Floral resource distribution and fitness consequences for two solitary bee species in agricultural landscapes

Floral resources are crucial for wild pollinators. Identifying the spatio-temporal floral resource use of wild pollinators and effects of resource distribution on their development might help to promote them and their pollination services to crops in agricultural landscapes.We established populations of Osmia cornuta and Osmia bicornis, two solitary wild bees, in 24 agricultural landscapes with varying floral resource availability. Based on their pollen use, we mapped the landscape-scale distribution of the visited plants, estimated pooled specific floral resource availabilities and measured its effects on reproductive output.Woody semi-natural habitats such as hedgerows provided the majority of pollen sources for both Osmia species. Pollen use differed strongly between the two species. The offspring of both Osmia increased with availability of pooled specific pollen resources. In accordance with their preferred pollen types, offspring of O. cornuta increased with increasing cover of trees and shrubs of the Rosaceae family, and that of O. bicornis with increasing cover of Papaver rhoeas, Ranunculus acris and Quercus spp. as well as with the proximity to oilseed rape. In spite of their specific responses to pollen resources, the offspring of both species decreased with the distance to forest. The floral resource availability did not significantly affect the proportion of adult females and the weight of the offspring. As forest does not appear to be a main foraging habitat for both species, the benefit of forest proximity indicates an additional role of forest in addition to food availability.Specific flowering plants and forests should thus be conserved and enhanced to maintain and support O. cornuta, O. bicornis and likely other wild bee populations in agricultural landscapes. The combined information of land cover and detailed floral resource availability gives a deeper understanding into population processes in agricultural landscapes.     

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.     

Limitations on reproductive success in endemic <Emphasis Type="Italic">Aquilegia viscosa</Emphasis> (Ranunculaceae) relative to its widespread congener <Emphasis Type="Italic">Aquilegia vulgaris</Emphasis>: the interplay of herbivory and pollination

Plant reproduction can be strongly affected by herbivory and different features of pollination ecology, such as pollinator visitation rates and capacity for self-pollination. The purpose of this study is to compare the relative impact of herbivory and pollination on maternal reproductive success in endemic Aquilegia viscosa and its widespread congener Aquilegia vulgaris. We conducted herbivore exclusion experiments in two populations of each species in 2 different years and showed that the maternal fertility of A. viscosa was significantly more limited by floral predation and pre-dispersal seed predation than its widespread congener. In the absence of herbivory, A. viscosa retained significantly lower maternal fertility than A. vulgaris. Experimental pollinations in an insect-free glasshouse showed that the two species have an equal seed/ovule ratio both in the absence of pollinators and in the presence of non-limiting outcross pollination. Pollinator visitation rates were significantly higher in populations of A. vulgaris than in populations of A. viscosa. In addition, path analyses showed that spur length, an important trait for pollinator attraction in Aquilegia, and, indirectly sepal and petal width, contribute positively to the seed/ovule ratio in A. vulgaris, but not in A. viscosa. These results indicate that maternal fertility of endemic A. viscosa is strongly reduced by flower and seed predation despite low rates of pollinator visitation, and that pollen or resource limitation in the wild may further reduce maternal fertility. Finally, floral trait variation appears to be decoupled from fertility variation in endemic A. viscosa, which possibly constrains the evolution of reproductive traits in this species.     

The reproductive biology of Stellera chamaejasme (Thymelaeaceae): A self-incompatible weed with specialized flowers

It has profound implications for species invasions and fitness, how weeds achieve reproductive success. We present the first study on the reproductive biology of Stellera chamaejasme, a toxic perennial weed which is abundant in the alpine meadows on the eastern Tibetan Plateau of China since the 1960s. S. chamaejasme has synchronogamy and herkogamous flowers, which last for about 11 days. Flowers can be yellow, white or reddish. Our results suggest that this species is self-incompatible and depends entirely on pollen vectors for seed production. All the pollinators observed were Lepidopteran species; however, the pollinator assemblage was significantly different among populations and likely independent on floral colour. Reproductive success and invasion of the weed might be a complex function of floral traits, breeding system and pollination ecology. Generally, plants with generalized pollination system are more invasive. However, our results suggest that specialized flowers and self-incompatibility in the present case may not prevent invasion of the weed.     

