Implications of a long-term, pollinator-mediated selection on floral traits in a generalist herb

Background and Aims

The phenotypic selection of a diverse insect assemblage was studied on a generalist plant species (Paeonia broteroi) in ten flowering seasons, with tests for whether visitor preferences for plants with larger flowers eventually translated into significant differences among plants in visitation rates, seed production, seed mass, seed germination and seedling survival.

Methods

Selection gradients were used to assess if selection on flower size contributed to explain differences in visitation rates, seed production and seed mass. First, independent analyses were carried out for each season; then for the ten season as a whole. Seedling emergence and survival were assessed by generalized linear models.

Key Results

Directional selection was found on flower size through visitation rates and seed production, and stabilizing selection through seed mass. Thus, larger flowers were more visited, and produced more, but lighter seeds, than smaller flowers. The results suggest a conflicting selection on flower size through seed number and size. Floral integration found in the study populations was larger than that in populations of a distant region. Finally, seed size did not influence seedling emergence and survival; thus, any advantages of seed size may be constrained under natural conditions before plants become reproductive individuals.

Conclusions

Plants with larger flowers may be benefited by producing more lighter seeds than fewer heavier ones, as they may contribute disproportionately to the seed bank, and have better chances that any descendant could eventually recruit. However, it seems unlikely that differences in flower size and integration found among populations in different regions could have been originated by rapid evolutionary change. First, because of the conflicting selection described; second, because of the remarkably low seedling survival found under natural conditions. Consequently, the influence of pollinator selection alone does not seem to explain differences in flower size and integration.Key words: Paeonia broteroi, long-term selection, conflicting selection, flower size, seed production, generalist pollination     

Flower thermoregulation facilitates fertilization in Asian sacred lotus

Background and Aims

The thermoregulatory flower of the Asian sacred lotus (Nelumbo nucifera) can maintain a relatively stable temperature despite great variations in ambient temperature during anthesis. The thermoregulation has been hypothesized to offer a direct energy reward for pollinators in lotus flowers. This study aims to examine whether the stable temperature maintained in the floral chamber influences the fertilization process and seed development.

Methods

An artificial refrigeration instrument was employed to cool flowers during the fertilization process and post-fertilization period in an experimental population. The effect of temperature on post-pollination events was also examined by removing petals in two field populations.

Key Results

Treatments with low floral temperature did not reduce stigma receptivity or pollen viability in undehisced anthers. Low temperature during the fertilization period significantly decreased seed set per flower but low temperature during the phase of seed development had no effect, suggesting that temperature regulation by lotus flowers facilitated fertilization success. Hand-pollination treatments in two field populations indicated that seed set of flowers with petals removed was lower than that of intact flowers in north China, where ambient temperatures are low, but not in south China, confirming that reducing the temperature of carpels did influence post-pollination events.

Conclusions

The experiments suggest that floral thermoregulation in lotus could enhance female reproductive success by facilitating fertilization.Key words: Nelumbo nucifera, Asian sacred lotus, beetle-pollination syndrome, fertilization process, post-pollination events, pollen viability, stigma receptivity, thermoregulation     

The relative importance of architecture and resource competition in allocation to pollen and ovule number within inflorescences of Hosta ventricosa varies with the resource pools

Background and Aims

Allocation of resources to floral traits often declines distally within inflorescences in flowering plants. Architecture and resource competition have been proposed as underlying mechanisms. The aim of the present study is to assess the relative importance of resource competition and architectural effects in pollen and ovule production on racemes of Hosta ventricosa, an apomictic perennial herb.

Methods

Combinations of two defoliation treatments (intact and defoliated) and two fruit-set treatments (no-fruit and fruit) were created, and the roles of architecture and resource competition at each resource level were assessed.

Key Results

Pollen and ovule number per flower increased after defoliation, but pollen to ovule ratio per flower did not change. Pollen, ovules and the pollen to ovule ratio per flower declined distally on racemes at each resource level. In the intact treatment, fruit development of early flowers did not affect either pollen or ovule number of late flowers. In the defoliated treatment, fruit development of early flowers reduced both pollen and ovule numbers of late flowers due to over-compensation caused by defoliation. Late flowers on defoliated fruit racemes produced less pollen than intact fruit racemes but the same number of ovules; therefore, the reduction in pollen number was not caused by over-compensation. In addition, the fruit-set rate of early flowers during flowering was higher in intact racemes than in defoliated racemes.

