Geographic variation in reproductive success of Stenocereus thurberi (Cactaceae): Effects of pollination timing and pollinator guild

? Premise of the study: It has been proposed that species of columnar cacti from dry tropical areas depend on bats for their reproduction, whereas species from dry subtropical areas are also pollinated by other species. To test this hypothesis, we examined the effects of pollinator guild and of variation in time and space on the reproductive success of a widespread species. ? Methods: Changes in fruit set, seed set, and pollinator activity through time were recorded in three widely separated populations of Stenocereus thurberi. Breeding system and sources of pollination limitation were determined by controlled pollinator exclusions in each population. ? Key results: Significant differences were found in the timing of activity and in the effectiveness of pollinators among sites. In the northern and central populations, reproductive success depends on bats, whereas in the southern population a combination of pollinators was more effective. No difference between open and hand cross-pollination treatments was found in the northern and central populations, which suggests no pollen limitation. However, significant differences were detected in the southern population, which indicates temporal differences in pollinator abundance or arrival time. ? Conclusions: Local variation in pollinator assemblages and reproductive success could greatly affect the evolution of pollination systems. The pattern of generalist pollination in the southernmost populations and specialized pollination in the central and northern populations contradicts the hypothesis of latitudinal variation. In the absence of nocturnal pollinators, the accumulated nectar can sustain visits by diurnal pollinators, a bet-hedging strategy that increases the chances of fruit set in some populations.     

Reproduction of beetle‐pollinated Anaxagorea dolichocarpa (Annonaceae) is resilient to habitat disturbance in rainforest fragments

Strong evidence exists that fragmentation negatively affects pollination and plant reproduction, but little research has been conducted with regards to tropical trees. Specifically, effects of forest fragmentation on reproduction of plants with beetle‐pollinated flowers are poorly understood, and there are no data on the impact of fragmentation on reproduction in the structurally important tropical family Annonaceae. We examined the relationship between fragment size, pollinator abundance and seed set of beetle‐pollinated Anaxagorea dolichocarpa (Annonaceae) in a disturbed Brazilian Atlantic rainforest. Flower and fruit production and abundance of pollinators were quantified over ten months in three large (306–388 ha) and three small (6–14 ha) forest fragments. We recorded per flower pollinator abundance, resulting fruit set (fruits per flower) and seed set (monocarps per fruit) for a total of 209 individually marked flowers, and compared pollinator abundance in 186 flowers across all fragments. Flower and fruit production differed among fragments, but were similar for the combined large and small fragments. Between 64.8% (large fragments) and 66.3% (small fragments) of flowers received at least one pollinator. We found no significant difference in pollinator numbers between large and small fragments, and no correlation between pollinator abundance and fruit and seed set. A single visitor had a high probability of pollinating a flower. We conclude that 1) fragment size had no influence on pollinator number and plant reproductive success, and 2) generalist behavior of the pollinating beetles mitigate the risk of pollination failure for the reproductively specialized plant. However, further research may yet reveal genetic impoverishment of populations in small fragments due to restricted pollinator movements.     

Flower herbivory and seed predation in Silene vulgaris (Caryophyllaceae): effects of pollination and phenology

Pollination and seed predation were studied in Silene vulgaris populations during two seasons, one with much lower pollinator abundance than the other. Among the pollinators, noctuid moths of the genus Hadena also acted as seed predators. Nectar-foraging female moths oviposited in flowers, and their larvae consumed flowers and seed capsules.
Despite a lower percentage of pollinated flowers in the year of low pollinator abundance, similar numbers of flowers set fruit in both years, because fewer flower buds and flowers were eaten by Hadena larvae during the year of low pollinator visitation. The number of seed capsules preyed upon was also lower in the year with low pollinator abundance, resulting in a higher seed set. The positive correlation between the percentage of pollinated flowers and the percentage of seed capsules destroyed was also observed when comparing flowers opening in different parts of the season.
Early flowering plant individuals had the same pollination success but suffered higher seed predation than late flowering ones. Selection for maximized pollination success through synchronous flowering, is probably the main reason for the compressed flowering period in 5. vulgaris , but the high level of predation early in the season may further increase the reproductive success of synchronous flowering individuals.     

