The maintenance of mixed mating by cleistogamy in the perennial violet Viola septemloba (Violaceae)

The production of both potentially outcrossed (chasmogamous) and obligately self-fertilized (cleistogamous) flowers presents a clear exception to the prediction that the only evolutionarily stable mating systems are complete selfing and complete outcrossing. Although cleistogamy has evolved repeatedly, the reason for its stability is not known for any species. We tested the hypothesis that the production of cleistogamous and chasmogamous flowers by a perennial violet constitutes adaptive phenotypic plasticity. We manipulated the season of flowering for each flower type and determined fruit set and the germination percentage of seeds produced by cleistogamous and chasmogamous flowers to test the hypothesis that adaptive plastic response to seasonal environmental variation makes mixed mating stable. Cleistogamous flowers had greater fruit set in all seasons and produced seeds with germination percentages as great as or greater than those from chasmogamous flowers. The consistent advantage of cleistogamous flowers is clearly not consistent with a role of adaptive plastic response to seasonal variation. The biomass cost of seed production by chasmogamous flowers was nearly three times that for cleistogamous flowers. Explaining why chasmogamous flower have not been eliminated by natural selection requires that this difference be balanced by an advantage to chasmogamous flowers that has not yet been identified.     

Evidence for selective fruit abortion in Banksia spinulosa (Proteaceae)

Fruit set is consistently low in the genus Banksia (Proteaceae). One explanation for low fruit:flower ratios is that excess flowers allow plants to selectively abort inferior progeny thereby increasing the average quality of the seed crop. We examined whether self-pollinated fruits were aborted selectively in a partially self-compatible population of Banksia spinulosa. Inflorescences were divided in half and pollinated with cross pollen on both sides, self pollen on both sides or cross pollen on one side and self pollen on the other. Seed production was reduced significantly by 38% on fully self-pollinated compared to fully cross-pollinated inflorescences, indicating partial self-incompatibility or inbreeding depression. On inflorescences pollinated with both cross and self pollen, selfed seed set was reduced by 63% compared to crossed seed set. On the selfed half of these inflorescences, fewer fruits and seeds were produced, and more fruits had aborted seeds than on randomly selected halves of fully self-pollinated inflorescences. These results suggest that self-fertilized embryos were developed in the absence of outcrossed progeny but were selectively aborted when outcrossed progeny were present on inflorescences. To determine the amount of outcrossing occurring in the study population, outcrossing rates (t) of naturally pollinated plants were examined by electrophoretic assessment of progeny arrays. Values of I did not differ significantly from one, indicating complete outcrossing. High outcrossing, despite the high probability of self pollen receipt under natural conditions, is consistent with the conclusion that selective abortion occurs in B. spinulosa.     

THE RELATIVE COSTS OF SELF- AND CROSS-FERTILIZED SEEDS IN IMPATIENS CAPENSIS (BALSAMINACEAE)

By using a generally applicable technique that involves monitoring the development and survivorship of flowers and seed capsules, I estimated the material and energetic costs of producing self- and cross-fertilized seeds in Impatiens capensis. All flowers and fruits on six plants were censused intensively for the two-month period of reproduction. Cleistogamous (selfing) flowers ripened seed in about 24 days, compared to about 36 days for the chasmogamous (outcrossing) flowers. In terms of dry weight, selfed seeds cost about two-thirds as much as outcrossed seeds: 12.4 versus 18.4 mg dry weight per seed. When adjusted to the currency of calories, and including an independent estimate of pollen and nectar production in outcrossing flowers, I estimate the costs to be about 65 and 135 calories per selfed or outcrossed seed. Sources of error include the accuracy of the estimates of flower and fruit weight, and possible differences among the developmental stages in respiratory costs. The cost discrepancy implies that outcrossed seeds should possess a countervailing fitness advantage large enough to offset their greater energetic cost.     

