treatment. Fruit set values among untreated plants varied from 32 to 54%, with a mean value of 41.8%. Seeds derived from self-pollination had a narrower window of opportunity for germination compared to seeds derived from nocturnal pollination. Self-pollinated seeds had lower germination, rate of germination or lag time in response to light, osmotic potential and heat shock treatments, compared to other pollination treatments, revealing an inbreeding cost. Overall, our results show that L. yerbabuenae is the likely major pollinator of the studied A. colorata population. However, under pollinator limitation A. colorata may produce seeds by autonomous self-pollination, at a cost expressed as lower germination.     

Floral traits mediate the vulnerability of aloes to pollen theft and inefficient pollination by bees

Background and Aims

Pollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral and inflorescence characteristics mediate the pollination role of pollen collection by bees.

Methods

For ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology.

Key Results

Species varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape.

Conclusions

Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.     

The relative importance of birds and bees in the pollination of Metrosideros excelsa (Myrtaceae)

Exclusion experiments were used to assess the effect of different pollinator groups on outcrossing and seed production in Metrosideros excelsa. The main study site was Little Barrier Island, New Zealand where indigenous bird and native solitary bees are the main flower visitors. Our results showed that native birds were more important pollinators of M. excelsa than native bees. Seed production was much higher in open pollination than in two exclusion experiments where either birds were excluded and native bees only had access to flowers, or where all pollinators had been excluded. The number of fertile seeds per capsule was 45% higher after open pollination than in treatments with bee visitation only and 28% higher than in treatments where all flower visitors were excluded. Estimated outcrossing rates were significantly higher (t_m = 0.71) for open pollination in the upper canopy (>4 m above‐ground level) where bird visitation is presumed to be more frequent than for a treatment with native bee access only (t_m = 0.40). Our results also suggest that a large proportion of seeds (66%) arise from autonomous self‐pollination when all pollinators are excluded. In four trees of a modified mainland population with predominantly introduced birds and a mixture of introduced and native bees there was no decrease in seed production for the treatment allowing bee access only, indicating that – in contrast to native bees – honeybees may be more efficient pollinators of M. excelsa. Observation of the foraging behaviour of both groups of bees showed that native bees contact the stigma of flowers less frequently than honeybees. This is likely to be a consequence of their smaller body size relative to honeybees.     

Floral biology of Echinopsis chiloensis ssp. chiloensis (Cactaceae): Evidence for a mixed pollination syndrome

In this study the flower biology of Echinopsis chiloensis ssp. chiloensis, a columnar cactus occurring in Central Chile, is investigated, in particular its pollination syndrome, its visitors, their frequencies and behaviors and their pollination efficiencies. As statements on floral anthesis of this species are contradictory, this study also intended to elucidate both its beginning and duration. A pollinator exclusion study of a total of 162 flowers from 12 plants was conducted at one of the two study sites. Fruit and seed production as well as seed viability were documented to evaluate pollinator efficiencies.     

Covariation of flower traits and bird pollinator assemblages among populations of Kniphofia linearifolia (Asphodelaceae)

Globally, bird-pollinated plants can be separated into two groups, one consisting of species pollinated by specialist nectarivores, and the other of plants pollinated by occasional nectarivores. There are marked differences in nectar properties among the two groups, implying that there has been pollinator-mediated selection on these traits. This raises the possibility that variation in bird assemblages among populations of a plant species could lead to the evolution of intraspecific variation in floral traits. We examined this hypothesis in Kniphofia linearifolia, a common and widespread plant in southern Africa. Although bees are common visitors to flowers of this species, exclusion of birds from inflorescences led to significant reductions in seed set, indicating that the species is primarily bird-pollinated. We showed that bird pollinator assemblages differ markedly between five different populations of K. linearifolia, and that variation in flower morphology and nectar properties between these populations are associated with the dominant guild of bird visitors at each population. We identified two distinct morphotypes, based on corolla length, nectar volume and nectar concentration, which reflect the bird assemblages found in each type. Further work is needed to establish if a natural geographic mosaic of bird assemblages are the ultimate cause of differentiation in floral traits in this species.     

