Removal and insertion of pollinia in flowers of Oxypetalum (Asclepiadaceae) in southeastern Brazil

Pollinarium removal and pollinium insertion of seven Oxypetalum species (O. alpinum var. alpinum, O. appendiculatum, O. banksii subsp. banksii, O. jacobinae, O. mexiae, O. pachyglossum and O. subriparium) were recorded in Vi?osa. Minas Gerais. They presented a tendency of one or two pollinarium removals and one pollinium insertion (single insertion), except O. appendiculatum. In this species, mainly, two pollinia of the same pollinarium were inserted per stigmatic chamber (double insertion), resulting exceptionally in 6-10 inserted pollinia in a flower, an unusual occurrence among the Asclepiadaceae. No association between removal and insertion was found, e.g., O. subriparium and O. banksii subsp. banksii had the highest pollinarium removal (1.78 and 1.45, respectively) and one of the lowest pollinium insertions (0.02 in both species), per flower. Oxypetalum mexiae showed the lowest pollinarium removal and pollinium insertion per flower (0.09 and 0.01, respectively) among the studied species and other Asclepiadaceae. Oxypetalum subriparium, O. banksii subsp. banksii and O. mexiae might be having reproductive limitations. Pollinarium removal and pollinium insertion per flower of the studied species varied from site to site, similarly to what was recorded for other Asclepiadaceae.     

Variation in stigma morphology – How does it contribute¶to pollination adaptation in Pedicularis (Orobanchaceae)?

 Stigma morphology in Pedicularis and its role in pollination have not been evaluated. This paper reports for the first time the relationship between stigma morphology and other aspects of floral design. 17 species of Pedicularis representing all the different types of corolla structure in the genus were studied in the field and in the laboratory. Results indicate that the stigmatic surface morphology of Pedicularis species is related to the type of corolla structure occurring in the species. Pollination studies in eight Pedicularis species demonstrated adaptation between stigmatic surface morphology and other aspects of floral biology. Species with larger, more corrugated stigmatic surfaces always possess a subsidiary construction (“tooth” or “beak”) on the galea. In addition, they have relatively longer flower tubes, higher stigmatic pollen loads, bigger pollen size and a shorter blooming period for individual flowers. Finally, the authors discuss the evolutionary significance of floral design in genus Pedicularis. Received April 8, 2002; accepted June 11, 2002 Published online: December 11, 2002     

青阳参花部特征及其传粉适应性

对青阳参花（Cynanchum otophyllum）部综合特征、访花昆虫种类、访花行为及传粉过程进行了研究,结果表明,青阳参花结构复杂,两个子房基部离生、花柱联合与雄蕊形成合蕊柱,柱头表面被邻近花药的侧翼紧密包围形成5个柱头腔。青阳参的花粉形成独特的花粉块,一次传粉过程可以转运大量的花粉。东方蜜蜂（Apis cerana）是青阳参的主要传粉昆虫,其传粉包括两个过程：（1）当蜜蜂的口器或足插入着粉腺的槽口后借助蜜蜂的力量将花粉块从花上拔起;（2）当蜜蜂再次访花时将携带的花粉块插入其中一个柱头腔。花粉块里面的花粉粒住柱头腔中萌发出花粉管,然后沿着花柱道向下生长最后进入子房。在整个花期仡粉保持有相对较高的生活力,而其柱头可授性则在7天后逐渐降低。     

Reproductive biology of Colletia spinosissima (Rhamnaceae) in Argentina

 We studied the reproductive behavior of Colletia spinosissima Gmel. at six sites in two regions of Argentina during the 1998–2000 flowering seasons. Flowering takes place in winter. The flowers are homogamous, fragrant, and entomophilous. There was evidence of self-incompatibility and cross-incompatibility between nearby partners. Fruit set of open-pollinated flowers (13.3%) seemed pollen-limited, since low fecundity was consistently associated with small stigmatic pollen loads and with low numbers of successful pollen tubes, and fecundity increased with abundance of efficient pollinators. Reproductive success (percent ovules becoming viable seeds) was 3.6. It is hypothesized that homogamy was established in Colletia because in an impoverished pollination environment – such as the one likely prevailing when this lineage evolved – achieving pollen export and receipt in a single pollinator visit has selective advantages. A large stigma and an extragynoecial compitum would compensate for the homogamy-driven, increased pollen-stigma interference. Received March 8, 2001 Accepted July 19, 2001     

The pollination of a self-incompatible, food-mimic orchid, Coelogyne fimbriata (Orchidaceae), by female Vespula wasps

Background and Aims

The study of specialized interactions between species is crucial to our understanding of processes in evolutionary ecology due to their profound effect on life cycles and diversification. Obligate pollination by a single wasp species is rare in Orchidaceae except in species with sexually deceptive flowers that are pollinated exclusively by male insects. The object of this study was to document pollination of the food-deceptive flowers of Coelogyne fimbriata, a species pollinated exclusively by female wasps.

