Ontogeny of the translator inAsclepiadaceae s.str.

The ontogeny of the translator is analysed inFockea edulis, Pachycarpus dealbatus, andCalotropis procera. The three species show a similar basic pattern in the construction of the corpusculum. Each side wall consists of two originally separate flanks. In the primitive state (Fockea), these flanks remain separate throughout their development, only the two inner flanks fuse along their base. In the derived state, the two flanks on each side fuse to form solid side walls, and the corpusculum develops a floor by additional secretion. The primitive state of divided flanks is recapitulated in early ontogeny. InFockea, the pollinia are attached apically and dorsally to the outer flanks by means of unmoulded adhesive material; in more advanced translators, caudicles are inserted as new elements between the outer flanks and the pollinia. The caudicles are attached to the lower part of the corpusculum. Within the four tribes of the family, many species have retained parts of the primitive structure in their corpuscula. Flanks divided in the lower part of the corpusculum and caudicles attached dorsally to the outer flanks seem to be common inMarsdenieae andStapelieae. InAsclepiadeae andGonolobeae the side walls are solid throughout the length of the corpusculum. — The question of cellular mechanisms underlying the formation of the translator is discussed. In all probability, the main factor responsible for the complex structure of the corpusculum is a differential secretory activity of the glandular field. This, in turn, points to differential genetic activity as a regulating factor of the ontogeny of the translator.     

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.     

The consequences of rewardlessness in orchids: reward-supplementation experiments with Anacamptis morio (Orchidaceae)

Pollinators are expected to respond to low reward availability in an inflorescence by visiting fewer flowers before departure, thus potentially causing reduced visitation, but also reduced geitonogamous selfing. I tested this hypothesis using Anacamptis morio, an orchid that does not reward its pollinators. Supplementation of inflorescences with artificial nectar did not result in an increase in fruit set on supplemented inflorescences compared to control inflorescences and tended to reduce pollinia removal. Supplementation resulted in reduced fruit quality, but there was no evidence that this was as a result of inbreeding depression. Behavioral experiments showed that pollinating bumble bees, as predicted, visited more flowers on supplemented inflorescences. Bumble bees also deposited more self-pollen on supplemented inflorescences, but this was marginally significant. Bumble bee queens removed significantly more pollinia from control inflorescences, while Bombus terrestris and B. lucorum workers did not. I conclude that while pollinators behaved as predicted, there was weak evidence that pollinia removal, pollen deposition, and fruit set followed the predictions of the hypothesis. I argue that this was probably because some pollinators were more efficient at removing and depositing pollen on control inflorescences, while others were not.     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

ROLES OF POLLEN AND OVARY AGE IN FOLLICLE PRODUCTION OF THE COMMON MILKWEED ASCLEPIAS SYRIACA

Using cross-pollination experiments, pollinia viability of the common milkweed Asclepias syriaca L. (Asclepiadaceae) did not change significantly over the five-day life of the flowers, but stigma receptivity decreased over threefold over this period. During most of the season pollinators inserted over 90% of their pollinia into flowers during the first two days following anthesis. However, pollinators were scarce during the first few days of flowering, and they inserted over 60% of their pollinia into flowers three or more days old. This behavior potentially decreased outcrossing success and increased the relative male contribution of the flowers. Pollinator visitation patterns may play an important role in determining the functional sexuality of these plants.     

Honey bees are the dominant diurnal pollinator of native milkweed in a large urban park

