Pollination mechanisms and pollinators are reported for a total of 137 species (75% of the non-abiotically pollinated flora) as they occur at three altitudinal levels (subandean scrub: 2,200-2,600 m; cushion-plant zone: 2,700-3,100 m; subnival feldfield: 3,200-3,600 m) in the Andean (alpine) zone on the Cordon del Cepo (33⚬17′S) in central Chile as part of community oriented research in reproductive biology in the high temperate Andes of South America. Only around 4% of the species studied faile
d to be visited by potential pollinators. Hymenopterans (principally bees) are important pollinators of 50% of the biotically pollinated flora, butterflies of 24% and flies of 46%. Other vectors include beetles, moths, and hummingbirds. An estimated 17% of the flora is anemophilous. Bee species-richness, specialist feeding, and melittophily reach maxima in the subandean scrub; thereafter, bees diminish rapidly in number, with bees pollinating only 13% of the subnival flora as contrasted with 68% of the subandean flora. Although fly and butterfly species-richness also decline with increasing altitude, the proportions of species pollinated by these vectors actually increases. High-altitude populations of melittophilous species with broad altitudinal ranges are invariably serviced by fewer bee species as compared with lower populations. The rich bee fauna at the lower end of the Andean zone in central Chile appears to have resulted from upward colonization from that of the subtending lowland Mediterranean sclerophyllous woodland vegetation. Altitudinal variation in pollination spectra is discussed in relation to contrasting life history characteristics and different modes of thermoregulation in the insect groups involved.
Altitudinal changes in breeding and pollination systems of tropical montane plants were studied in 13 species of Espeletia growing in the Venezuelan Andes from 2,000 to 4,300 m. Hand pollination tests showed that all species were strongly self-incompatible. The four species found only above 4,000 m had up to 10% median seed set in self-pollinated heads, which was significantly higher than the lower elevation species. Nine species were insect-pollinated, with large bees the major pollinator g
roup. An endemic p?ramo hummingbird, Oxypogon guerinii, was an important visitor of E. schultzii in three populations examined. Experimental bagging experiments showed that the four high elevation species were wind-pollinated, further evidenced by the lack of pollinator visits and markedly different capitulum morphologies. Open-pollinated seed set in two wind-pollinated species, E. spicata and E. timotensis, was strongly dependent on the population's flowering density, which varied significantly from year to year. The shift from insect to wind pollination in Espeletia can be related to the low pollinator availability at high elevations in the Andes, protection of the capitula from snow and daily frosts, and the extremely long flowering periods of individual heads.
This study explores the effects of emergence time and reproductive phenology on seed number, seed size, and seedling survival in a population of the alpine buttercup, Ranunculus adoneus. Phenology in this snow bowl population is structured by snow depth. Plants in late melting interior portions of the bowl emerged and flowered 3 to 4 wk after those in early melting zones at the bowl perimeter during the summers of 1988 and 1989. Flowering time differences of buttercups across the bowl were consi
stent from one year to the next. In 1988, late flowering plants tended to set fewer seeds than early flowering ones; in 1989 no decrease in seed number accompanied flowering date. Path analysis showed that equal fecundity in early and late emerging portions of the bowl population during 1989 resulted from balancing spatial and temporal constraints on seed production. Spatial aspects of habitat quality improved toward the interior of the bowl, but temporal regimes deteriorated in these late melting sites. In both 1988 and 1989 seed size declined with delays in flowering. Path analysis of 1989 data showed that because of reduced time for seed growth, plants in late melting portions of the bowl set smaller seeds than those in earlier melting zones. Differences in seed size due to parental phenology are likely to influence fitness in snow buttercups. Under natural conditions, seedlings from large seeds (≥0.65 mg) have sixfold higher survival than do those from smaller seeds (<0.65 mg). We conclude that seedling recruitment may be infrequent in late-melting portions of the snow bowl due to delayed parental phenology.
Flowering phenology, fruit set, and pollinator frequency of the ericaceous alpine dwarf shrub, Rhododendron aureum, were compared among three quadrats arranged along a snowmelt gradient on a slope. Bagging and self-pollination experiments showed that R. aureum was physiologically self-compatible, but pollinator visitation highly enhanced fruit set. Depending on the lateness of snowmelt, flowering time varied from mid-June to late July among quadrats. Pollinator visitation increased as the flower
ing season progressed, and fruit set was significantly higher in the quadrat with latest snowmelt. Emasculation experiments suggested that later flowering might promote cross-pollination. Thus, later flowering was advantageous for effective pollination. On the other hand, later-flowering plants often failed to set fruits because of the onset of autumn frost and snow before fruit maturation.
Surface features, anatomy, and ultrastructure of the floral nectary of Eccremocarpus scaber (Bignoniaceae), pollinated predominantly by the largest-known hummingbird (Patagona gigas gigas), were studied together with nectar sugar content and secretion rate. The annular disk nectary comprises epidermis, secretory and ground parenchyma with intercellular spaces, and branched vascular bundles terminating in the secretory parenchyma where only phloem is found. Amyloplasts and vacuoles increase in si
ze throughout development, the latter becoming sites of organelle degradation. Transferlike cells in nectary phloem and P-proteinlike fibrillar material in phloem parenchyma were observed. Flowers produced around 32 μl of nectar (mostly after anthesis) with 11 mg of sugar composed of fructose, glucose, sucrose, and maltose in a ratio of 0.34:0.32:0.17:0.17. Morphological studies as well as the presence of maltose and glucose in nectar suggest storage of the originally phloem-derived sugars as starch with its subsequent hydrolysis. The low sucrose/hexose ratio (0.25) and high nectary secretion force (nectar per flower biomass) observed places E. scaber close to large-bodied bat-pollinated plants. A hypothesis based on nectar origin and nectar secretion is advanced to explain pollinator-correlated variation in sucrose/hexose ratio.
Pollination by deceit in Myristica insipida, a beetle‐pollinated nutmeg, was hypothesized to operate on intersexual differences in flower production and longevity, producing a daily fluctuation between floral display maxima and minima. Sticky traps were used to continuously monitor beetle activity. Flower production and naturally occurring intersexual differences in display were recorded. Male and female trees flowered in synchrony producing daily display maxima at 1800–0600 and display minima a
t 1400–1800. Rewarding male trees produced three times the number of flowers of female trees, but the greater longevity of female flowers reduced the intersexual difference in display maxima to a factor of two. There was no intersexual difference in display minima. Beetles were demonstrated to be sensitive to differences in both maximum and minimum displays on rewarding male trees, a necessary prerequisite for directional selection on display size. Beetle captures were significantly higher at male trees during floral display maxima, and no intersexual differences in capture rate occurred during floral display minima. However, capture rates at male trees did not decline as predicted, and the pattern of captures was consistent with crepuscular activity. Beetle captures at male and female trees were lower from 1800 to 0600 and 1000 to 1400, and higher from 0600 to 1000 and 1400 to 1800, but the differences were only significant at female trees. These data suggests that foraging errors are numerous, frequent, and the result of overall foraging activity.
An understanding of the process of submarine pollination should provide insight into the evolutionary and reproductive ecology of the marine angiosperms (seagrasses). The flow around the reproductive organs of the seagrass Zostera marina L. (Potamogetonales) was, therefore, examined in a flow chamber. The phenological emergence of flowers during (1) pollen capture and (2) pollen release, and by fruit during (3) seed release, led to a reduction in flow rate toward the inflorescence. This change i
n flow due to floral emergence was associated with a 50% increase in the fluid shear stress [tau = (2.2 _ 0.3) x 10-3 Pa for an immature flower vs. tau = (3.1 _ 0.5) x 10-3 Pa for a receptive flower]. The Reynolds number (Re) and fluid shear stress around inflorescences and infructescences were comparable, indicating a dynamic similarity in the processes of pollen capture and fruit dehiscence [Re = 47 _ 5, tau = (1.6 _ 0.3) x 10-3 Pa for inflorescences; Re = 38 _ 5, tau = (1.3 _ 0.1) x 10-3 Pa for infructescences]. These results indicate that the emergence of reproductive organs leads to changes in fluid shear stress, which will affect the release, transport, and capture of particles including pollen. Theoretical considerations of these observations using aerosol-filtration theory suggest that pollen capture in Z. marina occurs through direct interception of pollen by stigmas.
In most experimental hybridizations between oilseed rape (Brassica napus) and weedy B. campestris, either intra- or interspecific pollen has been applied to individual flowers. Under field conditions, however, stigmas will often receive a mixture of the two types of pollen, thereby allowing for competition between male gametophytes and/or seeds within pods. To test whether competition influences the success of hybridization, pollen from the two species was mixed in different proportions and appl
ied to stigmas of both species. The resulting seeds were scored for paternity by isozyme and randomly amplified polymorphic DNA analysis. Using data on the proportion of fully developed seeds and the proportion of these seeds that were hybrids, a statistical model was constructed to estimate the fitness of conspecific and heterospecific pollen and the survival of conspecific and heterospecific zygotes to seeds. B. campestris pollen in B. napus styles had a significantly lower fitness than the conspecific pollen, whereas no difference between pollen types was found in B. campestris styles. Hybrid zygotes survived to significantly lower proportions than conspecific zygotes in both species, with the lowest survival of hybrid zygotes in B. napus pods. This is in contrast to the higher survival of hybrid seeds in B. napus than in B. campestris pods when pollinations are made with pure pollen. Altogether, the likelihood of a foreign pollen grain producing a seed was much lower on B. napus than on B. campestris. In addition, pods on B. napus developed to a lower extent the more heterospecific pollen was in the mix, whereas this had no effect on B. campestris.
Flow chamber observations of the filamentous pollen of Zostera marina L. (Potamogetonales) revealed that pollen rotated and moved toward inflorescences where they were captured by stigmas. The mechanics of this abiotic pollination process were examined and found to be related to the flow environment around emergent flowers. The translational movement of pollen was imparted by the advection of the fluid (e. g., pollen kinetic energy, K, ranged from 0.8 × 10−−14 to 2.4 × 10−−14 J, and the average
K of the fluid was ≈ 0.7 × 10−−14 J), while the rotational motion was imparted by the fluid shear stress (τ) within the velocity gradient (e. g., pollen shear stress, σt = ωµ where ω is the rotational velocity and µ is the dynamic viscosity, ranged from 3.4 × 10−−14 to 26 × 10−−14 Pa, and the average fluid shear stress was τ ≈ 10 × 10−−4 Pa; Ackerman, 1997, American Journal of Botany 84: 1099–1109). These results indicate that there is a greater potential for pollination by filamentous pollen relative to spherical pollen. Functionally, while spherical pollen needs to be directly upstream from stigmas to be captured, filamentous pollen need only be in the vicinity of inflorescences and flowers to be captured by stigmas. Thus, in addition to direct interception on stigmas, filamentous pollen can be captured while they rotate past flowers or when they are redirected through the velocity gradient towards flowers. Filamentous pollen is an adaptation to submarine pollination in seagrasses.
We document the pollination biology and mating systems of Neobuxbaumiamezcalaensis and Neobuxbaumia macrocephala, two Mexican giant columnarcacti. These two species form mixed forests in the western Tehuacan Valley,Mexico. The flowers of both N. mezcalaensis and N. macrocephala exhibitnocturnal anthesis, are self-incompatible, and are pollinated primarily bythree species of nectar-feeding bats (Choeronycteris mexicana,Leptonycteris curasoae, and Leptonycteris nivalis). Neobuxbaumiamezcalaensis i
s androdioecious, a breeding system that appears to beuncommon among Cactaceae. Neobuxbaumia macrocephala is hermaphroditic. Wehypothesize that columnar cacti show a geographical dichotomy in floralbiology specialization that probably can be related to predictability inpollinator abundance.
We investigated the functional significance of raised black spots on the ray florets of Gorteria diffusa (Asteraceae) in South Africa. Field observations showed that G. diffusa is pollinated by a small bee-fly, Megapalpus nitidus (Bombyliidae), which is strikingly similar to the raised spots that occur on some of the ray florets. Removal of the spots resulted in a significant decrease in the rate of fly visits to capitula, but did not significantly affect seed set. Replacement of the spots with
simple ink spots also significantly reduced the rate of pollinator visits, suggesting that flies respond to details in the structure of the spots. Investigations using scanning electron microscopy showed that the spots of G. diffusa consist of a complex of different cell types. Differences in epidermal sculpturing may partly explain the UV reflectance pattern of these spots, which is similar to that of the flies. Male flies are strongly attracted to the spots, as well as to other flies sitting in the capitula, although female flies also visit the capitula. We conclude that the spots of G. diffusa mimic resting flies, thereby eliciting mate-seeking and aggregation responses in fly pollinators. Similar dark spots have evolved in unrelated South African Gazania, Dimorphotheca, and Pelargonium species pollinated by bee-flies.
The floral biology, reproductive system, and visitation behavior of pollinators of four species of columnar cacti, Stenocereus griseus, Pilosocereus moritzianus, Subpilocereus repandus, and Subpilocereus horrispinus, were studied in two arid zones in the north of Venezuela. Our results support the hypothesis that Venezuelan species of columnar cacti have evolved toward specialization on bat pollination. Additional information on the floral biology of a fifth species, Pilosocereus lanuginosus, wa
s also included. All species showed the typical traits that characterize the pollination syndrome of chiropterophily. All species but Pilosocereus moritzianus were obligate outcrossers. Nectar and pollen were restricted to nocturnal floral visitors. Two species of nectar-feeding bats, Leptonycteris curasoae Miller and Glossophaga longirostris Miller, were responsible for practically all the fruit set in these cacti. Frequency of bat visitation per flower per night was highly variable within and between species of cactus, with average frequencies varying between 27 and 78 visits • flower21 • night21. In general terms, the pattern of floral visitation through the night was significantly correlated with the pattern of nectar production and nectar sugar concentration for all species of cactus. Under natural pollination, fruit:flower ratios varied from 0.46 in Subpilocereus repandus to 0.76 in Stenocereus griseus. The efficiency of bat pollination in terms of seed:ovule ratio was high in all species, varying between 0.70 and 0.94.
Pollination in the genus Pedilanthus is commonly effected by hummingbirds. Pollination by vespid wasps in Pedilanthus is documented for the first time based on observations of Pachodynerus nasidens and Eumenes americanus pollinating Pedilanthus diazlunanus. An hypothesis concerning the mechanism by which hummingbird pollination was replaced by insect pollination in Pedilanthus is advanced based on observations of insects on P. bracteatus.
We studied the patterns of adaptive radiation in Disa, a large orchid genus in southern Africa. A cladogram for 27 species was constructed using 44 morphological characters. Pollination systems were then mapped onto the phylogeny in order to analyze pathways of floral evolution. Shifts from one pollination system to another have been a major feature of the evolutionary diversification of Disa. Unlike many plant genera that are pollinated mainly by a single group of insects, radiation in Disa has
encompassed nearly all major groups of pollinating insects; in all, 19 different specialized pollination systems have been found in the 27 species included in this analysis. Another striking pattern is the repeated evolution of broadly similar pollination systems in unrelated clades. For example, butterfly-pollinated flowers have evolved twice; showy deceptive flowers pollinated by carpenter bees, twice; long-spurred flowers pollinated by long-tongued flies, four times; night-scented flowers pollinated by moths, three times; and self-pollination, three times. This suggests that a few dominant pollinator species in a region may be sufficient to generate diversification in plants through repeated floral shifts that never retrace the same pathways.
