Fruit abortion in the Chihuahuan-Desert endemic cactus <Emphasis Type="Italic">Opuntia microdasys</Emphasis>

Resource and pollen limitation, as well as pollen/ovule incompatibility, have been proposed as causes to explain fruit abortion. To assess whether abortion in Opuntia microdasys was due to resource and/or pollen limitation and could therefore be reversed fruit set and seed set were studied using controlled pollination experiments on 60 plants that had been randomly assigned a combination of watering and fertilization treatments. On the other hand, to test whether fruit abortion was irreversible, due to pollen/ovule incompatibility, we examined the reproductive biology of the species. This included observations on floral phenology, nectar production, flower visitors, numbers of pollen grains and ovules, and self-pollination experiments. Results showed that O. microdasys is a fully self-incompatible species and its floral biology and the activity of the main pollinator allow constant deposition of incompatible pollen onto stigmas, which may contribute to fruit abortion. Reproductive success was limited by nutrients and pollen, but the fruit set increased only by 58%, compared to 47% of the control, after the experimental addition of pollen, nutrients and water. The magnitude of pollen and resource limitation suggests that similar levels of abortion will be present in good as well as in bad years. Selfing as well as incompatibility between ramets from the same clone and between closely related plants seem plausible candidates to explain the large proportion of fruit abortion, and experimental cross pollination between genotypes identified through molecular markers are necessary to fully understand the considerable abortion rate that remains unexplained after pollen and resource addition. Interestingly, the possible reason why the abortion of energetically expensive fruits has not been eliminated by natural selection is that the aborted fruits are propagules able to root and produce new plants with the same genotype of the mother. Abortion would have a dramatic effect on cross-fertilized genotypes because they result in zero fitness, but it would have a positive effect on the fitness of the maternal genotype because a clonal offspring is produced. Evidently, the exact fitness consequences to the maternal plant will depend on the differences in survival and reproduction of these different offspring types.     

Reproductive biology of Ranunculus weyleri (Ranunculaceae), a narrowly endemic plant from the Balearic Islands with disjunct populations

Ranunculus weyleri is a species endemic to Mallorca Island (western Mediterranean basin) that is currently threatened due to its reduced geographical distribution and disjunct, isolated populations. It is a small, perennial herb that inhabits rocky environments, together with other endemic plants with creeping life-forms. In the present study, for the first time the reproductive biology of the species was investigated in terms of its floral phenology (floral stages and times of formation), pollen/ovule (P/O) ratios, and fruit and seed setting under natural and pollen manipulation experiments, assessing the breeding system, pollination vectors and pollen limitations. Cross-pollinations among populations were performed to evaluate the levels of compatibility between populations. In addition, the floral displays and the reproductive outputs of four wild populations were determined. The observed high P/O ratios, results of the hand-pollination experiments, and flower traits indicate that this species is predominantly allogamous. Combinations of both insects (belonging to Coleoptera, Diptera, Lepidoptera and Hymenoptera) and wind (ambophily) constitute the main pollination vectors, and pollen limitation was observed. Interpopulation crossing was effective, and exogamic depression was not present, as evidenced by the fecundity levels. The predation of the floral stems by herbivores (mainly goats) severely limits the reproductive success of this endemic species in wild populations. Finally, several ways of action are proposed for R. weyleri conservation.     