Pollinator‐mediated interactions between cultivated papaya and co‐flowering plant species

Many modern crop varieties rely on animal pollination to set fruit and seeds. Intensive crop plantations usually do not provide suitable habitats for pollinators so crop yield may depend on the surrounding vegetation to maintain pollination services. However, little is known about the effect of pollinator‐mediated interactions among co‐flowering plants on crop yield or the underlying mechanisms. Plant reproductive success is complex, involving several pre‐ and post‐pollination events; however, the current literature has mainly focused on pre‐pollination events in natural plant communities. We assessed pollinator sharing and the contribution to pollinator diet in a community of wild and cultivated plants that co‐flower with a focal papaya plantation. In addition, we assessed heterospecific pollen transfer to the stigmatic loads of papaya and its effect on fruit and seed production. We found that papaya shared at least one pollinator species with the majority of the co‐flowering plants. Despite this, heterospecific pollen transfer in cultivated papaya was low in open‐pollinated flowers. Hand‐pollination experiments suggest that heterospecific pollen transfer has no negative effect on fruit production or weight, but does reduce seed production. These results suggest that co‐flowering plants offer valuable floral resources to pollinators that are shared with cultivated papaya with little or no cost in terms of heterospecific pollen transfer. Although HP reduced seed production, a reduced number of seeds per se are not negative, given that from an agronomic perspective the number of seeds does not affect the monetary value of the papaya fruit.     

Patterns of pollinator-generated fruit set inOrchis morio (Orchidaceae)

Reproductive success of a deceptive species,Orchis morio, was investigated in relation to floral display, vertical position of fruits and dispersion pattern. A caging experiment confirmed the necessity of the presence of insect vectors for fruit development. In two Czech populations studied in 1997, insect-pollinated plants had on average 27.3 and 38% fruit set, compared to 96.7% for flowers pollinated by hand. Floral display (number of flowers per spike) was expected to positively affect the probability of pollinator visit. However, fruit set (number of fruits per spike) did not increase linearly with increasing floral display, but rose sigmoidally and approached an asymptotic value. Floral display enhanced reproductive success (fruit/flower ratio) up to a certain number of flowers (11–14), above which the relationship started to decrease and only increased the probability of production of at least one fruit. We also studied the effect of flower position within an inflorescence on the probability of being pollinated and we counted the number of seeds per capsule. A rapid decline in fruit set from the bottom to the top of the inflorescence observed probably reflects the behaviour of pollinators. The number of seeds per one capsule varied from 550 to 12 270. In addition, individuals growing in large clumps or isolated had lower reproductive success than those growing in small clumps. This result implies the existence of an intraspecific competition for pollinators and on the other hand inability of isolated individuals to attract a large number of pollinators.     

Enhancement of fruit quality in Capsicum annum through pollination by Hypotrigona gribodoi in Kakamega,Western Kenya

A study was carried out in Kakamega forest, in the western region of Kenya, to evaluate the effectiveness of the stingless bee Hypotrigona gribodoi (Magretti, 1884) on the pollination of green pepper. Three treatments were applied and consisted of self‐pollination, pollination by feral pollinators in the open field and pollination by H. gribodoi in a net cage. The differences in fruit yield and seed quality were compared among treatments. Flowers pollinated by H. gribodoi produced the heaviest fruits with the highest seed numbers followed by feral pollinators and lastly self‐pollinated flowers. Moreover, seeds were significantly bigger in size in fruits resulting from flowers pollinated by H. gribodoi compared to fruits obtained from self‐pollinated flowers or flowers pollinated by feral insects. We, therefore, conclude that H. gribodoi is an efficient pollinator of green pepper in the tropical region of East Africa.     