In eastern North America, the field milkweed, Asclepias syriaca L. (Asclepiadaceae), is used in planting schemes to promote biodiversity conservation for numerous insects including the endangered monarch butterfly, Danaus plexippus (Linnaeus) (Nymphalidae). Less is known about its pollinators, and especially in urban habitats where it is planted often despite being under increasing pressure from invasive plant species, such as the related milkweed, the dog‐strangling vine (DSV), Vincetoxicum rossicum (Kleopow) Barbar. (Asclepiadaceae). During the A. syriaca flowering period in July 2016, we surveyed bees in open habitats along a DSV invasion gradient and inspected 433 individuals of 25 bee species in 12 genera for pollinia: these were affixed to bees that visited A. syriaca for nectar and contain pollen packets that are vectored (e.g., transferred) between flowers. Of all bees sampled, pollinia were found only on the nonindigenous honeybee, Apis mellifera (43% of all bees identified), as well as one individual bumblebee, Bombus impatiens Cresson. Pollinia were recorded from 45.2% of all honeybees collected. We found no relationship between biomass of DSV and biomass of A. syriaca per site. There was a significant positive correlation between A. syriaca biomass and the number of pollinia, and the proportion vectored. No relationship with DSV biomass was detected for the number of pollinia collected by bees but the proportion of vectored pollinia declined with increasing DSV biomass. Although we find no evidence of DSV flowers attracting potential pollinators away from A. syriaca and other flowering plants, the impacts on native plant–pollinator mutualisms relate to its ability to outcompete native plants. As wild bees do not appear to visit DSV flowers, it could be altering the landscape to one which honeybees are more tolerant than native wild bees.     

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

Background and Aims

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

Methods

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

Key Results

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

Conclusions

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

Components of pollination effectiveness in Psychotria suerrensis,a tropical distylous shrub

In this paper I report components of effectiveness for pollinators of a tropical distylous shrub, Psychotria suerrensis (Rubiaceae), which is visited by a variety of bees, wasps, and butterflies, and by two species of hummingbirds. In the field, I measured the following components of effectiveness: frequency of visits, evenness of visits across plants, and diurnal pattern of visits. I also used flight-cage experiments to compare pollentransfer abilities of euglossine bees and heliconiid butterflies. Euglossine bees visited more frequently, visited earlier in the day, and visited a higher proportion of plants in the population than did other taxa. In flight cage experiments, bees and butterflies transferred similar amounts of pollen overall, but bees transferred significantly more inter-morph (compatible) pollen. For each component measured, euglossine bees appeared to be the most effective pollinators.     

The influence of distinct pollinators on female and male reproductive success in the Rocky Mountain columbine

Although there are many reasons to expect distinct pollinator types to differentially affect a plant's reproductive success, few studies have directly examined this question. Here, we contrast the impact of two kinds of pollinators on reproductive success via male and female functions in the Rocky Mountain columbine, Aquilegia coerulea . We set up pollinator exclusion treatments in each of three patches where Aquilegia plants were visited by either day pollinators (majority bumble bees), by evening pollinators (hawkmoths), or by both (control). Day pollinators collected pollen and groomed, whereas evening pollinators collected nectar but did not groom. Maternal parents, potential fathers and progeny arrays were genotyped at five microsatellite loci. We estimated female outcrossing rate and counted seeds to measure female reproductive success and used paternity analysis to determine male reproductive success. Our results document that bumble bees frequently moved pollen among patches of plants and that, unlike hawkmoths, pollen moved by bumble bees sired more outcrossed seeds when it remained within a patch as opposed to moving between patches. Pollinator type differentially affected the outcrossing rate but not seed set, the number of outcrossed seeds or overall male reproductive success. Multiple visits to a plant and more frequent visits by bumble bees could help to explain the lack of impact of pollinator type on overall reproductive success. The increase in selfing rate with hawkmoths likely resulted from the abundant pollen available in experimental flowers. Our findings highlighted a new type of pollinator interactions that can benefit a plant species.     

The floral biology ofThelymitra epipactoides (Orchidaceae), and the implications of pollination by deceit on the survival of this rare orchid

Thelymitra epipactoides has a highly variable visual display achieved through polychromatic flowers and variable inflorescence size, bearing between 7 and 31 flowers, which attract foraging polylectic bees. Only bees of the genusNomia were observed carrying pollinia and successfully pollinating the orchid. The genusNomia contains polylectic, pollen gathering species that store pollen in both the crop and scopa on the hind legs. The absence of a reward for the bees indicates the orchid is relying on deception to attract visitors. The relationship of deception to mimicry is discussed. Once on the flower, tactile, visual and possibly olfactory stimuli direct bees to the false anther formed by the voluminous column wings, where morphological adaptations of the flower ensure that the pollinarium is deposited on the gaster of the bee to effect pollination. — The lack of seed set observed on the Victorian coast appears to be due to the absence of pollinators from the heath and grassland communities in which the orchid grows. This may well be a consequence of the reduced number of plants flowering in the community (a result of the elimination of fire at these sites), thus not maintaining a floral community attractive to potential pollinators.     