Methods

Field observations and experiments were conducted in two populations of C. fimbriata. Floral phenology was recorded, and functional floral architecture was measured. Insect visitors to flowers were observed from 2005 to 2007. Bioassay experiments were conducted to check whether the floral odour attracted pollinators. Natural (insect-mediated) rates of pollinarium removal, pollinium deposition on stigmas, and fruit set were recorded. To determine the importance of cross-pollination, the breeding system was assessed via controlled, hand-pollination experiments.

Key Results

Two populations of C. fimbriata with fragrant, nectarless flowers are pollinated by females of the same Vespula species (Vespidae, Hymenoptera). Experiments on wasps show that they crawl towards the source of the odour. The flowering period appears to coincide with an annual peak in Vespula colony expansion when additional workers forage for carbohydrates. Rates of pollinarium removal (0·069–0·918) and pollinium deposition on stigmas (0·025–0·695) are extremely variable. However, fruit set in C. fimbriata is always low (0·014–0·069) and appears to be based on self-incompatibility coupled with intraclonal (geitonogamous) deposition of pollinia.

Conclusions

Coelogyne fimbriata and Steveniella satyrioides are now the only orchid species known to have food-deceptive flowers that are pollinated exclusively by eusocial, worker wasps. In C. fimbriata, floral odour appears to be the major attractant. Sub-populations may go through flowering seasons when pollinators are abundant or infrequent, but fruit set always remains low because the obligate pollinator does not often appear to transfer pollinaria between intercompatible genets.Key words: Coelogyne fimbriata, Vespula wasps, food deception, floral odour, pollinarium removal, pollinium deposition, self-incompatibility     

One for all and all for one: retention of colour‐unchanged old flowers increases pollinator attraction in a hermaphroditic plant

• Long‐lived flowers increase pollen transfer rates, but these entail high water and carbon maintenance costs. The retention of pollinated and reward‐free old flowers enhances pollinator visitation to young receptive flowers by increasing floral display size. This mechanism is associated with acropetal inflorescences or changes in flower colour and openness, but the retention of unchanging solitary flowers remains overlooked.
• We examined pollination‐dependent variation in floral longevity and determined stigmatic receptivity, pollen viability and pollen removal rates among flower ages in Kielmeyera regalis, a Neotropical savanna shrub. We also evaluated the effects of floral display size on pollinator visitation rates. Lastly, we determined whether old flowers are unvisited and exclusively increase pollinator attraction to young flowers through flower removal experiments.
• Regardless of pollination treatment, flowers lasted fully open with no detectable physical changes for 3 days. Over time, stigmas remained receptive but >95% of pollen was removed. Pollinator visitation significantly increased with floral display size and intermediate percentages (15–30%) of newly opened flowers. Accordingly, the retention of reward‐free and unvisited old flowers increased young flower–pollinator interaction.
• Our results reveal the importance of a prolonged floral longevity in increasing pollinator attraction toward newly opened receptive flowers without changes in flower colour and form. We conclude that the retention of pollinated, reward‐free and unvisited colour‐unchanged old flowers in K. regalis is a strategy that counteracts the water use costs associated with the maintenance of large flowers with increased mate opportunities in a pollen‐limited scenario.

     

Quantification of trehalose in biological samples with a conidial trehalase from the thermophilic fungus Humicola grisea var. thermoidea

The partially-purified, thermally-stable trehalase from conidia of Humicola grisea was highly specific for trehalose and was free of potentially interfering activities. The enzyme was fully stable when stored in solution at -15°C for at least 6 months. This preparation could be used to quantify trehalose from 0.05 to 1.25 mol/ml either in carbohydrate mixtures or in complex biological materials.M.J. Neves, H.F. Terenzi and J.A. Jorge are with Departmento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirao Preto, Universidade de São Paulo, 14040-901-Ribeirão Preto, São Paulo, Brazil; F.A. Leone is with Departamento de Quimica, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, 14040-901-Ribeirão Preto, São Paulo, Brazil.     