Conclusions

In flowering plants, the relative importance of architecture and resource competition in allocation to pollen and ovules may vary with the resource pools or the overall resource availability of maternal plants.     

Did Drosera evolve long scapes to stop their pollinators from being eaten?

Background and Aims

Insectivorous plants frequently display their flowers on the ends of long racemes. Conventional wisdom is that long racemes in insectivorous plants have evolved to provide spatial separation between flowers and traps, which consequently prevents pollinators from being captured. However, it is also possible that long racemes evolved for better seed dispersal or to make flowers more visible to pollinators.

Methods

Two sympatric insectivorous plants with identical pollinators were studied: Drosera cistiflora, with an upright growth form but a short raceme; and Drosera pauciflora, with a basal rosette of traps and a very long raceme. If long racemes evolved to protect their pollinators then D. cistiflora should capture more pollinators than D. pauciflora. However, if long racemes evolved to attract pollinators then taller flowers should receive more pollination visits than shorter flowers.

Key Results

Examination of D. pauciflora and D. cistiflora traps revealed that no pollinators were captured by either species, suggesting that long racemes did not evolve to protect pollinators from being captured. Experimental manipulations of flower height in D. cistiflora showed that experimentally shortened plants received significantly fewer pollination visits than plants which were taller in stature.

Conclusions

Long scapes in Drosera and non-insectivorous plants probably evolved due to similar selective pressures such as pollinator attraction.     

Mutual reproductive dependence of distylic Cordia leucocephala (Cordiaceae) and oligolectic Ceblurgus longipalpis (Halictidae, Rophitinae) in the Caatinga

Background and Aims

The close relationship between distylic Cordia leucocephala and the bee Ceblurgus longipalpis, both endemic to the Caatinga, north-east Brazil, was investigated, emphasizing reproductive dependence, morphological adaptations of the partners, and pollen flow.

Methods

In the municipality of Pedra, in the Caatinga of Pernambuco, the breeding system and reproductive success of C. leucocephala, its interaction with flower visitors and inter- and intramorph pollen flow were determined.

Key Results

The bee Ceblurgus longipalpis, the unique flower visitor and effective pollinator of self-incompatible Cordia leucocephala, presents morphological features adapted to exploit hidden pollen and nectar in the long and narrow corolla tubes. Pollen of low-level anthers is collected with hairs on prolonged mouthparts and pollen of high-level anthers with clypeus, mandibles, and labrum, showing pollen removal from both levels with the same effectiveness. In both morphs, this results in similar legitimate, i.e. intermorph cross-pollen flow. Illegitimate pollen flow to stigmas of pin flowers, however, was much higher than to stigmas of thrum flowers. Moreover, more illegitimate pollen was transported to stigmas of pin and less to those of thrum flowers when compared with legitimate pollen flow.

Conclusions

The study reveals a one-to-one reproductive inter-dependence between both partners. Data indicate that this relationship between bee species and plant species is one of the rare cases of monolecty among bees. Monotypic Ceblurgus longipalpis, the only rophitine species of Brazil, evolved prolonged mouthparts rare among short-tongued bees that enable them to access pollen from flowers with short-level anthers hidden for bees of other species, and nectar at the base of the flower tube.     

Stigmatic fluid aids self-pollination in Roscoea debilis (Zingiberaceae): a new delayed selfing mechanism

Background and Aims

Delayed selfing is the predominant mode of autonomous self-pollination in flowering plants. However, few delayed selfing mechanisms have been documented. This research aims to explore a new delayed selfing mechanism induced by stigmatic fluid in Roscoea debilis, a small perennial ginger.

Methods

Floral biology and flower visitors were surveyed. The capacity of autonomous selfing was evaluated by pollinator exclusion. The timing of autonomous selfing was estimated by emasculation at different flowering stages. The number of seeds produced from insect-pollination was assessed by emasculation and exposure to pollinators in the natural population. The breeding system was also tested by pollination manipulations.