RELATIVE POLLINATOR EFFECTIVENESS AND EVOLUTION OF FLORAL TRAITS IN SPATHIPHYLLUM FRIEDRICHSTHALII (ARACEAE)

We studied the relative effectiveness of different pollinators of Spathiphyllum friedrichsthalii Schott for 15 months on Barro Colorado Island, Panama. Pollen-foraging stingless bees (Apidae: Trigona) made 87% of floral visits. Experiments showed that these bees pollinate flowers, and correlations of fruit- and seed-set with visitation frequencies and floral contact times suggested that they were responsible for the majority of seeds produced. Fifteen species of fragrance-foraging, male euglossine bees (Apidae: Euglossini) collectively accounted for a small portion of seed-set in fewer than 27% of the inflorescences. Neocorynura (Halictidae) were pollen thieves and were unimportant as pollinators. We propose that euglossine and stingless bees differentially influence outcrossing rates and the evolution of floral traits of S. friedrichsthalii. Foraging behavior of male euglossines should allow for more long-distance pollen flow whereas stingless bees are likely to promote near-neighbor and geitonogamous pollinations. We discuss why the prolonged male phase of anthesis in this protogynous species may be maintained through pollination by stingless bees rather than male euglossines. Furthermore, although the floral fragrance is attractive to many species of male euglossines, it attracts few individuals. This condition may represent an intermediate step in the evolution of predominant pollination by male euglossines.     

Pollinator visitation closely tracks diurnal patterns in pollen release

Premise

Animal-pollinated plants face a high risk of pollen loss during its transfer. To limit the negative effect of pollen losses by pollen consumption and heterospecific transfer, plant species may adjust and stratify their pollen availability during the day (i.e., “schedule” their pollen presentation) and attract pollinators in specific time frames.

Methods

We investigated diurnal patterns of pollen availability and pollinator visitation in three coflowering plant species: Succisa pratensis with open flowers and accessible pollen, pollinated mainly by pollen-feeding hoverflies; Centaurea jacea with open flowers and less accessible pollen, pollinated mainly by pollen-collecting bees; and Trifolium hybridum with closed flowers and pollen accessible only after the active opening of the flower, pollinated exclusively by bees.

Results

The three plant species differed in the peak pollen availability, tracked by the visitation activity of their pollinators. Succisa pratensis released all pollen in the morning, while pollinator activity was still low and peaked with a slight delay. In contrast, C. jacea and T. hybridum had distinct pollen presentation schedules, peaking in the early afternoon. The pollinator visitation to both of these species closely matched their pollen availability.

Conclusions

Stratifying pollen availability to pollinators during the day may be one of several mechanisms that allow coflowering plants to share their pollinators and decrease the probability of heterospecific pollen transfer.     

Diurnal butterfly pollination in the orchid Habenaria radiata

Habenaria is one of the largest orchid genera, with its main centers of diversity in Brazil, eastern Asia and central and southern Africa. In spite of its diversity, the pollinators of few Habenaria spp. have been studied, while most published reports indicate that Habenaria spp. are mainly pollinated by moths. Here we report that the diurnal skipper Parnara guttata can be at least one of the pollinators of Habenaria radiata. Skippers have generally been thought of as nectar thieves, transferring few pollen grains. However, orchids transmit their pollen grains in large groups within pollinia. Such innovation can probably render the skipper a pollinator of H. radiata.     