广西蜘蛛抱蛋(Aspidistra retusa)的传粉生物学研究

蜘蛛抱蛋属植物的花部结构丰富多样,且常常贴地表开花,但其传粉生物学研究鲜有报道。广西蜘蛛抱蛋(Aspidistra retusa)是广西特有植物,为揭示该物种的自然传粉机制,在蜘蛛抱蛋属世界分布和分化中心的广西境内,开展了广西蜘蛛抱蛋开花物候、访花昆虫及繁育系统的野外观测研究。结果表明:广西蜘蛛抱蛋的花部结构中雄蕊高于雌蕊,但自交不亲和,为异花传粉;双翅目瘿蚊属成虫(Cecidomyia sp.,Cecidomyiidae,Diptera)为其专性传粉者,作为回报,广西蜘蛛抱蛋为瘿蚊提供理想的交配场所,同时提供大量的花粉供雌瘿蚊取食以补充生殖所需能量,并为受精卵的孵化以及幼虫的发育提供舒适的环境。该研究用实地现场观察第一手数据验证了前人有关蜘蛛抱蛋属植物瘿蚊传粉的推测,为中国蜘蛛抱蛋属植物传粉生物学研究的首次报道,然而蜘蛛抱蛋属植物花部结构极具多样性,应存在更多样的传粉方式,还有待更深入的调查研究。     

Floral morph composition and pollen limitation in the seed set of <Emphasis Type="Italic">Nymphoides indica</Emphasis> populations

We investigated the floral morph composition and seed set of the populations in a heterostylous aquatic plant, Nymphoides indica O. Kuntze (Menyanthaceae). The bias of floral morph ratio in the populations was negatively correlated with the seed set. In the populations with biased floral morph ratio, the pollen on the stigma of the common morphs were dominated by pollen of the same morph, resulting in reduced seed set. The addition of pollen of the opposite morph on the stigmas of the common morph resulted in an increase of fruit and seed set rates. The intermorph pollination rate and the fruit and seed set rates of Nymphoides indica were highest when two floral morphs were close in the populations. Various insect species including Lepidoptera, Diptera and Hymenoptera visited the flowers. However, geitonogamous pollination was common because of the short flying distance of the pollinators. These results indicate that the seed set of Nymphoides indica is limited by the shortage of compatible pollens and that the success of intermorph pollination is influenced by spatial distribution of the two floral morphs. As Nymphoides indica is endangered, conservation of the populations with high fecundity is proposed by us.     

宁夏枸杞的花粉呈现式样及交配系统研究北大核心CSCD

被子植物的花回馈、雌雄蕊时空分离特征和花粉呈现式样等花部特征及传粉者效率会影响雄性适合度和有性繁殖过程。宁夏枸杞(Lycium barbarum)是中国宁夏、新疆、内蒙等干旱和半干旱地区分布的特有种。该研究对新疆喀什地区宁夏枸杞自然种群的花部综合征、花粉呈现式样、花回馈、传粉者行为与交配方式的关系进行观察及统计分析,以探讨其花粉逐步呈现的适应性及其在提高雌雄繁殖过程中的意义。结果表明:(1)宁夏枸杞的单花寿命为(4.07±0.15) d,而雄性持续时间(0.07±0.01 d)比雌性持续时间短(4±0.01 d);花寿命内雌蕊长度比雄蕊长,属柱头探出式异位类型;花粉呈现式样为不完全逐步呈现。(2)花寿命不同阶段花回馈间存在显著差异。(3)意大利蜜蜂(Apis mellifera)、熊蜂(Bombus sp.)、食蚜蝇(Syrphidae sp.)是宁夏枸杞在自然居群的主要访花者,其中熊蜂和食蚜蝇是主要传粉者,但传粉者效率低,属于高移出低沉积类型。(4)宁夏枸杞的花粉胚珠比(2 448.11±448.32)及授粉实验结果均表现出兼性异交特征;自发自交及人工自花授粉花的座果率及结籽率很低,属于自交不亲和类型;自然传粉花的座果率及结籽率比人工去雄异花低,存在其较高的花粉限制(40.71%)。研究发现,宁夏枸杞花部综合征表现出雌雄异位和花粉不完全逐步呈现式样,这是避免雌雄功能及雄蕊各花药间的干扰、减少花粉同步移出及保障其雄性适合度的有效途径,但自然居群的传粉者种类限制、传粉效率低以及自交不亲性是导致宁夏枸杞花粉限制及降低雌性繁殖成功的主要原因。     

High specialisation in the pollination system of Mandevilla tenuifolia (J.C. Mikan) Woodson (Apocynaceae) drives the effectiveness of butterflies as pollinators