Key Results

Autonomous self-pollination occurred after flowers wilted. The stigmatic fluid formed a globule on the stigma on the third day of flowering. The enlarged globule seeped into the nearby pollen grains on the fourth flowering day, thus inducing pollen germination. Pollen tubes then elongated and penetrated the stigma. Hand-selfed flowers produced as many seeds as hand-crossed flowers. There was no significant difference in seed production between pollinator-excluded flowers and hand-selfed flowers. When emasculated flowers were exposed to pollinators, they produced significantly fewer seeds than intact flowers. Visits by effective pollinators were rare.

Conclusions

This study describes a new form of delayed autonomous self-pollination. As the predominant mechanism of sexual reproduction in R. debilis, delayed self-pollination ensures reproduction when pollinators are scarce.     

Floral structure and development in the monoecious palm Gaussia attenuata (Arecaceae; Arecoideae)

Background and Aims

Sexual dimorphism, at both the flower and plant level, is widespread in the palm family (Arecaceae), in contrast to the situation in angiosperms as a whole. The tribe Chamaedoreeae is of special interest for studies of the evolution of sexual expression since dioecy appears to have evolved independently twice in this group from a monoecious ancestor. In order to understand the underlying evolutionary pathways, it is important to obtain detailed information on flower structure and development in each of the main clades.

Methods

Dissection and light and scanning electron microscopy were performed on developing flowers of Gaussia attenuata, a neotropical species belonging to one of the three monoecious genera of the tribe.

Key Results

Like species of the other monoecious genera of the Chamaedoreeae (namely Hyophorbe and Synechanthus), G. attenuata produces a bisexual flower cluster known as an acervulus, consisting of a row of male flowers with a basal female flower. Whereas the sterile androecium of female flowers terminated its development at an early stage of floral ontogeny, the pistillode of male flowers was large in size but with no recognizable ovule, developing for a longer period of time. Conspicuous nectary differentiation in the pistillode suggested a possible role in pollinator attraction.

Conclusions

Gaussia attenuata displays a number of floral characters that are likely to be ancestral to the tribe, notably the acervulus flower cluster, which is conserved in the other monoecious genera and also (albeit in a unisexual male form) in the dioecious genera (Wendlandiella and a few species of Chamaedorea). Comparison with earlier data from other genera suggests that large nectariferous pistillodes and early arrest in staminode development might also be regarded as ancestral characters in this tribe.     

Female reproductive success decreases with display size in monkshood, Aconitum kusnezoffii (Ranunculaceae)

Background and Aims

Reduction in female fitness in large clones can occur as a result of increased geitonogamous self-fertilization and its influence through inbreeding depression. This possibility was investigated in the self-compatible, bee-pollinated perennial herb Aconitum kusnezoffii which varies in clone size.

Methods

Field investigations were conducted on pollinator behaviour, flowering phenology and variation in seed set. The effects of self-pollination following controlled self- and cross-pollination were also examined. Selfing rates of differently sized clones were assessed using allozyme markers.

Key Results

High rates of geitonogamous pollination were associated with large display size. Female fitness at the ramet level decreased with clone size. Fruit and seed set under cross-pollination were significantly higher than those under self-pollination. The pre-dispersal inbreeding depression was estimated as 0·502 based on the difference in seed set per flower between self- and cross-pollinated flowers. Selfing rates of differently sized clones did not differ.

Conclusions

It is concluded that in A. kusnezoffii the negative effects of self-pollination causing reduced female fertility with clone size arise primarily from a strong early-acting inbreeding depression leading to the abortion of selfed embryos prior to seed maturation.Key words: Early-acting inbreeding depression, Aconitum kusnezoffii, clone size, female reproductive success, geitonogamy     

Preformation and distribution of staminate and pistillate flowers in growth units of Nothofagus alpina and N. obliqua (Nothofagaceae)

Background and Aims

The distribution and differentiation times of flowers in monoecious wind-pollinated plants are fundamental for the understanding of their mating patterns and evolution. Two closely related South American Nothofagus species were compared with regard to the differentiation times and positions of staminate and pistillate flowers along their parent growth units (GUs) by quantitative means.