Mechanisms of pollen deposition by insect pollinators

Summary Studies of pollen dispersal in insect-pollinated plants have often documented highly leptokurtic patterns of pollen deposition that can increase the likelihood of long-distance mating. To examine potential causes of highly leptokurtic deposition, we introduce four functions that arise when (1) the duration of pollinator visits to pollen sources is limited, (2) the rate of pollen deposition varies randomly among pollinators and/or among visits, (3) the rate of pollen deposition changes monotonically over time or (4) pollen is carried in layers or compartments on the pollinator's body that differ in deposition rate. Maximum likelihood techniques were used to fit deposition functions to data obtained from honey bees (Apis mellifera L.) visiting mustard plants (Brassica campestris L.) that contained a marker gene. Each of the alternative leptokurtic functions fit the experimental data better than a simple exponential function and the best-fit function predicted a mean pollen dispersal distance more than three times greater than the exponential. We argue that studies of pollen deposition need to test a broader range of deposition models to assess outcrossing distance in plant populations accurately.     

The structure and function of orchid pollinaria

Cohesive masses of pollen known as pollinia have evolved independently in two plant families — Orchidaceae and Asclepiadaceae. Yet, the bilateral symmetry of orchids has allowed a greater degree of specialization in pollination systems and a much greater diversity in the morphology of pollinaria — units comprising the pollinia(um) together with accessory structures for attachment to the pollinator. Pollinaria differ in the degree of cohesion of pollen in the pollinium, which may be soft, sectile (comprised of sub-units known as massulae) or hard. A single hard pollinium may contain more than a million pollen grains, yet pollen:ovule ratios in orchids are several orders of magnitude lower than in plants with powdery pollen due to the lack of wastage during transport to the stigma. Attachment of pollinia to the pollinator is usually achieved by means of a viscidium that adheres most effectively to smooth surfaces, such as the eyes and mouthparts of insects and beaks of birds. The stalk connecting a pollinium to the viscidium may be comprised of a caudicle (sporogenous in origin) and/or a stipe (derived from vegetative tissue), or be lacking altogether. Caudicles and stipes may undergo a gradual bending movement 20 s to several hours after withdrawal from the flower, the main function of which appears to be to reduce the possibility of geitonogamous pollination. Other mechanisms that promote outcrossing and pollen export in orchids include pollen carryover (achieved by sectile or soft pollinia), temporary retention of the anther cap, protandry and self-incompatibility (rare among orchids). Pollinaria ensure that large pollen loads are deposited on the stigma, thus enabling the fertilization of the large numbers of ovules in the flowers of Orchidaceae. Pollinaria also ensure efficient removal of pollen from the anther, minimal pollen wastage during transit, and a high probability of deposition on conspecific stigmas.     

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.     

Pollination genetics of hybridization in sympatric populations of asclepias exaltata and a. syriaca (asclepiadaceae)

Enzyme electrophoresis of pollinia and seeds was used to identify pollinia on pollinators, interspecific pollinations, and hybrid seed production in sympatric populations of Asclepias exaltata and A. syriaca. The frequency of mixed pollinia loads on pollinators was low. Only 4.1% (N= 169 pollinia) and 8% (N= 244 pollinia) of the pollinia collected from pollinators of A. syriaca and A. exaltata, respectively, were identified as pollinia from the other species. Natural levels of interspecific pollination, 3.6% (N = 166 pollinia) on A. syriaca and 3.1% (N = 228 pollinia) on A. exaltata, were typically lower than the number of foreign pollinia carried by pollinators. Hybrid seeds were identified in only two of 208 A. syriaca fruits and one of 178 A. exaltata fruits. Hybrid seeds were largely underdeveloped in the single A. exaltata fruit. High fruit set (34.7%) and near normal seed set following hand-pollination of A. syriaca with A. exaltata pollinia suggest that hybridization is more likely between these taxa when A. syriaca is the maternal parent. Our findings indicate that the likelihood of hybridization between A. exaltata and A. syriaca is remote. Nevertheless, introgressive hybridization has been documented in these species, which suggests that hybridization remains a strong evolutionary force even for species that infrequently interbreed.     

