两种生境对四川黄龙沟少花鹤顶兰有性繁殖成功的影响

四川黄龙沟森林和钙化滩流地两种生境中分布着大量的少花鹤顶兰,而且生长在这两种生境中的少花鹤顶兰有性繁殖成功存在着差异,目前尚不清楚这种差异是否是由生境差异所致,及其影响有性繁殖成功的机制。通过对黄龙沟少花鹤顶兰两种生境主要环境因子的调查和两种生境中的花部特征和传粉情况的测量,结果表明森林生境中林木盖度、土壤厚度、土壤含水量、土壤有机质、土壤全氮以及空气相对湿度比钙化滩流地中的高,而灌木盖度、草本盖度、苔藓盖度、空气温度、相对光照强度、土壤温度、全钙以及pH则是钙化滩流地中更高。而两种生境中少花鹤顶兰的形态特征和有性繁殖成功之间也存在明显的差异。在森林生境中的个体与钙化滩流地中的个体相比,最大叶长和最大叶宽更大,花朵数更多,距长也更长,单花和花序寿命更长。但花粉移走率和结实率却是钙化滩流地中的更高。两种生境中温度和光照的差异可能是导致两种生境中少花鹤顶兰花粉移走率和结实率差异的重要原因。     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Reproductive biology of an alpine orchid Phaius delavayi

The genus Phaius is distributed mainly in pantropical areas, and most species reported are autogamous. In contrast, Phaius delavayi, an alpine plant endemic to temperate zones of China, is strictly pollinator dependent. Only female workers and males of the bumblebee species B. hypnorum and B. lepidus are effective pollinators. The pollinaria are deposited on top of the head or the prothorax of the pollinators. The different depositions of pollinaria are regarded as an adaptation of the chamber dimensions of the flower to the size of the pollinator. The natural fruit set reported here (24.6–37.1%) suggests that pollination is successful in the deceptive P. delavayi. The yellow perianth and sweet floral scent of P. delavayi are important cues in attracting pollinators, while flowering time and mass population are associated with the process of “learning and avoidance” of the rewardless flowers. This orchid, therefore, employs a deceptive pollination strategy with diverse pollinators and pollinaria depositions rather than autogamy as do its congeners.     

Plant Parenthood in Milkweeds: A Direct Test of the Pollen Donation Hypothesis

Abstract For hermaphroditic plant species whose fruit production is limited by maternal resources, the "pollen donation hypothesis" views large floral displays as an adaptation to enhance the probability of fathering seeds on other plants. This hypothesis has frequently been used to describe the evolution of large floral displays in milkweeds ( Asclepias ). Most tests of the pollen donation hypothesis, however, have used indirect measures, such as flower production or pollen removal, to estimate male reproductive success. To test the pollen donation hypothesis directly, we performed a paternity analysis and determined the number of seeds sired by individual genotypes in a natural population of poke milkweed, A. exaltata , in southwestern Virginia. Seeds sired (male success) and seeds produced (female success) were significantly correlated with flower number per plant (for male success: r = 0.32, P < 0.05; for female success: r = 0.66, P < 0.0001). Functional gender of plants that reproduced both as males and females (N = 17) was not correlated with flower number per plant ( r = 0.35, P>0.05), indicating that plants with large floral displays did not reproduce primarily as males. Percent fruit-set and seed number per fruit were higher in 1986, when levels of pollinarium removal also were higher. Furthermore, several umbels that experienced high pollinator activity selectively matured fruits that contained many seeds. We argue that the evolution of large floral displays in milkweeds is the result of selection to increase overall reproductive success rather than male reproductive success alone.     

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.