The effect of timing of pollination on the mating system and fitness ofKalmia latifolia (Ericaceae)

Plant mating systems are known to vary within species and some immediate ecological factors have been found to be associated with the geographic distribution of selfing. The environmental condition of the maternal plant may influence the production of selfed seed relative to outcrossed seed. This study investigated the effect of late pollination on the mating system of Kalmia latifolia, a long-lived perennial shrub. A 2 × 2 experimental design was used to determine whether reproductive success declines over the course of the flowering season and whether there was an interaction between pollination time (early vs. late in the season) and pollen type (self-fertilized vs. outcrossed). An interaction of this sort would indicate context-dependent fitness of selfed seeds compared to outcrossed seeds and, thus, show an ecological influence over a plant's mating system. Relative fitness was assessed in terms of female reproductive success. Timing of pollination did not affect abortion of outcrossed seeds; however, delay in pollination increased abortion of selfed seeds by 34.7%. Thus, it appears that plants selectively aborted selfed seeds rather than outcrossed seeds and this selection was more intense at the end of the season. An ecological factor such as time of pollination may affect the mating system of K. latifolia.     

Plants,position and pollination: Planting arrangement and pollination limitation in a revegetated eucalypt woodland

The spatial arrangement of plants is a key determinant of pollination services in natural ecosystems. Despite this, plant arrangements are rarely considered an important characteristic of revegetated communities and this may be limiting successful pollination dynamics in these re‐created systems. We assessed if aggregated South Australian Blue Gum (Eucalyptus leucoxylon, F. Muell) had greater pollination success than more dispersed trees by measuring fruit set, seed production and germination in a revegetated eucalypt woodland. The proportion of buds that developed into fruit (fruit set) was similar between aggregated (five flowering Blue Gum within 30 m) and dispersed (no flowering Blue Gum within 30 m) trees (<14%). Aggregated and dispersed trees produced a similar number of seeds per fruit in 2015, when fewer trees flowered and those that flowered produced fewer flowers (ca 8 seeds/fruit). In 2016, when flowering was more abundant, aggregated trees produced more seeds per fruit than dispersed trees (12 seeds/fruit compared to 4 seeds/fruit). Despite differences in seed number, there was no difference in the quality of seed produced, indicating outcrossed pollen was received by both aggregated and dispersed trees. Although outcrossing was likely, this did not prevent pollination limitation, with the addition of outcrossed pollen increasing fruit and seed set for trees in both arrangements. Consequently, we propose that planting individuals of the same species (conspecifics) in clumps, rather than intermixing species, can increase seed set in revegetated eucalypts, particularly in good flowering years. However, aggregated plantings will be most effective if the needs and foraging behaviour of pollinators are also considered, to attract and sustain pollinators in revegetated systems, thereby minimizing the risk of pollination limitation.     

Cleistogamy in the rare high Andean perennial herb Cryptantha capituliflora (Boraginaceae)

 Axial and apical flowers of Cryptantha capituliflora were analyzed with regard to morphology and pollen tube growth to assess the occurrence of cleistogamy. Although intermediate floral forms do occur, cleistogamous flowers were significantly smaller than chasmogamous flowers, had fewer anthers, and showed a distinctive stigmatic surface. Chasmogamous flowers can be cross-pollinated. Nevertheless, the growth of self-pollen tubes in few chasmogamous buds jointly with flower characters suggests that these flowers can probably produce fruits through autonomous selfing. The mean seed number per fruit did not differ between fruits from chasmogamous and cleistogamous flowers. Cleistogamous flowers were only observed in axial inflorescences, which are completely covered by the leaf. Other species of section Cryptantha also show the same trend, with cleistogamous flowers located in the lower half of the stems. This pattern is discussed in relation to dissimilarities in the outcrossing opportunities between flower types within the plant. Received May 22, 2002; accepted November 14, 2002 Published online: March 20, 2003     

The role of pollination level on the reproduction of females and hermaphrodites in the gynodioecious plant Gypsophila repens (Caryophyllaceae)