The ability to produce seeds when pollinators or potential mates are scarce is thought to be one of the main advantages of self-fertilization in flowering plants. However, whether autonomous selfing increases seed set in natural populations has seldom been tested, and even fewer studies have evaluated the advantage of selfing across a gradient of pollen availability. This study examines the fertility consequences of autonomous selfing in Aquilegia canadensis (Ranunculaceae), a shortlived, spring
-flowering perennial typically found in small, patchy populations on rock outcrops. We used a pollinator exclusion experiment to confirm reports that A. canadensis has a well-developed capacity for autonomous selfing resulting from incomplete protogyny and close proximity of stigmas and anthers during dehiscence. Flowers excluded from pollinators set 87% as many seeds per carpel (X¯ 6 1 SE 5 7.1 6 1.4 seeds) as hand-pollinated flowers (8.1 6 1.3 seeds), and seed production in unpollinated flowers correlated negatively with the distance between stigmas and anthers (r 5 20.46). Autonomous selfing could be potentially valuable in providing reproductive assurance because only 2.7 6 0.5 pollen grains were deposited on each stigma before anther dehiscence, compared to 134.1 6 17.9 pollen grains by the end of anther dehiscence. However, prevention of autonomous selfing by anther removal before dehiscence did not decrease seed set, even for plants at low plant densities where outcross pollen may have been in short supply. Emasculated flowers set as many seeds per carpel (9.3 6 0.9) as intact flowers (8.4 6 1.1), indicating that sufficient cross pollen is deposited for full seed set. These results do not support the hypothesis that autonomous selfing by A. canadensis has been selected because it provides reproductive assurance.
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 d
eclines 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.
The Turonian flora from Sayreville New Jersey includes one of the world's most diverse assemblages of Cretaceous angiosperm flowers. This flora is made even more interesting by its association with a large insect fauna that is preserved by charcoalification as well as in amber. Floral diversity includes numerous representatives of Magnoliidae, Hamamelididae, Rosidae, Dilleniidae, and Asteridae (Ericales sensu lato). Included are hypogynous, five‐merous flowers with uniseriate hairs on the pedice
ls and stamens in bundles most frequently borne opposite the petals. There is considerable variation in filament length, and some filaments are branched. On some anthers, strands of residue, suggesting the former presence of a liquid of unknown nature, partially occlude the apparent zone of dehiscence. In other cases, open anthers are fully occluded by an amorphous substance. pollen is rarely found associated with anthers, but is common on stigmatic surfaces. pollen is prolate and tricolporate with reticulate micromorphology. The superior syncarpous ovary is five‐carpellate with axile/intruded parietal placentation and numerous anatropous ovules/carpel. Ovary partitions have closely spaced, parallel ascending channels (secretory canals?), and there are apparent secretory canals/cavities in receptacles, sepals, and petals. Individual stigmas are cuneiform with a central groove and eccentrically peltate. Styles are short and fused. In aggregate, the stigmas form a secondarily peltate stigma. Seeds have a reticulate sculpture pattern, a pronounced raphe, and funicular arils with sculpture similar to the seeds. phylogenetic analyses of several data matrices of extant taxa place this fossil in a monophyletic group with the modern genera Garcinia and Clusia within the Clusiaceae. As such, these fossils represent the earliest fossil evidence of the family Clusiaceae. Some modern Clusiaceae are notable, in particular, for their close relationship with meliponine and other highly derived bee pollinators; the fossil flowers share several characters that suggest a similar mode of pollination. This possibility is consistent with other floral and insect data from the same locality.
Abstract In their classic study, Alston and Turner (American Journal of Botany, vol. 50, 159-173, 1963) documented extensive hybridization among four morphologically distinct Baptisia species native to East Texas. While Alston and Turner found putative F1 hybrids in great numbers, they found no evidence of backcrossing. In this study prezygotic and postzygotic reproductive barriers between two of these species, B. leucophaea and B. sphaerocarpa, were investigated and found to be quite weak. Flow
ering times overlap and bumble bees were observed visiting both species and intermediate hybrids. While pollinator constancy in flights between B. leucophaea and B. sphaerocarpa was moderately strong, significant levels of constancy were not observed in flights involving hybrids and either parental species. Thus, backcrossing was not impeded by pollinator behavior. Further, hybrid pollen was highly stainable (93.5%) and able to effectively set seed in crossing experiments with both parental species. Pollinator behavior was compared in experimental populations with and without hybrid ramets and found to differ between these two treatments. Hybrids were found to facilitate pollinator movement between species. In total, these results suggest that reproductive isolation is not responsible for the rarity of backcrossing in naturally hybridizing B. leucophaea and B. sphaerocarpa populations.
Wind pollination was experimentally demonstrated in Linanthus parviflorus (Polemoniaceae), a predominantly beeflypollinated, self-incompatible annual. Seed set in plants enclosed in mesh tents that excluded pollinators but allowed airborne pollen flow provided evidence for wind pollination, and the extent of seed set due to wind pollination was compared to that in open-pollinated controls and pollen-supplemented treatments. Additional controls were included to test for possible confounding effec
ts of the mesh tent. Mean seed number in open-pollinated plants was 72.8–81.1% of that in pollensupplemented plants, while wind pollination alone produced 49.5–52.2%, a smaller but substantial proportion of seed set with pollen supplementation. Further evidence for wind pollination was found in a comparison of sites differing in the extent of wind exposure in two populations of L. parviflorus. Airborne pollen counts were higher in exposed sites than in protected sites, and the difference was marginally significant. Seed set was significantly pollen limited in protected sites, but not in exposed sites. Taken together, the data suggest that wind pollination provides some reproductive assurance in this obligately outcrossing species. Wind pollination is hypothesized to represent an alternative to selfing as an evolutionary solution to the problem of temporal or spatial variation in pollination visitation.
Pollination-related and time-dependent floral morphological changes occur in a diverse set of angiosperm taxa and appear to be particularly common in species occupying resource-limited environments. In deer weed (Lotus scoparius), such floral modifications include a color change from yellow to orange and a folding of the banner petal down over the keel. These changes are rapidly induced by pollination, but will also occur much more slowly without pollination. Orange flowers typically lack nectar
and pollen. We examined the reproductive success of these plants to test the hypothesis that retention of orange flowers increases pollinator visitation rate and fruit set while reducing costs to the pollinators. All of the common species of bee pollinators that visited deer weed easily distinguished between yellow and orange flowers at close range and preferentially probed yellow flowers. Retention of orange flowers by these plants resulted in a higher frequency of pollinator visits and a higher fruit set per flower than plants that lacked orange flowers. The number of flowers visited by each pollinator was lower on plants with a mixture of yellow and orange flowers, suggesting that the presence of orange flowers may reduce selfing. The possible selective pressures involved in the evolution of these mechanisms and their relation to stressful environments are also discussed.
Differences in pollen tube growth rates (certation) between heterospecific (foreign) and conspecific pollen may strongly influence whether hybrid offspring are produced after mixed pollen loads are delivered to a stigma. For both members of a sympatric species pair, Hibiscus moscheutos and H. laevis, pollination by pure loads of foreign pollen resulted in fruit set that was not significantly different from conspecific pollination, indicating that pure loads of foreign pollen could readily result
in hybrid offspring. However, the number of seeds per fruit from pure foreign pollinations was significantly less than that of pure conspecific pollination. Simultaneous mixed pollination resulted in a proportion of hybrid seeds (detected by an electrophoretic marker enzyme) that was significantly lower than expected based upon the capacity of foreign pollen to effect fertilization when applied in pure pollinations. After these 50/50% pollen mixtures were applied to stigmas, 8.0 and 7.4% hybrids were produced when H. moscheutos and H. laevis were the ovule parents, respectively. For these Hibiscus species, pollen competition appears to function as a barrier to hybridization that is of moderate intensity compared with similar barriers occurring between other recently studied sympatric species pairs.
Nearly forty years ago R. L. Berg proposed that plants with specialized pollination ecology evolve genetic and developmental systems that decouple floral morphology from phenotypic variation in vegetative traits. These species evolve separate floral and vegetative trait clusters, or as she termed them, “correlation pleiades.” The predictions of this hypothesis have been generally supported, but only a small sample of temperate‐zone herb and grass species has been tested. To further evaluate this
hypothesis, especially its applicability to plants of other growth forms, we examined the patterns of phenotypic variation and covariation of floral and vegetative traits in nine species of Neotropical plants. We recognized seven specific predictions of Berg's hypothesis. Our results supported some predictions but not others. Species with specialized pollination systems usually had floral traits decoupled (weak correlation; Canna and Eichornia) or buffered (relationship with shallow proportional slope; Calathea and Canna) from variation in vegetative traits. However, the same trend was also observed in three species with unspecialized pollination systems (Echinodorus, Muntingia, and Wedelia). One species with unspecialized pollination (Croton) and one wind‐pollinated species (Cyperus) showed no decoupling or buffering, as predicted. While species with specialized pollination usually showed lower coefficients of variation for floral traits than vegetative traits (as predicted), the same was also true of species with unspecialized or wind pollination (unlike our prediction). Species with specialized pollination showed less variation in floral traits than did species with unspecialized or wind pollination, as predicted. However, the same was true of the corresponding vegetative traits, which was unexpected. Also in contrast to our prediction, plants with specialized pollination systems did not exhibit tighter phenotypic integration of floral characters than did species with generalized pollination systems. We conclude that the patterns of morphological integration among floral traits and between floral and vegetative traits tend to be species specific, not easily predicted from pollination ecology, and generally more complicated than R. L. Berg envisaged.
Next Section Abstract Pollination biology, breeding system, and floral phenology of the columnar cactus Stenocereus stellatus were studied in wild, wild managed in situ and cultivated populations of central Mexico, in order to examine whether these aspects have been modified under domestication and whether they determine reproductive barriers between wild and manipulated individuals. Individuals of both wild and manipulated populations are self-incompatible, indicating that artificial selection
has not modified the breeding system. Their pollination biology is also similar. Anthesis is mainly nocturnal, with a peak of nectar production between 0200 and 0400 when the stigma presents maximum turgidity. Nocturnal visitors are the effective pollinators. Nearly 75% of flowers exposed for nocturnal pollination set fruit, while none of the flowers exposed for diurnal pollination produced fruits. The bats Leptonycteris curasoae, L. nivalis, and Choeronycteris mexicana (Glossophaginae) are the most likely pollinators, and their time of foraging is synchronized with the time of nectar production and stigma receptivity in S. stellatus. Bats potentially move pollen over a considerable distance, so there is apparently no spatial isolation to prevent pollen exchange between wild and cultivated populations. Phenological studies showed that there are also no apparent temporal barriers. However, manual cross pollination failed between some domesticated and wild phenotypes, suggesting that gene flow between wild and cultivated populations might be limited by pollen incompatibility.
The pollinators of 29 ginger species representing 11 genera in relation to certain floral morphological characteristics in a mixed-dipterocarp forest in Borneo were investigated. Among the 29 species studied, eight were pollinated by spiderhunters (Nectariniidae), 11 by medium-sized Amegilla bees (Anthophoridae), and ten by small halictid bees. These pollination guilds found in gingers in Sarawak are comparable to the pollination guilds of neotropical Zingiberales, i.e., hummingbird-, and euglos
sine-bee-pollinated guilds. Canonical discriminant analysis revealed that there were significant correlations between floral morphology and pollination guilds and suggests the importance of plant-pollinator interactions in the evolution of floral morphology. Most species in the three guilds were separated on the plot by the first and second canonical variables. Spiderhunter-pollinated flowers had longer floral tubes, while Amegilla-pollinated flowers had wider lips than the others, which function as a platform for the pollinators. Pistils and stamens of halictid-pollinated flowers were smaller than the others. The fact that gingers with diverse morphologies in a forest with high species diversity were grouped into only three pollination guilds and that the pollinators themselves showed low species diversity suggests that many species of rare understory plants have evolved without segregating pollinators in each pollination guild.
Plant populations vary in density both naturally and as a consequence of anthropogenic impacts. Density in turn can influence pollination by animals. For example, plants in dense populations might enjoy more frequent visitation if pollinators forage most efficiently in such populations. We explored effects of plant density on pollination and seed set in the larkspur Delphinium nuttallianum and monkshood Aconitum columbianum. At our site in the Colorado Rocky Mountains, flowers of D. nuttallianum
are pollinated primarily by queen bumble bees, solitary bees, and hummingbirds, whereas those of A. columbianum are pollinated primarily by queen and worker bumble bees. We found that the quantity of pollination service to both species (pollinator visitation rate and pollen deposition) was at best weakly related to density. In contrast, seed set declined by approximately one-third in sparse populations relative to nearby dense populations. This decline may stem from the receipt of low-quality pollen, for example, inbred pollen. Alternatively, sparsity may indicate poor environmental conditions that lower seed set for reasons unrelated to pollination. Our results demonstrate the value of simultaneously exploring pollinator behavior, pollen receipt, and seed set in attempting to understand how the population context influences plant reproductive success.
Pollination ecology of an emergent tree species, Shorea (section Mutica) parvifolia (Dipterocarpaceae), was studied using the canopy observation system in a lowland dipterocarp forest in Sarawak, Malaysia, during a general flowering period in 1996. Although the species has been reported to be pollinated by thrips in Peninsular Malaysia, our observations of flower visitors and pollination experiments indicated that beetles (Chrysomelidae and Curculionidae, Coleoptera) contributed to pollination o
f S. parvifolia in Sarawak. Beetles accounted for 74% of the flower visitors collected by net-sweeping, and 30% of the beetles carried pollen, while thrips accounted for 16% of the visitors, and 12% of the thrips carried pollen. The apical parts of the petals and pollen served as a reward for the beetles. Thrips stayed inside the flower almost continuously after arrival, and movements among flowers were rare. Fruit set was significantly increased by introduction of beetles to bagged flowers, but not by introduction of thrips. Hand-pollination experiments and comparison of fruit set in untreated, bagged, and open flowers suggested that S. parvifolia was mainly outbreeding.
Lactoris fernandeziana, monotypic in its family, is endemic to the cloud forests of Robinson Crusoe Island. Although there has been considerable study of the relationships of Lactoris, as a rare species and as a putative primitive paleoherb, little is known of its reproductive biology. Knowledge of the latter is essential for effective conservation programs. The species is gynomonoecious. The overall proportion of flowers is ∼1 female:1 hermaphrodite. The inconspicuous semipe
ndulous green flowers, usually in mixed-gender inflorescences, do not produce rewards. Hermaphrodite flowers are herkogamous and protogynous. Pollen grains are shed from the extrorse anthers in permanent dry tetrads. There is a mean of 12879 tetrads per hermaphrodite flower. Both flower types bear an average of ∼18 ovules. The P/O (pollen/ovule) ratios imply facultative or obligate xenogamy, but hand pollinations show that Lactoris is self-compatible. No floral visitors were ever observed, but stigmata of open-pollinated flowers bore tetrads, and 64% of such styles had pollen tubes. Flowers enclosed in large mesh (1 mm) bags bore similar numbers of tetrads and pollen tubes. Thus, we conclude that Lactoris is anemophilous, a syndrome perhaps reflected by the P/O ratio. Low genetic diversity (isozymes and DNA) supports selfing and implies limited distance wind pollen dispersal. The small size of the island, the ± 1000 extant Lactoris plants, coupled with anemophily, self-compatibility, and pendant flower position, have yielded a geitonogamous system with high seed set and low genetic diversity. If inbreeding depression is expressed, it is in seed germination and seedling vigor, for Lactoris is very difficult to cultivate. For this species, effective conservation practices need to focus on habitat preservation and promotion of outcrossing.
