Pollination biology of the Galápagos endemic, Tournefortia rufo-sericea (Boraginaceae)

Tournefortia rufo-sericea is an endemic member of the Galápagos angiosperm flora. Although not uncommon within the archipelago, its status is presently listed as vulnerable and, as such, a complete knowledge of its pollination biology may prove useful in preventing its decline. Pollination experiments, flower-visitor observations, nectar and fluorescence studies, as well as pollen : ovule ratio and pollen size studies were included in this investigation. The small, white flowers of this species set fruit via open pollination (81%), autonomous self-pollination (80%), diurnal pollination (80%), and nocturnal pollination (85%). Ants are the most common visitor to the flowers of this species, primarily during the day, whereas beetles and moths make visits at night. A small amount of nectar is presumably produced, as moths are often seen probing the corollas. Experiments with fluorescent dust did not support interflower pollen movement. The pollen : ovule ratio was 4972 : 1, which suggests xenogamy, and the mean pollen size was 21 µm. Despite the pollen : ovule ratio, it appears that this species exhibits a breeding strategy of facultative autogamy, in which the majority of flowers set fruit via autonomous self-pollination, but limited outcrossing may occur when pollinators are available. Conservation efforts for T. rufo-sericea should focus on the control or eradication of exotic species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 21–31.     

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

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

Reproductive ecology of the endemic Lecocarpus pinnatifidus (Asteraceae) in an isolated population in the Galápagos Islands

Lecocarpus pinnatifidus is an endemic member of the Asteraceae occurring on only one island in the Galápagos archipelago. The capitula are large with female ray florets and male disc florets. They are self-compatible but this study suggests fruit set is pollen limited. Visits from Xylocopa darwini and other larger insect pollinators are rare, and small insects seem to be the main pollinators. Small insects carry few pollen grains and most likely mediate self-pollinations. Self-compatibility and seed set after selfing are the most common reproductive strategy in the Galápagos Islands and L. pinnatifidus seemingly fits well into this group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 171–180.     

Noncorrelated evolution between herbivore- and pollinator-linked features in Aristolochia chilensis (Aristolochiaceae)

The effect of mutualists (i.e. pollinators) and antagonists (i.e. herbivores) can have non-additive effects on plant fitness. This is often interpreted as evidence for correlated evolution on a suite of traits leading to an increase and decrease of the interaction of plants with mutualists and antagonists, respectively. This situation has been found to prevail in plants that have large floral and fruit displays but are not limited by pollinators for seed set. We suggest the alternative hypothesis, where plants limited by pollinators for seed set (e.g. deceit-pollinated plants) exhibit additive effects of pollinators and herbivores on fitness (i.e. noncorrelated evolution). Using a 2 × 2 factorial design, we tested this hypothesis by solely and simultaneously evaluating the effects of pollinators and the single herbivore, Battus polydamas archidamas , on female reproductive success of Aristolochia chilensis . Plants exposed to herbivores presented 2.6-fold greater herbivory than plants that excluded them. In addition, plants exposed to pollinators showed strong limitation by pollinators for seed set compared with other plants of the genus Aristolochia. However, only pollinators had a significant effect on fruit and seed set because plants that excluded them did not set fruits or seeds. Furthermore, herbivores and pollinators exerted additive effects on fruit and seed production. Collectively, these results indicate that herbivore- and pollinator-linked traits in A. chilensis exhibit noncorrelated evolution.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 239–245.     

Pollination biology of a night‐flowering Galápagos endemic,Ipomoea habeliana (Convolvulaceae)

Ipomoea habeliana is an endemic, night‐flowering member of the Galápagos flora. Pollination experiments, flower‐visitor observations, nectar sampling, pollen transfer, and pollen to ovule ratio and pollen size studies were included in this project. The large, white flowers of this species set fruit via open pollination (55%), autonomous autogamy (51%), facilitated autogamy (91%), cross‐pollination (80%), diurnal open pollination (60%) and nocturnal open pollination (60%). Fruit set is pollen‐limited. Ants, beetles, crickets and hawk moths regularly visit the flowers. Ants are the most frequent visitors, but hawk moths are the only effective pollinators. Nectar is available throughout the night, but is most abundant early in the evening when hawk moth visits are most frequent. Experiments with fluorescent dust demonstrate intra‐ and inter‐plant pollen movement by hawk moths. Although this species is adapted for hawk moth pollination, it readily sets fruit via autonomous autogamy when no visits are made. Thus, it is concluded that it is facultatively xenogamous. Additional support for this conclusion is provided by the pollen to ovule ratio of 1407 and by the fact that the plants grow in a region that has few or no faithful pollinators. Conservation efforts for I. habeliana should include hand pollinations, which could significantly increase seed set. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 11–20.     