Autogamy in Boronia falcifolia: consequences for inbreeding and persistence in fragmented coastal heaths

Coastal heath is becoming increasingly fragmented, resulting in reduced pollinator abundance affecting species viability. The reproductive ecology and inbreeding of a common coastal heath species, Boronia falcifolia (Rutaceae), was investigated. Pollination observations and experiments were carried out at one site, and inbreeding was assessed using genetic markers at eight populations. Four pollination treatments were assessed: cross-pollination, self-pollination, autogamous pollination and open (insect) pollination. Boronia falcifolia was found to be highly self-compatible, frequently setting seeds from autogamous pollination. There were no significant differences between the autogamous, self and cross pollen treatments for any measures of reproductive success (pollen tube growth, fruit set, swollen locules and seed production). Insect activity resulted in fewer seeds per fruit than hand cross pollination, although pollinator visits were few. All populations studied were highly inbred. Autogamous self-pollination is widespread in B. falcifolia and this combined with its ability to vegetatively regenerate enhances its potential for survival when pollinators are few.     

Generalist passerine pollination of a winter-flowering fruit tree in central China

Background and Aims

Winter-flowering plants outside the tropics may experience a shortage of pollinator service, given that insect activity is largely limited by low temperature. Birds can be alternative pollinators for these plants, but experimental evidence for the pollination role of birds in winter-flowering plants is scarce.

Methods

Pollinator visitation to the loquat, Eriobotrya japonica (Rosaceae), was observed across the flowering season from November to January for two years in central China. Self- and cross-hand pollination was conducted in the field to investigate self-compatibility and pollen limitation. In addition, inflorescences were covered by bird cages and nylon mesh nets to exclude birds and all animal pollinators, respectively, to investigate the pollination role of birds in seed production.

Results

Self-fertilization in the loquat yielded few seeds. In early winter insect visit frequency was relatively higher, while in late winter insect pollinators were absent and two passerine birds (Pycnonotus sinensis and Zosterops japonicus) became the major floral visitors. However, seed-set of open-pollinated flowers did not differ between early and late winter. Exclusion of bird visitation greatly reduced seed-set, indicating that passerine birds were important pollinators for the loquat in late winter. The whitish perigynous flowers reward passerines with relatively large volumes of dilute nectar. Our observation on the loquat and other Rosaceae species suggested that perigyny might be related to bird pollination but the association needs further study.

Conclusions

These findings suggest that floral traits and phenology would be favoured to attract bird pollinators in cold weather, in which insect activity is limited.     

Generalization versus Specialization in Pollination Systems: Visitors,Thieves, and Pollinators of Hypoestes aristata (Acanthaceae)

Many recent studies have suggested that the majority of animal-pollinated plants have a higher diversity of pollinators than that expected according to their pollination syndrome. This broad generalization, often based on pollination web data, has been challenged by the fact that some floral visitors recorded in pollination webs are ineffective pollinators. To contribute to this debate, and to obtain a contrast between visitors and pollinators, we studied insect and bird visitors to virgin flowers of Hypoestes aristata in the Bamenda Highlands, Cameroon. We observed the flowers and their visitors for 2-h periods and measured the seed production as a metric of reproductive success. We determined the effects of individual visitors using 2 statistical models, single-visit data that were gathered for more frequent visitor species, and frequency data. This approach enabled us to determine the positive as well as neutral or negative impact of visitors on H. aristata’s reproductive success. We found that (i) this plant is not generalized but rather specialized; although we recorded 15 morphotaxa of visitors, only 3 large bee species seemed to be important pollinators; (ii) the carpenter bee Xylocopa cf. inconstans was both the most frequent and the most effective pollinator; (iii) the honey bee Apis mellifera acted as a nectar thief with apparent negative effects on the plant reproduction; and (iv) the close relationship between H. aristata and carpenter bees was in agreement with the large-bee pollination syndrome of this plant. Our results highlight the need for studies detecting the roles of individual visitors. We showed that such an approach is necessary to evaluate the pollination syndrome hypothesis and create relevant evolutionary and ecological hypotheses.     

Effects of natural and artificial pollination on fruit and offspring quality

Worldwide, many crops rely on insect pollination. Insufficient pollination can reduce fruit and seed set by directly reducing pollen deposition, and can also affect offspring quality, such as growth rate and resistance to herbivores, by limiting outcrossing opportunities. Both effects are important in fruit agroecosystems where fruit size and the quality of seeds for re-planting are dependent on sufficient pollination. We experimentally manipulated pollination of the cape gooseberry, Physalis peruviana L. (Solanaceae), to test the effects of honey and bumble bee pollination compared to manual outcrossing and autonomous self-pollination on fruit and offspring characteristics. Compared to manual and self-pollination, bee pollination increased fruit size, seed set and germination rates, supporting the hypothesis that sufficient pollination increases plant fitness. Interestingly, plant growth rate and herbivore resistance were significantly and marginally greater in manually outcrossed plants compared to self-pollinated offspring, suggesting that inbreeding reduces offspring quality. Herbivore resistance and plant growth did not differ between one honeybee visit and self-pollination suggesting that multiple pollinator visits are needed to prevent inbreeding events. Our data suggest that the quantity and quality of pollen deposited by bee visitation can significantly alter ecologically and economically relevant traits in this agroecosystem.     