     

Functional differentiation in pollination processes among floral traits in Serapias species (Orchidaceae)

Floral displays, influencing attractiveness to insects, increase the number of pollinator visits and the efficiency of each visit in terms of pollen exchange and thus affect the plant reproductive success. Here, we conducted an in situ manipulation experiment to investigate whether the floral modifications affect reproductive success in natural orchid populations of Serapias lingua and Serapias vomeracea. We estimated male and female reproductive success of three treatment groups, disassembly of floral tube, cutting of lip, and painting of the callus surface, in terms of pollinaria removed/deposited and fruit production. Results revealed that phenotypic modification had opposite effects on reproductive success of two examine species. Indeed, reproductive success was significantly increased by the detached of the petals and sepals, and decreased, due to callus painting and lip removal, in S. lingua. On the contrary, unmanipulated plants of S. vomeracea showed significantly higher value of pollinaria removed and deposited and fruit set than manipulated ones. The differences between S. lingua and S. vomeracea agree to the different pollination strategy of examined species. S. vomeracea shows shelter imitation strategy, and thus, the disassembly of tunnel‐like corolla does not allow the insects to use the flower as a refuge, while S. lingua is a sexually deceptive orchid and therefore the opening of the flower made more visible callus (visible at a greater distance) increasing the pollinators attraction. This study provides evidence that pollinators were largely sensitive to the experimental modification of the flower phenotype, which is consistent with the presence of significant selection on individual floral characters. Our experimental investigations of the effects of variation in display on pollinator visitation provide insights into the evolution of floral morphology in orchid with shelter imitation strategy.     

Phenotype–fitness relationships and pollen-transfer efficiency of five orchid species with different pollination strategies

Premise

Deceptive pollination, a fascinating mechanism that independently originated in several plant families for benefiting from pollinators without providing any reward, is particularly widespread among orchids. Pollination efficiency is crucial in orchids due to the aggregated pollen in a pollinarium, which facilitates pollen transfer and promotes cross-pollination as pollinators leave after being deceived.

Methods

In this study, we compiled data on reproductive ecology from five orchid species with different pollination strategies: three deceptive-strategy species (shelter imitation, food deception, sexual deception), one nectar-rewarding species, and one shelter-imitation but spontaneously selfing species. We aimed to compare the reproductive success (female fitness: fruit set; male fitness: pollinarium removal) and pollination efficiency of species representing these strategies. We also investigated pollen limitation and inbreeding depression among the pollination strategies.

Results

Male and female fitness were strongly correlated in all species but the spontaneously selfing species, which had high fruit set and low pollinarium removal. As expected, pollination efficiency was highest for the rewarding species and the sexually deceptive species. Rewarding species had no pollen limitation but did have high cumulative inbreeding depression; deceptive species had high pollen limitation and moderate inbreeding depression; and spontaneously selfing species did not have pollen limitation or inbreeding depression.

Conclusions

Pollinator response to deception is critical to maintain reproductive success and avoid inbreeding in orchid species with non-rewarding pollination strategies. Our findings contribute to a better understanding of the trade-offs associated with different pollination strategies in orchids and highlight the importance of pollination efficiency in orchids due to the pollinarium.

     

A Sexually Dimorphic Corolla Appendage Affects Pollen Removal and Floral Longevity in Gynodioecious Cyananthus delavayi (Campanulaceae)

The floral traits of bisexual flowers may evolve in response to selection on both male and female functions, but the relative importance of selection associated with each of these two aspects is poorly resolved. Sexually dimorphic traits in plants with unisexual flowers may reflect gender-specific selection, providing opportunities for gaining an increased understanding of the evolution of specific floral traits. We examined sexually dimorphic patterns of floral traits in perfect and female flowers of the gynodioecious species Cyananthus delavayi. A special corolla appendage, the throat hair, was investigated experimentally to examine its influences on male and female function. We found that perfect flowers have larger corollas and much longer throat hairs than female flowers, while female ones have much exerted stigmas. The presence of throat hairs prolonged the duration of pollen presentation by restricting the amount of pollen removed by pollen-collecting bees during each visit. Floral longevity was negatively related to the rate of pollen removal. When pollen removal rate was limited in perfect flowers, the duration of the female phases diminished with the increased male phase duration. There was a weak negative correlation between throat hair length and seed number per fruit in female flowers, but this correlation was not significant in perfect flowers. These results suggest that throat hairs may enhance male function in terms of prolonged pollen presentation. However, throat hairs have no obvious effect on female function in terms of seed number per fruit. The marked sexual dimorphism of this corolla appendage in C. delavayi is likely to have evolved and been maintained by gender-specific selection.     