Effect of pollination on floral longevity and costs of delaying fertilization in the out-crossing Polygala vayredae Costa (Polygalaceae)

Background and Aims

The effect of pollination on flower life span has been widely studied, but so far little attention has been paid to the reproductive consequences of delayed pollination in plants with long floral life spans. In the present study, Polygala vayredae was used to answer the following questions. (1) How does male and female success affect the floral longevity of individual flowers? (2) How does delaying fertilization affect the female fitness of this species?

Methods

Floral longevity was studied after experimental pollinations involving male and/or female accomplishment, bagging and open pollination. The reproductive costs of a delay in the moment of fertilization were evaluated through fruit set, seed–ovule ratio and seed weight, after pollination of flowers that had been bagged for 2–18 d.

Key Results

Senescence of the flowers of P. vayredae was activated by pollen reception on the stigmatic papillae, while pollen removal had no effect on floral longevity. Nonetheless, a minimum longevity of 8 d was detected, even after successful pollination and pollen dissemination. This period may be involved with the enhancement of male accrual rates, as the female accomplishment is generally achieved after the first visit. Floral life span of open-pollinated flowers was variable and negatively correlated with pollinator visitation rates. Delayed pollination had a major impact on the reproductive success of the plant, with fruit set, seed–ovule ratio and seed weight being significantly diminished with the increase of flower age at the moment of fertilization.

Conclusions

A strong relationship between pollination and floral longevity was observed. Flowers revealed the ability to extend or reduce their longevity, within some limits, in response to the abundance of efficient pollinators (i.e. reproductive fulfilment rates). Furthermore, with scarce or unpredictable pollinators, a long floral life span could maintain the opportunity for fertilization but would also have reproductive costs on production of offspring. Reduced female fitness late in the flower''s life could shift the cost–benefit balance towards a shorter life span, partially counteracting the selection for longer floral life span potentially mediated by scarce pollination services.Key words: Delayed pollination, endemic species, flower longevity, life span, pollen limitation, pollination, pollinator scarcity, Polygala vayredae, Polygalaceae, reproductive consequences, secondary pollen presentation     

Hanging by a coastal strand: breeding system of a federally endangered morning-glory of the south-eastern Florida coast, Jacquemontia reclinata

Background and Aims

Coastal development has led to extensive habitat destruction and the near extinction of the beach clustervine, Jacquemontia reclinata (Convolvulaceae), an endangered, perennial vine endemic to dune and coastal strand communities in south-eastern Florida. We examined the breeding system of this rare species, and observed visitors to its flowers, as part of a larger effort to document its status and facilitate its recovery.

Methods

Reproductively mature experimental plants were grown from seed collected from wild plants in two of the largest remaining populations. Controlled hand pollinations on potted plants were conducted to determine the level of compatibility of the species and to investigate compatibility within and between populations. Seeds from the hand pollinations were planted in soil, and they were monitored individually, recording time to seed germination (cotyledon emergence). Wild plants were observed in several of the remaining populations to determine which species visited the flowers.

Key Results

Hand pollination and seed planting experiments indicate that J. reclinata has a mixed mating system: flowers are able to set fruit with viable seeds with self-pollen, but outcross pollen produces significantly greater fruit and seed set than self-pollen (≥50 % for crosses vs. <25 % for self-pollinations). Visitors included a wide array of insect species, primarily of the orders Diptera, Hymenoptera and Lepidoptera. All visitors captured and examined carried J. reclinata pollen, and usually several other types of pollen.