In gynodioecious plant species, females are expected to have more resources available for maturing seeds because pistillate flowers are smaller, do not produce pollen, and are thus less costly that perfect flowers. The potential female advantage arising from more abundant resources is, however, likely to vary depending on whether seed production is limited by resource or pollen availability. Here we experimentally investigated the influence of pollen and resource limitation on female advantage in a gynodioecious species using two levels of pollination. Total seed production of females was always greater than that of hermaphrodites: females produced more flowers and more fruits that contained similar numbers of seeds of similar mass. Under low pollination, female and hermaphrodite plants allocated resources to increased flower production rather than to increased seed size or quality. We did not detect any influence of pollen or resource limitation on female advantage, which remained similar under low (= abundant resources) and full pollination. Outcrossed fruits performed better than selfed fruits when the same plant received both selfed and outcrossed pollen on different flowers. These differences were not greater under high pollination, possibly because resources available for each fruit did not differ between our pollen intensity treatments.     

Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae)

The reproductive biology of an exotic species will affect its ability to become naturalized and invasive in non-native habitats. Rhododendron ponticum is an ecologically damaging exotic weed in the British Isles, which spreads predominantly by seed. I investigated how inbreeding and outcrossing affect seed production and germination in a wild population of this species in Ireland. Experimental manipulations revealed low fruit and seed set when insects were excluded from flowers, suggesting that this species has limited capability for spontaneous autogamy. Hand-pollination treatments showed that, although flowers are self-compatible (with self and same plant pollen), higher levels of seed set occur following outcrossing (xenogamy). There was no significant difference in rate of germination of seeds from inbred or outcrossed treatments. The addition of xenogamous pollen to open flowers did not increase fruit or seed set, suggesting that flowers in this population are not pollen limited: native generalist pollinators, mainly bumblebees ( Bombus spp.), are providing an adequate pollinator service. This work demonstrates that outcrossing increases seed set and pollinators are required to facilitate this. Hence, generalist native pollinators can promote invasion by exotic plants. Native pollinators can clearly play an important part in alien species invasion.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 373–381.     

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.     

Differential outcrossing rates in dispersing and non-dispersing achenes in the heterocarpic plant Crepis sancta (Asteraceae)

The association between mating systems and dispersal in plants has been studied mostly in cleistogamous species where, generally, seeds produced by cleistogamous (selfed) flowers are less dispersed than seeds produced by chasmogamous (potentially outcrossed) flowers. In heterocarpic Asteraceae, non-dispersing fruits (achenes) are produced at the periphery of the capitulum (outer florets) whereas dispersing achenes are produced by inner florets in the same capitulum. Since all the florets are protandrous, the outer floret developing first are in female phasis when anthesis of inner florets takes place. Thus, outer florets can be potentially selfed by the inner florets of the same capitulum whereas the latter must be pollinated by flowers of other capitula. Therefore outer florets should be more inbred than inner florets. To test this hypothesis, we measured the natural outcrossing rate in outer and inner florets using allozymes in three populations of the heterocarpic Crepis sancta. The results showed that the outcrossing rate was highest for non-dispersed achenes. Moreover, among the outcrossed achenes within a capitulum it was observed that the number of paternal parents of non-dispersing achenes was higher than for dispersing achenes. The pattern observed was therefore the opposite to the pattern of cleistogamous plants and contradicts the putative pollination mechanism we proposed for Asteraceae. The results agree with the predictions of sib competition theory which considers that outcrossing may minimize competitive interactions among relatives (sibs) falling near the mother plant. Higher outcrossing rate in outer florets could also occur because pollinators are more attracted to these florets.     

THE INFLUENCE OF THE MATING SYSTEM ON SEED SIZE VARIATION IN CRINUM ERUBESCENS (AMARYLLIDACEAE)

We present evidence that extreme seed size variation in fruits of Crinum erubescens (range: 0.1 to 66.5 g per seed) occurs when mating pairs are inbred, either from selfing or biparental inbreeding. Several relatively uniform seeds of intermediate size are produced when pollen from several pollen donors is applied simultaneously to a flower. Selfed fruits and some fruits pollinated with a single pollen donor produce both large and small seeds, although selfed fruits produce fewer seeds than outcrossed fruit. These results are contrary to the hypothesis that variation in seed size is attributable to either pollen competition or differential allocation of maternal resource to seeds of different genotypes.     