四川黄龙沟少花鹤顶兰繁殖成功特征

少花鹤顶兰(Phaiu delavayi)是多年生、多次结实的具有克隆能力的地生兰科植物,是中国特有种。目前少花鹤顶兰开花物候、花寿命、繁殖成功等生物学特征尚不清楚,而这些资料是开展少花鹤顶兰进化和保护生物学研究的基础。利用2005-2007年四川黄龙沟少花鹤顶兰的调查数据,对少花鹤顶兰繁殖成功特征进行了研究。结果表明黄龙沟少花鹤顶兰每个花序的平均花朵数为3(1-7)朵。叶和花的形态指标在年季间无显著变异。2005-2007年3年间的开花物候没有明显的差异,花期始于6月中旬,于7月底至8月初结束,持续约6-7周,80%-90%的花在7月初的开花高峰期开放,属花集中开放模式。花寿命与是否授粉密切相关,成功授粉的花寿命比没有授粉的花寿命短。少花鹤顶兰是自交亲和的,但其结实完全依赖于两种熊蜂的传粉,不存在自动自花授粉和无融合生殖现象。柱头可授性和花粉活力可维持15 d左右。繁殖成功率年季间的变化很大,并表现出逐年下降的趋势,花粉移走率为18%-51%,自然结实率为10%-36%。繁殖成功下降的趋势可能与黄龙沟喷洒农药有关。     

Floral visual signal increases reproductive success in a sexually deceptive orchid

Sexually deceptive orchids mimic signals emitted by female insects in order to attract mate-searching males. Specific attraction of the targeted pollinator is achieved by sex pheromone mimicry, which constitutes the major attraction channel. In close vicinity of the flower, visual signals may enhance attraction, as was shown recently in the sexually deceptive orchid Ophrys heldreichii. Here, we conducted an in situ manipulation experiment in two populations of O. heldreichii on Crete to investigate whether the presence/absence of the conspicuous pink perianth affects reproductive success in two natural orchid populations. We estimated reproductive success of three treatment groups (with intact, removed and artificial perianth) throughout the flowering period as pollinaria removal (male reproductive success) and massulae deposition (female reproductive success). Reproductive success was significantly increased by the presence of a strong visual signal??the conspicuous perianth??in one study population, however, not in the second, most likely due to the low pollinator abundance in the latter population. This study provides further evidence that the coloured perianth in O. heldreichii is adaptive and thus adds to the olfactory signal to maximise pollinator attraction and reproductive success.     

Does flower color variation matter in deception pollinated<Emphasis Type="Italic"> Psychilis monensis</Emphasis> (Orchidaceae)?

Negative frequency dependent selection (FDS) had been proposed as a mechanism for the maintenance of the high levels of variability in floral traits of rewardless plants. Thus far the evidence has been equivocal for discontinuous traits. We experimentally tested the FDS hypothesis for continuous variation in flower color of Psychilis monensis, a rewardless, epiphytic orchid of Mona Island, Puerto Rico. P. monensis flowers all year long with a peak in June and July. Plants are self-incompatible. Individual flowers last 9–15 days if unpollinated. Over a 9-month observation of an unmanipulated population, a high percentage of plants showed pollinarium removals (79%) and fruit initiation (63%), but the actual percentage of flowers visited was very low (12%) and final fruit set was as low as the lowest recorded for orchids (2.4%). In a FDS experiment using a Latin Square design, we manipulated flower color in three populations. Over 50% of the variation in either male or female reproductive success was explained by time and site with no significant effect of treatment except as part of a three-way interaction of time × site × treatment. Paired comparisons with the controls gave equivocal results for both male and female measures of reproductive success. Major community changes had occurred during the FDS experiment with flower activity falling dramatically and by the third run of the experiment, only P. monensis was in flower. Coincidentally, the numbers of effective visits increased with time, presumably as pollinators became less discriminating in search of new food resources. Thus, negative frequency dependent selection is either sporadic or non-existent. Reproductive success was statistically related to locality and date, which was reflected in the flowering phenology of the local communities. High natural levels of color variation may be more influenced by drift than selection.     