Variation in Pollen-Donor Composition among Pollinators in an Entomophilous Tree Species,Castanea crenata,Revealed by Single-Pollen Genotyping

Background

In plants, reproductive success is largely determined by the composition of pollen (i.e., self-pollen and outcross-pollen from near and distant pollen-donors) transported as a result of pollinator foraging behavior (e.g., pollen carryover). However, little evidence is available on how and to what extent the pollen carryover affects the pollen-donor composition and on which insect taxa are effective outcross-pollen transporters under field conditions. In this study, we explored roles of foraging behavior of insect pollinators on pollen-donor composition and subsequent reproductive success in a woody plant.

Methods

We performed paternity analyses based on microsatellite genotyping of individual pollen grains found on diurnal pollinators (i.e., bumblebee, small bee, fly, small beetle, and honeybee) visiting Castanea crenata trees.

Results

The outcross-pollen rate was highest in bumblebees (66%), followed by small bees (35%), flies (31%), and small beetles (18%). The effective number of pollen donors, representing pollen carryover, was greater in bumblebees (9.71) than in flies (3.40), small bees (3.32), and small beetles (3.06). The high percentages of pollen from outside the plot on bumblebees (65.4%) and flies (71.2%) compared to small bees (35.3%) and small beetles (13.5%) demonstrated their longer pollen dispersal distances.

Conclusions

All of the diurnal insects carried outcross-pollen grains for long distances via pollen carryover. This fact suggests that a wide range of insect taxa are potential outcross-pollen transporters for the self-incompatible C. crenata.     

Division of labour within flowers: heteranthery,a floral strategy to reconcile contrasting pollen fates

In many nectarless flowering plants, pollen serves as both the carrier of male gametes and as food for pollinators. This can generate an evolutionary conflict if the use of pollen as food by pollinators reduces the number of gametes available for cross‐fertilization. Heteranthery, the production of two or more stamen types by individual flowers reduces this conflict by allowing different stamens to specialize in ‘pollinating’ and ‘feeding’ functions. We used experimental studies of Solanum rostratum (Solanaceae) and theoretical models to investigate this ‘division of labour’ hypothesis. Flight cage experiments with pollinating bumble bees (Bombus impatiens) demonstrated that although feeding anthers are preferentially manipulated by bees, pollinating anthers export more pollen to other flowers. Evolutionary stability analysis of a model of pollination by pollen consumers indicated that heteranthery evolves when bees consume more pollen than should optimally be exchanged for visitation services, particularly when pollinators adjust their visitation according to the amount of pollen collected.     

FLORAL BIOLOGY AND REGULATION OF SEED SET AND SEED SIZE IN THE LILY,CLINTONIA BOREALIS

We studied the reproductive ecology of Clintonia borealis, a clonal understory species, in eastern Ontario. Flowers are protogynous and require insect pollination for outcrossing and maximum seed set. Most pollination is done by nectar-gathering bumble bees. We recorded pollen deposition and flowering patterns in 1983. Flowering lasted for about 10 days in mid-June. Stigmatic pollen loads were lowest at the onset of blooming, reflecting the scarcity of male-phase flowers. Pollination increased significantly by peak bloom and subsequently remained high. However, since bees fly mostly between neighboring stems, much of the pollen transferred may have been geitonogamous. In 1983, seed set per flower was not increased by supplementing pollination or by reducing the number of fruits competing for resources per stem. Rather, flowers set more seeds (14%) only if both treatments were performed simultaneously. Seed size was increased by 25% when competitive fruits were removed and by 5% more when pollen was added under the removal treatment. Further work in 1984 showed that pollination effects may be related to changes in pollen source. Selfed flowers set fewer and smaller seeds than outcrossed ones when screened from insects and hand pollinated. Outcrossing distance had little effect on seed set or seed size. Thus, in C. borealis seed number and seed size are limited by a balance between maternal resource availability and the amount of outcrossing provided by pollinators.