Effects of anoxia on root ultrastructure of four neotropical trees

Summary. The root ultrastructure of seedlings grown in anaerobic conditions was investigated in four neotropical species: Sesbania virgata, Erythrina speciosa, Sebastiania commersoniana (all present in waterlogged or flooded areas), and Schizolobium parahyba (that occupies mainly dry areas). Anaerobiosis induced an increase in the size of mitochondria, dilatation of cristae and of the endoplasmic reticulum, and fragmentation or concentric arrangement of reticulum saccules. The ultrastructural alterations were reversible only for S. virgata and E. speciosa. The seedlings of S. parahyba and S. commersoniana were more sensitive to oxygen deprivation and presented extensive cell disruption. The results are discussed in terms of energy supply.Correspondence and reprints (present address): Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade Estadual de Londrina, P.O. Box 6001, 86051-970, Londriná, Paraná, Brazil.     

Morphological diversity and function of the stigma in Ficus species (Moraceae)

The stigma plays several roles such as pollen hydration and selection, and pollen tube nutrition. In the Ficus-fig wasp mutualism, stigmata have an additional, almost unknown, function by representing a physical interface for both plant and wasp reproduction. We used light and electron microscopy to compare the detailed morphology of the stigmata of nine Ficus species of different sections and with different pollination modes and sexual expressions. Figs were collected at the stage when the stigmata were receptive for pollination. Stigmata in actively pollinated monoecious species have well developed papillae concentrated on the adaxial surface exposed towards the fig cavity. Conversely, the passively pollinated monoecious species have the whole surface of the stigmata covered by somewhat smaller papillae. In both actively and passively pollinated monoecious species these features are consistent, irrespective of style length. In all actively pollinated gynodioecious species, the stigmata of pistillate flowers were tubular or infundibuliform whereas in almost all actively pollinated monoecious species (except F. racemosa) the stigmata were filiform, with one branch or two asymmetric branches. In gynodioecious species the short-styled flowers in “male” figs show a limited receptive surface with small papillae, while the stigmata of long-styled flowers in “female” figs are covered by papillae that extend down the sides of the style, increasing the stigmatic surface. In actively pollinated species, stigmata are cohesive, forming a common surface for pollen tube germination (= synstigma). The synstigma arrangement was quite variable: lax, cohesive or very cohesive, with entanglement by stigmatic papillae and stylar trichomes. Entanglement by stylar trichomes is common in gynodioecious species. The synstigma arrangement did not correlate with phylogeny or breeding system. This study is the first to report a very loose synstigma in actively pollinated monoecious Ficus species. Our analyses revealed that, in Ficus, the synstigma is functionally analogous to an extra-gynoecial compitum. Comparative studies will be required to test further hypotheses about the evolutionary determinants of such variation.     

Post-pollination changes in the floral organs of two <Emphasis Type="Italic">Cymbidium</Emphasis> species

It was observed that the unpollinated flowers of Cymbidium pendulum (Roxb.) Sw. and C. aloifolium (L.) Sw. stayed fresh for 20 and 18 d, respectively, but attained senescence in 8 and 7 d, respectively, after pollination. The higher content of total soluble sugars, reducing sugars and free amino acids was observed in all the floral organs of pollinated flowers than in unpollinated ones. Pollination also up-regulated the activity of hydrolytic (α-amylase, β-amylase, invertase) and proteolytic enzymes (proteases) in floral organs. Amongst floral organs, the lip and perianth possessed highest contents of metabolites. Application of auxin inhibitor (0.25 μM triiodobenzoic acid) and ethylene inhibitor (0.25 μM AgNO₃) to the pollinated flowers partially prevented the process of senescence.     

Cleistogamy in the rare high Andean perennial herb Cryptantha capituliflora (Boraginaceae)

 Axial and apical flowers of Cryptantha capituliflora were analyzed with regard to morphology and pollen tube growth to assess the occurrence of cleistogamy. Although intermediate floral forms do occur, cleistogamous flowers were significantly smaller than chasmogamous flowers, had fewer anthers, and showed a distinctive stigmatic surface. Chasmogamous flowers can be cross-pollinated. Nevertheless, the growth of self-pollen tubes in few chasmogamous buds jointly with flower characters suggests that these flowers can probably produce fruits through autonomous selfing. The mean seed number per fruit did not differ between fruits from chasmogamous and cleistogamous flowers. Cleistogamous flowers were only observed in axial inflorescences, which are completely covered by the leaf. Other species of section Cryptantha also show the same trend, with cleistogamous flowers located in the lower half of the stems. This pattern is discussed in relation to dissimilarities in the outcrossing opportunities between flower types within the plant. Received May 22, 2002; accepted November 14, 2002 Published online: March 20, 2003     

