Population size,pollinator visitation and fruit production in the deceptive orchid Calypso bulbosa

It has been proposed that in non-rewarding animal-pollinated plants the pollination intensity should decrease with increasing population size and should increase with increasing local abundance of reward-producing plants. To test these hypotheses, we examined how population size, local abundance of Salix caprea, and tree cover were related to pollen removal and fruit production in 16 populations of the deceptive, early-flowering and bumblebee-pollinated orchid Calypso bulbosa in northern Sweden in 3 consecutive years. To determine whether fruit production was limited by pollinator visitation, supplemental hand-pollinations were performed in three populations in 3 years. Finally, to examine whether increased fruit production was associated with a reduction in future flower production, vegetative growth or survival, supplemental hand-pollination was repeated for 5 years in one population. The levels of pollen export, pollen deposition, and fruit set of C. bulbosa varied considerably among years and among populations. The proportion of plants exporting pollen was negatively related to population size, and positively related to density of S. caprea and to tree cover in 1 of the 3 years. In the other 2 years, no significant relationship was detected between proportion of plants exporting pollen and the latter three variables. In no year was there a significant relationship between fruit set and population size, density of S. caprea and tree cover. There was substantial among-year variation in the extent to which fruit production was limited by insufficient pollen deposition and in the amount of weather-induced damage to flowers and developing fruits. Fruit set was consistently higher in hand-pollinated than in open-pollinated plants, but this difference was statistically significant in only one of 3 years. Supplemental hand-pollination in 5 consecutive years increased cumulative fruit production 1.8 times, but did not affect flower production, plant size, or survival. Tree cover was negatively correlated with the incidence of frost damage in 1 year. The results indicate that life-time seed production may be pollen limited in C. bulbosa, and that variation in population size and local abundance of the early-flowering, nectar-producing S. caprea can only partly explain the extensive variation in pollinator visitation among populations of this species.     

Low legitimate pollen flow in distylic Turnera hermannioides (Passifloraceae) and its consequences on fruit and seed set

Turnera hermannioides is a ruderal distylic subshrub, native to NE-Brazil. In the Catimbau National Park, situated within the semi-arid Caatinga region, we studied the pollination ecology of this species, emphasizing (1) effective pollinators; (2) characteristics of short- and long-styled flowers; (3) intra- and intermorph pollen flow; and (4) fruit and seed set. Short and long-styled morphs differ in pollen size and ornamentation, stigmatic surface, style and stamen length and nectar production. The flowers are obligate intermorphic outcrossers and depend on animals for pollination. The flowers of T. hermannioides attracted insect visitors of 25 species, among them butterflies, beetles, but mainly bees. Polylectic bees, such as Apis mellifera, stingless bees, and solitary Callonychium brasiliense were the most frequent visitors and the principal pollinators. The frequency of visits, however, was very low (on average 1.9 visits/flower/day), resulting in a low and unbalanced legitimate pollen flow, i.e. a much lower number of pollen grains from short-styled flowers reaching stigmas of long-styles than vice versa, which in turn compromised fruit and seed set. Surprisingly, Protomeliturga catimbaui and P. turnerae, both oligolectic on flowers of the Turneraceae-clade of the Passifloraceae s.l., were very rare flower visitors and did not contribute significantly to the pollination of Turnera hermannioides. We discuss how the low visitation rate and the inefficiency of pollinators (i.e. inefficiency in promoting legitimate pollination) compromise pollen flow and, consequently, fruit and seed set in this species.     

Physical site characteristics limit pollination and fruit set in the dioecious hydrophilous species,Vallisneria americana

Experimental and observational studies of the submersed, freshwater macrophyte Vallisneria americana Michx. revealed that depth, wind and wave exposure, and current velocity may all influence fruit set. In this dioecious species, long-pedunculate female flowers are pollinated by free-floating male flowers at the water surface. Average fruit set in the natural populations studied varied from zero to 97% of the flowers observed. With increasing water depth in New York and Pennsylvania lakes, female plants continued to flower, though these flowers were unable to reach the surface, and consequently, did not set fruit. Fruit set was also lower in relatively open sites exposed to wind and waves, presumably because male flowers do not remain in the vicinity of female flowers long enough for effective pollination. This was particularly striking at a site with low male flower densities, but fruit set was increased to 100% at that site by confining the floating male flowers within a field enclosure. In a river, fruit set was negatively correlated with surface current velocity, and was reduced to zero in current velocities greater than 0.30 m · s^–1. Fruit set in V. americana appears to be restricted or precluded by physical environmental conditions in a variety of sites.     