Reproductive biology of Boswellia serrata, the source of salai guggul, an important gum-resin

Detailed studies were carried out on the phenology, floral biology, pollination ecology and breeding system of Boswellia serrata Roxb. (Burseraceae) the source of 'salai guggul'. The trees remain leafless during the entire period of flowering and fruiting. The inflorescence is a terminal raceme and produces up to 90 bisexual, actinomorphic flowers. On average a flower produces 10 044 ± 1259 starch-filled pollen grains. About 85% of the fresh pollen grains are viable; the pollen to ovule ratio is 3348 : 1. The stigma is of the wet papillate type. The style is hollow with three flattened stylar canals filled with a secretion product. The stylar canals are bordered by a layer of glandular canal cells. The inner tangential wall of the canal cells shows cellulose thickenings. The ovary is trilocular and bears three ovules, one in each locule. Flowers offer nectar and pollen as rewards to floral visitors. The giant Asian honey bee ( Apis dorsata ) and A. cerana var. indica (Indian honey bee) are the effective pollinators. The species is self-incompatible and the selfed pollen tubes are inhibited soon after their entry into the stigma. Self-pollen tubes develop a characteristic 'isthmus' as a result of enlargement of the tube soon after emergence through the narrow germ pore. Cross-pollinated flowers allowed normal pollen germination and pollen tube growth, and resulted in fruit- and seed-set. Under open pollination fruit-set was only about 10%. Although manual cross-pollinations increased fruit set, it was only up to about 20%. Low fruit set appears to be the result of inadequate cross-pollination and other constraints, presumably limitation of available nutrients.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 73–82.     

How flower biology and breeding system affect the reproductive success of the narrow endemic Polygala vayredae Costa (Polygalaceae)

Polygala vayredae is a narrow endemic species from the oriental pre-Pyrenees. Despite its conservation status and rarity, no information is available on its reproductive biology. As the flower is the structure directly involved in pollinator attraction, its morphological and functional traits have major effects on the reproductive success of the plant. In this work, the flower biology and breeding system of P. vayredae were studied to evaluate how they affect the reproductive outcome in natural populations. Flower morphology, flower rewards, and male and female functioning throughout the lifespan of the flower were assessed. Pollination experiments, involving pollinator exclusion and pollen from different sources, were conducted, and the pollen ovule index was determined. Female fitness and the occurrence of pollen limitation were assessed in three natural populations over 2 years by observing the presence of pollen on the stigma, pollen tube development, and fruit production. Polygala vayredae flowers are elaborate and long-lived with nectar rewards. The floral traits are well adapted to xenogamy and entomophily, which are in accordance with the observed breeding system and auto-incompatibility system. No mechanism of reproductive assurance was observed and P. vayredae strictly depends on pollinators to set fruit. Low fruit production was observed in the studied populations, which was largely the result of scarce, unreliable, and/or inefficient pollinators and poor pollen quality. In addition, available resources may be a limiting factor. The reproductive strategy of P. vayredae prevents inbreeding depression by a self-incompatibility system, which in years of scarce pollinators is overcome by the plant habit.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 67–81.     