Methods

Two samples of GUs that had extended in the 2004–2005 growing season were taken in 2005 and 2006 from trees in the Lanín National Park, Patagonia, Argentina. For the first sample, axillary buds of the parent GUs were dissected and the leaf, bud and flower primordia of these buds were identified. The second sample included all branches derived from the parent GUs in the 2005–2006 growing season.

Key Results

Both species developed flowering GUs with staminate and/or pistillate flowers; GUs with both flower types were the most common. The position of staminate flowers along GUs was similar between species and close to the proximal end of the GUs. Pistillate flowers were developed more distally along the GUs in N. alpina than in N. obliqua. In N. alpina, the nodes bearing staminate and pistillate flowers were separated by one to several nodes with axillary buds, something not observed in N. obliqua. Markovian models supported this between-species difference. Flowering GUs, including all of their leaves and flowers were entirely preformed in the winter buds.

Conclusions

Staminate and pistillate flowers of N. alpina and N. obliqua are differentiated at precise locations on GUs in the growing season preceding that of their antheses. The differences between N. alpina and N. obliqua (and other South American Nothofagus species) regarding flower distribution might relate to the time of anthesis of each flower type and, in turn, to the probabilities of self-pollination at the GU level.Key words: Branch, bud, growth unit, Markovian models, Nothofagus alpina, N. obliqua, Patagonian forests, pistillate flower, preformation, staminate flower     

Floral closure induced by pollination in gynodioecious Cyananthus delavayi (Campanulaceae): effects of pollen load and type, floral morph and fitness consequences

Background and aims

Pollination-induced floral changes, which have been widely documented in flowering plants, have been assumed to enhance the plant''s reproductive success. However, our understanding of the causes and consequences of these changes is still limited. Using an alpine gynodioecious species, Cyananthus delavayi, we investigated the factors affecting floral closure and estimated the fitness consequences of floral closure.

Methods

The timings of floral closure and fertilization were determined. The effects of pollen load, pollen type (cross- or self-pollen) and floral morph (female or perfect flower) on the occurrence of floral closure were examined. Ovule fertilization and seed production were examined to investigate the causes and consequences of floral closure. Flowers were manipulated to prevent closing to detect potential benefits for female fitness.

Key Results

Floral closure, which could be induced by a very low pollen load, occurred within 4–7 h after pollination, immediately following fertilization. The proportion of closed flowers was influenced by pollen load and floral morph, but not by pollen type. Floral closure was more likely to occur in flowers with a higher proportion of fertilized ovules, but there was no significant difference in seed production between closed and open flowers. Those flowers in which closure was induced by natural pollination had low fruit set and seed production. Additionally, seed production was not influenced by closing-prevented manipulation when sufficient pollen deposition was received.

Conclusions

The occurrence of floral closure may be determined by the proportion of fertilized ovules, but this response can be too sensitive to ensure sufficient pollen deposition and can, to some extent, lead to a cost in female fitness. These results implied that the control of floral receptivity by the recipient flowers does not lead to an optimal fitness gain in C. delavayi.     

Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome

Background and Aims

If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.

Methods

Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.

Key Results

Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.

Conclusions

This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.     

Is Floral Longevity Influenced by Reproductive Costs and Pollination Success in Cohniella ascendens (Orchidaceae)?

Background and Aims

Although studies have shown that pollen addition and/or removal decreases floral longevity, less attention has been paid to the relationship between reproductive costs and floral longevity. In addition, the influence of reproductive costs on floral longevity responses to pollen addition and/or removal has not yet been evaluated. Here, the orchid Cohniella ascendens is used to answer the following questions. (a) Does experimental removal of flower buds in C. ascendens increase flower longevity? (b) Does pollen addition and/or removal decrease floral longevity, and does this response depend on plant reproductive resource status?

Methods

To study the effect of reproductive costs on floral longevity 21 plants were selected from which we removed 50 % of the developing flower buds on a marked inflorescence. Another 21 plants were not manipulated (controls). One month later, one of four flowers on each marked inflorescence received one of the following pollen manipulation treatments: control, pollinia removal, pollination without pollinia removal or pollination with pollinia removal. The response variable measured was the number of days each flower remained open (i.e. longevity).