Breeding systems in a temperate mediterranean-type climate montane sclerophyllous forest in central Chile

The frequency of dioecy in the predominantly biotically-pollinated native flora of a temperate montane sclerophyllous forest in central Chile, 33^oS, is determined. Experimental crosses and other tests were performed on a taxonomically diverse set of annual herbs, perennial herbs and woody species to detect genetic self-incompatibility, spontaneous selfing capacity and obligate agamospermy.
The overall frequency of dioecy in the community is 9%.Dioecy is unequally represented among life-forms, increasing in frequency with greater longevity: 0% in annual herbs; 2% in perennial herbs; 17% in shrubs; 57% in trees. Thirty-eight percent of 37 hermaphrodite species proved to be genetically self-incompatible. Self-incompatibility, like dioecy, increases in frequency with longevity: 0% in annual herbs; 50% in perennial herbs; 80% in shrubs. The self-compatible species showed a wide range of breeding habits from facultative outcrossing to strong autogamy. However, in most self-compatible species hand self-pollination increased fruit and seed set over spontaneous selfing. Obligate agamospermy was not detected among hermaphrodite specieS. Absolute and relative fecundity were strongly correlated with the degree of compatibility. However, reduced fecundity in obligate outbreeding species could not be unequivocally attributed to pollinator limitation.
For equivalent life-forms, the montane sclerophyllous forest exhibits proportionately more obligate outbreeding species than some tropical forests and other temperate communities. The strong tendency for obligate outcrossing in the Chilean community is seen as a result of interplay between life-history constraints and the intense abiotic and biotic pressures that are characteristic of the mediterranean-type climate community.     

Effect of pollination time,the hour of daytime,pollen storage temperature and duration on pollen viability,germinability, and fruit set of date palm (Phoenix dactylifera L.) cv "Deglet Nour"

Success artificial pollination with viable pollen is crucial process in the production chain of date palms. This study evaluated the impact of pollen storage temperature and duration, pollination time following spathe cracking, and the hour of daytime on pollen viability, germinability, fruit set and yield of 'Deglet Nour' date palm cultivar. In in vitro tests, fresh pollen showed the maximum viability (96.3%) and germination (85%) but it decreased thereafter upon the storage temperature (28, 4 and ?30 °C) and duration (3, 6, 9 and 12 months). In this respect, pollen stored at ?30 °C retained highest viability and germinability followed by those stored at 4 and then at 28 °C. In filed experiments, fruit set was 85, 75, 65, and 45% with pollination using fresh pollen, or pollen stored at ?30, 4 and 28 °C, respectively. Fruit set was 95%, 75%, and less than 50%, for pollination performed on the same day of spathe cracking, 6 and 12 days later, respectively. The highest fruit set percentage and yield/bunch were obtained with pollination performed between 12.0 pm and 15.0 pm in contrast to 8.0–11.0 am or 16.0–17.0.     

Quantitatively determined self-incompatibility. IV. Pollination and seed set in Borago officinalis

Summary The outcomes of sequential double pollination, mixed double pollination and single pollination are compared. Single pollination leads to lower seed set than double pollination. Systematic differences between female genotypes are shown to be possible. It is also shown that failure to set seed is generally due to pre-zygotic maternal factors.     

Low fruit set in the abundant dioecious tree Clusia hilariana (Clusiaceae) in a Brazilian restinga

Clusia hilariana (Clusiaceae), a dioecious tree, is an important nurse plant in the open, patchy restinga vegetation of coastal sandy plains in northern Rio de Janeiro (SE-Brazil). Although highly abundant locally, we found low production of fruit and viable seeds in open-pollinated as compared to hand-pollinated flowers. Reward from flowers of both sexes was resin, which was produced in higher quantities in male flowers. Flower visitors, including honeybees, were infrequent. We argue that low sexual reproductive success was due to inefficient pollen transport, and that vegetative propagation may contribute significantly to the high abundance of plants of this species.     