Nectar robbing improves male reproductive success of the endangered Aconitum napellus ssp. lusitanicum

Nectar robbery is usually thought to impact negatively on the reproductive success of plants, but also neutral or even positive effects have been reported. Very few studies have investigated the effects of nectar robbing on the behaviour of legitimate pollinators so far. Such behavioural changes may lead to the reduction of geitonogamy or to increased pollen movement. We simulated nectar robbing in experimental sites as well as in natural populations of Aconitum napellus ssp. lusitanicum, a rare plant pollinated by long-tongued bumblebees. In an experimental setup, we removed the nectaries of 40 % of the flowers, which is similar to rates of robbing observed in wild populations. Patches of plants with experimentally robbed flowers were compared with control patches containing plants with untreated flowers. We observed pollinator behaviour, mimicked male reproductive success (pollen dispersal) using fluorescent dye, and measured female reproductive success (seed set). The main legitimate visitors were bumblebees while honeybees were often observed robbing nectar. They did so by “base working”, i.e. sliding between tepals. Bumblebees tended to visit fewer flowers per plant and spent less time per single flower when these had been experimentally robbed. This change in behaviour consequently increased the proportion of flowers visited by bumblebees in patches with robbed flowers. Fluorescent dye mimicking pollen flow was dispersed larger distances after pollinators had visited patches with robbed flowers compared to control patches. Average seed set per plant was not affected by nectar robbing. Our results demonstrated that A. napellus does not suffer from nectar robbery but may rather benefit via improved pollen dispersal and thus, male reproductive success. Knowledge on such combined effects of behavioural changes of pollinators due to nectar robbery is important to understand the evolutionary significance of exploiters of such mutualistic relationships between plants and their pollinators.     

Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation

 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002     

新疆产药用植物黑果枸杞有性生殖产出差异的繁殖生态学研究

由于黑果枸杞（Lycium ruthenicum）果实具有重要的保健价值,近年来倍受关注,但其野生资源正遭受着前所未有的采伐压力。目前关于影响黑果枸杞果实产量的有性生殖过程研究较少,本实验对新疆库尔勒地区6个不同生境的野生黑果枸杞居群开展了繁殖生态学观察,并对各居群的访花频率、访花昆虫种类、柱头花粉落置、胚珠数、座果率、结实率和单株果实产量等指标进行了比较分析。结果表明,访花频率与结实率、座果率具有显著正相关关系,说明传粉者是限制黑果枸杞有性生殖的关键因素之一;传粉者种类越丰富的居群其有性生殖产出越高,揭示了传粉昆虫的多样性和丰富度是影响黑果枸杞繁殖成功的重要限制因子。此外,和静县华侨农场和焉耆县霍拉山河沟的黑果枸杞居群生殖产出明显与其传粉成功率相偏离,进一步分析表明,风沙侵蚀和人为采伐对植株的破坏是影响这两个居群果实产量的重要原因。本研究结果对于野生黑果枸杞居群的保护及其人工规模栽培提供了重要的参考依据。     

Pollination and reproduction enhance the invasive potential of an early invader: the case of Lythrum salicaria (purple loosetrife) in South Africa

The potential of an alien plant to spread rapidly and colonize new habitat may be related to the mode of reproduction and the ability to attract pollinators. Most studies focus on widespread invasive plants, in which pollinators are rarely limiting. Here, we assess the ability of a recent invader in South Africa, the tristylous Lythrum salicaria to self-reproduce and whether this can explain the delay between introduction and spread. This study was conducted in one of the largest known populations (a total of 7 populations in South Africa) of L. salicaria in the Liesbeek river in the fynbos biome. We assessed the importance of pollinators and autonomous selfing in L. salicaria by comparing seed set between pollinator excluded and naturally pollinated flowers. Overall, 5 pollinators (4 native and 1 alien) were recorded with Cape honeybees and Africa Monarch butterflies the most prominent. Seed and fruit set were significantly higher in open pollinated flowers compared to pollinator excluded flowers. Also, seed and fruit set in pollinator excluded flowers were higher in long and medium morphs compared to short morphs. Germination was high for seeds from pollinator, but also from pollinator excluded treatments. This shows that L. salicaria in South Africa is self-compatible to some extent, but it is frequented by pollinators, significantly increasing seed production. Despite L. salicaria being tristylous, all 3 morphs are present in South Africa and with a huge seed production, this species has the potential to become a major invader of rivers and wetlands in South Africa.