The bilobed stigma of many species in the order Scrophulariales closes in response to touch by an animal pollinator. In hummingbird-pollinated bush monkey flower, Mimulus aurantiacus (Scrophulariaceae), closure is rapid, occurring within seconds of tactile stimulus. We investigated the proximate causes of stigma closure and subsequent reopening in M. aurantiacus, as well as potential costs and benefits of stigma closure for female fitness. Stigma closure is elicited by both touch and pollen, but
closure in response to pollen is much slower, requiring 0.5–1.5 h. Stigmata reopen within 2.5–4.5 h if touch, but no pollen, is applied. Upon receipt of pollen, most stigmata remain closed for the remaining lifetime of the flower, even if less pollen is received than is needed for full seed set. Those stigmata that do reopen after pollination generally require between 20 and 28 h to do so, much longer than for unpollinated stigmata. Reopening after pollination appears to be a response to low seed set rather than to low pollen load. Natural pollination of stigmata manipulated to prevent closure shows that closure does not increase capture of pollen or seed set. In fact, closure reduces the average pollen load received by flowers. Despite this, there is no evidence that stigma closure has any negative effect on female fitness in terms of seed set or germinability. Hypotheses for the adaptive significance of stigma closure are discussed. Understanding proximate causes of stigma closure and reopening is essential in the evaluation of these hypotheses.
We examined the function of floral traits associated with buzz pollination through studies of Rhexia virginica (Melastomataceae) in the Muskoka region of Ontario, Canada. Controlled pollinations demonstrated that the species is self-compatible, but dependent on insects for pollen transfer. Bumble bees made 82 and 90% of observed insect visits to R. virginica in 1996 and 1997, respectively, and effectively buzzed flowers. Buzz pollination did not appear to be highly “specialized”
since various species of bumble bee were capable of pollination, and pollen transfer efficiency appeared to be relatively low. Experimental manipulations provided little support for the hypothesis that the yellow color of melastome anthers mimics abundant pollen, thereby deceiving pollinators to visit regardless of whether most pollen has been removed. Fruit set averaged 52.6% among populations, owing largely to infrequent pollinator visits and pollen limitation. Flowers of R. virginica were infertile after a single day of anthesis, but petals were subsequently maintained for 1–2 d and stamens underwent a color change from bright yellow to red. Second-day flowers may function to increase floral display size and hence fertility, without a concomitant increase in pollen discounting. Studies of bumble bee foraging behavior and correlates of seed set provided indirect support for this hypothesis.
We investigated patterns of fruit and seed production on inflorescences of a population of Pancratium maritimum in northwest Spain over a 2-yr period. Initial findings showed that the earliest opening flowers on an inflorescence are more likely to set fruit and produce more seeds than later opening flowers and that this pattern is maintained throughout the flowering season. Supplementary pollination and flower-removal experiments were performed to investigate whether the observed pattern is attr
ibutable (a) to variation in pollen receipt, (b) to sequestration of resources by the earliest flowers on an inflorescence, and/or (c) to "architectural" limitations on the fruit/seed production of later flowers. Supplementary pollination did not improve fruit or seed production by late flowers in either of the 2 yr of study. In flower-removal experiments, the remaining flowers showed improved fruit set and mean number of seeds per flower, by comparison with flowers in the same position on control inflorescences. When all flowers except the latest third were removed, these showed fruit set and seed production similar to those of early flowers on control inflorescences. These results strongly suggest that the observed within-inflorescence patterns of fruit and seed production in P. maritimum are mainly attributable to competition for resources (i.e., explanation b), though other adaptive explanations cannot be ruled out.
I studied the pollination biology of two closely related species of agave, Agave palmeri and A. chrysantha (Agavaceae), which exhibit several chiropterophilous (bat-pollinated) traits. Floral studies, floral visitor observations, and pollination studies were conducted over four summers at six different sites to examine floral traits and determine the relative importance of diurnal vs. nocturnal pollinators. Agave chrysantha appears to have developed minor shifts in several floral characters that
enhance diurnal pollination. Although floral shifts towards diurnal pollination were fewer in A. palmeri, stigmas were diurnally receptive and copious floral rewards were available in the morning, indicating that some adaptations exist to allow for multiple pollinators. Differences in fruit and seed set between naturally day- and night-pollinated umbels for both species were either not significant or significantly higher in day-pollinated plants. Bats were not important pollinators of A. chrysantha, and the mutualistic relationship between A. palmeri and the lesser long-nosed bat was found to be asymmetric. "Bat-adapted" floral traits appear to be flexible enough to respond to the climatic and pollinator unpredictability experienced by agaves at the northern edge of their distribution. This variability may be a more important factor affecting evolution of floral characters than a particular pollinator.
Next Section Abstract A previously undescribed pollination system involving a monoecious tree species, Artocarpus integer (Moraceae), pollinator gall midges, and fungi is reported from a mixed dipterocarp forest in Sarawak, Borneo. The fungus Choanephora sp. (Choanephoraceae, Mucorales, Zygomycetes) infects male inflorescences of A. integer, and gall midges (Contarinia spp., Cecidomyiinae, Diptera) feed on the fungal mycelia and oviposit on the inflorescence. Their larvae also feed on the myceli
a and pupate in the inflorescence. The gall midges are also attracted by female inflorescences lacking mycelia, probably due to a floral fragrance similar to that of male inflorescences. Because of the sticky pollen, dominance of Contarinia spp. in flower visitors, and pollen load observed on Contarinia spp. collected on both male and female inflorescences, Artocarpus integer is thought to be pollinated by the gall midges. Although several pathogenic fungi have been reported to have interactions with pollinators, this is the first report on a pollination mutualism in which a fungus plays an indispensable role. The pollination system described here suggests that we should be more aware of the roles fungi can play in pollinations.
The selective maintenance of gynodioecy depends on the relative fitness of the male-sterile (female) and hermaphroditic morphs. Females may compensate for their loss of male fitness by reallocating resources from male function (pollen production and pollinator attraction) to female function (seeds and fruits), thus increasing seed production. Females may also benefit from their inability to self-fertilize if selfing and inbreeding depression reduce seed quality in hermaphrodites. We investigated
how differences in floral resource allocation (flower size) between female and hermaphroditic plants affect two measures of female reproductive success, pollinator visitation and pollen receipt, in gynodioecious populations of Geranium richardsonii in Colorado. Using emasculation treatments in natural populations, we further examined whether selfing by autogamy and geitonogamy comprises a significant proportion of pollen receipt by hermaphrodites. Flowers of female plants are significantly smaller than those of hermaphrodites. The reduction in allocation to pollinator-attracting structures (petals) is correlated with a significant reduction in pollinator visitation to female flowers in artificial arrays. The reduction in attractiveness is further manifested in significantly less pollen being deposited on the stigmas of female flowers in natural populations. Autogamy is rare in these protandrous flowers, and geitonogamy accounts for most of the difference in pollen receipt between hermaphrodites and females. Female success at receiving pollen was negatively frequency dependent on the relative frequency of females in populations. Thus, two of the prerequisites for the maintenance of females in gynodioecious populations, differences in resource allocation between floral morphs and high selfing rates in hermaphrodites, occur in G. richardsonii.
Competition among pollen grains for the chance to fertilize ovules typically involves two stages: arrival times on stigmas and/or the growth of pollen tubes through styles. In a previous study of Hibiscus moscheutos, we found that individual pollen donors often differed in pollen tube competitive ability. Here we determined whether short delays in pollen arrival time altered the average success of “fast” and “slow” pollen donors when both types of pollen experienced the same delays. Hand-pollina
tion experiments were carried out using four pairs of pollen donors that differed in competitive ability. We allowed delays of 15 or 30 min between the first and second pollen donor and then determined seed paternity using allozyme markers. The second donor typically sired fewer seeds than pollen that arrived earlier, but, contrary to expectation, “faster” pollen did not always sire significantly more seeds than “slower” pollen when each was applied after delays of the same duration. In two of the four pairs of donors, differences that were seen following simultaneous pollinations disappeared when each type of pollen was applied following identical delays of 15 or 30 min. This unexpected response suggests that the dynamics of pollen tube competition are more complex than anticipated.
Pollination ecology of three Durio species, D. grandiflorus, D. oblongus, and D. kutejensis (Bombacaceae), was studied in a lowland dipterocarp forest in Sarawak, Malaysia, during a peak flowering period when at least 305 species of plants bloomed in 1996. Durio has been reported to be pollinated by bats in Peninsular Malaysia. However, my observations of flower visitors and pollination experiments indicated that two species, D. grandiflorus and D. oblongus, were pollinated by spiderhunters (Nec
tariniidae) and that the other species, D. kutejensis, was pollinated by giant honey bees and bats as well as birds. Hand-pollination experiments showed that all three species were obligate outbreeders. A resource limitation in fruit production was suggested. The former two species were visited only by spiderhunters, and the bagged flowers that were opened for animal visitors only at night bore no fruit, while those that were opened only during the day bore fruits, at comparable fruiting ratios to open pollination. Durio kutejensis was observed to be visited by giant honey bees, birds, and bats at different times of day, and three series of bagged experiments that exposed the flowers to animal visitors at different times of day bore fruits at a comparable ratio to open-pollination.
This paper examines the hypothesis that nectar robbing can affect plant reproductive success either positively or negatively. To this end, I investigated various aspects of the pollination ecology of a population of the herb Anthyllis vulneraria subsp. vulgaris in northwest Spain over 5 yr. By observing floral visitors, I found that the most important pollinator species was the long-tongued bee Anthophora acervorum, which accounted for ∼45% of recorded insect visits. However, just over 45% of vi
sits were by the nectar-robbing bumble bees Bombus terrestris and B. jonellus. Although the incidence of robbing differed considerably over 5 yr of study, the frequency in every season was very high (66.4–76.5% of robbing) except for 1997 (0% robbing). Despite this high frequency of robbing, robbed flowers had a higher probability of setting fruit than nonrobbed flowers in all years of the study (mean: 82.0 vs. 51.0%; excluding 1997). This increased fruit set in robbed flowers is directly related to bumble bee behavior because the robbers' bodies came into contact with both the anthers and stigmas while robbing. Thus, the robbers effect pollination. These results suggest that the effect of nectar robbers on plant reproductive success is dependent both on the robbers' behavior and on flower/inflorescence structure. The importance of nectar-robbing bumble bees on the reproductive success of A. vulneraria and its yearly high frequency suggest that the relationship between robbers and this plant is part of a successful long-term mutualism.
Nuphar comprises 13 species of aquatic perennials distributed in the temperate Northern Hemisphere. The European species N. lutea and N. pumila in Norway, the Netherlands, and Germany are pollinated by bees and flies, including apparent Nuphar specialists. This contrasts with reports of predominant beetle pollination in American N. advena and N. polysepala. We studied pollination in N. ozarkana in Missouri and N. advena in Texas to assess whether (1) there is evidence of pollinator shifts associ
ated with floral-morphological differences between Old World and New World species as hypothesized by Padgett, Les, and Crow (American Journal of Botany 86: 1316–1324. 1999) and (2) whether beetle pollination characterizes American species of Nuphar. Ninety-seven and 67% of flower visits in the two species were by sweat bees, especially Lasioglossum (Evylaeus) nelumbonis. Syrphid fly species visiting both species were Paragus sp., Chalcosyrphus metallicus, and Toxomerus geminatus. The long-horned leaf beetle Donacia piscatrix was common on leaves and stems of N. ozarkana but rarely visited flowers. Fifteen percent of visits to N. advena flowers were by D. piscatrix and D. texana. The beetles' role as pollinators was investigated experimentally by placing floating mesh cages that excluded flies and bees over N. advena buds about to open and adding beetles. Beetles visited 40% of the flowers in cages, and flowers that received visits had 69% seed set, likely due to beetle-mediated geitonogamy of 1st-d flowers. Experimentally outcrossed 1st-d flowers had 62% seed set, and open-pollinated flowers 76%; 2nd-d selfed or outcrossed flowers had low seed sets (9 and 12%, respectively). Flowers are strongly protogynous and do not self spontaneously. Flowers shielded from pollinators set no seeds. A comparison of pollinator spectra in the two Old World and three New World Nuphar species studied so far suggests that the relative contribution of flies, bees, and beetles to pollen transfer in any one population depends more on these insects' relative abundances (and in the case of Donacia, presence) and alternative food sources than on stamen length differences between Old World and New World pond-lilies.
Dichogamy is very common in flowering plants and is widely thought to reduce pollen‐pistil interference, especially self‐pollination. Yet, the functional significance of dichogamy has rarely been studied. We investigated the nature and functioning of dichogamy in eastern Ontario populations of Aquilegia canadensis, a highly selfing columbine previously described as protogynous. We then manipulated flowers to determine whether increased protogyny would reduce self‐fertilization. Contrary to previ
ous reports, A. canadensis is not dichogamous. Controlled pollinations in a greenhouse showed that pollen tubes generally begin to develop after anther dehiscence. Although stigmas can collect pollen early in floral development, naturally pollinated flowers collected from four populations had few pollen grains on stigmas and almost no pollen tubes in styles until after anther dehiscence. Limited pollen deposition before anther dehiscence was also associated with low nectar availability and limited sepal expansion. Because inbreeding depression is strong in this species, selection may favor increased protogyny if it reduces selfing. We tested this hypothesis by comparing the level of selfing in flowers rendered protogynous by the removal of the first 19 (of 39) anthers to develop, with nonprotogynous control flowers. Contrary to expectations, protogyny did not reduce selfing. Our results emphasize the importance of detailed field observations and manipulative experiments in understanding the nature and functional significance of dichogamy.
In a study of sexual reproduction in long-lived semelparous plants, we observed Agave macroacantha in the tropical desert of Tehuacán-Cuicatlán, Mexico, describing duration of flowering, flower phenology, and nectar production patterns. We also performed two manipulative experiments evaluating (a) the seed production efficiency of different crossing systems (selfing, cross-pollination, apomixis, and control), and (b) the effect of different pollinators (diurnal exposure to pollinators, nocturnal
exposure, exclusion, and control) on the seeds produced. Flowering occurred from early May to late July and had a mean duration of 29 days in the individual rosettes. The flowers were protandrous; anthesis occurred in the afternoon of the third day after floral opening, and the pistils matured in the afternoon of the fifth day. The stigmas remained receptive from dusk to the following morning. Pollination was mostly allogamous. Nectar was produced principally during the night, from the first stages of floral aperture until the stigmas wilted and flowering ceased. The flowers were visited during the day by hymenoptera, butterflies, and hummingbirds and during the night by bats and moths. Only the nocturnal visitors, however, were successful pollinators. Agave macroacantha is extremely dependent on nocturnal pollinators for its reproductive success.
We did a series of observational studies and manipulative experiments on the guild of nocturnal visitors of Agave macroacantha, including (1) a description of the hourly patterns of visits by moths and bats, (2) an evaluation of the relative contribution of bats and moths to flowering success, and (3) an evaluation of the pollination efficiency of the different bat species. Scapes exposed to moths but excluded to bats yielded ∼50% fewer fruits than those exposed to both pollinator groups. Flower
s exposed to the bat species Leptonycteris curasoae showed similar fruiting success to those exposed to Choeronycteris mexicana and to those exposed to the whole nocturnal visitor guild. However, the fruits originated from flowers pollinated by Leptonycteris curasoae yielded significantly more seed than those exposed to Choeronycteris mexicana or to the whole pollinator guild. It is concluded that Agave macroacantha is extremely dependent on nocturnal pollinators for its reproductive success and that bats are especially important for successful pollination. Some of these pollinators are migratory and have been reported to be steadily declining. A continuing decline in the populations of pollinators may impede the successful sexual reproduction of the plant host and may put the long‐term survival of this agave species under risk.