Effects of pollen quality and quantity on pollen limitation in Crataegus monogyna (Rosaceae) in NW Spain

Pollen limitation occurs when plants produce less fruits and/or seeds than they would with adequate pollen receipt. If the addition of cross-pollen to stigmas increases fruit/seed production, it is interpreted as an evidence of pollen limitation. Much of the limitation may be associated with the quality rather than quantity of pollen; however, most studies do not discriminate between the two, which may lead to misinterpretation of the results. We studied the effects of quality and quantity of pollen on the reproduction of a northern Spanish population of Crataegus monogyna. The treatments included self- and cross-pollination, and supplementation to open and bagged flowers. The response variables considered were number of pollen grains per stigma, pollen tubes per style, and initial and final fruit set. In the Cantabrian range, C. monogyna requires insect pollinators to set fruit and is partially self-incompatible. We found that the number of pollen tubes did not differ between cross- and self-pollination treatments; however, self-pollinated flowers set less fruits than flowers that received pure cross-pollen or were supplemented with both cross- and self-pollen. The experimental design allowed us to infer qualitative rather than quantitative pollen limitation. Comparison of the number of pollen grains and tubes, and initial and final fruit set among pollination treatments suggested post-zygotic embryo selection against selfed progeny.     

Pollen-limited seed set in pseudogamous blackberries (Rubus L. subgen. Rubus)

Summary Pseudogamous blackberry species are polyploid and usually exhibit meiotic irregularities causing severe reduction in pollen viability. When species means were compared, relative seed set was strongly correlated with the number of good pollen grains produced per flower divided by the number of ovules (good-pollen/ovule ratio).Partial correlation analysis for percentage viable pollen and relative seed set, controlling for good-pollen/ovule ratio, revealed an almost significant relationship, presumably because meiotic irregularities have a similar effect on both pollen viability and the viability of meiotically derived embryo sacs. Seed germination, on the other hand, showed no relationship with relative seed set or with pollen viability.The automatic selfing capacity exhibited by most of these blackberry species does not appear to be related to either pollen production or relative seed set.     

Effects of varying pollen load on fruit set,seed set and seedling performance in apple and pear

Two apple crosses and one pear cross were carried out using three different pollen densities. In one of the apple crosses, both fruit and seed set decreased when a mixture of pollen and powder (diluted pollen treatment) was used for pollination. Also, seedlings from the undiluted treatment were more vigorous than those originating from pollination with diluted pollen. In the other apple cross, however, no effect on fruit set, seed set or growth in the first year was found. Mean diameter after the second year differed among treatments, but did not seem to be related to pollen density. For pear, the number of good seeds decreased and the number of deaf, probably unfertilized, seeds increased with decreasing pollen density. However, there was no apparent significant effect of pollen density on seedling growth. With respect to scab resistance and mildew attack in both apple crosses, no differences among treatments were found. The results indicate that gametophytic competition may occur in apple crosses, but that it is probably weaker in pear.     

花粉和资源限制对青阳参坐果率的影响

以生长于自然种群的青阳参植株为材料进行补充授粉试验,以人工种植的青阳参植株为材料进行资源限制试验,探讨花粉和资源限制对青阳参坐果率的影响.结果表明:自花授粉、天然授粉和异株异花授粉植株的结实率分别为1.7％、2.25 ％和24％,花粉来源对青阳参坐果存在显著影响;补充施肥和不补充施肥植株的坐果率分别为5.12％和3.1...     

Sexual dimorphism and fruit production in a dioecious understory tree,Ilex opaca Ait.