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.     

Paternal effects on seed mass in Arabidopsis thaliana

Who is in control of seed size, and do some fathers sire bigger seeds than others? We used isogenic male-sterile genotypes of the Arabidopsis thaliana accessions Col and Ler. By fertilising flowers side-by-side with either pollen from the same accession ('self-pollination') or pollen from another accession (outcrossing), we compared, on the same mother plant, seed set of flowers that were very similar in resource status. Some paternal genotypes had a significant effect on seed mass, with the most extreme father siring seeds 15.3% heavier than seeds resulting from 'self-pollination'. There was no correlation between seed mass of paternal parents and the seeds they sired. We discuss the evolution of seed size as a tug-of-war between parent and offspring.     

Analysis of mating system and genetic structure in the endangered,amphicarpic plant,Lewton’s polygala (<Emphasis Type="Italic">Polygala lewtonii)</Emphasis>

Polygala lewtonii is a federally endangered, amphicarpic plant with a mixed mating system and three types of flowers: (1) aboveground, chasmogamous flowers (i.e., open-pollinated; CH), (2) aboveground, cleistogamous flowers (i.e., closed, selfing; CL) and (3) CL flowers on belowground stems (amphicarpy). Aboveground seeds are ant-dispersed, whereas belowground seeds are spaced across the length of the rhizome. Here, we collected individuals of P. lewtonii at both range-wide and fine geographic scales and genotyped them at 11 microsatellite loci. We analyzed patterns of genetic diversity and structure to understand: (1) the predominant mating system (selfing or outcrossing), (2) the movement of pollen and seeds across the landscape, and (3) the optimal strategy to conserve the full range of genetic variation. P. lewtonii reproduces predominantly by selfing or bi-parental inbreeding, but reproduction occurred through each of the three flower types. Some individuals produced by selfing/inbreeding were tightly clustered spatially, and were likely produced either by belowground flowers or by aboveground flowers with limited seed dispersal. Other selfed/inbred individuals were spatially separated (maximum of 15 m), and were likely produced by aboveground flowers followed by seed dispersal by ants. Fine-scale patterns of genetic structure indicate that some gene flow is occurring among aboveground CH flowers but both pollen and outcrossed seeds are moving limited distances (maximum of 0.5 km). Because genetic variation is structured at a fine spatial scale, protecting many populations is necessary to fully conserve the genetic variation in P. lewtonii. Conservation seed banking, if accompanied by research on seed germination requirements, may also contribute to the effective protection of genetic variation in P. lewtonii.     

Elevation-related variation in the population characteristics of distylous Primula nivalis affects female fitness and inbreeding depression

The population characteristics of distylous species are highly sensitive to stochastic natural selection pressure.Therefore,populations growing under different environmental conditions may vary in floral morph ratios,potentially affecting female fitness and leading to inbreeding depression.However,the variation in offspring quality among populations as a result of inbreeding depression is poorly understood in distylous species.This study investigates variations in plant density,seed mass,seed viabilityfemale fitness,and post-dispersal inbreeding depression in both sexual morphs(long-styled and shortstyled plants)of the distylous Primula nivalis that were subjected to different pollination treatments along an elevational gradient from 1657 to 2704 m a.s.l.Population characteristics(morph plant density and ratio)and fruit set were significantly affected by sexual morph and elevation.Plant density and fruitset frequencies were lower for short-styled than for long-styled plants at 2704 m a.s.l.The seeds from the cross-pollinated flowers of both morphs were higher in quality than those of self-pollinated flowers.The female fitness of seeds from cross-pollinated flowers of both morphs was higher than that of seeds from open-pollinated and self-pollinated flowers.The female fitness of seeds from long-styled flowers was higher than that of seeds from short-styled flowers at all elevations.Inbreeding depression increased with elevation among plants with short-styled flowers but not among those with long-styled flowers.Variation in the elevation-dependent mating system might influence female fitness and affect inbreeding depression in both floral morphs.In conclusion,the low quality of seeds from short-styled flowers at high elevations might decrease short-styled flower frequency,affecting population characteristics.     