The pollination ecology of Pedicularis rex subsp. lipkyana and P. rex subsp. rex (Orobanchaceae) from Sichuan, southwestern China

Pedicularis is one of few genera for which pollination ecology has been studied extensively. Although over half of the species of Pedicularis are found in the mountains of southwestern China, pollination ecology has been studied there on a few species only. The present paper reports pollination ecology of Pedicularis rex subsp. lipskyana and P. rex subsp. rex from Sichuan, southwestern China. The two subspecies are nectariferous, with sugar content 28% in P. rex lipskyana and 22% in P. rex, respectively. The flowers of the two subspecies are adapted to pollination by bumblebee workers. Bumblebees are the primary and effective pollinators although honeybees were also observed pollinating P. rex subsp. lipskyana at one studied population. Usually bumblebees entered the corolla tube from the right side in an upright or a nearly half-inverted position to imbibe nectar located at the base of the corolla tube. The stigma usually contacts the side region of the thorax and abdomen or occasionally directly contacted pollen loads, leading to pollination of flowers. Much less commonly, bumblebees foraged on flowers of P. rex subsp. rex in an inverted position for pollen, which pollinated flowers sternotribically. We suggest that pollination mechanism is closely associated with floral morphology. Some important differences were found from a previous study on P. rex in floral morphology and the primary pollination mechanism.     

Adaptive differentiation in floral traits in the presence of high gene flow in scarlet gilia (Ipomopsis aggregata)

Plant–pollinator interactions are thought to be major drivers of floral trait diversity. However, the relative importance of divergent pollinator‐mediated selection vs. neutral processes in floral character evolution has rarely been explored. We tested for adaptive floral trait evolution by comparing differentiation at neutral genetic loci to differentiation at quantitative floral traits in a putative Ipomopsis aggregata hybrid zone. Typical I. aggregata subsp. candida displays slender white tubular flowers that are typical of flowers pollinated by hawkmoths, and subsp. collina displays robust red tubular flowers typical of flowers pollinated by hummingbirds; yet, hybrid flower morphs are abundant across the East Slope of the Colorado Rockies. We estimated genetic differentiation (F_ST) for nuclear and chloroplast microsatellite loci and used a half‐sib design to calculate quantitative trait divergence (Q_ST) from collection sites across the morphological hybrid zone. We found little evidence for population structure and estimated mean F_ST to be 0.032. Q_ST values for several floral traits including corolla tube length and width, colour, and nectar volume were large and significantly greater than mean F_ST. We performed multivariate comparisons of neutral loci to genetic correlations within and between populations and found a strong signal for divergent selection, suggesting that specific combinations of floral display and reward traits may be the targets of selection. Our results show little support for historical subspecies categories, yet floral traits are more diverged than expected due to drift alone. Non‐neutral divergence for multivariate quantitative traits suggests that selection by pollinators is maintaining a correlation between display and reward traits.     

The pollination of Habenaria rhodocheila (Orchidaceae) in South China: When butterflies take sides

Habenaria is one of the largest terrestrial genera in the family Orchidaceae. Most field studies on Habenaria species with greenish–white and nocturnal scented flowers are pollinated by nocturnal hawkmoths and settling moths. However, H. rhodocheila presents reddish flowers lacking a detectable scent and fails to fit the moth pollination syndrome. We investigated the pollinators, breeding system, and functional traits of H. rhodocheila in South China and found that two diurnal swallowtail butterflies Papilio helenus and Papilio nephelus (Papilionidae) were the effective pollinators. When butterflies foraged for nectar in the spur, the pollinia became attached between the palpi. A triangular projected median rostellar lobe was found at the entrance (sinus) of the spur of H. rhodocheila. This lobe divided the spur opening into two entrances forcing butterflies to enter their proboscides through the left or right side. When the projection of median rostellar lobe was removed, the site of pollinium attachment changed to the eyes of the butterflies, leading to a higher rate of pollinium removal but lower rate of pollinium deposition. Our quartz glass cylinder choice experiment suggested that visual rather than olfactory cues provided the major stimuli for butterflies to locate these flowers. Hand pollination experiments suggested this species was self‐compatible but pollinator‐dependent. However, the proportion of seeds with large embryos produced in self‐pollinated fruits was significantly lower than in cross‐pollinated fruits, indicating a significant inbreeding depression. Unlike many other orchid species, fruit set was higher than rates of pollinium removal, indicating a high level of pollination efficiency in a species with friable pollinia. Shifts from moth to butterfly pollination in the genus Habenaria parallel other orchid lineages providing insights into the potential for pollinator‐mediated floral trait selection.     