Conclusions

Remnant populations of beach clustervine will have greater reproductive success not only if floral visitor populations are maintained, but also if movement of either pollen or seed takes place between populations. Restoration efforts should include provisions for the establishment and maintenance of pollinator populations.Key words: Breeding system, conservation, beach clustervine, Jacquemontia reclinata, Convolvulaceae, endangered species, floral visitors, coastal dunes, pollination, reproductive biology, Florida, Caribbean, bees, butterflies     

So many visitors and so few pollinators: variation in insect frequency and effectiveness governs the reproductive success of an endemic milkwort

Plant–pollinator interactions are one of the most important and variable mutualisms having major implications for plant fitness. The present study evaluates the interactions between an endemic milkwort, Polygala vayredae, and its floral visitors by studying the temporal variability, foraging behaviour and effectiveness of floral visitors in three populations during three consecutive years. The flowers were visited by a diverse array of insects, totalling 24 different species. However, only four species were effective pollinators, depositing pollen on stigmas after one visit, while the remaining species behaved as nectar robbers, secondary nectar robbers or nectar thieves and were completely ineffective for pollination. Among the effective pollinators, two groups with distinct foraging behaviours were observed: the nectar collecting long-tongued bees Bombus pascuorum and Anthophora sp. and the pollen collectors Eucera longicornis and Halictus sp. No significant differences were observed among pollinators in their efficiency in pollen deposition on stigmas, but significant differences were observed in the foraging behaviour between nectarivorous and pollen collectors. Variation in the abundance and assemblage of floral visitors was observed at the temporal scale and among populations, with the effective pollinators being generally scarce. Consequently, the reproductive outcome in this species was low and significantly variable among populations and years. The results highlight the importance of studying floral visitor effectiveness when determining pollinator assemblages.     

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.     

Abnormalities in sexual development and pollinator limitation in Michelia coriacea (Magnoliaceae), a critically endangered endemic to Southeast Yunnan,China

Michelia coriacea (Magnoliacae) is a critically endangered tree, endemic to Southeast Yunnan province, China. Most of the individuals in the extant populations normally bear flowers, but fruit set and fertile seed production rates were only 6.7% and 0.2%, respectively. To identify possible causes of reproductive barriers, the following studies were carried out: examination of stamen and pistil development; experiments using controlled pollination; observations of behavior and frequency of floral visitors in both cultivated and natural populations. The results revealed that about 60% of pollen was abnormal and approx. 70% of ovules had delayed development. Hand pollination could effectively enhance the fruit set (F_4,25=35.139, P<0.0001) and seed set per fruit (F_4,25=85.022, P<0.0001). Both cultivated and wild M. coriacea had an extremely low frequency of floral visitors. Some beetles, a few species of Andrenidae and some Bombus sp. are likely to be the effective pollinators. The fruit set and seed set per fruit from controlled self-pollination and cross-pollination were significantly different (P<0.05), and thus it is inferred, that inbreeding depression may be a contributing factor in the very low seed production. It appears that low seed set in M. coriacea is due to a combination of factors: abnormalities in pollen and ovules, low number of effective pollinators, and inbreeding depression.     

Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads

Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self‐compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self‐compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.     

The ant-pollination system of Cytinus hypocistis (Cytinaceae), a Mediterranean root holoparasite

Background and Aims

The genus Cytinus is composed of rootless, stemless and leafless parasites whose flowers are only visible during the reproductive period when they arise from the host tissues. Most of the taxa occur in Madagascar and South Africa, where mammal pollination has been suggested for one species. There is only one species in the Mediterranean region, and its pollination system has been unknown. Here, a long-term field observation study is combined with experimental pollination treatments in order to assess the pollination biology and reproductive system in the Mediterranean species Cytinus hypocistis.

Methods

Field studies were carried out in six populations in southern Spain over 4 years. Temporal and spatial patterns of variation in the composition and behaviour of floral visitors were characterized. Pollen loads and pollen viability were observed, and exclusion and controlled-pollination treatments were also conducted.