We studied the relative role of inflorescence traits, flowering synchrony, and pollination context for infructescence and fruit initiation in two Spanish populations of Arum italicum, a species in which inflorescences are the pollination unit. In this species, a specialized inflorescence organ, the appendix, is important for pollinator attraction. However, the short floral longevity and the production of mostly one inflorescence per plant make its pollination potentially dependent on strong flow
ering synchrony and on external factors not controlled by the plant (the pollination context). The flowering period in both sites lasted >3 mo. Day-to-day variation in simultaneous antheses was high, and 11–50% of antheses occurred on days during which no pollen donor was present. Inflorescence traits, flowering synchrony, and between-plant distance all influenced infructescence and fruit initiation, but their relative importance differed between sites. In one large population, infructescence initiation was positively related to inflorescence traits; in a smaller population infructescence initiation increased with the number of donor inflorescences. In both sites, percentage of fruits initiated per infructescence was dependent on a combination of inflorescence traits, flowering synchrony, and between-plant distance. Plants producing 2–4 inflorescences had higher probability of infructescence initiation and overlapped their antheses with more plants than single-inflorescence ones.
Dichogamy is one of the most widespread floral mechanisms in flowering plants and is thought to have evolved to reduce interference between pollen import and export within flowers, especially self‐pollination. Self‐pollination between flowers may also be reduced if dichogamy is synchronous among flowers on an inflorescence. The analysis of dichogamy at both levels requires that the sexual phases of individual flowers be defined functionally in terms of pollen deposition and removal. We conducted
morphological and functional analyses to investigate the degree of dichogamy within flowers and the synchronicity of dichogamy between flowers within inflorescences in an emergent, aquatic monocot, flowering rush (Butomus umbellatus). Based on daily observations of the development of marked flowers, data on the schedule of anther dehiscence within flowers, and repeat surveys of floral sex ratios in three populations, individual flowers appear to be strictly protandrous. On average, each flower spends ∼1 d in each of male and female phases with an intervening 1‐d neuter phase during which there is no available pollen in anthers and stigmas are not yet exposed to receive pollen. Morphological criteria used to delimit the beginning and end of each of these three sex phases were validated by quantifying the temporal schedule of pollen removal from anthers and pollen deposition on stigmas. Experimental pollinations showed that the morphological changes marking the end of female phase are hastened by pollen deposition. At the umbel level, synchronous development within sequential cohorts of flowers reduced overlap of male and female sexual phases between flowers. On average (±1 SE), 72 ± 3% of flowers completed their female phase while no other flowers on the same umbel were in male phase. Computer simulations of umbel development showed that this value is significantly higher than expected if the timing of flower development within umbels was random (30 ± 1%). Surveys of floral sex ratios in three populations revealed that 87% of umbels were either unisexual male or female at any given time. Pollinators usually visited more than one flower in sequence when foraging on umbels, suggesting that synchronous dichogamy may be an adaptation to avoid geitonogamy. The adaptiveness of both flower‐ and umbel‐level dichogamy is also suggested because both traits are expressed to a lesser extent in obligately clonal, triploid populations, where flowers do not make seeds and hence floral adaptations are not maintained by natural selection.
The function of stamen dimorphism in the breeding system of the alpine shrub Rhododendron ferrugineum was studied in two populations in the French Alps. This species has pentameric flowers with two whorls of stamens: an inner whorl of five long stamens and an outer whorl of short stamens. We studied the development of stamens from buds to mature flowers (measurement of the filament, anther, and style lengths at five successive phenological stages) and compared the size and position of reproducti
ve organs at maturity in control and partially emasculated flowers (removal of long-level stamens) to determine whether the presence of long-level stamens constitutes a constraint for the development of the short-level ones. Stamen dimorphism can be observed early in stamen development, from the bud stage of the year prior to flowering. At this early stage, meiosis had already occurred. Emasculation of the long-level stamens induced the short-level ones to grow longer than in normal conditions. We also performed seven pollination treatments on ten randomly chosen individuals in each population, and the number of seeds following each treatment was recorded. Results from these treatments showed that R. ferrugineum produced spontaneous selfed seeds in the absence of pollinators. However, no seed was produced when short-level stamens were emasculated and pollinators excluded, suggesting that long-level stamens are not responsible for selfing in the absence of pollinators and that reproductive assurance is promoted by short-level stamens.
Androdioecy is a rare sexual system in nature, as predicted theoretically. Among the androecious species reported so far, Castilla elastica (Moraceae) is unique in that flowers are unisexual and staminate and pistillate flowers on cosexual plants are produced on different inflorescences. In addition, inflorescence structure of staminate inflorescences on males and staminate and pistillate inflorescences on cosexes is markedly different. Staminate inflorescences on males are bivalvate, while stam
inate inflorescences on cosexes are "fig-like" and urceolate. Pistillate inflorescences are discoidal. The difference may reflect different roles and requirements of the three inflorescences in pollination and protection from herbivores. This study reports thrips pollination of C. elastica, demonstrated by a pollinator introduction experiment. Thrips pollination of the species may be an example of mutualism originating from plant-herbivore interactions. In the Moraceae, shifts from simple herbivores on flowers to pollinators might have occurred many times, evolving into diverse pollination systems including the fig-fig wasp mutualism. The family, of which little is known about pollination systems, provides interesting and unique opportunities to study evolution of pollination systems and roles of nonpollinating associates of inflorescences.
Next Section Abstract We quantified pollinator visit behavior, pollen receipt and export, and changes in allele and genotype frequencies from initial Hardy- Weinberg conditions in experimental arrays of two color morphs of snapdragons (Antirrhinum majus) visited by freely foraging bumble bees (Bombus appositus and B. flavifrons). The number of pollen grains received by a flower depended most on the number of pollinator visits to the flower, whereas the number of grains exported was best predicte
d by the total time pollinators spent inside the flower. The pattern of mating generally was assortative with respect to color, as bees tended to overvisit one color or the other within a foraging bout. In arrays where nectar was augmented in one color, the augmented color received both more visits and longer visits. Allele and genotype frequencies in offspring samples were in accord with qualitative expectations based on the pollinator observations, demonstrating that pollinators can directly influence the evolution of single-locus floral traits, at least under simplified experimental conditions.
We compared the extent of pollen limitation on female reproductive success of Salix lanata L., an entirely insect-pollinated willow, and S. lapponum L., which is 50 : 50% insect : wind pollinated (ambophilous). Supplemental hand-pollination significantly increased seed number per fruit by nearly 50% in the insect-pollinated willow, but had no significant impact on seed number in the dually pollinated species. Fruit set was not affected by the treatment in either of the species. These results dem
onstrate that pollen limitation on reproductive success is most pronounced in the species that depends entirely on insects for pollination. In general, pollinator visitation was highest to S. lapponum, but bumble bees were only observed on S. lanata, suggesting that the quantity and quality of pollinator visitation differed between the species. Our results empirically support the hypothesis that a dual pollination strategy is most effective in alpine environments with low and infrequent pollinator activity and high wind speeds.
The reproductive assurance hypothesis posits that selection favors self-pollination in flowering plants where mates and/or pollinators are scarce. A corollary is that self-pollinating populations are expected to be superior colonizers of mate- and pollinator-scarce environments. The California annual Clarkia xantiana includes outcrossing populations (ssp. xantiana) and autogamously self-pollinating populations (ssp. parviflora). Outcrossing is ancestral, and the subspecies have parapatric distri
butions with a narrow contact zone. We tested aspects of the reproductive assurance hypothesis by examining geographic and subspecies variation in the densities of mates and pollinators (native bees) and the density dependence of pollinator visitation and pollen receipt. Plant and flower densities, pollinator density, and pollinator visitation rates were lowest in the region of exclusively self-pollinating populations. Pollinator assemblages there lacked Clarkia-associated pollinator taxa that were common elsewhere. Self-pollinating populations in the contact zone generally had densities and visitation rates intermediate between allopatric self-pollinating populations and outcrossing populations. Visitation rate and pollen receipt increased significantly with plant density. These findings suggest that selection for reproductive assurance influenced the origin of self-pollination and/or that reproductive assurance influenced the geographic distribution of self-pollination. Geographic variation in pollinator assemblages may have generated variation in the value of reproductive assurance.
Autonomous self‐pollination may be considered as a mechanism enhancing plant reproductive success when plant access to pollen sources may limit seed production. We have studied the relationship between geographical patterns of variation in pollinator service to Helleborus foetidus and self‐pollination ability in three widely spaced regions in the Iberian Peninsula. As could be expected from its early flowering period, pollinator visitation rates to both plants and flowers of H. foetidus were ver
y low at all sites. Pollinator composition remained consistent among regions, but there was significant variation among regions in pollinator service. Despite the low visitation rates, fruit set did not appear to be pollen limited in any of the study areas, which may be explained by the long duration of flowers (up to 20 d). When pollinators were excluded experimentally, fruit set decreased significantly, but substantial levels of self‐pollination occurred at all regions. Autonomous self‐pollination levels were lowest in the two regions with lowest pollinator service and highest in the region with highest pollinator service. This disagreement between our results and the expectations derived from the reproductive assurance hypothesis may reflect a nonequilibrium situation of the northern H. foetidus populations in relation to their current pollinating environment.
Field studies in the semiarid Succulent Karoo region of South Africa showed that flowers of Massonia depressa (Hyacinthaceae) are visited at night by at least four rodent species, including two gerbil species. Live-trapped rodents were found to carry Massonia pollen on their snouts; they also had large quantities of Massonia pollen in their feces as a result of grooming their fur. Visits by insects to the flowers were infrequent at one site and apparently absent at another site. Plants enclosed
in large-mesh wire cages, which excluded rodents but not insects, set very few seeds relative to open controls. Our initial hypothesis of rodent-pollination in M. depressa was based on the striking similarity of its flowers to those of unrelated, rodent-pollinated Protea species. Convergent traits include dull-colored and very robust flowers situated at ground level, a strong yeasty odor, and secretion of copious amounts of sucrose-dominant nectar during the evening when rodents are active. Despite having a low sugar concentration (∼20%), the nectar of M. depressa is almost 400 times as viscous as an equivalent sugar solution. The jelly-like constituent in the nectar may discourage robbing by insects, while also facilitating lapping by rodents. Our findings illustrate the utility of floral syndromes for generating testable predictions about pollination systems.
Genetic improvement of willows through traditional breeding can be facilitated by pollen collection and storage so that female flower receptivity need not be synchronized with pollen shed for breeding. Two experiments were completed to test the effectiveness of various organic solvents for willow pollen collection. In the first experiment, seven pollen collection treatments and an untreated control were tested with two willow clones. The other experiment tested three treatments that showed promi
se in the initial experiment and an untreated control with eight willow clones. Toluene and carbon tetrachloride were effective for pollen extraction, with average pollen germination percentages that were >15%, but both chemicals reduced pollen viability by 10–20% compared with an untreated control based on in vitro germination tests. Pollen extracted with carbon tetrachloride or toluene was successfully used in controlled pollination, and >100 new families were produced with this technique. Pollen viability remained high after 18 mo of storage at −20°C. Based on our results, toluene is the preferred solvent for future willow pollen extractions because it is as effective as carbon tetrachloride, is not a known carcinogen, and is less expensive.
Enantiostyly is a form of directional asymmetry in plants in which the style is deflected away from the main axis of the flower, either to the left or right side. In Wachendorfia (Haemodoraceae), a small genus of insect-pollinated geophytes restricted to the Cape Province of South Africa, populations are usually polymorphic for asymmetry. Here we investigate dimorphic enantiostyly in the four species of Wachendorfia to determine whether variation in their reproductive systems influences the main
tenance of this genetic polymorphism. Experimental field pollinations of W. paniculata and W. thyrsiflora indicated higher fertility for cross- than for self-pollinations, whereas in W. brachyandra, these types of pollination produced similar levels of fertility. Outcrossing rates were highest in W. paniculata (t = 0.78–0.98), with W. brachyandra (t = 0.39–0.79) and W. thyrsiflora (t = 0.76) exhibiting mixed mating systems. Outcrossing rates in two populations of W. parviflora varied from mixed mating (t = 0.61) to predominant selfing (t = 0.07). Population style-morph ratios ranged from 1 : 1 in outcrossing W. paniculata to monomorphism in selfing W. parviflora and clonal W. thyrsiflora. In W. brachyandra, a species with delayed selfing, morph ratios were usually biased. The maintenance of enantiostyly in Wachendorfia appears to be strongly influenced by levels of disassortative mating and the balance between sexual and clonal recruitment.
Flowers that are open for >12 h may be visited by both diurnal and nocturnal pollinators. I compared the effectiveness (measured as seed production and pollen movement distance) of diurnal and nocturnal pollinators of Silene alba, a species whose flowers open in evening but close by midmorning the following day. By bagging flowers either during evening hours or during daylight hours or both day and night, I compared seed production caused by diurnal and nocturnal pollinators. Flowers exposed onl
y to nocturnal visitors (mostly sphingid and noctuid moths) produced significantly more seeds than flowers exposed only to diurnal visitors (bees, flies, and wasps). Fluorescent dye applied to anthers moved significantly further and to more stigmas at night than during the day. In both measures of pollination effectiveness, nocturnal‐visiting moths are better pollinators of S. alba than are the diurnal‐visiting bees, flies, and wasps. These data support the hypothesis that floral phenology is an adaptation to expose flowers to the most effective pollinators.
The floral longevity of unpollinated, hand self-, and hand cross-pollinated flowers was compared in two varieties of Impatiens hypophylla, which contrasts with their mating systems. When flowers were emasculated and hand-pollinated every day after anthesis, their longevity was reduced. In the absence of emasculation and hand pollination, the staminate phase of the flowers of both varieties was 1 d longer. After the staminate phase, flowers of the outcrossing variety dropped their androecium, exp
osing the stigma and initiating the pistillate phase, which lasted for ∼2 d. In contrast, flowers of the mixed-mating variety self-pollinated autonomously and then terminated their flowering. Under great seasonal variation in the pollinator visitation rate, which was observed in their natural populations, the outcrossing variety should maximize expected outcross success through the phenology of floral sex phases, whereas the mixed-mating variety self-pollinated ovules that were not outcrossed within the staminate phase. Based on these results, I suggest that the autonomous self-pollination in I. hypophylla induced differences both in the selfing coefficient and in floral longevity between the varieties.
Discussion about thrips (Thysanoptera) as main pollinators has been controversial in the past because thrips do not fit the preconception of an effective pollinator. In this study, we present evidence for thrips pollination in the dioecious pioneer tree genus Macaranga (Euphorbiaceae). Macaranga hullettii is pollinated predominantly by one thrips species, Neoheegeria sp. (Phlaeothripidae, Thysanoptera). As a reward for pollinators, the protective floral bracteoles function as breeding sites for
thrips and trichomal nectaries on the adaxial surface of the floral bracteoles provide alimentation. Flowering phenology of both staminate and pistillate trees was highly synchronized within 3-4 wk periods. In contrast to pistillate trees, staminate trees start to breed the thrips inside the developing inflorescences ∼2 wk before anthesis. Breeding of Neoheegeria sp. in the laboratory indicates that the thrips development is completed within ∼17 d. Thus, staminate trees offer breeding sites for one thrips generation until the onset of pollen presentation. Intraspecific pollen transfer by thrips was proved by pollen loads of thrips taken from receptive pistillate inflorescences of M. hullettii. Bagging experiments of different mesh sizes showed that seed set reached almost the level of open-pollinated flowers when exclusively tiny insects like thrips were able to enter the net bags, but no apomictic seed set occurred when no insect access was given to the flowers.