     

Effects of habitat disruption on the activity of nectarivorous bats (Chiroptera: Phyllostomidae) in a dry tropical forest: implications for the reproductive success of the neotropical tree Ceiba grandiflora

In the tropical dry forest of the central Pacific coast of Mexico the pollination and reproductive success of the bombacaceous tree Ceiba grandiflora was negatively affected by habitat disruption. Two of the three bat species that function as effective pollinators for this species ( Glossophaga soricina and Musonycteris harrisoni) visited flowers found in trees in disturbed habitats significantly less than trees found in undisturbed habitats. A similar pattern was observed for the effective bat pollinator, Leptonycteris curasoae; however the difference was not significant. The three nectarivorous bats that functioned as effective pollinators of C. grandiflora also visited flowers to exclusively feed on pollen by biting or pulling off an anther (see Fig. S1 of Electronic Supplementary Material). The number of pollen grains deposited on stigmas from flowers in undisturbed areas was significantly greater than from flowers in disturbed habitats. The greater visitation rate and the greater number of pollen grains deposited on flowers from trees in undisturbed forest resulted in a significantly greater fruit set for trees in these areas. Our study demonstrates the negative effect that habitat disruption has on bat pollinators in tropical dry forest ecosystems and documents the negative consequences for the plants they pollinate.     

Pollinator activity and pollination success of Medicago sativa L. in a natural and a managed population

Medicago sativa L. is an important cash crop in the arid region of northwest China. Pollinator activity is an essential aspect of pollination success, but the relationships between pollinator visitation rate and seed set still need further study of M. sativa. We investigated the following characteristics of M. sativa in natural and managed populations: floral traits, pollinator activity, and breeding system. Our results indicated the management could affect the number of flowers produced; however, there was no detectable effect on the seed set per flower. We found the percentage of seeds among pollinated flowers in the managed population was significantly higher than that in the natural population. Moreover, the increase in the proportion of pollinated flowers could significantly increase seed set per flower, and pollinator visitation rate was the important limiting factor for seed set in both populations. Andrena lebedevi Popov was found to be the most frequent pollinator in both populations. Outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. Our study suggested that proper management (artificial selection) could promote pollination success of M. sativa.     

Variation in sex ratio, morph-specific reproductive ecology and an experimental test of frequency-dependence in the gynodioecious Kallstroemia grandiflora (Zygophyllaceae)

An enduring puzzle in gynodioecious species is the great variation in female frequency seen among populations. We quantified sex ratio in 44 populations of gynodioecious Kallstroemia grandiflora. Then, we measured pollinator visitation, pollen deposition, autonomous selfing rate and pollen limitation of females. Finally, using experimental populations, we tested whether female fitness responds to the frequency of female plants. We found broad variability in sex ratio among populations (0-44% female). Hermaphrodite flowers received more pollinator visits and pollen grains than females, and bagged hermaphrodite flowers produced fruits. However, we found no evidence of pollen limitation in females. In experimental populations, female plants showed no evidence of frequency-dependent pollinator visitation, fruit set, seed set or total seed mass. These results do not support frequency-dependent variation in fitness as a major mechanism affecting female frequencies in K. grandiflora. Within the context of this study, pollinators are abundant and pollinator movement appears to operate at a large enough scale to overcome the potential reproductive disadvantages of producing solely female flowers.     

Effects of local density on pollination and reproduction in Delphinium nuttallianum and Aconitum columbianum (Ranunculaceae)

Plant populations vary in density both naturally and as a consequence of anthropogenic impacts. Density in turn can influence pollination by animals. For example, plants in dense populations might enjoy more frequent visitation if pollinators forage most efficiently in such populations. We explored effects of plant density on pollination and seed set in the larkspur Delphinium nuttallianum and monkshood Aconitum columbianum. At our site in the Colorado Rocky Mountains, flowers of D. nuttallianum are pollinated primarily by queen bumble bees, solitary bees, and hummingbirds, whereas those of A. columbianum are pollinated primarily by queen and worker bumble bees. We found that the quantity of pollination service to both species (pollinator visitation rate and pollen deposition) was at best weakly related to density. In contrast, seed set declined by approximately one-third in sparse populations relative to nearby dense populations. This decline may stem from the receipt of low-quality pollen, for example, inbred pollen. Alternatively, sparsity may indicate poor environmental conditions that lower seed set for reasons unrelated to pollination. Our results demonstrate the value of simultaneously exploring pollinator behavior, pollen receipt, and seed set in attempting to understand how the population context influences plant reproductive success.     