Key Results

Cytinus hypocistis is a self-compatible monoecious species that relies on insects for seed production. Ants were the main visitors, accounting for 97·4 % of total floral visits, and exclusion experiments showed that they act as true pollinators. They consistently touched reproductive organs, carried large pollen loads and transported viable pollen, although the different ant species observed in the flowers differed in their pollination effectiveness. The abundance of flying visitors was surprisingly low, and only the fly Oplisa aterrima contributed to fruit production and cross-pollination.

Conclusions

Mutualistic services by ant are essential for the pollination of Cytinus hypocistis. Although this parasite does not exhibit typical features of the ‘ant-pollination syndrome’, many other characteristics indicate that it is evolving to a more specialized ant-pollination system. The striking interspecific differences in the pollination systems of Mediterranean Cytinus (ant-pollinated) and some South African Cytinus (mammal-pollinated) make this genus an excellent model to investigate the divergent evolution of pollination systems in broadly disjunct areas.Key words: Ant, breeding system, Cytinus hypocistis, Cytinaceae, insects, flies, Mediterranean Basin, parasitic plant, pollination, Rafflesiaceae     

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.

     

Domestication of cardamom (Elettaria cardamomum) in Western Ghats, India: divergence in productive traits and a shift in major pollinators

Background and Aims

Elettaria cardamomum, a highly priced spice, is native to the Western Ghats of South India. Wild populations still occur in isolated patches in their natural habitats; however, much of today''s commercial product comes from cultivated sources. There is no information on domestication-related traits of this species; the main objective of this study was to compare wild and cultivated populations of cardamom in terms of vegetative and reproductive features in order to identify domestication syndromes and to examine whether the two populations have developed reproductive barriers.

Methods

Two wild populations and five cultivated plantations were used for the present study. Vegetative and floral traits, flowering phenology, pollination biology and breeding systems of wild and cultivated populations were compared. Effective pollinators amongst floral visitors were identified by confirming pollen transfer as well as by fruit set following their visit to virgin flowers. Manual pollinations were carried out in order to study the breeding systems of the two populations and reproductive barriers, if any, between them.

Key Results

Several productive traits including the number of branches, number of inflorescences, and total number of flowers per clump, number of flowers that open each day, the duration of flowering, the length of the flower and the amount of nectar per flower are significantly greater in cultivated cardamom. The principal pollinators in wild cardamom are solitary bees, Megachile sp. and two species of Amegilla, whereas those in cultivated cardamom are the social bees Apis dorsata, A. cerana and Trigona iridipennis. Both the wild and cultivated populations are self-compatible and there are no reproductive barriers between the two populations.

Conclusions

Domestication in cardamom has brought about significant changes in vegetative and reproductive traits and a shift in effective pollinators from native solitary bees to social bees. The shift in pollinators seems to be due to the availability of a large number of flowers for prolonged periods in cultivated cardamom that can attract and sustain social bees, rather than due to co-evolution of the flower and the pollinator.Key words: Elettaria cardamomum, wild cardamom, domestication, Amegilla sp., Apis cerana, Apis dorsata, Megachile sp., pollination efficiency, solitary bees, social bees     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Female and male fitness of a sexually deceptive orchid with a narrow distribution area: from phenotypic traits to spatial distribution patterns

• The Orchidaceae family presents one of the most extravagant pollination mechanisms: deception. While many studies on reproductive success have been performed on food‐deception orchids, less have been performed on sexually deceptive orchids. Here, we focused on Ophrys balearica P. Delforge, an endemic orchid of the Balearic Islands, to study its reproductive ecology, the spatio‐temporal variation of its reproductive success and the individual (floral display and geospatial position) and population parameters (patch size, shape and density) that affect its reproductive success.
• We performed hand‐pollination experiments, along with the recording of floral display parameters and GPS position of over 1,100 individuals from seven populations in two consecutive years. We applied, for the first time, GIS tools to analyse the effects of individual’s position within the population on the reproductive success. Reproductive success was measured both in male (removed pollinia) and female (fruit set) fitness.
• The results confirm that this species is pollinator‐dependent and mostly allogamous, but also self‐compatible. This species showed high values for the cumulative inbreeding depression index and high pollen limitation. Male fitness was almost equal to female fitness between years and populations, and reproductive success exhibited huge spatio‐temporal variation.
• Although we did not find strong correlations between floral display and reproductive success, patches with low‐plant density and individuals in the external portion of the population showed significantly higher plant fitness. These findings must be considered in conservation actions for endangered orchid species, especially considering that most orchids are strongly dependent on pollinators for their species’ fitness.