Pollination effectiveness and pollinator importance in a population of Heterotheca subaxillaris (Asteraceae)

 Assessing the relative contributions to seed set for each of a plant species’ floral visitors provides an indication of the relative influence of these visitors on the plant’s reproductive success. This study examined pollinator activity and seed set in a population of Heterotheca subaxillaris, a species that exhibits a floret dimorphism (heads bearing disk and ray florets), and that is visited by both generalist foragers and specialist bees. Visits by nine bee genera and one genus of skipper were recorded in the study population. During the period of study, these insects varied in their relative abundance, in their foraging activity on a head, and in their pollination effectiveness. The pattern of pollination effectiveness shown by the different pollinators was similar for both floret types, although seed set was higher overall for ray florets. Pollinator importance, calculated as the product of pollination effectiveness and relative abundance, was dictated by a pollinator’s relative abundance. The single specialist bee species observed in the study population proved to have neither higher pollination effectiveness nor higher pollinator abundance than other pollinators. This would suggest that H. subaxillaris is not under strong selective pressure to co-specialize with its specialist visitor. Received: 21 December 1995 / Accepted: 20 June 1996     

Status of pollinators and their efficiency in coffee fruit set in a fragmented landscape mosaic in South India

The region of Kodagu, South India, comprises a fragmented landscape with a high density of remnant forest patches dispersed within a mosaic dominated by shaded coffee agro-forests. We evaluated the role of self, wind and insect pollination to Coffea canephora production in this landscape. The giant Asian honeybee, Apis dorsata, which nests in remnant forests, was the main pollinator of coffee (accounting for 58% of the floral visitors). The proportion of flowers that developed into fruits was highest when hand cross-pollinated (44%), followed by open- (insect and wind combined; 33%) and wind- (22.1%) pollination treatments. Pollination by bees therefore increases fruit production by 50% over that achieved by wind. Self-pollination (1.7%) and no pollination (1%) treatments produced very low fruit set, emphasizing the importance of cross-pollination in C. canephora. Unlike measures of pollination success, initial fruit set (five weeks after flowering) proved an unreliable proxy for final fruit set. Size of adjoining forest fragments (mostly 0.3–20 ha, with a few exceeding 200 ha) positively influenced pollinator visitation to coffee flowers, but distance to such fragments had no influence on pollination. This study demonstrates the importance of cross-pollination for crop production in C. canephora, the important contribution that pollinating insects make to coffee production, and the benefits of relatively large forest fragments within the landscape mosaic to support insect pollinators of coffee. A comparison of pollinator composition to that of 100 years ago indicated that coffee pollination services remained intact despite changes in pollinator community composition.     

Relative contributions to seed production by floral visitors of slickspot peppergrass, Lepidium papilliferum (Brassicaceae)

Assessing the relative contributions to seed production made by different types of floral visitors is fundamental to understanding the evolution of floral morphology and the influence that particular pollinator taxa have on plant fitness and reproduction. This 3-year study examined the pollinator activity and the seed production in three populations of Lepidium papilliferum, a threatened mustard endemic to sagebrush-steppe habitat in southwest Idaho. Relative amounts of time visitor taxa spent foraging on flowers, visitation rates (number of flowers visited per unit time during a foraging bout), and pollination effectiveness (fruit set per single visit to a virgin flower) were recorded for each of 12 insect taxa that visited L. papilliferum flowers. Relative contributions to seed production were calculated as the product of relative interaction frequencies (the relative number of flowers visited by each taxon—the “quantity” component of pollination) and pollination effectiveness (fruit set per single visit to a virgin flower—the “quality” component of pollination). Despite the superiority of some insect taxa in terms of pollination effectiveness, estimates of relative pollinator contributions to seed production were influenced primarily by an insect taxon’s interaction frequency with flowers. Pollinator assemblages varied widely both spatially and temporally, which suggest that L. papilliferum is not under strong selective pressure to shift from its generalist pollination strategy toward greater specialization. For this threatened plant, reliance on a diverse assemblage of insect pollinators may serve as an important buffer against disruption in reproductive success caused by fluctuations in population sizes of individual pollinator taxa.     