Butterfly pollination in the tropics is considered somewhat effective or solely effective in a few plant species. In the present study, we tested the hypothesis that Mandevilla tenuifolia (Apocynaceae), which has floral attributes associated with psychophily, has strategies adapted to pollination by butterflies, restricting other floral visitors and making these insects act as efficient pollinators. We analysed the floral and reproductive biology of M. tenuifolia, as well as the frequency and efficiency of its flower visitors. M. tenuifolia is an herb whose flowers have strong herkogamy and secondary pollen presentation on the style head, which corresponds to 60.4% of pollen on the anthers. Flower longevity and the long period of receptivity of the stigmatic region associated with the large amount of pollen removed in the first visits suggest that flowers remain functionally female during part of anthesis. Butterflies, mainly of the families Nymphalidae and Pieridae, are the only pollinators of M. tenuifolia. Despite being self‐compatible, M. tenuifolia depends on biotic vectors for fruit production. A non‐significant difference in fruit set between controlled treatments and natural conditions suggests that the pollinators are efficient. The inclination resulting from the landing of butterflies on flowers, together with flower morphology, guiding the insect proboscis inside the floral tube, as well as the frequency and efficiency of butterfly visits, are evidence of the close relationship between butterflies and M. tenuifolia, and also of the efficiency of these insects as pollinators.     

Influence of reproductive traits on pollination success in two <Emphasis Type="Italic">Daphne</Emphasis> species (Thymelaeaceae)

Taxonomically related species can differ in a number of reproductive traits, which may translate into a differential mating system and pollination success. Here we compare two hermaphroditic insect-pollinated Daphne species (D. rodriguezii and D. gnidium) which differ in distribution (island endemic vs. mediterranean) and floral traits (long- vs. short-tube corolla). We investigated their mating system and pollen limitation by means of hand-pollination experiments and quantified the diversity and abundance of flower visitors by direct observations. Plant size and five reproductive traits (flower production, proportion of viable anthers, pollen production, flower tube length and tepal area) were studied to assess how they contribute to reproductive success, measured as proportion of pollen grains germinated per stigma and fruit set. Selfing was very low and pollen limitation existed in both species, though was higher in D. rodriguezii probably due to the scarcity of flower visitors. The low fruit set in both species suggests that most of the pollen grains found on stigmas are self-pollen. Pollinators appeared to favour some floral traits (specifically, flower tube length or tepal area) in both species, although flower crop in D. rodriguezii was the only reproductive trait influencing fruit set. In both species, the highest variability in reproductive traits and pollination success was within individuals. Our findings suggest that despite both species showed similar mating system, dependency on outcrossing pollen and selection of floral traits, pollen limitation was higher in D. rodriguezii, probably as a higher proportion of self-pollen arrives to its stigmas.     

Many to flower, few to fruit: the reproductive biology of Hamamelis virginiana (Hamamelidaceae)

Hamamelis virginiana flowers from late September to late November. In 1977, we began studying the reproductive biology of this eastern North American arborescent shrub by examining floral phenology and rewards, pollen-ovule ratios, breeding system, pollination, pollinator and resource limitation, and seed dispersal. The homogamous, self-incompatible flowers emit a faint odor, bear nectar with sucrose ratios typical of bee- and fly-pollinated flowers, and produce abundant sticky pollen. Flowers were visited infrequently by insects representing six orders. Flies were the most common floral visitors, specifically members of the genus Bradysia, but small bees also carried high percentages of Hamamelis pollen. Despite high pollen/ovule ratios (11?445 grains/ovule), bees and flies are likely pollinators, as experiments indicate wind pollination is less likely. Pollen quantity and resource availability did not appear to limit reproductive output, but pollen quality did. Tests of >40?000 flowers showed natural fruit set to be <1%. The flowering time, breeding system, and clumped distribution of plants, likely due in part to limited seed dispersal, combine to yield this remarkably low fruit set. Because all other species of Hamamelis flower from late winter to early summer, it may be that H. virginiana evolved a fall flowering phenology to avoid competition for pollinators with the closely related H. vernalis.     