GENETIC CONSEQUENCES OF OUTCROSSING IN THE CLEISTOGAMOUS ANNUAL,IMPATIENS CAPENSIS. II. OUTCROSSING RATES AND GENOTYPIC CORRELATIONS

The genetic consequences of a plant's mating system depend on both the degree of outcrossing and the genetic relationship between mates. We examined the electrophoretic genotypes of seeds derived from cleistogamous (CL) and chasmogamous (CH) flowers in six populations of the facultatively cleistogamous annual, Impatiens capensis. Multilocus estimates of the outcrossing rates for the strongly protandrous CH flowers ranged from 0.29 to 0.71 and were higher than estimates based on single-locus data. Such results suggest that the CH flowers experience variable levels of both geitonogamous self-fertilization and biparental inbreeding. A new and generally applicable technique based on the relative level of inbreeding within progeny groups provided direct estimates of the correlation between the genotypic values of outcrossed mates. These correlations varied widely among populations and contributed up to half of the inbreeding observed among the CH progeny. Such biparental inbreeding biases estimates of the outcrossing rate based on the mixed-mating model downward and influences mating-system evolution by decreasing the “cost of meiosis.”     

The Effect of Pollen Source vs. Flower Type on Progeny Performance and Seed Predation under Contrasting Light Environments in a Cleistogamous Herb

Dimorphic cleistogamy is a specialized form of mixed mating system where a single plant produces both open, potentially outcrossed chasmogamous (CH) and closed, obligately self-pollinated cleistogamous (CL) flowers. Typically, CH flowers and seeds are bigger and energetically more costly than those of CL. Although the effects of inbreeding and floral dimorphism are critical to understanding the evolution and maintenance of cleistogamy, these effects have been repeatedly confounded. In an attempt to separate these effects, we compared the performance of progeny derived from the two floral morphs while controlling for the source of pollen. That is, flower type and pollen source effects were assessed by comparing the performance of progeny derived from selfed CH vs. CL and outcrossed CH vs. selfed CH flowers, respectively. The experiment was carried out with the herb Ruellia nudiflora under two contrasting light environments. Outcrossed progeny generally performed better than selfed progeny. However, inbreeding depression ranges from low (1%) to moderate (36%), with the greatest value detected under shaded conditions when cumulative fitness was used. Although flower type generally had less of an effect on progeny performance than pollen source did, the progeny derived from selfed CH flowers largely outperformed the progeny from CL flowers, but only under shaded conditions and when cumulative fitness was taken into account. On the other hand, the source of pollen and flower type influenced seed predation, with selfed CH progeny the most heavily attacked by predators. Therefore, the effects of pollen source and flower type are environment-dependant and seed predators may increase the genetic differences between progeny derived from CH and CL flowers. Inbreeding depression alone cannot account for the maintenance of a mixed mating system in R. nudiflora and other unidentified mechanisms must thus be involved.     

Reproductive versatility in legumes: the case of amphicarpy in Trifolium polymorphum