Flowering phenology and reproductive biology of Drosera anglica (Droseraceae)

Prompted by the sparse knowledge of the reproductive biology of carnivorous plants, compared with studies of their trapping habits, we investigated the flowering phenology and pollination biology of Drosera anglica Huds. in two fens in mid-western Canada. Seed set and germination were used to compare the effectiveness of a series of pollination treatments, including single insect visits to virgin flowers. Flowers opened during mid-morning but closed by early afternoon, and exhibited pseudo-cleistogamic behaviour in cool, overcast weather. D. anglica was found to be self-compatible, and able to self-pollinate and self-fertilize. Geitonogamy was an uncommon mode of self-reproduction because plants typically possessed a lone inflorescence upon which a single, short-lived flower opened, a few days before the next bud reached anthesis. Insect visits to the fragrance-lacking, nectarless flowers, chiefly by flies (Diptera: Syrphidae), were infrequent (one visit per 1 h 40 min of observation), and the low frequency of seed set and low numbers of seeds per fruit in pollination treatments involving insects, suggest the species does not rely on insects to effect pollination. Self-pollination, with or without the aid of a vector (insects, wind) was as effective as natural pollination; ultimately, autogamy is chiefly responsible for natural seed set. Thus, the species exhibits characteristics of facultative autogamy.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 417–426.     

Pollination by fungus gnats in four species of the genus Mitella (Saxifragaceae)

The first example of pollination by fungus gnats in the eudicots is reported. The genus Mitella (Saxifragales) is characteristically produces minute, inconspicuous, mostly dull-coloured flowers with linear, sometimes pinnately branched, petals. To understand the function of these characteristic flowers, we studied the pollination biology of four Mitella species with different floral traits and different sexual expression: dioecious M. acerina , gynodioecious M. furusei var. subramosa , and hermaphroditic M. stylosa var. makinoi and M. integripetala. Flower-bagging experiments showed that wind pollination did not occur in the dioecious and gynodioecious species. Two years of observations of flower visitors at six study sites in Japan revealed that the principal pollinators of all four Mitella were specific species of fungus gnats (Mycetophilidae), which landed on the flowers with their long spiny legs settling on the petals. Characteristically, numerous pollen grains were attached to the fungus gnats in specific locations on the body. Although, on average, 1.3–2.6 fungus gnats visited each inflorescence per day, the fruit set of both bisexual and female flowers exceeded 63%. These results suggest that fungus gnats are highly efficient pollinators of Mitella spp., and that Mitella flowers are morphologically adapted to pollination by fungus gnats.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 449–460.     

Flowering in a bamboo, Melocanna baccifera (Bambusoideae: Poaceae)

Melocanna baccifera (Roxburgh) Kurtz ex Skeels, a species of bamboo introduced to Sri Lanka from India, flowered and set fruit during 2001–2002. Culms that flowered and set fruit died. The incidence of flowering is significant in that flowering took place close to the predicted mast flowering in 2007. At the onset of flowering, inflorescences were predominantly staminate. But later in 2002, bisexual and pistilate flowers also developed leading to fruiting. Both protandry and protogyny were observed in the bisexual florets. Floral characters indicated that the species was mainly out-crossing. Although anther dehiscence released pollen and stigmas were exerted, pollination of stigmas was inefficient. The few stigmas that were naturally pollinated showed limited pollen tube growth. However, fruit set took place. Fruits were very rarely seen to germinate naturally. Many fruits were devoid of an embryo, indicating that parthenocarpy and/or apomixis may have taken place. Excised embryos germinated in vitro .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 287–291.     

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

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

Invasive Africanized honey bee impact on native solitary bees: a pollen resource and trap nest analysis

Little is known of the potential coevolution of flowers and bees in changing, biodiverse environments. Female solitary bees, megachilids and Centris , and their nest pollen provisions were monitored with trap nests over a 17-year period in a tropical Mexican biosphere reserve. Invasion by feral Apis (i.e. Africanized honey bees) occurred after the study began, and major droughts and hurricanes occurred throughout. Honey bee competition, and ostensibly pollination of native plants, caused changes in local pollination ecology. Shifts in floral hosts by native bees were common and driven by plant phylogenetics, whereby plants of the same families or higher taxa were substituted for those dominated by honey bees or lost as a result of natural processes. Two important plant families, Anacardiaceae and Euphorbiaceae, were lost to competing honey bees, but compensated for by greater use of Fabaceae, Rubiaceae, and Sapotaceae among native bees. Natural disasters made a large negative impact on native bee populations, but the sustained presence of Africanized honey bees did not. Over 171 plant species comprised the pollen diets of the honey bees, including those most important to Centris and megachilids (72 and 28 species, respectively). Honey bee pollination of Pouteria (Sapotaceae) plausibly augmented the native bees' primary pollen resource and prevented their decline. Invasive generalist pollinators may, however, cause specialized competitors to fail, especially in less biodiverse environments.  No claim to original US government works. Journal compilation © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 152–160.     