Summary This study suggested that sexual selection is potentially an important factor in the maintenance of dioecy in the American holly, Ilex opaca (Aquifoliaceae). Sexual dimorphisms in flower production and phenology were highly significant in this understory tree. On average, individual males produced 7.4 times as many flowers as did female trees. Staminate flowers lasted only a single day, whereas pistillate flowers lasted 3–4 days, during which they showed no significant decline in their ability to produce fruit after pollination. Individual male trees opened their flower buds asynchronously during the season, maximizing the number of days they were in flower. Individual females opened their buds more synchronously, maximizing their floral display at one point in time. Females produced fruits in numbers that were somewhat less than proportional to their flower production. Fruit development was initiated from only 38.9% and 69.5% of pistillate flowers in 1987 and 1988, respectively. By the time of ripening, an average female had lost 62.3%, 24.3%, and 11.1% of its initial fruit crop in 1986, 1987, and 1988, respectively. The proportion of fruit lost in 1986 was independent of the number of fruit that initially began development. In 1988, artifically supplementing pollen to a large number of flowers failed to increase either fruit or seed production relative to control branches with unsupplemented flowers. This suggested that resource levels were likely more important than pollen availability in limiting female reproductive success. These observations on I. opaca were consistent with the expectations for a population in which male reproductive success continues to benefit from continued pollinator service and female reproductive success does not.     

Exploring the limiting causes of fruit production in the tropical tree Ipomoea wolcottiana Rose (Convolvulaceae)

Path analysis was used to evaluate a group of variables related to fruit production in the tropical tree Ipomoea wolcottiana (Convolvulaceae). The variables were tree size (trunk diameter), available phosphorus in the soil, floral phenotypic sex expression (i.e. gynoecium biomass allocation), flower production, fruits initiated, and length of the corolla and diameter of its opening. The path model explained 79.9% of the total variation, and the production of mature fruits was related in large part by the number of fruits initiated and by phenotypic sex expression. In turn, fruits initiated were related to corolla tube diameter and to a lesser extent to corolla tube length. The model could explain a low percentage of the variation (18.5%) of the number of fruits initiated, but once a fruit was initiated it had a high probability of maturing. Phenotypic sex expression related negatively to the number of flowers produced and positively, to almost the same degree, to available phosphorus in the soil. Tree size had a positive effect on flower production but a negligible effect on fruits initiated and matured. Soil phosphorus affected positively, although weakly all variables. The relationships between tree size, number of flowers and phenotypic sex expression suggested that abundant flowering in this species functions more for pollen donation than its reception.     

A new contribution to the Holocene vegetation history of the West African Sahel: pollen from Oursi,Burkina Faso and charcoal from three sites in northeast Nigeria

A pollen diagram from Oursi in Burkina Faso is compared with anthracological (charcoal analysis) results from three sites in northeast Nigeria (Konduga, Gajiganna, Lantewa). The present-day vegetation at all four sites is Sahelian or Sahelo-Sudanian and under heavy human impact. At Oursi, a closed grassland with only few trees and almost no Sudanian elements can be reconstructed for the middle Holocene. At the Nigerian sites, on the other hand, Sudanian woody plants were present during this period. We assume that the Sahel was not a uniform zone during the middle Holocene but rather a mosaic of different vegetation types according to local site conditions. In the light of these results, a simple model of latitudinally shifting vegetation zones is not applicable. Around 3000 B.P. the closed grassland at Oursi was opened by agro-pastoral activities, and at Gajiganna, plants characteristic of pasture lands can be directly related with the presence of cattle. Human impact seems to have been the dominant factor in the vegetation history of the Sahel from 3000 B.P. until today, masking possible effects of climatic change.     