The dispersion and capture of differently shaped particles within a Zostera marina L. (eelgrass; Zosteraceae) bed were examined to understand submarine pollination and other dispersals. During periods of moderate flow in the canopy, the capture rate of “spherical” (the shape of ancestral pollen) and “filamentous” (the shape of eelgrass pollen) particles was greater for particles released at the top of the canopy (3.07 and 4.53% × 10−5 cm−2 of collector; i.e., percentage of particles captured nor
malized to collector area) and greater for filamentous than for spherical particles (4.51% × 10−5 cm−2 vs. 2.01% × 10−5 cm−2). Estimates of the horizontal P (Joseph‐Sendner diffusion velocity) and the vertical diffusivity (Gaussian K) of filamentous particles were small (P ≈ 4 × 10−4 m/s; K ≈ 10−4 m2/s) compared to theoretical values that do not consider plant canopies. These findings support the concept that eelgrass canopies modify the fluid dynamics (i.e., reduced turbulent mixing) within their canopies. These results indicate that 1000–10 000 Z. marina pollen are required to pollinate a single flower. Similarly, it was estimated that under some conditions, the probability of particle impaction on eelgrass vegetation approaches certainty. These results provide insight into the evolution of filamentous pollen and submarine pollination, as well as dispersal and other mass transport phenomena within macrophyte canopies.
In small, fragmented populations of self‐incompatible plant species, genetic drift and increasingly close relationships between plants may restrict the number of genetically different pollen donors, the availability of compatible mates, and the opportunity for pollen competition and selection. These restrictions may reduce the siring success or increase the probability of inbreeding depression in the offspring. To test if this was the case, we hand‐pollinated maternal plants in small and large p
opulations of the rare, endemic plant Cochlearia bavarica (Brassicaceae) with pollen from one, three, or nine donors from the same population or with nine donors from a different population. In one additional population of intermediate size, maternal plants were hand‐pollinated with ten donors located at a distance of 1, 10, 100, or 1000 m. We then recorded seed and offspring characters. On average, offspring from small populations were smaller than normal and fewer survived to maturity. Increasing the number of pollen donors had a positive effect on reproductive success in small and large populations, but at the highest pollen diversity this occurred at the expense of slightly reduced offspring fitness. Because the total amount of transferred pollen was held constant, these effects could not be attributed to increasing pollen load. Rather, the increasing pollen diversity may have increased the chances of selecting a particularly “good” donor for fertilization—an example of a sampling effect of diversity. Pollen from outside a population or from 10–100 m away resulted in higher reproductive success and greater offspring size. Effects of population size and pollination treatments on reproductive success and offspring fitness were additive. Apparently, there is no obvious size threshold above which the potential of inbreeding depression can be ignored in C. bavarica.
Recent ecological research has revealed that the Sonoran Desert columnar cactus Lophocereus and the pyralid moth Upiga virescens form an obligate pollination mutualism, a rare but important case of coevolution. To investigate the phylogenetic origins of this unusual pollination system, we used molecular sequence data to reconstruct the phylogeny of the four taxa within the genus Lophocereus and to determine the phylogenetic position of Lophocereus within the North American columnar cacti (tribe
Pachycereeae). Our analysis included Lophocereus, six Pachycereus species, Carnegiea gigantea, and Neobuxbaumia tetetzo within the subtribe Pachycereinae, and Stenocereus thurberi as an outgroup within the Stenocereinae. Extensive screening of chloroplast and mitochondrial genomes failed to reveal sequence variation within Lophocereus. At a deeper phylogenetic level, however, we found strong support for the placement of Lophocereus within Pachycereus as sister group to the hummingbird‐pollinated P. marginatus. We discuss possible hypotheses that may explain the transition from bat pollination (ancestral) to moth and hummingbird pollination in Lophocereus and P. marginatus, respectively. Additional phylogenetic analyses suggest that the genus Pachycereus should be expanded to include Lophocereus, Carnegiea, Neobuxbaumia, and perhaps other species, whereas P. hollianus may need to be excluded from this clade. Future study will be needed to test taxonomic distinctions within Lophocereus, to test for parallel cladogenesis between phylogroups within Lophocereus and Upiga, and to fully delineate the genus Pachycereus and relationships among genera in the Pachycereinae.
While a considerable amount of attention has been devoted to the effects that increased ultraviolet-B (UV-B) radiation has on vegetative plant growth and physiological function, the impact that UV-B may have on plant fitness has been the focus of fewer studies, with attention given primarily to a few crop species. Further, the possible interactions between UV-B and additional potential stresses found in natural environments have rarely been studied experimentally. Because the reported effects of
increased UV-B on plant growth and fitness have been highly variable, studies that focus on factors that may lead to these differences in results are important for the formulation of accurate predictions about future plant success under varying UV-B levels. We examined the effects of UV-B dose and intraspecific competition on growth, phenology, pollen production, pollination success, fruit and seed production, and offspring quality in two species of Phacelia. Increased UV-B was neutral or beneficial for all traits, while competition was neutral or detrimental. There were no significant interactions between UV-B and competition in the parental generation. Phacelia campanularia offspring were unaffected by parental competition, but derived indirect beneficial effects on germination, growth, and fitness traits from parental enhanced UV-B.
We investigated the floral and pollination biology of two monoecious root holoparasites, Balanophora kuroiwai and B. tobiracola (Balanophoraceae), in the subtropical forests of southern Japan. Both species secrete nectar from extrafloral nectaries distributed among the flowers, which is mainly consumed by ants, cockroaches, and pyralid moths. Pollen grains were found attached to the bodies of these insects. Pyralid moths of the genera Assara and Nacoleia were observed laying eggs on the inflores
cences of B. kuroiwai. In both Balanophora species, pyralid larvae were found feeding on vegetative tissue without exploiting the seeds, and adults emerged from the fruited infructescences. In B. kuroiwai, we assessed pollination success under different experimental conditions by estimating the percentage of styles that had pollen tubes reaching the ovules. This revealed that: (1) the plants were at least sporophytically self‐compatible; (2) they were generally pollinated within an inflorescence (geitonogamy); (3) outcrossing occurred, but the rates varied greatly among inflorescences; and (4) ants were probably responsible for the geitonogamy. While ants and flightless cockroaches were the most likely contributors to geitonogamous self‐pollination, we consider pyralid moths to be the most likely cross‐pollinators of Balanophora species. This is a new example of pollination mutualism involving a plant and its pollinating parasite.
Species of Collinsia and Tonella, the two sister genera of self‐compatible annuals that constitute tribe Collinsieae, show extensive variation in floral size and morphology and in patterns of stamen and style elongation during the life of the flower (anthesis). We used a nuclear ribosomal ITS phylogeny, independent contrasts, and phylogenetically corrected path analysis to explore the patterns of covariance of the developmental and morphological traits potentially influencing mating system. Larg
e‐flowered taxa maintain herkogamy (spatial separation of anthers and stigmas) early in anthesis by differential elongation of staminal filaments, which positions each of the four anthers at the tip of the “keel” upon dehiscence. Small‐flowered taxa do not show this pattern of filament elongation. The styles of large‐flowered taxa elongate late in the 2–5 d of anthesis, resulting in late anther‐stigma contact and delayed self‐pollination. Anther‐stigma contact and self‐pollination occur early in anthesis in small‐flowered species/populations. Thus, we found complex covariation of morphological and developmental traits that can be interpreted as the result of multitrait adaptation for early selfing and high levels of autogamy, delayed selfing and higher levels of outcrossing, or intermediate levels of outcrossing. Continuous variation in these traits suggests the operation of continuous variation in selective optima or the combined effects of divergent selection and phylogenetic inertia.
Flower stalks of Pulsatilla cernua, an early spring herb in north temperate Asia, changed position from erect to pendulous and back to erect during 6–10 d anthesis. We tested three possible explanations for this movement. Our results showed that (1) this movement is unlikely to be a mechanism to attract pollinators or enhance pollen output, because no pollinator preference was observed between erect and pendulous flowers and we found no buzz-pollination in this species; (2) hand self-pollination
yielded higher seed set than open pollination in the field, but spontaneous selfing rarely occurred. Among open-pollinated flowers, seed set was depressed by emasculation, indicating that in the presence of insects, self-pollen provided reproductive assurance in this protogynous and self-compatible species. However, the change in flower orientation cannot be explained as reproductive assurance in that even self-pollination largely depended on pollinator visits rather than gravity. (3) A pollen germination experiment indicated that pollen damage by water is serious in this species. We deduced that the bending of the flower stalk during anthesis was to avoid rain damage to pollen grains in this species. During the 3–6 d period of pollen presentation, the petals elongated and were covered with unwettable hairs. Together with flower stalk movement, this was enough to protect the organs inside the flower from rain. This movement of the flower stalk seems to be important to maintain pollen viability in a rainy habitat with a scarcity of pollinators.
Floral longevity is assumed to reflect a balance between the benefit of increased pollination success and the cost of flower maintenance. Flowers of Kalmia latifolia (Ericaceae), mountain laurel, have a long duration and can remain viable up to 21 d if unpollinated. I experimentally tested whether this long duration increases pollination success by clipping stigmas to reduce functional floral longevity to 3-4 d. Clipping stigmas decreased fruit set from 65% to only 10%. Flowers with natural life
spans were not pollination-limited, demonstrating that long floral duration ensured female reproductive success. The long floral duration of K. latifolia was unique in this site (the Great Swamp, Rhode Island, USA). Coflowering shrubs in summer had a mean floral life span of 3.4 d. Spring-flowering species had significantly longer mean floral durations (7.2 d). These duration differences may reflect seasonal variation in pollinator availability. However, K. latifolia flowers in summer, when its bumble bee pollinators are abundant but it is a poor competitor for bees because its flowers produce little nectar. The long floral duration allows K. latifolia to outlast coflowering competitors and attract sufficient pollinators. I hypothesize that the long floral duration of K. latifolia functions as a mechanism for competitive avoidance and reproductive assurance.
The majority of flowering plants require animals for pollination, a critical ecosystem service in natural and agricultural systems. However, quantifying useful estimates of pollinator visitation rates can be nearly impossible when pollinator visitation is infrequent. We examined the utility of an indirect measure of pollinator visitation, namely pollen receipt by flowers, using the hummingbird-pollinated plant, Ipomopsis aggregata (Polemoniaceae). Our a priori hypothesis was that increased polli
nator visitation should result in increased pollen receipt by stigmas. However, the relationship between pollinator visitation rate and pollen receipt may be misleading if pollen receipt is a function of both the number of pollinator visits and variation in pollinator efficiency at depositing pollen, especially in the context of variable floral morphology. Therefore, we measured floral and plant characters known to be important to pollinator visitation and/or pollen receipt in I. aggregata (corolla length and width and plant height) and used path analysis to dissect and compare the effect of pollinator visitation rate vs. pollinator efficiency on pollen receipt. Of the characters we measured, pollinator visitation rate (number of times plants were visited multiplied by the mean percentage of flowers probed per visit) had the strongest direct positive effect on pollen receipt, explaining 36% of the variation in pollen receipt. Plant height had a direct positive effect on pollinator visitation rate and an indirect positive effect on pollen receipt. Despite the supposition that floral characters would directly affect pollen receipt as a result of changes in pollinator efficiency, corolla length and width only weakly affected pollen receipt. These results suggest a direct positive link between pollinator visitation rate and pollen receipt across naturally varying floral morphology in I. aggregata. Understanding the relationship between pollinator visitation rate and pollen receipt may be of critical importance in systems where pollinator visitation is difficult to quantify.
Reproductive assurance is frequently used to explain the evolution of selfing but has become controversial from lack of evidence. We studied the pollination system of the near carnivorous plant genus Roridula and showed that reproductive assurance is important in this system. Hemipterans have a digestive mutualism with Roridula and have been implicated in pollination but flowers show adaptations to hymenopteran pollination. We deduce that hemipterans are the primary pollinators of Roridula becau
se seed set is significantly reduced when hemipterans are excluded from the flowers. Using allozyme electrophoresis, we show that hemipterans are responsible for mostly selfed progeny. Although bees still pollinate Roridula on very rare occasions, their exclusion does not affect seed set. The complicated floral structures that occur in Roridula most likely evolved as adaptations for bee pollination. Resident hemipterans facilitate selfing by Roridula, and this acts as a reproductive assurance mechanism because it increases seed production and ensures that plants still reproduce in the absence of more motile, outcrossing pollinators.
Paniculate agaves from tropical deserts depend on nectar‐feeding bats for their reproduction, while species from extratropical areas near the northern limit of Leptonycteris curasoae are pollinated by diurnal and nocturnal insects and birds. Agave angustifolia is a paniculate agave with a broad distribution in Mexico whose range coincides with the distribution of L. curasoae, while A. subsimplex has a narrow distribution in Sonora within the range of nectar‐feeding bats. We studied the pollinati
on biology of A. angustifolia and A. subsimplex in northwestern Mexico to evaluate the relative importance of bat pollination in a region where L. curasoae is seasonal. Flower visitors included a wide variety of animals, with bats having greater visitation rates in A. angustifolia. A pollinator‐exclusion experiment revealed that bats were responsible for most of the control fruit set in A. angustifolia, whereas for A. subsimplex, diurnal and nocturnal visitors were equally effective. Overall, our data indicate that in central Sonora, A. angustifolia depends on nectar‐feeding bats for its sexual reproductive success, while A. subsimplex relies on both diurnal and nocturnal pollinators. Given the contrasting distribution ranges of paniculate agaves, our results seem to support the trend from specialization to generalization along the continuum of tropical to extratropical deserts.
We evaluated the role of floral scents in the reproductive success of Alocasia odora C. Koch (Araceae). Alocasia odora is pollinated by its specific pollinators, Colocasiomyia alocasiae (Okada) and C. xenalocasiae (Okada) (Diptera: Drosophilidae). These flies use the spadix of A. odora as breeding sites. The appendix, which is at an upper part of the spadix and is the most attractive region, attracted these pollinators by emitting volatiles, although the male zone of the inflorescence was also a
ttractive. The number of flies attracted was positively correlated with appendix size. During the pistillate phase of the protogynous spadix, attracted flies aggregated in the lower part (female zone) to mate, lay eggs, and perhaps obtain nutrients. The flies moved to the upper part (male zone) of the spadix by the tightening of the constriction separating the upper and lower parts, and then the staminate phase started. This movement of the flies on the spadix promotes outcrossing of A. odora. Removal of the appendix or the whole upper part of the spadix resulted in much reduced fruit set, suggesting that the absence of the scent‐producing region leads to insufficient pollination because of reduced pollinator attraction.
The role of geitonogamy in the evolution of inflorescence design is not well understood. The plant's dilemma hypothesis proposes that evolution of larger inflorescences is driven by selection for greater pollinator attraction, but constrained by higher rates of geitonogamy experienced by larger inflorescences. Here we investigate the role of geitonogamy on fruit set in natural populations of Asclepias speciosa. We compared fruit set from three pollination treatments: (1) inflorescences bagged be
fore and after receiving 6 hand outcross pollinia (Bag), (2) inflorescences unbagged and receiving 6 hand outcross pollinia (Open), and (3) naturally pollinated inflorescences (Control). The Bag and Open treatments initiated significantly more fruits than the Control. Bag aborted significantly fewer fruits than Open or Control. Fruit set was significantly higher in Bag than Open, and Open had significantly higher fruit set than Control. From these results, we conclude that (1) high rates of geitonogamy significantly increase fruit abortion and reduce fruit set in natural populations of A. speciosa and (2) natural populations are compatible pollen limited. Both findings are consistent with the plant's dilemma hypothesis.