     

Assessment of distyly syndrome in Primula palinuri Petagn. a rare species living on maritime vertical cliffs

The distinctive habitat of vertical cliffs provides refuges for many relict plant species. Primula palinuri Petagn. is the only maritime and Mediterranean species of the genus Primula. It only occurs on the cliffs of a narrow area of southern Italy and is classified as being at risk of extinction. Given that only fragmentary information on the biology and ecology of P. palinuri is available, we studied flower morphology, breeding system and other reproductive traits of this species to furnish data for the evaluation of its conservation status. We assessed that plants of P. palinuri develop either long-styled or short-styled flowers and both morphs are equally represented in the studied populations. Reciprocal position of anthers and stigmas is complemented by differences in pollen–ovule ratios, pollen size and exine ornamentation. In both morphs, about 90 % of open-pollinated flowers developed mature capsules and the average number of seeds per capsule was around sixty. Hand pollination experiments showed that reproductive success was high (92.86 and 100 % fruit set) following inter-morph crossings, but no fruit develops after same-morph pollination or selfing. We have ascertained that full distyly syndrome (distinct floral morphs, intra-morph incompatibility and morphs equally represented in the populations) occurs in P. palinuri. However, a further reduction of the severely fragmented range, any rapid change of the environmental conditions due to human impact on coastal areas or stochastic events of plant mortality may alter the proportion of the two morphs, affecting the fragile conservation status of the species.     

Effects of conspecific and heterospecific floral density on the pollination of two related rewarding orchids

Variation in within-population floral density can affect interactions between plants and pollinators, resulting in variable pollen export for plants. We investigated the effects of conspecific and heterospecific floral densities on pollination success both of two related, self-compatible, nectar-rewarding orchid species in Ireland, Spiranthes romanzoffiana (rare and listed as endangered) and its congener, S. spiralis (more abundant and not of conservation concern). Floral densities, insect visitation rates, and orchid pollen transport were recorded in multiple quadrats in four populations of both orchid species over their flowering season. We found that conspecific and heterospecific co-flowering plant density affected pollination in both orchid species. For S. romanzoffiana, higher heterospecific density increased pollen removal. For S. spiralis, higher conspecific visitation increased pollen removal and increased heterospecific density decreased pollen deposition. In addition, increased conspecific density increased pollen deposition in both species. This study shows that plants may interact to facilitate or compete for different components of the pollination process, namely; pollinator attraction, pollen removal and deposition. Such interactions have immediate consequences for endangered plant species, as increases in both conspecific and heterospecific coflowering density may ameliorate the negative effects of rarity on pollination, hence overall reproductive success.     

Anthecology and reproductive system of Mourera fluviatilis (Podostemaceae): Pollination by bees and xenogamy in a predominantly anemophilous and autogamous family?