Effects of habitat isolation on pollinator communities and seed set

Destruction and fragmentation of natural habitats is the major reason for the decreasing biodiversity in the agricultural landscape. Loss of populations may negatively affect biotic interactions and ecosystem stability. Here we tested the hypothesis that habitat fragmentation affects bee populations and thereby disrupts plant-pollinator interactions. We experimentally established small ”habitat islands” of two self-incompatible, annual crucifers on eight calcareous grasslands and in the intensively managed agricultural landscape at increasing distances (up to 1000 m) from these species-rich grasslands to measure effects of isolation on both pollinator guilds and seed set, independently from patch size and density, resource availability and genetic erosion of plant populations. Each habitat island consisted of four pots each with one plant of mustard (Sinapis arvensis) and radish (Raphanus sativus). Increasing isolation of the small habitat islands resulted in both decreased abundance and species richness of flower-visiting bees (Hymenoptera: Apoidea). Mean body size of flower-visiting wild bees was larger on isolated than on nonisolated habitat islands emphasizing the positive correlation of body size and foraging distance. Abundance of flower-visiting honeybees depended on the distance from the nearest apiary. Abundance of other flower visitors such as hover flies did not change with increasing isolation. Number of seeds per fruit and per plant decreased significantly with increasing distance from the nearest grassland for both mustard and radish. Mean seed set per plant was halved at a distance of approximately 1000 m for mustard and at 250 m for radish. In accordance with expectations, seed set per plant was positively correlated with the number of flower-visiting bees. We found no evidence for resource limitation in the case of mustard and only marginal effects for radish. We conclude that habitat connectivity is essential to maintain not only abundant and diverse bee communities, but also plant-pollinator interactions in economically important crops and endangered wild plants. Received: 7 May 1999 / Accepted: 19 July 1999     

Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres

A widely accepted paradigm for speciation in tropical forests, the refuge theory, requires periodic habitat fragmentation driven by global climatic fluctuations to provide conditions for allopatric speciation. This implies that comparative species richness in refugia is due to loss of diverse communities in areas affected by climatic cycles. In this study we compare distribution patterns of bird and plant taxa which we consider to be of either deep phylogenetic lineages or recent radiations. It is demonstrated that lowland areas which have been postulated as Pleistocene refugia are dominated by species which represent lineages of pre-Pleistocene age. Since variations in species richness within these forest tracts reflect currently apparent environmental variables which might be considered to determine carrying capacity, we do not need to postulate that richness is the result of changes in forest cover in the past. Recently diversified taxa of plants and birds are found mainly at the periphery of the main rain forest blocks and in habitat islands outside them. Here, peak concentrations of young restricted-range species are often congruent with clusters of old and biogeographically relictual species. It is suggested that this reflects special intrinsic environmental properties of these areas, in the form of long-term environmental stability caused mainly by persistent orographic rain or mist. In this case, richness is not necessarily due to extinction outside these areas. Stability not only enables survival of relictual taxa, but also promotes morphological differentiation of radiating taxa, leading to aggregates of taxa of restricted distribution.     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

Effect of pollination intensity on fruit and seed set in cacao (Theobroma cacao L.)

We studied the functional relationship between pollination intensity and fruit survival as well as the number of seeds per pod in the tropical tree Theobroma cacao L. on a Forastero Upper-Amazon clone (UPA 409) in Ivory Coast. Cutting the style 24 h after pollination allowed for counting the number of pollen grains deposited on a stigma without affecting fruit set and seed development. Forty-three pollen grains were necessary to reach 50% of maximum fruit set 28 days after pollination. Above 115 pollen grains, the proportion of developing ovaries reached a maximum of 88% 28 days after pollination and 75% at maturity. With fewer than 238 pollen grains per stigma, there was a close relationship between pollination intensity and number of seeds per pod; the pollenseed ratio increased from 1.61 to 3.81 for PI increasing from 30 to 238 pollen grains. For higher pollination intensities, the average number of seeds per pod reached a maximum of 58. The relationship between pollination intensity and seed content was modelled. Results are consistent with the hypothesis that ovules attracted pollen tubes in a similar way regardless of whether or not they had already been reached by another pollen tube.     

侧柏和北美香柏(柏科)的传粉机制(英)

观察了侧柏（Phaycladusorientalis（L．）Franco）和北美香柏（ThujaoccidentalisL．）散粉后花粉进入珠孔的过程。在散粉期,这两种植物的胚珠均分泌出传粉滴。当花粉落到传粉滴上后,引起传粉滴表面的形状发生改变或减弱胚珠的继续分泌,使得该传粉滴蒸腾加快,导致其比未授粉的传粉滴明显收缩。观察结果表明：不同植物的花粉导致侧柏传粉滴的收缩速率不尽相同。其中,与侧柏亲缘关系较近的植物花粉引起传粉滴的收缩速率和侧柏自身花粉引起的传粉滴收缩速率相似;反之,收缩速率变慢。侧柏传粉滴的收缩可能主要是由于花粉减弱胚珠分泌的结果。