Geographical variation in floral traits of the tree tobacco in relation to its hummingbird pollinator fauna

Nicotiana glauca , a hummingbird pollinated plant, exhibits geographical variation in several floral traits. We examined whether geographical differentiation occurred for different flower characters and if this differentiation could be explained, at least in part, by the existence or abundance of different hummingbird species in the respective pollinator assemblages. The comparison between five populations showed significant variation in six floral traits and two female fitness measures. The traits that better discriminated between populations were corolla length and corolla width. There were metric correlations between corolla length and style length in all populations studied and, in four of the five populations, both corolla length and width were also correlated. Among plants in each population, seed weight was correlated positively and significantly with style exertion, suggesting that fruit quality is dependent on the degree of cross pollination. Assemblages of hummingbirds differed between populations in species composition, visitation frequencies, and bill length. Linear regression involving bill length of the more frequent hummingbird pollinators and corolla length yielded positive and significant relationships. Thus, there appears to be an adjustment between pollinators and flowers traits that have high incidence in the among population variation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 657–667.     

Why be a rewarding trap? The evolution of floral rewards in Arum (Araceae), a genus characterized by saprophilous pollination systems

Plant species of the genus Arum typically have lure-and-trap pollination systems that are saprophilous (i.e. attracting flies or beetles searching for breeding sites in decaying organic matter). They have been assumed to always attract and trap their pollinators by deception because the inflorescences provide unsuitable breeding grounds for pollinators. The present study explored the possibility that one species, Arum creticum Boiss. & Heldr., which has yellow, sweet-smelling inflorescences, rewards its pollinators and that this increases its success in attracting pollinators over its close relative, Arum idaeum Coust. & Gadoger. The relationship between rewards provided, floral structure, insect attraction, and pollen import and export was examined in two naturally occurring sympatric populations of A. creticum , A. idaeum , and their natural hybrids. The results showed that plants providing more pollen were visited by larger numbers of females of a mining bee Lasioglossum marginatum Brullé as well as adults and nymphs of a hemipteran bug Dionconotus cruentatus creticus Heiss. In A. creticum , L. marginatum was found to be a better outcrossing vector than D. cruentatus in areas where both pollinators occurred because L. marginatum individuals carried more pollen on their bodies and travelled greater distances between inflorescences, thus increasing the potential for outcrossing. The finding that floral rewards may result in increased fitness, compared to nonrewarding systems that rely on attracting saprophilic pollinators, suggests that it is possible for rewarding species to evolve from saprophilous systems.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 257–268.     

Floral phenology and sex expression in functionally monoecious Rhoiptelea chiliantha (Rhoipteleaceae)

The reproductive ecology of wind-pollinated gynomonoecious species, in which the individual plant produces both female (pistillate) and perfect flowers, has rarely been studied. We examined the floral phenology and reproductive traits in Rhoiptelea chiliantha , described as gynomonoecy, to understand the adaptive significance of this sexual system. This species is a rare tree native to south-western China and northern Vietnam. The flowers are characterized by an anemophilous pollination syndrome, but no insects were observed foraging on them. Perfect flowers have larger tepals but smaller stigmas than female flowers, indicating flower size dimorphism. Floral ratios of female to perfect flowers are stable in different individuals and populations. On individual plants, perfect flowers open first, followed by female flowers, with a 1-week interval. Perfect flowers are protogynous with a 3.7-day interval (neuter phase) between the female phase (1.5 days) and expanded male phase (8.2 days). Both female and perfect flowers exhibit pronounced synchrony in flowering at the levels of inflorescences and individuals. However, flowers on different individuals show asynchronicity in timing of initial blooming. Tracking the process from pollination to fruit maturation, we found that female flowers contributed almost exclusively to seed production, but perfect flowers were sterile (functionally males). Therefore, this plant is functionally monoecious. This finding resolved a puzzle on the occurrence of female flowers in this plant, because previous reports described female flowers as being sterile. As the sex phases were completely separate between individuals, the pattern of floral phenology may ensure that outcrossing strongly predominates.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 145–151.     