The bees don't know and the flowers don't care: the effect of heterospecific pollen on reproduction in co‐occurring Thysanotus species (Asparagaceae) with similar flowers

Species integrity relies on the maintenance of reproductive isolation, particularly between closely related species. Further, it has been hypothesized that the presence of heterospecific pollen on flower stigmas adversely affects plant reproduction with increasing effect in closely related species. Using two pairs of co‐occurring buzz‐pollinated Thysanotus spp. in the Mediterranean climate region of Perth, Western Australia, we quantified the effect of heterospecific pollen on fruit and seed set. We first determined the mating systems of the two focal species using self‐ and outcross pollen, followed by separate treatments with heterospecific pollen within each species pair. Of the two species receiving pollen, Thysanotus triandrus had a mixed mating system, but with significantly lower fruit and seed set from self‐pollen relative to outcross pollen. Thysanotus tenellus was autogamous with no difference in fruit or seed set between autogamous, self‐ or outcross pollinations. Heterospecific pollen had no effect on fruit or seed set of either focal species. These outcomes point to post‐pollination reproductive isolation, consistent with a floral morphology that causes low specificity of pollen placement and thus a poor capacity for pre‐pollination discrimination. Negative effects of heterospecific pollen, therefore, do not appear to play a role in the reproduction in this group of buzz‐pollinated flowers.     

Pollen–ovule ratios are strongly correlated with floral reciprocity,in addition to sexual system,in Mussaenda (Rubiaceae)

Heterostyly is a floral polymorphism that increase inter‐morph pollen transfer and promote disassortative mating. Breakdown of heterostyly has happened many times, either leading to dioecy or monomorphism. Mussaenda is a genus with diverse sexual systems including distyly, dioecy, floral monomorphism, and homostyly, making it an ideal system to study the evolution of floral traits and their relationship with sexual system shifts. Here, floral traits and pollen–ovule (P/O) ratios were estimated and used to test hypotheses about the relationships among P/O ratios, floral trait evolution, and sexual system shifts. Our results revealed that there is no significant difference of reciprocity in upper level sexual organs between species with different sexual systems. The reciprocity indices of the lower level sexual organs, however, are smaller in species with functional dioecy than in those with distyly. P/O ratios in dimorphic Mussaenda species were relatively lower than in monomorphic outcrossing species, but did not differ significantly between species with distyly and functional dioecy. Populational P/O ratios were negatively correlated with reciprocity index in the dimorphic species. We suggest that the loss of function in lower level organs in species with functional dioecy has resulted from less strict reciprocity. The relationship between P/O ratios and reciprocity indices strongly support that efficient disassortative pollination of distylus flowers may have promoted evolution towards low P/O ratios.     

Pollination biology and reproductive success in four Brazilian species of Gomesa (Orchidaceae: Oncidiinae): Specific pollinators,but high pollen loss and low fruit set

Four species of Gomesa (Gomesa flexuosa, Gomesa ranifera, Gomesa cornigera, and Gomesa riograndensis), a horticulturally important, albeit scarcely studied orchid genus, were investigated. Pollination biology was studied through fieldwork in Rio Grande do Sul in Southern Brazil during two flowering seasons. Breeding systems were studied through controlled pollinations in cultivated plants. All studied species secrete floral oils through elaiophores and are exclusively pollinated by native, oil-collecting bees of the Centridini or Tapinotaspidini tribes. Pollinarium reconfiguration after withdrawal, a feature likely promoting cross-pollination, was recorded in G. cornigera and G. riograndensis. All studied species are pollinator-dependent and either predominantly or fully self-incompatible, that is, unable to set fruit following self-pollination. The used indexes of pollination efficiency consistently scored values below 1, indicating that in all studied species less than one flower is pollinated per pollinarium removed. Accordingly, pollen loss was high, ranging from 40% to 75%. Furthermore, inflorescences in natural populations displayed low fruit sets (below 13%). Gomesa flexuosa presented the highest values of pollinarium removal (36.24%) and deposition (15.34%). Significantly, this is the only studied species bearing a tabula infraestigmatica, a column swelling that bees hold to stabilize themselves while gathering the floral oils. Since pollinators were observed in all studied species spending considerable time (up to 3 min 46 s) and visiting several (up to 19) flowers, we propose that the observed low fruit set is the result of a combination of self-incompatibility, pollinator-mediated self-pollination, and pollen loss.     