We examined the breeding system and pollination of Chamaecrista keyensis Pennell (Fabaceae: Caesalpinioideae) and the effects of urban edge and mosquito control on reproduction of this rare endemic herb of the Lower Florida Keys. Controlled hand‐pollination treatments were applied to plants in the field. Although C. keyensis flowers are self‐compatible, they are not capable of automatic selfing. Inbreeding depression was observed in both seed set and percentage seed germination. Bees of seven ge
nera were observed visiting C. keyensis flowers during the peak flowering season (June to July). Only Xylocopa micans and Melissodes spp. may be effective pollinators for C. keyensis, as they were the only bees that “buzz pollinate” this species, which has poricidal anther dehiscence. Chamaecrista keyensis received substantially more visits by X. micans, but fewer visits from Melissodes spp. in urban‐edge vs. forest sites. Aerial mosquito spraying may exacerbate the existing pollinator limitation suffered by C. keyensis by reducing the number of visits by the buzz‐pollinating bees. Individuals of C. keyensis at urban edges produced fewer seeds per fruit than did individuals in a pristine forest mainly because of greater insect seed predation.
The contribution of a pollinator toward plant fitness (i.e., its “effectiveness”) can determine its importance for the plant's evolutionary ecology. We compared pollinators in a population of Asclepias incarnata (Apocynaceae) for several components of pollinator effectiveness over two flowering seasons to evaluate their importance to plant reproduction. Insects of the order Hymenoptera predominate in A. incarnata pollination, but there appears to be no specialization for pollination within this
order. Pollinators varied significantly in nearly every component of effectiveness that we measured, including pollen load, removal and deposition of pollen, pollination efficiency (deposition/removal), flower‐handling time, and potential for geitonogamy (fractional pollen deposition). The visitation rate of pollinators also varied significantly between years and through time within years. Pollination success and percentage fruit‐set of unmanipulated plants in the population also varied significantly between years, and pollination success varied among sample times within years. Most components of effectiveness were weakly correlated, suggesting that the contributions of visitor species toward pollination varied among effectiveness components. Mean flower‐handling time, however, was strongly correlated with several components, including pollen removal and deposition, pollination efficiency, and fractional pollen deposition. These findings highlight the significance of pollination variability for plant reproduction and suggest that time‐dependent foraging behaviors may play an important role in determining pollinator effectiveness.
Xerophyllum tenax is a mass‐flowering, nectarless herb in which self‐pollination is unavoidable as anthers shed pollen onto the three, receptive stigmatic ridges attached to each pistil within a few hours after expansion of the perianth. We compared the pollination system with reproductive success in this species through controlled, hand‐pollination experiments. Ovaries of flowers sampled from unbagged inflorescences were visited by pollen‐eating flies (primarily members of the family Syrphidae)
, beetles (primarily Cosmosalia and Epicauta spp.), and small bees, and produced normal‐sized capsules and mature seeds. Ovaries of flowers from inflorescences bagged to prevent insect pollination produced small capsules containing undeveloped or no seeds. Epifluorescence analyses suggest that 0.95 of the uncovered flowers are cross‐pollinated by insects with pollen tubes penetrating style and ovary tissue. Flowers show a “leaky” but early‐acting self‐incompatibility system. While hundreds of pollen tubes germinate on each stigmatic surface following self‐pollination, few pollen tubes penetrate the stigmatic surface and none penetrate the ovary. In contrast, when stigmas are cross‐pollinated by hand with pollen from a second inflorescence pollen tubes were seen penetrating style and ovary. Self‐incompatibility in X. tenax parallels that of some species of Trillium, a sister genus within the Melanthiaceae.
This paper reports obligate seed-parasitic pollination mutualisms in Breynia vitis-idea and B. fruticosa (Phyllanthaceae). The genus Breynia is closely related to Glochidion and Gomphidium (a subgenus of Phyllanthus), in which pollination by species-specific, seed-parasitic Epicephala moths (Gracillariidae) have been previously reported. At night, female Epicephala moths carrying numerous pollen grains on their proboscises visited female flowers of B. vitis-idea, actively pollinated flowers, and
each subsequently laid an egg. Examination of field-collected flowers indicated that pollinated flowers of B. vitis-idea and B. fruticosa almost invariably had Epicephala eggs, suggesting that these moths are the primary pollinators of the two species. Single Epicephala larvae consumed a fraction of seeds within developing fruit in B. vitis-idea and all seeds in B. fruticosa. However, some of the fruits were left untouched, and many of these had indication of moth oviposition, suggesting that egg/larval mortality of Epicephala moths is an important factor assuring seed set in these plants. The overall similarity of the specialized floral structure among Breynia species may indicate that this pollination system is fairly widespread within the genus.
We investigated the origin of stylar polymorphisms in Narcissus, which possesses a remarkable range of stylar conditions and diverse types of floral morphology and pollination biology. Reconstruction of evolutionary change was complicated by incomplete resolution of trees inferred from two rapidly evolving chloroplast regions, but we bracketed reconstructions expected on the fully resolved plastid-based tree by considering all possible resolutions of polytomies on the shortest trees.
Stigma-height dimorphism likely arose on several occasions in Narcissus and persisted across multiple speciation events. As proposed in published models, this rare type of stylar polymorphism is ancestral to distyly. While there is no evidence in Narcissus that dimorphism preceded tristyly, a rapid transition between them may explain the lack of a phylogenetic footprint for this evolutionary sequence. The single instances of distyly and tristyly in Narcissus albimarginatus and N. triandrus, respectively, are clearly not homologous, an evolutionary convergence unique to Amaryllidaceae. Floral morphology was likely an important trigger for the evolution of stylar polymorphisms: Concentrated-changes tests indicate that a long, narrow floral tube may have been associated with the emergence of stigma-height dimorphism and that this type of tube, in combination with a deep corona, likely promoted, or at least was associated with, the parallel origins of heterostyly.
The pollination biology of the cactus Pachycereus pecten-aboriginum was studied in a tropical location in western Mexico (ca. 18° N latitude) to compare with data from a northern population (ca. 28° N latitude). Throughout this range, the nectar-feeding bat Leptonycteris curasoae is resident within the tropics but migratory in its northern range. The hypothesis was tested that if a predictable bat presence has been an important force in the evolution of pollination systems in columnar cacti, P.
pecten-aboriginum will have a specialized pollination system within the tropics and a generalized pollination system in northern populations. In both areas, pollination experiments showed that P. pecten-aboriginum has a self-incompatible, hermaphroditic breeding system. In the tropical area, flowers open at night and close early in the morning. Nectar is secreted only during the night, and flowers are exclusively pollinated by three species of nectar-feeding bats, indicating a specialized pollination system. In contrast, anthesis and nectar secretion in northern populations occur during the night and day, allowing visitation and effective pollination by both nocturnal and diurnal pollinators. This study provides evidence of divergence mediated by pollinator predictability (resident vs. migrant), through shifts from short to long anthesis and nectar production periods from southern to northern populations.
Traits associated with self-pollination are common in island plants. This pattern could simply reflect the vestige of selection during colonization. Alternatively (or in addition), the ability to self-pollinate may provide a reproductive assurance benefit in established island plant populations due to inferior island pollinator service. To test these alternatives I studied an introduced plant (Nicotiana glauca; Solanaceae) on the California mainland and on two Channel Islands colonized at differ
ent times (approximately 30 and 100 yr ago). I compared these populations in terms of (1) capacity for self-pollination (self-compatibility, autogamy, stigma-anther distance, and incidence of a crumpled floral morph) and (2) current selection for the ability to self-pollinate (pollinator service by hummingbirds and the effect of emasculation on reproductive success). In general, island plants exhibited a higher capacity for self-pollination than mainland plants, especially on the most recently colonized island. However, island plants were not visited less frequently or more variably, nor did I detect current selection for selfing on islands. This supports the hypothesis that selfing traits in island plants are the product of a filter to successful establishment during colonization and not of selection for selfing in established island populations.
Flowers of columnar cacti are animal-pollinated, often displaying a chiropterophylic syndrome. This study examined if the columnar cactus Stenocereus queretaroensis, a tropical species endemic to western Mexico, is bat-pollinated, by studying its pollination biology and the foraging behavior of potential pollinators. Flowers were produced in winter through spring, peaking in April. Anthesis was nocturnal, and stigma and anther turgidity began around 2200 hours. Production of nectar secretion and
highest sugar concentration and energy supply were nocturnal, peaking between 2200 and 2400 hours. Manual auto-pollination and exclusion experiments showed that self-pollination yielded no fruits, while nocturnal pollinators resulted in high fruit set and seed set compared to diurnal pollination treatments. The nectar-feeding bat Leptonycteris curasoae (Phyllostomidae) was the main nocturnal pollinator with the highest effective pollination. Peak bat visitation coincided with peaks in nectar production. The high abundance of L. curasoae throughout the 4-yr study, suggests that it is a seasonally reliable pollinator for this columnar cactus. While pollination syndromes have been increasingly called into question in recent years, this study suggests that at least for this system, there is a fairly close fit between pollinator and pollination syndrome.
Phylogenetic studies are often hampered by the independent evolution of characters that may potentially obscure relationships. The adaptive significance of the nectar spur and its evolution within the Mexican lobeliads (Campanulaceae) is considered here. The taxonomic delimitations of Heterotoma from the Mexican species within the genera Lobelia and Calcaratolobelia were tested. Independent molecular data were gathered to determine whether the Mexican spurred lobeliads should be treated as disti
nct genera. The internal transcribed spacer (ITS) region from 18–26S nuclear rDNA and chloroplast DNA from the 3′ trnK intron were sequenced from 14 representative species. Our data suggest that Heterotoma, as originally conceived, is a good evolutionary unit within Lobelia and that the presence of a nectar spur is an important morphological character that can be used in defining phylogenetic position. This study also suggests that morphological changes associated with hummingbird pollination have evolved more than once in the Mexican lobeliads, from small blue-flowered, insect-pollinated relatives.
The identity and behavior of pollinators are among the main factors that determine the reproductive success and mating system of plants; however, few studies have directly evaluated the relationship between pollinators and the breeding system of the plants they pollinate. It is important to document this relationship because the global decline in pollinators may significantly affect the breeding systems of many animal-pollinated plants, particularly specialized systems. Ceiba pentandra is a trop
ical tree that has chiropterophilic flowers and a variable breeding system throughout its distribution, ranging from fully self-incompatible, to a mixed system with different degrees of selfing. To determine if regional differences in pollinators may result in regional differences in the outcrossing rate of this species, we used systematic observations of pollinator behavior in two tropical life zones and high-resolution genetic analysis of the breeding system of populations from these two regions using microsatellites. We found a predominantly self-incompatible system in regions with high pollinator visitation, while in environments with low pollinator visitation rates, C. pentandra changed to a mixed mating system with high levels of self-pollination.
Arisaema serratum possesses a pitfall-trap flower pollination system. However, little is known about the efficiency and pattern of pollen movement in A. serratum. Thus, the aims of this study are to (1) determine the paternal parents of the seeds and (2) elucidate pollen movement in a natural population. Paternity analysis using microsatellite markers was performed. Seeds were collected from a natural population of A. serratum in 2001 at Horigane, Japan. Small midges became trapped in female spa
the tubes during the flowering period. We found that (1) seeds in a fruit were fertilized by multiple sires; (2) seeds sired by a paternal parent were either clumped, exclusively, or randomly distributed on the spadix, depending on the parent; (3) to a great extent, a few males contributed as sires; (4) distance from a female was not a factor in the inequality of reproductive success among males; (5) male reproductive success was not correlated with its size. We conclude that pollen carryover and the trap-flower pollination system are likely to result in multiple paternity and inequality in male success.
Complementary field and laboratory tests confirmed and quantified the pollination abilities of Tranes sp. weevils and Cycadothrips chadwicki thrips, specialist insects of their respective cycad hosts, Macrozamia machinii and M. lucida. No agamospermous seeds were produced when both wind and insects were excluded from female cones; and the exclusion of wind-vectored pollen alone did not eliminate seed set, because insects were able to reach the cone. Based on enclosure pollination tests, each wee
vil pollinates an average 26.2 ovules per cone and each thrips 2.4 ovules per cone. These pollinators visited similar numbers of ovules per cone in fluorescent dye tests that traced insect movement through cones. Fluorescent dye granules deposited by Cycadothrips were concentrated around the micropyle of each visited ovule, the site of pollen droplet release, where pollen must be deposited to achieve pollination. In contrast, Tranes weevils left dye scattered on different areas of each visited ovule, indicating that chance plays a greater role in this system. Each weevil and 25 thrips delivered 6.2 and 5.2 pollen grains, respectively, on average, to each visited ovule per cone, based on examination of dissected pollen canals. In sum, the pollination potential of 25 Cycadothrips approximates that of one Tranes weevil.
Next Section Abstract Tacca, a genus of tropical herbs, possesses near black flowers, conspicuous involucral bracts and whisker-like filiform bracteoles. These features have been assumed to function as a “deceit syndrome” in which reproductive structures resemble decaying organic material attracting flies that facilitate cross-pollination (sapromyiophily). We investigated pollination and mating in Tacca chantrieri populations from SW China to evaluate this assumption. Contrary to this expectatio
n, populations were highly selfing. Pollinator visitation was infrequent and bagged flowers set abundant seed. Pollen loads on stigmas indicated autonomous self-pollination, some of which occurred prior to flower opening. The seed set of inflorescences with bracts and bracteoles removed was not significantly different from unmanipulated inflorescences, suggesting that these structures play a limited role in pollinator attraction, at least at our study sites. Pollen : ovule ratios averaged 49, a value expected in a highly selfing species. Selfing rates estimated in four populations using allozyme markers averaged 0.86 (range 0.76–0.94), thus corroborating this inference. Our results indicate that despite considerable investment in extravagant display, populations of T. chantrieri are highly selfing. We propose several hypotheses to resolve this paradox and argue that future studies of pollination syndromes would benefit by investigation of both pollination and mating biology.
Gynodioecy is a dimorphic breeding system in which hermaphrodite and female individuals coexist in populations. Sex ratio and gender-relative lifetime seed production determine the stability of gynodioecy, and both genetic and ecological factors may influence these parameters. I analyzed the consequences of variation in population sex ratio and site elevation for the relative pollination success of female and hermaphrodite individuals of Daphne laureola in southern Spain, where previous studies
failed to detect female fecundity advantages at two mid-elevation sites. Pollination success, estimated as stigmatic pollen loads, number of pollen tubes per style, and percentage of fertilized flowers, was higher for hermaphrodites than females in populations with 20–56% females. Furthermore, female quantitative disadvantage in pollination success increased with elevation, suggesting that the higher availability of pollen due to the increased proportion of hermaphrodites could not mitigate the negative effect that other factors associated with elevation apparently had on pollination. Supplemental hand pollinations showed that female seed production was pollen limited in populations with a proportion of females >50%, although both pollination success and natural fruit set of females in these sites were the highest recorded.
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 i
n 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.