Reproductive strategies of Strobilanthes kunthianus, an endemic, semelparous species in southern Western Ghats, India

Strobilanthes kunthianus (Acanthaceae) is a semelparous species with synchronized flowering and mast seeding once in 12 years. As semelparous plants have only one chance to reproduce, they are expected to develop effective strategies to prevent reproductive failure. The reproductive strategies of S. kunthianus were investigated by studying the floral traits, pollination biology, and breeding system that are critical for reproductive success. The species exhibits a series of floral traits: (1) gregarious flowering attracts a large number of Apis cerana indica , the major pollinator; (2) the stigma is sensitive to touch by the pollinator; in fresh flowers, the receptive surface faces the entry path of the incoming bee, facilitating pollen deposition; as an immediate response, the stigma curves backwards moving the receptive surface away from the path of the exiting bee, thus preventing autogamy and interference in pollen transfer; (3) flowers remain fresh for 2 days with receptive stigma and nectar and pollen reward. These traits render the species 100% pollination efficient to ensure a high seed set. As the species is self-compatible, the prevailing high degree of geitonogamous pollinations does not interfere with fruit set. The evolution of the adaptive floral traits has facilitated mast seeding in the species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 155–163.     

Self‐incompatibility and post‐fertilization maternal regulation cause low fecundity in Aegle marmelos (Rutaceae)

Outbreeding confers an evolutionary advantage, and flowering plants have evolved a variety of contrivances for its maximization. However, neither fruit set nor seed set is realized to its fullest potential for a variety of reasons. The causes of low flower to fruit and seed to ovule ratios were investigated in naturally occurring bael trees (Aegle marmelos) at two sites for three seasons. The study established that the mass effect of synchronized flowering attracted a variety of insect pollinators to the generalist flowers; Apis dorsata was the most efficient pollinator. The seed to ovule ratio was low despite high natural pollination efficiency (c. 2400 pollen per stigma). Although pollination‐induced structural and histochemical changes in the style allowed many (9.5 ± 2.1) pollen tubes to grow, only cross‐pollen tubes could grow through the style. Gametophytic self‐incompatibility, manifested in the stylar zone, resulted in a significantly slower growth rate of self‐pollen tubes. The occurrence of obligate self‐incompatibility, coupled with increased self‐pollen deposition (geitonogamy), caused a significant number of flowers to abort. Fruit retention in the trees declined from 40% to 12% as a result of abortion of fruits at different stages of development. The number of mature fruits on a tree was negatively correlated (r = ?0.82) with their size. It is inferred that low natural fecundity in A. marmelos is primarily a result of obligate self‐incompatibility and strong post‐fertilization maternal regulation of allocation of resources to the developing fruits. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 572–585.     

Floral biology and pollination mechanisms of four Mexico-endemic Fuchsia species with contrasting reproductive systems