Reproductive biology of Alpinia blepharocalyx (Zingiberaceae): another example of flexistyly

The floral biology and breeding system of Alpinia blepharocalyx were studied in Yunnan province, southwest China. Our results indicate that populations of A. blepharocalyx have two floral morphs, which differ in flowering behaviour: the cataflexistylous morph in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes curved under the anther at afternoon, and the anaflexistylous morph in which the receptive stigma is curved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen in the afternoon; the flowering period of both floral morphs is 12 hours; the stylar movements in the two floral morphs are synchronous, and they have similar traits to those found in other Alpinia species previously reported. Also, on average, cataflexistylous flowers are larger than anaflexistylous flowers, especially the labellum and corolla tube length; moreover, the P/O ratio of the two floral morphs is significantly different: the cataflexistylous morph has more pollen grains and fewer ovules than the anaflexistylous morph; the evolutionary significance of this comparison is discussed. Cataflexistylous flowers were observed to produce a lower concentration of nectar than that of anaflexistylous flowers before 11:00 a.m., but they then have higher nectar concentration; cataflexistylous flowers always have more nectar than anaflexistylous flowers. Flowers of A. blepharocalyx were visited by fewer insects at a lower frequency, including honey bees (Apis cerana cerana) and two species of carpenter bees (Xylocopa spp.). No significant differences were found in the fruit set among the hand-pollination treatments regardless of selfing (geitonogamy and autogamy) or crossing (intermorph and intramorph); but all hand-pollination treatments exhibited much higher fruit set than the controls; meanwhile, no fruit set occurred in the unpollinated bagged plants. Pollen tube growth was examined using fluorescence microscopy following self, intramorph, and intermorph hand pollinations at different times of the day; the pollen tubes of both floral morphs have the same growth rate and the capability to reach the ovary. Both the results of pollen tube growth and fruit sets under different pollination conditions show that A. blepharocalyx is self-compatible and dependent upon insects for pollination. The distinctive flexistyly mechanism of A. blepharocalyx is likely a floral mechanism that promotes effective intermorph pollen transfer.     

Reproductive ecology of Ocotea catharinensis,an endangered tree species

• Ocotea catharinensis (Lauraceae) is an endangered tree species from the Brazilian Atlantic Rainforest. Currently, little is known about the reproductive ecology of this species. Aiming to propose conservation measures, we described aspects related to phenology, floral biology, pollination, seed dispersal and mating system of O. catharinensis.
• We conducted phenological observations in 62 individuals for 2 years. In one reproductive event, we evaluated nectar production, stigmatic receptivity and pollen viability. Floral visitors were observed, identified and classified on a scale of pollination effectiveness. Seed dispersers were observed and identified using camera traps. Finally, the mating system was evaluated through pollen/ovule ratios, experimental pollination treatments and genetic analysis with molecular markers.
• Ocotea catharinensis presented a supra‐annual fruiting pattern with a substantial reduction of reproducing individuals from bud phase to ripe fruit phase. Several mechanisms prompting cross‐fertilisation were identified, such as attractive, herkogamic and protogynic flowers. The main floral visitors and pollinators were from the Diptera order, and all seed dispersers were birds. The species presented a predominantly outcrossed mixed mating system with significant selfing rate (17.3%).
• Although based on restricted evidence, we hypothesised that selfing is an escape mechanism for situations unfavourable to cross‐fertilisation. Specifically, for the studied population selfing is a response to reduced population size, which is caused by the non‐reproduction of all potentially reproductive individuals and by past exploitation events. Therefore, conservation efforts should be able to enhance population sizes, as well as prevent overexploitation.