The aerodynamics of wind pollination selects for an intimate relation between form and function in anemophilous plants. Inflorescence architecture and floral morphology vary extensively within the Poaceae, but the functional implication of this variation remains largely unknown. Here we quantify associations between floret, culm, and inflorescence characteristics for 25 grass species in Kananaskis, Alberta, Canada, and consider whether different architectures may implement unique mechanisms to a
id pollination. The species cluster into four categories defined by all combinations of floret size (small vs. large) and inflorescence architecture (compact vs. diffuse). Species differed significantly for all 18 traits that we measured, with 12 traits differing only between floret‐size classes, three differing only between inflorescence types, and three differing among both (independently or by an interaction). Based on these morphological associations, we discuss the aerodynamic and functional consequences of each category for wind pollination. The independence of inflorescence and floral traits has probably allowed exploration of all possible combinations of inflorescence architecture and floret size during the evolution of the Poaceae.
In the cultivated cranberry (Vaccinium macrocarpon), reproductive stems produce 1–3 fruit even though they usually have 5–7 flowers in the spring. We undertook experiments to test the hypothesis that this was an adaptive life history strategy associated with reproductive effort rather than simply the result of insufficient pollination. We compared fruit production on naturally pollinated plants with those that were either manually pollinated or that were caged to exclude insects. Clearly, insect
s are necessary for the effective pollination of cranberry plants, but hand pollination of all flowers did not result in an increase in fruit number. Most of the upper flowers, which had significantly fewer ovules than did the lower flowers, aborted naturally soon after pollination. However, when the lower flower buds were removed, the upper flowers produced fruit. This suggests that the upper flowers may serve as a backup if the earlier blooming lower ones are lost early in the season. Furthermore, the late‐blooming flowers may still contribute to the plant's reproductive success as visiting pollinators remove the pollen, which could serve to sire fruit on other plants. These results are discussed in the context of their possible evolutionary and proximate causes.
Plants interact simultaneously with a diversity of visitors, including herbivores and pollinators. Correlations among traits associated with herbivory and pollination may constrain the degree to which plants can evolve in response to any one interactor. Using the distylous plant, Gelsemium sempervirens, we tested the hypothesis that traits typically associated with pollination (distyly) and herbivore resistance (secondary compounds) were phenotypically correlated and examined how these traits in
fluenced plant interactions with floral visitors. The flowers of G. sempervirens are visited by pollinators and a nectar robber, and the leaves and flowers express gelsemine, an alkaloid that is deterrent and sometimes toxic to visitors. Using an observational approach across five populations, we found the thrum floral morph (short-styled) expressed more leaf gelsemine than the pin morph (long-styled). Leaf gelsemine concentrations were positively correlated with flower gelsemine; however, there were no correlations between gelsemine and other floral morphological traits. Trait expression influenced pollination more so than robbing. Thrums received two times less pollen than pins. Moreover, across both morphs, pollen receipt was lower in plants that expressed higher levels of leaf gelsemine in two sites. These results imply that traits associated with pollination and herbivore resistance may not be independent.
or shifts, we traced the evolutionary trajectory of flower traits and visitors onto a phylogeny based on sequence data from ITS, waxy, and trnF/ndhJ DNA. Maximum-likelihood ancestral reconstruction of floral traits suggests that ancestral Schizanthus had a bee-pollination syndrome. The hummingbird syndrome evolved in S. grahamii, a high elevation species in the Andes. The moth syndrome evolved in the ancestor of three species that inhabit the Atacama Desert. Results of mapping flower visitors onto the phylogeny show that the shift from bee to hummingbird pollination concurred with a shift in pollinators as predicted by the syndromes. However, the same pattern was not found for the moth syndrome. Visits by moths were observed only in one of the three moth-syndrome species, and at a very low rate. This mismatch suggests either anachronic floral characters or maintenance of rare, imperceptible moth pollination backed up by capacity for autonomous selfing. Overall, results suggest that diversification of flower traits in Schizanthus has occurred in relation to pollinator shifts.
The long‐standing notion of pollination syndromes, which postulates that plants form recognizable groups according to pollinator type, has been challenged recently on the basis of apparent widespread generalization in pollination systems. As a test of the pollination syndrome concept, I examined the pollination biology of a group of 15 orchids that share a recognizable syndrome of floral features that includes yellow‐green coloration, oil secretion, pungent scent, shallow flowers, and a Septembe
r peak in flowering. The orchids occur in sympatry in the Cape Floral Region of South Africa. According to the pollination syndrome concept, the similar floral features of this group indicate a shared pollinator. To test this prediction, I observed pollinators on Pterygodium alatum, P. caffrum, P. catholicum, P. volucris, Corycium orobanchoides, and Disperis bolusiana subsp. bolusiana. They shared a single species of pollinator, the oil‐collecting bee, Rediviva peringueyi. Female bees collected oil from the lip appendage using modified front tarsi. The orchids reduce interspecific reproductive interference through differences in pollinarium length or the use of mutually exclusive pollinarium attachment sites on the body of the bee. The results are contrary to the expectation of generalization in pollination systems and suggest that pollinators play an important role in mediating selection on floral traits.
Stigma position is a key aspect of flower morphology that may influence pollination success and seed production. In the self-incompatible, tristylous herb, Lythrum salicaria, the stigma is positioned inside the corolla tube in the short-styled morph, but outside the corolla in the mid- and long-styled morphs. We performed controlled pollinations to determine whether style morphs differ in duration of stigma receptivity and in interference caused by self-pollen deposition. As predicted, flowers o
f the short-styled morph remained receptive for longer if not pollinated than did flowers of the other two style morphs. When deposition of self-pollen preceded compatible cross-pollination, seed set was higher in the mid-styled morph than in the other style morphs. Interference caused by self-pollen deposition was overall strongest in the short-styled morph, but the magnitude of differences among style morphs was contingent on the relative timing of self and compatible pollen receipt. The results indicate that both differences in duration of stigma receptivity and interference from self-pollen deposition may contribute to observed variation in seed production and pollen limitation among style morphs in natural populations of L. salicaria. An association between stigma position and duration of stigma receptivity should constrain the evolution of flower morphology.
The floral traits of plants with specialized pollination systems both facilitate the primary pollinator and restrict other potential pollinators. To explore interactions between pollinators and floral traits of the genus Burmeistera, I filmed floral visitors and measured pollen deposition for 10 species in six cloud forest sites throughout northern Ecuador. Nine species were primarily bat‐pollinated (84–100% of pollen transfer); another (B. rubrosepala) was exclusively hummingbird‐pollinated. Ac
cording to a principal components analysis of 11 floral measurements, flowers of B. rubrosepala were morphologically distinct. Floral traits of all species closely matched traditional ornithophilous and chiropterophilous pollination syndromes; flowers of B. rubrosepala were bright red, lacked odor, opened in the afternoon, and had narrow corolla apertures and flexible pedicels, which positioned them below the foliage. Flowers of the bat‐pollinated species were dull‐colored, emitted odor, opened in the evening, and had wide apertures and rigid pedicels, which positioned them beyond the foliage. Aperture width appeared most critical to restricting pollination; hummingbirds visited wide flowers without contacting the reproductive parts, and bats did not visit the narrow flowers of B. rubrosepala. Aperture width may impose an adaptive trade‐off that favors the high degree of specialization in the genus. Other floral measurements were highly variable amongst bat‐pollinated species, including stigma exsertion, calyx lobe morphology, and pedicel length. Because multiple species of Burmeistera often coexist, such morphological diversity may reduce pollen competition by encouraging pollinator fidelity and/or spatially partitioning pollinator's bodies.
Explaining the diversity of mating systems and floral forms in flowering plants is a long-standing concern of evolutionary biologists. One topic of interest is the conditions under which self-pollination can interfere with seed set for flowering plants with a self-incompatibility system. We investigated the effect of self-pollen interference for wild radish, Raphanus raphanistrum, which has sporophytic self-incompatibility. We performed pollinations and determined seed set for plants grown in th
e greenhouse, using pollen mixtures representing either self- with outcross-pollen or outcross-pollen alone. Stigmas were collected for a subset of pollinated flowers to determine the number of pollen grains applied. Average seed set for the self/cross (5.13 seeds/pollination) and cross treatments (5.09 seeds/pollination) did not differ significantly. Stigmatic pollen loads averaged around 700 grains, an amount close to observed natural pollen loads on R. raphanistrum. We concluded that for R. raphanistrum in natural populations, self-pollen is unlikely to interfere with outcross-pollen success. This study is the first to investigate effects of self-pollen interference on seed set for a homomorphic species with sporophytic self-incompatibility where rejection occurs at the stigmatic surface.
To increase our knowledge about mating‐system evolution, we need to understand the relationship between specific floral traits and mating system. Species of Collinsia (Plantaginaceae) vary extensively in mating system; this variation is associated with variation in floral morphology and development and with the timing of self‐pollination. Counterintuitively, large‐flowered, more outcrossing species tend to have delayed stigma receptivity, reducing the amount of time that the stigma is receptive
to cross‐pollination before autonomous self‐pollination. To understand how the timing of stigma receptivity is related to mating‐system evolution, we studied in detail the timing of both stigma receptivity and self‐pollination (anther–stigma contact) in two greenhouse‐grown populations of large‐flowered Collinsia heterophylla. Crosses on emasculated flowers at different stages of floral development always produced seeds, suggesting that cross‐fertilization can be effected by pollen arriving prior to physiological receptivity. Phenotypic and genetic variation within populations in the timing of stigma receptivity and anther–stigma contact was substantial, although slightly less for the contact. Despite strong interspecific and interpopulation correlations, we did not find an among‐genet phenotypic correlation between the traits. This indicates that each trait may respond independently to selection, and the trait association may be the result of correlational selection.
In distylous, self‐incompatible plants, clonal propagation, unbalanced floral morph frequencies, and reduced population size can interfere with the functioning of distyly by compromising legitimate intermorph pollinations, resulting in reduced reproductive output. Here, we examined the mating system and the impact of mate availability, population size, and spatial aggregation of morphs on reproductive output in the distylous, clonal, aquatic plant Hottonia palustris. Controlled pollinations unde
r greenhouse conditions detected no spontaneous selfing without the action of a pollen vector (autonomous autogamy) and demonstrated very low fruit and seed development after self‐pollination. Intermorph (legitimate) crossings resulted in high reproductive output in both floral morphs (long‐ and short‐styled individuals), whereas intramorph (illegitimate) crossings decreased fruit and seed development by more than 50%, indicating that the species has partial intramorph‐incompatibility. In natural populations, small population size and increasing deviation of floral morph frequencies negatively affected reproductive outcome. Individuals of the majority morph type developed significantly fewer fruit and seeds than individuals of the minority morph type. This rapid decline in fecundity was symmetrical, indicating that regardless of which morph was in the majority, the same patterns of negative frequency‐dependent mating occurred. Increasing spatial isolation between compatible morphs significantly reduced fruit and seed set in both morphs similarly. This study provides clear indications of frequency‐ and context‐dependent mating in natural populations of a distylous plant species.
Next Section Abstract Exposed nectar presentation is a key trait in flowers specialized for pollination by short-tongued insects. We investigated the pollination of Satyrium microrrhynchum, a rare South African orchid in which nectar is secreted as droplets on long floral hairs (“lollipop hairs”) at the mouth of a shallow labellum. Our observations indicate that this orchid is pollinated specifically by two insect species: a cetoniid beetle (Atrichelaphinus tigrina) and a pompilid wasp (Hemipeps
is hilaris). Both insects have short mouthparts and remove nectar from the hairs with sweeping motions of their mouthparts. Pollinaria become attached to the upper surface of their heads while they feed on the nectar. Beetles damage the hairs while feeding, which may explain the positive relationship between hair damage and pollination success in plants of S. microrrhynchum from populations where beetles were common. The orchid has cryptic green-yellow flowers with spectral reflectance similar to that of its leaves. The fragrance from plants in three populations, analyzed using gas chromatography coupled to mass spectrometry, was dominated by various terpenoids; linalool was the most abundant. Plants in different populations emitted similar compounds, but eugenol and derivatives of this compound were found in only one of the three populations. In an electrophysiological study (gas chromatography coupled to electroantennography), using antennae of A. tigrina, clear signals were elicited by some of the floral scent compounds.
Gene flow through pollen and seed dispersal is important in terms of population differentiation and eventually speciation. Seed and pollen flow are affected in turn by habitats and pollen vectors. We examined the effect of different pollinators and habitats on gene flow by comparing two species of Streptocarpus, using microsatellite and chloroplast RFLP markers. Populations of the forest-dwelling S. primulifolius were highly differentiated according to nuclear microsatellite data and had mutuall
y exclusive chloroplast haplotypes. This result is congruent with infrequent seed dispersal and limited between-population foraging by the long-tongued fly pollinator Stenobasipteron wiedemanni. In contrast, populations of S. dunnii growing in exposed crags had lower levels of population differentiation according to both nuclear and chloroplast data, congruent with a hypothesis of more effective between population seed dispersal and greater pollen-mediated gene flow due to the sunbird pollinator Nectarinia famosa. The population genetic behavior of these species is reflected in their taxonomy and phylogenetic position; S. primulifolius belongs to a taxonomically complex clade in which recent speciation is evident, while the clade containing S. dunnii is characterized by taxonomically well-defined species on longer phylogenetic branches. Our study shows that pollinator movements and seed dispersal patterns are a major determinant of the evolutionary trajectories of these species.
Sexual dimorphism may be especially pronounced in wind-pollinated species because they lack the constraints of biotically pollinated species that must present their pollen and stigmas in similar positions to ensure pollen transfer. Lacking these constraints, the sexes of wind-pollinated species may diverge in response to the different demands of pollen dispersal and receipt, depending on the magnitude of genetic correlations preventing divergence between sexes. Patterns of sexual dimorphism and
genetic variation were investigated for inflorescence traits in Schiedea adamantis (Caryophyllaceae), a species well adapted to wind-pollination, and compared to S. salicaria, a species with fewer adaptations to wind pollination. For S. adamantis, dimorphism was pronounced for inflorescence condensation and its components, including lateral flower number and pedicel length. Within sexes, genetic correlations between traits may constrain the relative shape of the inflorescence. Correlations detected across sexes may retard the evolution of sexual dimorphism in inflorescence structure, including features favoring enhanced dispersal and receipt of pollen. Despite genetic correlations across sexes, common principal components analysis showed that genetic variance–covariance matrices (G matrices) differed significantly between the sexes, in part because of greater genetic variation for flower number in hermaphrodites than in females. G matrices also differed between closely related S. adamantis and S. salicaria, indicating the potential for divergent evolution of inflorescence structure despite general similarities in morphology and pollination biology.
Next Section Abstract The African orchid flora has a high proportion of species with long-spurred white flowers. Few data exist to test the prediction that this floral syndrome pattern reflects an important role for hawkmoth pollination in the evolution and ecology of these orchids. The pollination biology of five aerangoid orchid species (Rangaeris amaniensis, Aerangis brachycarpa, A. confusa, A. thomsonii, and A. kotschyana) was investigated in Kenya. Four of these have long spurs (>10 cm) and
were pollinated by Agrius convolvuli and Coelonia fulvinotata. Aerangis confusa, which has relatively short spurs (ca. 4 cm), was pollinated by the short-tongued hawkmoths Hippotion celerio and Daphnis nerii. Nectar frequently filled the entire spur in some of the study species, even at anthesis. Sugar concentration of the nectar of four species was found to vary from ca. 1% at the mouth of the spur to 20% at the tip. Gradients were expressed more strongly in species with long, straight spurs. Species with spirally twisted spurs showed both steep and shallow nectar gradients. These gradients, previously unknown in plants, may function as a “sugar trail,” enticing long-tongued hawkmoths to probe deeply into spurs without incurring the cost of filling an entire spur with concentrated nectar. In addition, the most concentrated nectar is kept out of reach of short-tongued pollinators.