Comparing the reproductive success and pollination biology of an invasive plant to its rare and common native congeners: a case study in the genus <Emphasis Type="Italic">Cirsium</Emphasis> (Asteraceae)

Previous studies have examined an association between reproductive success and pollination biology of rare versus widespread species through pair-wise comparisons of native and invasive congeners or rare and common congeners. To determine the importance of reproductive success and pollination biology for an invasive thistle, Cirsium vulgare, we compared it in its invaded range to five, co-occurring native Cirsium species that range from rare to common. Native study species include C. fontinale var. fontinale, C. andrewsii, C. brevistylum, C. occidentale, and C. quercetorum. We compared all species’ reproductive success, insect visitation rate and composition, autonomous self-pollination, and level of pollen limitation in multiple populations. Species differed in their reproductive success; the invasive C. vulgare produced more flower heads per plant than most native species. C. vulgare attracted more visitors than its congeners. In addition, reproductive success and insect visitation significantly varied between populations within species, mainly due to aphid infestation in one population of C. occidentale. Unlike the rare species (C. fontinale and andrewsii), C. vulgare did not require a pollinator for high-levels of seed production. The remaining native species set fewer seeds than C. vulgare without a pollinator. However, differences in insect visitation and autonomous self-pollination did not lead to differences in pollen limitation across species or between populations. This result suggests that factors other than pollination biology determine the difference in reproductive success of these species. However, high levels of autonomous self-pollination and generalist insect visitation may allow the invasive C. vulgare to easily establish new populations from low numbers of propagules. Our study provides one contrast that should build towards a larger comparative analysis to examine general patterns in the relationship between reproductive success, pollination biology, rare and invasive species, and our ability to predict biological invasions in introduced species.     

Entomophily of the cloudberry (Rubus chamaemorus)

Cloudberry (Rubus chamaemorus L.), a dioecious perennial plant of boreal circumpolar distribution, is greatly prized for its berries. We crossed two treatments, pollinator exclusion and supplementary hand-pollination, to determine i) the relative importance of insects as pollinators, ii) if pollinator activity was a limiting factor for the sexual reproduction of the plant, and iii) the relative contribution of diurnal vs. nocturnal visitors to pollination. The activity of natural pollinators resulted in 97.5 and 88.5% fruit set, along with 76.7 and 62.5% seed set in 1998 and 1999, respectively. When insects were excluded, fruit-set dropped significantly to 18.4 (1998) and 12.8% (1999) and seed-set to 5.4 (1998) and 5.0% (1999) showing the importance of mid- and large-sized insects as pollinators. Natural levels of insect activity were sufficient to ensure complete pollination in both years as supplementary hand-pollination did not significantly increase either parameter in plots where pollinators had free access. Nocturnal insects may serve as pollinators (fruit-set = 41%), although they were less effective than diurnal pollinators (fruit-set = 93%).     

Pollination biology and breeding system in five nocturnal species of <Emphasis Type="Italic">Oenothera</Emphasis> (Onagraceae): reproductive assurance and opportunities for outcrossing

The capacity to produce seed, both by selfing and outcrossing, or mixed mating strategies, is considered a mechanism for overcoming unpredictable pollinator availability. In the present study, we investigate breeding system, insect visitations and the role of insect visitors in the pollination of five species of Oenothera subsect. Oenothera. Field experiments showed that autonomous selfing occurs at bud stage, prior to the opening of the flower. Control flowers showed similar seed set to hand-pollinated flowers, whereas emasculated flowers and those subject to open pollination set fewer seed. These results are consistent with the hypothesis that the investigated Oenothera exhibit a great capacity for autonomous selfing and that selfing is selected in order to provide reproductive assurance. Although flowers were visited mostly by nocturnal lepidopterans, these insects did not precipitate pollination and are thus considered nectar thieves. Conversely, analysis of pollen loads and behavior during foraging by diurnal insect visitors revealed that honeybees and bumblebees are the probable pollinators. We conclude that production of flowers capable of autonomous selfing at bud stage, followed by anthesis and opportunities for outcrossing, probably improves the invasive potential of these Oenothera in Europe, together with a rapid increase in their populations, even when pollinators are scarce.