Amphicarpy is a fascinating reproductive strategy, defined as fruit produced both below the soil surface and as aerial fruit on the same plant. Trifolium polymorphum is a grassland species subject to herbivory that combines amphicarpy with vegetative reproduction through stolons. Underground flowers have been described as obligate autogamous and aerial ones as self‐compatible allogamous, with aerial floral traits favouring cross‐pollination. In the present work we performed different pollination treatments on aerial flowers to analyse rates of pollen tube development and offspring fitness, measured as fruit set, seed production and germination percentage. This last variable was compared to that of seeds produced underground. No significant differences were found between fruit set in self‐ and cross‐pollinations. Seed production was higher in self‐pollinations, which is consistent with the higher rate of pollen tube development observed in self‐crosses. Spontaneous self‐pollination is limited in aerial flowers; thus pollen transfer by means of a vector is required even within the same flower. Germination tests showed that aerial seeds produced after self‐ and cross‐pollination did not differ in fitness, but underground seeds had higher germination percentage than aerial ones. Thus, we conclude that T. polymorphum has a mixed mating system. In grasslands with heavy grazing pressure, clonal propagation and underground seed production ensure persistence in the field. An intermediate level of selfing in aerial flowers ensures offspring, but morphological (herkogamy) and functional (dicogamy) floral traits maintain a window to incorporate genetic variability, allowing the species to tolerate temporal and spatial pressures.     

Sex expression and breeding strategy in <Emphasis Type="Italic">Commelina benghalensis</Emphasis> L.

This paper describes the results of a series of experiments conducted to unravel the patterns of sex expression and reproductive output in a fascinating species with high variation in sexuality. Commelina benghalensis L., an andromonoecious rainy season weed, bears male and bisexual flowers in axillary spathes of all the plants investigated. Bisexual flowers are of two types; chasmogamous (CH) and cleistogamous (CL). The former are borne on subaerial and the latter on subterranean shoots, in addition to those on aerial spathes. Three populations of the species, designated JU1, JU2 and JU3, were scanned for three consecutive years from 1996 to 1998, and the number and distribution of male, CH and CL flowers per plant were found to vary. The mere number of CH/CL flowers per plant is by itself not an accurate measure of mixed mating. It is necessary to confirm that CH flowers actually outcross and, if they do so, to what extent. Comparison of the pollen/ovule (P/O) ratio and percentage pollen germination on the stigmas of the CH and CL flowers have been used as indices of the pollination system. Confirmation of this was sought from the fruit and seed sets obtained after manual pollination of emasculated flowers with self- and cross-pollen. Results so obtained were compared with those of natural pollination. In the majority of CH flowers, the male and female reproductive phases (i.e. anther dehiscence and stigma receptivity) overlap, providing for self-pollination. However, two exceptions to this general behaviour were found in some plants of all the three populations. In some CH flowers, the female phase matures prior to anther dehiscence while in others, the anthers are sterile. Such plants, designated as variants 1 and 2, respectively, facilitate cross-pollination. While the CL flowers contribute to the production of selfed progeny, the variants of CH ones permit formation of outcrossed progeny, indicating a mixed mating strategy in C. benghalensis.     

Conservation implications of the reproductive ecology of Agalinis acuta (Scrophulariaceae)

Reproductive ecology of Agalinis acuta was investigated by examining potential for self-fertilization before and at anthesis, reproductive output from outcrossed vs. selfed matings, and effects of browsing, plant size, and conspecific plant density on seed and fruit production. These features of a plant species can provide indirect information pertinent to conservation such as patterns and maintenance of genetic diversity, risk associated with inbreeding depression, and changes in pollinator abundance or effectiveness. The species is self-compatible, with 97% of selfed flowers setting fruit; pollinators were not required for reproduction. However, seed set in self-pollinated fruits averaged 17-20% less than that in open-pollinated fruits. Geitonogamous and facilitated selfing are possible throughout anthesis and autonomous selfing is possible late in anthesis as corollas abscise. Delaying self-pollination until after outcrossing opportunities likely limits selfing rates and thus reduces risks associated with inbreeding but allows reproduction in absence of pollinators. Supplementing pollen on open-pollinated flowers yielded no additional seed set over controls. Neither early-season browsing of primary stems nor conspecific plant density had significant effects on number of fruits per plant, on fruit size, or on number of seeds from open-pollinated flowers. Currently, reproduction appears to be high (about 2400 seeds/plant), and future risks due to lack of genetic diversity are likely low.