Among the Cycadales (Cycadaceae and Zamiaceae), the Zamiaceae are known to be insect-pollinated. In contrast, the Cycadaceae are still considered wind-pollinated, although some doubt has been cast on several species, including Cycas revoluta. Using a large population of C. revoluta on Yonaguni Island (Okinawa, Japan), we performed exclusion experiments, documented insects from male and female cones, and analyzed the morphology of the apical part of the ovule to determine the pollination method o
f this species. Insect exclusion resulted in a notable reduction in seed set, except in a few individuals growing near male cones. The amount of airborne pollen was abundant within a 2-m radius of male cones but decreased markedly beyond this distance. Pollen grains of C. revoluta were found on the body of Carpophilus chalybeus (Nitidulidae, Coleoptera), one of a few species of insects collected from both male cones and female cones far from males. We conclude that C. revoluta relies on both wind (anemophily) and insect pollination (entomophily), although such anemophily is restricted to female trees growing within a 2-m radius of male trees. The nitidulids are not host specific to this cycad and primarily feed on plant tissue but serve as pollinators during pollen release. Cycas revoluta appears to be in an initial mode of animal pollination, as opposed to the host-specific insect pollination observed in most Zamiaceae.
The effectiveness of flower visitors as pollinators will determine their potential role as selective agents on flower traits. Pitcairnia angustifolia has floral characters that would fit pollination by long-billed hummingbirds, and they should be the most effective pollinators for this plant. To test this prediction, we characterized the behavior of visitors toward flowers and their pollination effectiveness. Coereba flaveola (bananaquits) was the most frequent flower visitor and acted as a prim
ary nectar robber; however, they pollinated incidentally and deposited pollen on stigmas. The endemic short-billed hummingbird Chlorostilbon maugaeus behaved as a secondary robber and did not pollinate flowers. As expected, the long-billed hummingbird, Anthracothorax viridis, was the most efficient visitor in terms of pollen deposition; however, it was the least frequent flower visitor. Introduced Apis mellifera (honeybees) were second in efficiency at depositing pollen and performed one third of the flower visits. Estimates of the expected rate of pollen deposition by each pollinator did not identify a single most effective pollinator. For P. angustifolia at least three flower visitors including an exotic bee and a nectar robber may be equally important to reproductive success. While these results limit our ability to make predictions on the role of hummingbird-pollination on current flower evolution, they do suggest the potential for pollination redundancy among flower visitors for P. angustifolia populations.
Narcissus triandrus is a tristylous daffodil from the Iberian Peninsula that has striking geographical variation in floral morphology and style‐morph ratios. Here, we investigate the relation between this variation and ecological factors to understand mechanisms governing morph ratios. We estimated morph ratios in 124 populations throughout the range of N. triandrus and measured 13 morphological traits in 35–78 populations. Sampling of morph ratios conducted over 2–10‐yr intervals demonstrated s
trong temporal stability. Variation in floral and vegetative traits enabled statistical prediction of morph‐frequency variation among populations. Latitudinal gradients in precipitation and temperature were correlated with plant and flower size, with larger flowers in northern populations associated with bumblebee visitation and stylar dimorphism. Flowers of the L‐ and M‐morphs differed significantly from the S‐morph in several size‐related characters, unlike other tristylous species. This pattern and the similarity in anther positions of the L‐ and M‐morphs suggest that the M‐morph of N. triandrus originated through genetic modifiers that shortened styles of the L‐morph. Our findings support the hypothesis that geographical variation in style‐morph ratios is largely governed by climatic gradients in the Iberian Peninsula, which influence the floral morphology and pollination biology of N. triandrus populations.
Despite evidence that both pollinators and nonpollinator agents of selection can shape the evolution of floral characters, there have been few attempts to compare the strengths and directions of selection from pollinators and other agents in the same study system. In this investigation of Leucanthemum vulgare, a self‐incompatible composite known for its conspicuous white rays, I obtained data from a ray removal experiment in the field and from a segregating F2 population in an experimental garde
n to assess the role of pollinator and nonpollinator selection as stabilizing factors on floral evolution in this species. Removal of all rays reduced the pollination success of heads by 31–35%, but did not significantly affect the level of infestation by larvae of the fly Tephritis neesii. Data from F2 plants indicated a potential for indirect selection on ray morphology, mediated through links between ray morphology and measures of vegetative size and plant vigor. The results of this study show that individuals of the normal, rayed phenotype have a clear selective advantage, in terms of both pollinator attraction and general plant vigor. Thus, there were no conflicting selection pressures between the pollinators and the other selective agents considered in this study.
The amount and genetic composition of pollen grains that are transported to fl owers infl uence the reproduction and fi tness of plants. Despite the importance of insect-pollination systems, an understanding of those systems is still lacking due to the absence of a genetic analysis of pollen grains that are transported to fl owers. We evaluated the pollination effi ciencies of bumblebees (Apidae, Bombus spp.), fl ower beetles (Scarabaeidae, subfamily Cetoniinae, Protaetia and Eucetonia sp.), and
small beetles (Lagriidae, Arthromacra sp.) that visited the fl owers of Magnolia obovata (Magnoliaceae) using quantitative (fl ower visitation frequency, amount of adherent pollen per insect) and qualitative (origin and genetic diversity of adherent pollen per insect) criteria. Most of the pollen adhering to bumblebees and small beetles was self-pollen. This result suggests that visitation by these insects may cause geitonogamous pollen fl ow and negatively affect the reproduction of M. obovata , causing inbreeding depression. In contrast, fl ower beetles transported large amounts of genetically diverse outcross pollen. Our results suggest that certain beetle species contribute quantitatively and qualitatively to the pollination of M. obovata . Direct genetic analysis of pollen grains will advance our understanding of plant mating systems and may shed light on the mutualism and coevolution of plants and fl ower visitors.
The buzz-pollinated genus Senna (Leguminosae) is outstanding for including species with monosymmetric flowers and species with diverse asymmetric, enantiomorphic (enantiostylous) flowers. To recognize patterns of homology, we dissected the floral symmetry character complex and explored corolla morphology in 60 Senna species and studied floral development of four enantiomorphic species. The asymmetry morph of a flower is correlated with the direction of spiral calyx aestivation. We recognized fiv
e patterns of floral asymmetry, resulting from different combinations of six structural elements: deflection of the carpel, deflection of the median abaxial stamen, deflection or modification in size of one lateral abaxial stamen, and modification in shape and size of one or both lower petals. Prominent corolla asymmetry begins in the earl-stage bud (unequal development of lower petals). Androecium asymmetry begins either in the midstage bud (unequal development of thecae in median abaxial stamen; twisting of androecium) or at anthesis (stamen deflection). Gynoecium asymmetry begins in early bud (primordium off the median plane, ventral slit laterally oriented) or midstage to late bud (carpel deflection). In enantiostylous flowers, pronouncedly concave and robust petals of both monosymmetric and asymmetric corollas likely function to ricochet and direct pollen flow during buzz pollination. Occurrence of particular combinations of structural elements of floral symmetry in the subclades is shown.
The co-occurrence of elaborate flowers visited by specific groups of pollinators and capacity for autonomous selfing in the same plant species has puzzled evolutionary biologists since the time of Charles Darwin. To examine whether autonomous selfing and floral specialization evolved in association, we quantified the autofertility level (AFI) in nine Schizanthus species characterized by a wide range of pollination specialization, revealing AFI values of 0.
02 to complete selfing. An independent contrasts analysis conducted on AFIs and number of functional pollinator groups showed that autonomous selfing evolved from an ancestral outcrossing system as plants became increasingly specialized (r = -0.82). To assess whether autonomous selfing together with specialization acts as a reproductive assurance mechanism, we estimated spatial and interannual variation in fruit set due to pollinator failure in two closely related high AndeanSchizanthus species differing in their specialization levels. Variation in pollinator failure rate was more pronounced and autonomous selfing increased fruit production over biotically assisted pollination in the more specialized species. Our study suggests that specialized pollination deems species more vulnerable to pollinator fluctuation thus promoting the evolution of delayed autonomous selfing.
The first three branches of the angiosperm phylogenetic tree consist of eight families with ∼201 species of plants (the ANITA grade). The oldest flower fossil for the group is dated to the Early Cretaceous (115-125 Mya) and identified to the Nymphaeales. The flowers of extant plants in the ANITA grade are small, and pollen is the edible reward (rarely nectar or starch bodies). Unlike many gymnosperms that secrete "pollination drops," ANITA-grade members examined thus far have a dry-type stigma.
Copious secretions of stigmatic fluid are restricted to the Nymphaeales, but this is not nectar. Floral odors, floral thermogenesis (a resource), and colored tepals attract insects in deceit-based pollination syndromes throughout the first three branches of the phylogenetic tree. Self-incompatibility and an extragynoecial compitum occur in some species in the Austrobaileyales. Flies are primary pollinators in six families (10 genera). Beetles are pollinators in five families varying in importance as primary (exclusive) to secondary vectors of pollen. Bees are major pollinators only in the Nymphaeaceae. It is hypothesized that large flowers in Nymphaeaceae are the result of the interaction of heat, floral odors, and colored tepals to trap insects to increase fitness.
This paper describes an open‐flower mutant, designated opf, that we discovered in a genetic screen of fast neutron bombardment mutants in an attempt at floral‐dip transformation of Melilotus alba (Fabaceae; white sweetclover), an alternative papilionoid legume host for Sinorhizobium meliloti. The opf mutant developed flowers with reflexed sepals and petals, thereby exposing the stamens and carpel, whereas wild‐type sweetclover inflorescences developed closed flowers where the young stamens and c
arpel remain covered during the early stages of flower development. Based on crosses with the wild type, the mutant segregated as a single, Mendelian recessive. Crosses were successful only when the opf mutant served as the female parent, suggesting that the mutant was male sterile. However, no obvious differences from wild‐type stamen development were observed in the opf mutant. The anther defect was due to indehiscence. However, as the plants approached the end of their life cycle, the frequency of selfing increased. We also investigated whether the opf mutant could be transformed via Agrobacterium tumefaciens floral‐dip infiltration because open flowers like those of Arabidopsis appear to be more readily transformable. However, similar to wild‐type M. alba, the opf mutant is refractory to floral‐dip transformation by Agrobacterium tumefaciens.
Evaluating the morphological relationships of angiosperm families that still remain unplaced in the current systems of classifi - cation is challenging because it requires comparative data across a broad phylogenetic range. The small neotropical family Metteniusaceae was recently placed within the lamiids, as sister to either the enigmatic Oncothecaceae or the clade (Boraginaceae + Gentianales + Lamiales + Solanales + Vahliaceae). We examined the development of two of the primary diagnostic trai
ts of Metteniusaceae, the moniliform anthers and the unilocular gynoecium. The gynoecium is 5-carpellate, and contains two ovules with a massive, vascularized integument. Late sympetaly and unitegmic ovules support placement of Metteniusaceae in the lamiids. The 5-carpellate gynoecium is consistent with a sister-group relationship between Metteniusaceae and Oncothecaceae. The gynoecium of Metteniusaceae is unusual in that it is monosymmetric throughout ontogeny, which indicates pseudomonomery; the fi ve carpel initials are congenitally fused by their margins and form a single locule; the two ovules develop from the two smallest and most poorly developed lateral carpels. Comparisons with other pseudomonomerous taxa allow us to propose division of the complex processes leading to pseudomonomery into eight characters, including carpel number and fusion, gynoecial symmetry, timing of carpel reduction, and number and position of nonfertile carpels.
Premise of the study: Flowering plants that rely on pollinators for most of their reproduction may experience unpredictable and inconsistent availability of effective pollinators throughout their reproductive lifetime. We investigated the reproductive ecology of two subspecies of the tufted evening primrose, Oenothera cespitosa, which occupy geographically and edaphically distinct habitats in western North America: O. cespitosa subsp. navajoensis inhabits sandstone soils on open sites or rocky s
lopes in the Colorado Plateau and O. cespitosa subsp. cespitosa grows in clay soils on talus slopes and exposed rocky ridges in the western Great Plains and northern Rocky Mountains of the United States.
Methods: Pollen augmentation and selfing experiments, floral visitor observations, and single‐visit effectiveness experiments were conducted over 4 years to examine the breeding system and spatiotemporal variation in pollinator behavior, assemblage, and abundance at different populations for each subspecies.
Key results: Both subspecies of O. cespitosa were self‐incompatible and pollen‐limited, suggesting that the relative abundance, effectiveness, and movement patterns of different insects as pollinators influenced the quality and quantity of seed production in these plants. Medium‐sized vespertine hawkmoths (Hyles lineata, Sphinx vashti) were effective pollinators when present, as were large matinal bees (Anthophora affabilis, A. dammersi, Xylocopa tabaniformis androleuca), whereas small oligolectic Lasioglossum bees primarily functioned as pollen thieves in the evening and morning.
Conclusions: These findings highlight the importance of variability of pollinator composition and abundance in the evolution of plant breeding systems and reproductive success at varying spatial and temporal scales.
The unique floral morphology of the South African orchid H. pulchra, with its twin meranthia, is best explained as an adaptation to pollination by oil‐collecting bees. Flowers consisting of meranthia (floral parts that function as single pollination units; commonly observed in garden Iris) are extremely rare among the angiosperms and their significance poorly understood. Unlike all other known examples of meranthia, the novel type described for H. pulchra is not bilabiate. All Huttonaea species
are unique in having twin petal sacs with glandular verrucae that secrete oil and are pollinated by Rediviva (Melittidae) oil‐collecting bees. But only Huttonaea pulchra has long and widely divergent petal claws that place the oil sacs well beyond the reach of a centrally positioned bee. The wide separation of these sacs forces the pollinator, R. colorata, to visit each side of the flower independently and effectively divides the flower into two meranthia. Molecular data indicate that the evolution of the Huttonaea‐type meranthium was dependent on the prior evolution of the oil flower/oil bee relationship. Meranthium evolution was also facilitated by the presence of oil in two separate structures (petal sacs) that were not physically constrained to remain in close proximity.
Premise of the study: We examined two accounts of the relationship between breeding system and life history variation in a clade of evening primroses (Oenothera, Onagraceae): (1) selection for reproductive assurance should generate an association between self‐compatibility and monocarpy and (2) phylogenetic conservatism leads to retention of breeding system and life history traits among closely related taxa.
Methods: We performed over 4000 hand pollinations under greenhouse conditions to determi
ne the compatibility of 10 Oenothera taxa (sections Anogra [17 taxa] and Kleinia [2 taxa)] for which breeding systems had not previously been reported. We used generalized linear mixed models to evaluate the influence of pollination treatment, parents, and population on fruiting success.
Key results: Among the taxa tested, six were self‐incompatible, two were variable in compatibility, and two were self‐compatible. We combined these data with published studies in Anogra and Kleinia and mapped breeding system and life history onto a published phylogeny.
Conclusions: We found no evidence for phylogenetic conservatism, but detected considerable evolutionary lability in both traits. Additionally, we found no evidence for a consistent relationship between breeding system and life history. Only eight of 19 taxa followed the predicted association between self‐incompatibility and polycarpy vs. self‐compatibility and monocarpy. Instead, many taxa have retained self‐incompatibility, regardless of monocarpy or polycarpy.