Key Results

The results showed significant flower bud removal and pollen manipulation effects on floral longevity; the interaction between these two factors was not significant. Flowers on inflorescences with previously removed flower buds remained open significantly longer than flowers on control inflorescences. On the other hand, pollinated flowers closed much faster than control and removed-pollinia flowers, the latter not closing significantly faster than control flowers, although this result was marginal.

Conclusions

The results emphasize the strong relationship between floral longevity and pollination in orchids, as well as the influence of reproductive costs on the former.Key words: Cohniella ascendens, floral longevity, flower bud removal, pollination, pollinia removal, reproductive costs     

Pollinators of the Rocky Mountain columbine: temporal variation, functional groups and associations with floral traits

Background and Aims

Pollinators together with other biotic and some abiotic factors can select for floral traits. However, variation in pollinator abundance over time and space can weaken such selection. In the present study, the variation in pollinator abundance over time and space was examined in populations of the Rocky Mountain columbine. The variation in three floral traits is described and correlations between pollinator type, functional pollinator groups or altitude and floral traits are examined.

Methods

Pollinator observations took place in six Aquilegia coerulea populations over 1–4 years and spur length, flower colour and sepal length were measured in 12 populations. Pollinator abundance, measured as visits per flower per hour, was compared among populations and years. Pollinators were grouped into two functional groups: pollen or nectar collectors. The following associations were examined: annual presence of hawkmoths and whiter flowers with longer spurs; the presence of Sphinx vashti and longer spurs; and higher altitudes and whiter flowers. The study looked at whether an increase in the proportion of hawkmoths in a population was associated with whiter and larger flowers with longer spurs.

Key Results

The abundance of different pollinator groups varied over time and space. Floral traits varied among populations. Higher altitude was correlated with bluer flowers. Whiter flowers were associated with the annual presence of hawkmoths. Populations visited by Sphinx vashti had longer spurs than populations visited only by Hyles lineata. Populations with greater percentage of nectar-collecting pollinators did not have whiter, larger flowers with longer spurs.

Conclusions

Despite the large variation in pollinator abundance over time and space, one species of bumble-bee or hawkmoth tended to predominate in each population each year. Future studies of Aquilegia coerulea should examine the specific influences of pollinators and the environment on flower colour and of hawkmoth species on spur length.Key words: Aquilegia coerulea, columbine, pollinator abundance, bumble-bee, hawkmoth, flower colour, spur length, functional pollinator group, altitude, floral trait     

Flower power: its association with bee power and floral functional morphology in papilionate legumes

Background and Aims

A test was made of the hypothesis that papilionate legume flowers filter pollinators according to their ability to exert strength to open flowers to access rewards. In addition, interactions with pollen vectors were expected to explain the structural complexity of the architecture of these flowers since operative flower strength may be determined by a combination of morphological traits which form part of an intrafloral functional module.

Methods

Six papilionate species were studied: Collaea argentina, Desmodium uncinatum, Galactia latisiliqua, Lathyrus odoratus, Spartium junceum and Tipuana tipu. Measurements were made of the strength needed to open keels and the strength that pollinators were capable of exerting. Morphological traits of all petals were also measured to determine which of them could be either mutually correlated or correlated with operative strength and moment of strength and participated in a functional module.

Key Results

It was observed that pollinators were capable in all cases of exerting forces higher and often several times higher than that needed to access floral rewards, and no association could be detected between floral operative strength and strength exerted by the corresponding pollinators. On the other hand, strong and significant correlations were found among morphometric traits and, of these, with operative strength and moment. This was particularly evident among traits of the keel and the wings, presumably involved in the functioning of the floral moveable mechanism.

Conclusions

Though visitors are often many times stronger than the operative strength of the flowers they pollinate, exceptionally weak bees such as Apis mellifera cannot open the strongest flowers. On the other hand, strong correlations among certain petal morphometric traits (particularly between the keel and wings) give support to the idea that an intrafloral module is associated with the functioning of the mechanism of these legume flowers. In addition, the highly significant correlations found across petals support the view of functional phenotypic integration transcending the ontogenetic organization of flower structure.