Molecular differentiation of two vicariant neotropical tree species, Plathymenia foliolosa and P. reticulata (Mimosoideae), inferred using RAPD markers

 Plathymenia foliolosa and P. reticulata are two tropical tree species economically important for their high quality wood. P. foliolosa occurs mainly in areas of Atlantic Forest (a tropical forest) whereas P. reticulata occurs mainly in the Cerrado region (a savanna vegetation), both Brazilian biomes classified as hotspots for conservation priorities. In the present study RAPD markers were used to differentiate P. foliolosa from the vicariant species P. reticulata. Using 10 random primers it was possible to obtain 84 markers, of which 22 (26.2%) were present in just one of the two species. Nine of the ten primers were successful in discriminating between the species through the detection of diagnostic markers or band frequency differences. One of the populations of P. foliolosa, located in a region where the two species co-occur, presented several markers that were characteristic of P. reticulata, suggesting the occurrence of gene flow between the two species. Received September 3, 2001; accepted August 19, 2002 Published online: November 22, 2002 Addresses of the authors: Daniela Rodrigues Lacerda (dlacerda@dedalus.lcc.ufmg.br), Maria Bernadete Lovato (lovatomb@icb.ufmg.br), Maria Dolores Porto Acedo (acedomd@icb.ufmg.br), Dept. Biologia Geral, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, CP: 486, Belo Horizonte, MG, 31270-901, Brasil. José Pires de Lemos Filho (lemos@icb.ufmg.br), Dept. Botanica, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, CP: 486, Belo Horizonte, MG, 31270-901, Brasil.     

The evolution of key functional floral traits in the early divergent angiosperm family Annonaceae

Potential key functional floral traits are assessed in the species‐rich early divergent angiosperm family Annonaceae. Pollinators (generally beetles) are attracted by various cues (particularly visual, olfactory, and thermogenic), with pollinators rewarded by nectar (generally as stigmatic exudate), heat, and protection within the partially enclosed floral chamber. Petals sometimes function as pollinator brood sites, although this could be deceptive. Annonaceae species are self‐compatible, with outcrossing promoted by a combination of protogyny, herkogamy, floral synchrony, and dicliny. Pollination efficiency is enhanced by pollen aggregation, changes in anthesis duration, and pollinator trapping involving a close alignment between petal movements and the circadian rhythms of pollinators. Most Annonaceae flowers are apocarpous, with syncarpy restricted to very few lineages; fertilization is therefore optimized by intercarpellary growth of pollen tubes, either by stigmatic exudate (suprastylar extragynoecial compitum) or possibly the floral receptacle (infrastylar extragynoecial compitum). Although Annonaceae lack a distinct style, the stigmas in several lineages are elongated to form “pseudostyles” that are hypothesized to function as sites for pollen competition. Flowers can be regarded as immature fruits in which the ovules are yet to be fertilized, with floral traits that may have little selective advantage during anthesis theoretically promoting fruit and seed dispersal. The plesiomorphic apocarpous trait may have been perpetuated in Annonaceae flowers as it promotes the independent dispersal of fruit monocarps (derived from separate carpels), thereby maximizing the spatial/temporal distance between seedlings. This might compensate for the lack of genetic diversity among seeds within fruits arising from the limited diversity of pollen donors.     

THE EVOLUTION OF INSECT-MEDIATED SELF-POLLINATION FROM AN OUTCROSSING SYSTEM IN CALYPTRIDIUM (PORTULACACEAE)

Calyptridium monospermum (Portulacaceae) comprises two biological races visited by different groups of insects. One race, pollinated by a bumblebee, Bombus vosnesenskii (Apidae), is outcrossing, while the other race is pollinated casually by a variety of other insects whose visits result in insect-mediated self-pollination and geitonogamy within the inflorescence. Insect-mediated selfing is also the mode of pollination in the closely related species, C. umbellatum. While all plants of both species are self-compatible, insect visitation is necessary for good seed set. Selfing populations of C. monospermum exhibit many floral adaptations for insect-mediated self-pollination which parallel those of the selfing C. umbellatum, while retaining vegetative characteristics typical of C. monospermum. These floral traits involve flower color and odor, style and stamen orientation, size of stigmatic surfaces, number of pollen grains produced, density of open flowers on inflorescences, and presence or absence of protogyny. Artifical pollination experiments show that these floral adaptations are effective in increasing the probability of insect-mediated self-pollination. Insect-mediated selfing in C. umbellatum and in some populations of C. monospermum is encouraged and probably made necessary by the inconstant pollinating behavior of the insects which visit these plants, as shown by the large percentage of non-Calyptridium pollen recovered from the bodies of insects captured while visiting these plants.     