The Podostemaceae are sessile hydrophytes restricted to habitats with waterfalls and seasonal flow variability. Mourera fluviatilis belongs to the family and has a disjunct distribution in the Amazon and northeastern Brazil. The following hypotheses were tested: (i) both autogamy and anemophily act on the pollination system of M. fluviatilis, and (ii) the highest reproductive success is achieved by cross-pollination. Natural populations were monitored in the rivers Pirangi and Ipojuca, northeastern Brazil. Floral visitors are the polylectic bees Apis mellifera, Trigona spinipes (Apidae), Augochlora sp. and Augochloropsis sp. (Halictidae), though only the latter three are effective pollinators. In experiments on the reproductive system and pollination there was a high percentage of fruit formation (>86%), both in treatment and control plants. Although the index of self-incompatibility is 0.86, there was a significant difference in the number of seeds produced among reproductive system treatments, and xenogamy produced more seeds. There was no significant difference in the number of penetrated ovules between control and manual cross-pollination (xenogamy) treatments, 24 h after pollination. However, there were differences among treatments, 48 h after pollination, except for the control and xenogamy treatments. The pollen tubes from cross-pollination treatments usually penetrate the micropyle in shorter time and in higher quantity. We found differences among pollination experiments, except for the control and non-emasculated treatments, and between emasculated treatments. In spite of the ambiguity of floral attributes (pedicel, filaments and anthers colors, sweet odor, reduced perianth, absence of nectar, small and dry pollen grains), wind pollination is less important for this species; what predominates instead are first allogamy mediated by bees and second self-pollination at the end of anthesis.     

Breeding system and bird pollination of Camellia pubipetala,a narrowly endemic plant from karst regions of south China

Camellia pubipetala is an endemic and endangered species with small and isolated populations occurring only in karst regions in Guangxi of south China. To understand the reproductive biology of C. pubipetala and its possible influences upon its endangered status, its breeding system and pollination ecology were studied in the Longhushan (LHS) and Longzhao (LZ) populations of this species. The flowering duration of the C. pubipetala populations spanned from late January to early April and anthesis of a single flower usually lasted 5–7 days. This species is homogamous, and the pollen and stigma are viable throughout anthesis. Each bagged flower could secrete 141.5 μL of nectar at a sugar concentration of 25.0% during anthesis. The observed high pollen/ovule ratio, and the results of hand-pollination experiments indicated that this species obligately outcrosses. Open pollination resulted in a significantly decreased fruit set (6.7%) and seed set (38.9%) compared to supplementary pollination treatment (23.3% and 64.7%, respectively), which is indicative of a pollen limitation in the process of pollination. The primary pollinator of C. pubipetala is the sunbird Aethopyga christinae and its visiting frequency is quite low, whereas the honeybee Apis cerana is only an occasional pollinator in wild populations. Low reproductive rates in C. pubipetala were found to be a consequence of few species of pollinators and their low visiting frequency. Pollen limitation may be a crucial factor that contributes to the endangered nature of this species. Artificial pollination and the release of pollinators are effective ways to increase the fruit and seed yield of this species.     

Pollinators,floral morphology and scent chemistry in the southern African orchid genus Schizochilus

Floral evolution often involves suites of traits, including morphology, colour and scent, but these traits are seldom analysed together in comparative studies. We investigated the associations between floral traits and pollination systems in Schizochilus, a southern African orchid genus with small nectar-producing flowers that has not been studied previously with respect to pollination biology. Field observations indicated the presence of distinct pollination systems in the four species which occur in the Drakensberg, including pollination by muscid flies in Schizochilus angustifolius, tachinid flies in Schizochilus zeyheri, various small flies in Schizochilus bulbinella and bees and wasps in Schizochilus flexuosus. Pollination success and pollen transfer efficiency clearly differed among the four species but were not correlated with the quantity of nectar rewards. Multivariate analysis of floral morphology and floral scent chemistry based on GC-MS data revealed significant differences among species as well as populations within species. The floral scent of S. angustifolius was dominated by the benzenoid compounds benzaldehyde and phenylacetaldehyde. Samples of one population of S. bulbinella were relatively similar to S. angustifolius but samples of another population were very distinct due to the occurrence of the nitrogen-containing compounds 3-methyl-butyl aldoxime (syn/anti) and the higher amounts of aliphatic esters, alcohols and acids. In contrast, the floral scent of S. flexuosus and S. zeyheri was characterized by high relative amounts of methyl benzoate. We conclude that Schizochilus has distinct, specialized pollination systems associated with subtle but significant variation in floral morphology and scent chemistry. We also caution that sampling of several populations may be required to characterize floral scent composition at the species-level in plants.