     

Breeding systems in Balsaminaceae in relation to pollen/ovule ratio,pollination syndromes,life history and climate zone

• Pollen/ovule (P/O) ratios are often used as proxy for breeding systems. Here, we investigate the relations between breeding systems and P/O ratios, pollination syndromes, life history and climate zone in Balsaminaceae.
• We conducted controlled breeding system experiments (autonomous and active self‐pollination and outcrossing tests) for 65 Balsaminaceae species, analysed pollen grain and ovule numbers and evaluated the results in combination with data on pollination syndrome, life history and climate zone on a phylogenetic basis.
• Based on fruit set, we assigned three breeding systems: autogamy, self‐compatibility and self‐incompatibility. Self‐pollination led to lower fruit set than outcrossing. We neither found significant P/O differences between breeding systems nor between pollination syndromes. However, the numbers of pollen grains and ovules per flower were significantly lower in autogamous species, but pollen grain and ovule numbers did not differ between most pollination syndromes. Finally, we found no relation between breeding system and climate zone, but a relation between climate zone and life history.
• In Balsaminaceae reproductive traits can change under resource or pollinator limitation, leading to the evolution of autogamy, but are evolutionary rather constant and not under strong selection pressure by pollinator guild and geographic range changes. Colonisation of temperate regions, however, is correlated with transitions towards annual life history. Pollen/ovule‐ratios, commonly accepted as good indicators of breeding system, have a low predictive value in Balsaminaceae. In the absence of experimental data on breeding system, additional floral traits (overall pollen grain and ovule number, traits of floral morphology) may be used as proxies.

     

Contrasting breeding systems in six species of Tillandsia L. (Bromeliaceae) from woody areas of Santa Fe Province: Argentina

The reproductive biology of six native Tillandsia species that co-occurred in woody areas of Santa Fe Province (Argentina) was studied for floral biology, breeding systems and reproductive success. Flower longevity, floral phases, hand self- and cross-pollinations or natural pollinations and fluorescence microscopy observations of post-pollination events were analysed. Also pollen/ovule (P/O) ratio and pre-emergent reproductive success (PERS) were compared between species. In T. aëranthos, T. ixioides and T. meridionalis, self-pollinated flowers mostly failed to form fruits, and fluorescence microscopy studies showed that in selfed pistils pollen tube growth was arrested in the style. It is likely that these species have homomorphic, gametophytic self-incompatibility. In contrast, T. capillaris, T. tricholepis and T. recurvata regularly formed fruits following autonomous self-pollination. Floral traits, the P/O ratios and PERS indices correlate with the breeding systems found in these two groups of species: the three selfing species have inconspicuous closed flowers, and they have low P/O ratios and high PERS values. In contrast, the three self-incompatible (SI) species have conspicuous flowers with relatively high P/O and low PERS values. We described here three new cases of SI in Tillandsia species subgenera Anoplophytum and confirm the autogamous and cleistogamous status in three species of Tillandsia subgenera Diaphoranthema.     

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

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

A preliminary study on pollination biology of Omphalogramma souliei Franch. (Primulaceae), a species endemic to China

The pollination biology of Omphalogramma souliei Franch., a species endemic to southwestern China, was investigated. Floral phenology, flower visitors, pollen/ovule ratio, attractants and rewards to the visitors were observed, measured, and recorded. Bagging experiments to exclude pollinators were carried out in the wild for two years. Our results revealed five important aspects of the reproductive biology of Omphalogramma souliei. 1) The pollen-ovule ratio was 1748±233. The breeding system was self-compatible, with facultative xenogamy. 2) The pollination syndrome is entomophily, and this species could not be pollinated by wind if the pollinators were unavailable. 3) Six insect species were observed visiting the flowers of Omphalogramma souliei in the wild, of them, three species of hymenoptera, Lasioglossum sp., Heriades parvula Cockerell and Micrapis florae Fabricius, are the principal floral visitors and effective pollinators. 4) The visual attractants to the visitors are floral color and shape, the large yellow anthers, and the rewards for visitors are pollen and nectar. 5) Cleistogamy may also occur, since the anthers of some flowers dehisced before opening of the corolla. The results of floral biology and pollination characters suggest that xenogamy predominated and autogamy played an assistant role in the evolution of reproduction and breeding system of O. souliei.     