Habitat dependent pollination success in a Cape orchid

Summary We examined the pollination success of Disa uniflora (Orchidaceae) in two habitats. Plants occurring in a rocky gorge were far more successful than plants occurring in an adjacent open valley. More than 55% of flowers in the gorge were pollinated and set fruit compared with less than 25% of flowers in the valley. These differences are explained by the preference of Meneris tulbaghia (Satyridae), the exclusive pollinator of the orchid, for rocky, sheltered habitats. Fruit set of hand-pollinated flowers did not differ significantly between the two habitats, indicating that resources did not account for the variation in fruiting success.     

Genetic rescue by distant trees mitigates qualitative pollen limitation imposed by fine‐scale spatial genetic structure

Restricted seed dispersal frequently leads to fine‐scale spatial genetic structure (i.e., FSGS) within plant populations. Depending on its spatial extent and the mobility of pollinators, this inflated kinship at the immediate neighbourhood can critically impoverish pollen quality. Despite the common occurrence of positive FSGS within plant populations, our knowledge regarding the role of long‐distance pollination preventing reproductive failure is still limited. Using microsatellite markers, we examined the existence of positive FSGS in two low‐density populations of the tree Pyrus bourgaeana. We also designed controlled crosses among trees differing in their kinship to investigate the effects of increased local kinship on plant reproduction. We used six pollination treatments and fully monitored fruit production, fruit and seed weight, proportion of mature seeds per fruit, and seed germination. Our results revealed positive FSGS in both study populations and lower fruit initiation in flowers pollinated with pollen from highly‐genetically related individuals within the neighbourhood, with this trend intensifying as the fruit development progressed. Besides, open‐pollinated flowers exhibited lower performance compared to those pollinated by distant pollen donors, suggesting intense qualitative pollen limitation in natural populations. We found positive fine‐scale spatial genetic structure is translated into impoverished pollen quality from nearby pollen donors which negatively impacts the reproductive success of trees in low‐density populations. Under this scenario of intrapopulation genetic rescue by distant pollen donors, the relevance of highly‐mobile pollinators for connecting spatially and genetically distant patches of trees may be crucial to safeguarding population recruitment.     

Reproductive success of Dactylorhiza incarnata ssp. incarnata (Orchidaceae): the effects of population size and plant visibility

Reproduction of plants pollinated solely by flower-visiting animals depends on the ability of the population and each of its flowering member to attract pollinators. Factors affecting the pollination of nectarless species differ somewhat from those affecting the pollination of rewarding species due to the avoidance behaviour of pollinators after visiting empty flowers.
We studied a non-mimic food-deceptive orchid, Dactylorhiza incarnata ssp. incarnata, in 16 populations in central Finland to examine if population properties and plant size affected reproductive success of plants.
We found that the number of flowering plants increased total pollinia removal and seed production of the population, but had no effect on reproduction of individual plants. Dactylorhiza incarnata occurred in open mires with virtually no rewarding species in the neighbourhood, but the distance from the forest edge did not have any effect on reproduction. However, increased variation in the number of flowers among within-population plants enhanced plant reproductive success. The increased variation probably hampered the learning process of flower visitors. Plant size affected reproductive success only in populations with a high general reproductive success indicating that in the rarely visited populations, pollination of individual plants is mainly affected by random effects.     

Extended flowering in a Mediterranean shrub: Seasonal variability in seed quality and quantity

Fluctuating conditions throughout the year and changes in floral display may promote shifts in pollinator activity and predator pressure, influencing female reproductive output, especially for extended flowering species under seasonal climates. In this regard, flowering and fruit production were tracked in 2 different years in 2 populations of Ononis tridentata in Central Spain. Total fruit production was estimated, and fruits were harvested to obtain primary fruit investment, pollination success, predation incidence, seed production, seed weight and germination rates. Ononis tridentata combined spring mass flowering with a steady long flowering period across the summer and fall. The fewer flowers that are produced in fall were successfully pollinated, and produced fruits that were subject to minimal predation pressure relative to spring fruits. Moreover, fall fruits contained a higher number of heavier seeds and showed higher germination rates than those of spring seeds. Fall reproductive output represent 10% of annual viable seeds and thus may act as an important complement to the main spring reproductive investment. Extended flowering could be interpreted as a “bet-hedging strategy” for enduring Mediterranean unpredictable and changing environmental conditions.