Aims The evolution of the outstanding variation of reproductive systems in angiosperms has been considered an important driver of lineage diversification. Closely related hermaphroditic and dioecious species with biotic pollination provide the opportunity to study and compare traits related to pollinator attraction and their consequences on reproductive components. A higher predictability of pollination syndromes is expected in dioecious species, which are dependent on pollinators, than in self-compatible hermaphroditic taxa. Dioecious species may suffer pollen limitation depending on the quality of floral rewards and the kind and abundance of pollinators, whereas no pollen limitation is expected in hermaphroditic species with autonomous self-pollination. Additionally, in the absence of pollen limitation, more or better seeds are expected in dioecious species, according to the sexual specialization hypothesis.Methods In natural populations of the hermaphroditic Fuchsia fulgens and Fuchsia arborescens and dioecious Fuchsia parviflora and Fuchsia obconica, all endemic to Mexico, we first described flower phenology, flower production and longevity and nectar volume and concentration. Then, we evaluated the correspondence between floral visitors and pollination syndromes. In hermaphrodite plants, we determined the level of herkogamy and the potential for autonomous self-pollination. Finally, we evaluated the effect of pollen limitation on fruit set and seed number and assessed seed germination for all species.Important findings In contrast to our prediction, dioecious species did not show a higher correspondence between pollination syndromes and floral visitors than did hermaphrodites; however, male flowers exhibited a higher correspondence than female flowers. No pollen limitation was detected in dioecious species, for which visitation rate did not differ between male and female flowers. The hermaphroditic F. fulgens showed pollen limitation for seed number, despite the presence of autonomous selfing. Fruit set from autonomous pollination was higher in F. arborescens, which showed a lower level of herkogamy compared with F. fulgens. Finally, dioecious species produced fewer but heavier seeds compared with hermaphrodite species. Although Fuchsia is classified as an outcrossing genus, both hermaphroditic species showed autonomous self-pollination. The heavier but lower number of seeds per fruit in dioecious species may be related to the more efficient resource allocation expected from sexual specialization. This could play an important role in the evolution of dioecy; however, a comparative phylogenetic approach is required to confirm this hypothesis.     

Positive effects of forest edges on plant reproduction: literature review and a case study of bee visitation to flowers of

Positive effects of fragmentation on plant reproduction are uncommon; in a literature review we found significant negative effects on fruit or seed set for 50 plant species, compared to 26 species showing no effect, and only nine affected positively. One of these is the declining New Zealand mistletoe Peraxilla tetrapetala (Loranthaceae), and here we investigate the mechanism of this positive effect. P. tetrapetala requires visits from native bird or bee pollinators to produce fruit. Fruit set was consistently pollen limited at several South Island sites because of a shortage of pollinators, but within a site at Lake Ohau, plants on forest edges had higher fruit set than those in the forest interior. Previous work showed that this difference was not caused by a shortage of resources in interior plants, but was associated with higher bird visitation rates to flowers on edges. In this study, we tested whether native bees also show a preference for edge flowers. At two sites (Ohau and Craigieburn) edge mistletoes had higher visitation by native bees (Hylaeus agilis and Leioproctus sp.) and higher fruit set. Some, but not all, of the higher visitation to edge flowers was explained by a preference amongst bees for flowers in direct sunshine. Therefore, P. tetrapetala experiences higher fruit set on edges because both of its main groups of pollinators (endemic birds and bees) visit edge flowers more often. The other eight published cases of positive effects of fragmentation on fruit set also all reported increased visitation rates by pollinators.     

Pollination of the heterostylous Galápagos native, <Emphasis Type="Italic">Cordia lutea</Emphasis> (Boraginaceae)

The hypothesis of this study was that in the Galápagos Islands, fruit and seed set via nocturnal pollination would exceed that of diurnal pollination due to greater insect activity at night typical of hot, arid regions. Cordia lutea, a heterostylous member of the Galápagos flora, was submitted to pollination experiments, visitor observations, nectar sampling, pollen transfer studies, pollen-ovule ratio studies, and pollen measurements. Flowers set fruit and seed via open pollination, autonomous autogamy, facilitated autogamy, facilitated cross-pollination, diurnal pollination, and nocturnal pollination. There was a significant difference in fruit set between flowers experiencing legitimate cross-pollinations (pin × thrum) and those experiencing all other pollination treatments except facilitated autogamy. There was no significant difference in seed set among any of the treatments, but there was a trend toward greater seed set for flowers experiencing open-pollination, legitimate cross-pollination, and nocturnal pollination. There was no significant difference in fruit set or seed set between flowers experiencing diurnal pollination and nocturnal pollination, although there was a trend toward greater seed set resulting from nocturnal pollination. Carpenter bees were the most effective diurnal pollinators, whereas moths were the most effective nocturnal pollinators. Of the two, moths are more efficient at transporting pollen from plant to plant. Results indicate that an overall low productivity of this species is due to pollen limitation exacerbated by nectar robbing. Cordia lutea exhibits a mixed mating system, producing a relatively low level of fruits through a combination of self- and cross-pollination, facilitated by the relatively few insects that are available.