Reproductive biology of Pachira aquatica Aubl. (Malvaceae: Bombacoideae): a tropical tree pollinated by bats,sphingid moths and honey bees

We investigated the reproductive biology, including the floral biology, pollination biology, breeding system and reproductive success, of Pachira aquatica, a native and dominant tropical tree of fresh water wetlands, throughout the coastal plain of the Gulf of Mexico. The flowers present nocturnal anthesis, copious nectar production and sugar concentration (range 18–23%) suitable for nocturnal visitors such as bats and sphingid moths. The main nocturnal visitors were bats and sphingid moths while bees were the main diurnal visitors. There were no differences in legitimate visitation rates among bats, moths and honey bees. Bats and honey bees fed mainly on pollen while moths fed on nectar, suggesting resource partitioning. Eight species of bats carried pollen but Leptonycteris yerbabuenae is probably the most effective pollinator due to its higher pollen loads. The sphingid moths Manduca rustica, Cocytius duponchel and Eumorpha satellitia were recorded visiting flowers. Hand pollination experiments indicated a predominant outcrossing breeding system. Open pollination experiments resulted in a null fruit set, indicating pollen limitation; however, mean reproductive success, according to a seasonal census, was 17 ± 3%; these contrasting results could be explained by the seasonal availability of pollinators. We conclude that P. aquatica is an outcrossing species with a pollination system originally specialized for bats and sphingid moths, which could be driven to a multimodal pollination system due to the introduction of honey bees to tropical America.     

THRIPS-MEDIATED SELF-POLLINATION OF TWO FACULTATIVELY XENOGAMOUS WETLAND SPECIES

Thrips and/or aphids played an important role in the self-pollination of two facultatively xenogamous herbs that inhabit wetlands in northwestern Iowa. In both Ranunculus sceleratus and Potentilla rivalis the fruit set and/or successful pollination of plants that were sprayed with malathion to kill thrips and aphids and caged to exclude typical flower visitors was substantially and significantly lower than that of open-pollinated and caged plants. We observed pollenbearing thrips and aphids on the flowers. The high fruit sets of emasculated flowers of R. sceleratus showed that insects moved pollen between flowers, and our observations of bees collecting pollen and moving between plants suggest that cross-pollination can occur in both species.     

Enantiostyly in Chamaecrista ramosa (Fabaceae‐Caesalpinioideae): floral morphology,pollen transfer dynamics and breeding system

Enantiostyly is a form of reciprocal herkogamy, in which floral morphs present reciprocal differences in the position of sexual elements, and occurs in monomorphic and dimorphic forms. This polymorphism maximises cross‐pollination and reduces self‐pollination, being very common within the subtribe Cassiinae (Fabaceae). Nevertheless, few studies have investigated the functionality of enantiostyly, particularly in this plant group. The present study aimed to investigate enantiostyly and its functionality in Chamaecrista ramosa, a monomorphic enantiostylous shrub, in an area of coastal vegetation in northeast Brazil. Pollen deposition and capture on the body of floral visitors, the relationship of these data with floral biology and breeding system, and morph ratio were evaluated. Pollen deposition and capture occurred in specific sites of the floral visitor body, showing the functionality of enantiostyly. The floral architecture, associated with the floral visitor behaviour, resulted in indirect pollen deposition on the floral visitor body. This occurred through a loop made by the pollen upon the inner petal surface, similar that generally reported for other Cassiinae. Chamaecrista ramosa is self‐compatible, although no fruit set was observed through spontaneous self‐pollination. The occurrence and number of floral morphs was similar within clumps. Enantiostyly seems to be advantageous for this species, as it results in efficient pollen capture and deposition, reduces the chances of autogamy and maximises intermorph pollen flow.