Post-pollination biochemical changes in the floral organs of<Emphasis Type="Italic">Rhynchostylis retusa</Emphasis> (L.) Bl. and<Emphasis Type="Italic">Aerides multiflora</Emphasis> Roxb. (Orchidaceae)

If left unpollinated, the flowers ofAerides multiflora (Roxb.) andRhynchostylis retusa (L.) Bl. can remain fresh for 17 and 24 d, respectively. However, they begin to wilt at 2 to 3 days after pollination (DAP) and 3 to 4 DAP, respectively, and become senescent at 5 DAP and 7 DAP, respectively. When measured at two developmental phases — Stage 1, start of wilting and Stage 2, progression to senescence — all the floral organs from pollinated flowers had higher contents of total soluble sugars, reducing sugars, and free amino acids than those from unpollinated flowers. A corresponding increase was noted in the activities of hydrolytic enzymes, i.e., α-amylase, β-amylase, and invertase, and proteolytic enzymes (proteases) in those organs. This indicated that signals related to pollination had up-regulated those activities, leading to a breakdown of complex molecules into simpler ones for mobilization. The amounts of sugars and enzyme activity were relatively greater in the pollinated flowers ofA. multiflora compared withR. retusa, and levels were always higher in the floral lips and perianths. When inhibitors of auxin (0.25 mM TIBA) or ethylene (0.25 mM AgNO₂) were applied to the pollinated flowers, their senescence was partially prevented, thus signifying hormonal involvement in governing the pollination-induced biochemical alterations normally found in those organs.     

Day–night fluctuations in floral scent and their effects on reproductive success in <Emphasis Type="Italic">Lilium auratum</Emphasis>

We examined the contribution of diurnal and nocturnal pollination to male and female reproductive success in Lilium auratum. Plants were bagged for either 12 h during the day or at night to allow either only nocturnal or only diurnal visitors to forage throughout the flowering period. We found that there was no significant difference in the seed:ovule ratio among diurnally pollinated, nocturnally pollinated, or control flowers. However, in terms of male reproductive success, it was more advantageous for the plants to be pollinated both diurnally and nocturnally: the numbers of pollen grains remaining in diurnally pollinated or nocturnally pollinated flowers were significantly greater than those in control flowers. The total amount of floral volatiles of L. auratum was significantly higher at night than during the day. The constituents of floral scent of all time series examined were mostly monoterpenoids, many of which serve as attractants for nocturnal hawkmoths. Such nocturnally biased floral scent emission of L. auratum might achieve male reproductive success by attracting nocturnal visitors, which may suggest that the relative contribution of floral scent in this species is biased towards male reproductive success.     

Effect of pollination success on floral longevity in the orchid Calypso bulbosa (Orchidaceae)

The lifespan of an individual flower is often affected by pollination success. Species differ regarding whether male function (pollen removal), female function (pollen deposition), or both trigger floral senescence. We studied senescence in the singleflowered, deceptive orchid Calypso bulbosa by manipulating the degree of male and female reproductive success. We found that deposition of any amount of pollen resulted in dramatic changes in shape and color within 4 d, whereas unmanipulated flowers and those that had had pollinia removed remained unchanged for 8-11 d after treatment. Selection may favor the reproductive function that is less easily satisfied as the trigger for senescence, because a flower that senesces after accomplishment of this function is likely to have already succeeded at the more easily satisfied one. Deceptive (i.e., rewardless) flowers are more likely to satisfy male than female function since the latter requires that a pollinator be fooled twice, first to pick up pollen and second to deposit it. A survey of naturally pollinated Calypso showed that male function, pollinium removal, was more likely to occur than female function, deposition (95% vs. 66% of visited flowers); thus floral senescence in Calypso is triggered by achievement of the function less likely to succeed. Studies of senescence triggers in species in which female function is more likely to be achieved than male are necessary to further test this hypothesis.