Which food‐mimic floral traits and environmental factors influence fecundity in a rare orchid,Calanthe yaoshanensis?

Orchid species that are food mimics produce fewer fruits than species offering rewards, but few studies have shown the impact of environmental factors (e.g. anthropogenic activity, frost and herbivores) on their reproductive success over several seasons. In this study, we focused on the sole population of the endangered Calanthe yaoshanensis as it secretes no nectar. We investigated its floral biology, fruit set rates and prevailing environmental factors over three seasons (2008–2010). Mechanical self‐pollination did not occur in C. yaoshanensis, but hand‐selfed and crossed flowers produced equal numbers of fruit. However, seed viability and embryo size were significantly higher in cross‐pollinated fruits maximizing embryonic fitness. Large hoverflies (Syrphidae) and Bombus patagiatus (gynes) were the only pollinarium vectors, but they often failed to disperse pollinaria. We interpret the temporary retention of the anther cap over the pollinarium as an adaptation lowering self‐pollination. Insect‐mediated rates of pollinarium removal were always higher than rates of pollinia deposition on stigmas. Over 3 years, natural rates of pollinarium removal differed significantly, whereas natural rates of fruit set were not significantly different (< 22%). Climate, herbivory and anthropogenic collections also inhibited some fruit set and maturation. Both biotic and abiotic factors appear to lower the fecundity of this endangered population. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 421–433.     

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     

Effects of floral display size on male and female reproductive success in Mimulus ringens

Background and Aims

The number of flowers blooming simultaneously on a plant may have profound consequences for reproductive success. Large floral displays often attract more pollinator visits, increasing outcross pollen receipt. However, pollinators frequently probe more flowers in sequence on large displays, potentially increasing self-pollination and reducing pollen export per flower. To better understand how floral display size influences male and female fitness, we manipulated display phenotypes and then used paternity analysis to quantify siring success and selfing rates.

Methods

To facilitate unambiguous assignment of paternity, we established four replicate (cloned) arrays of Mimulus ringens, each consisting of genets with unique combinations of homozygous marker genotypes. In each array, we trimmed displays to two, four, eight or 16 flowers. When fruits ripened, we counted the number of seeds per fruit and assigned paternity to 1935 progeny.

Key Results

Siring success per flower declined sharply with increasing display size, while female success per flower did not vary with display. The rate of self-fertilization increased for large floral displays, but siring losses due to geitonogamous pollen discounting were much greater than siring gains through increased self-fertilization. As display size increased, each additional seed sired through geitonogamous self-pollination was associated with a loss of 9·7 seeds sired through outcrossing.

Conclusions

Although total fitness increased with floral display size, the marginal return on each additional flower declined steadily as display size increased. Therefore, a plant could maximize fitness by producing small displays over a long flowering period, rather than large displays over a brief flowering period.     

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     

Analysis of trait–performance–fitness relationships reveals pollinator-mediated selection on orchid pollination traits

Premise

The role of pollinators in evolutionary floral divergence has spurred substantial effort into measuring pollinator-mediated phenotypic selection and its variation in space and time. For such estimates, the fitness consequences of pollination processes must be separated from other factors affecting fitness.

Methods

We built a fitness function linking phenotypic traits of food-deceptive orchids to female reproductive success by including pollinator visitation and pollen deposition as intermediate performance components and used the fitness function to estimate the strength of pollinator-mediated selection through female reproductive success. We also quantified male performance as pollinarium removal and assessed similarity in trait effects on male and female performance.

Results

The proportion of plants visited at least once by an effective pollinator was moderate to high, ranging from 53.7% to 85.1%. Tall, many-flowered plants were often more likely to be visited and pollinated. Given effective pollination, pollen deposition onto stigmas tended to be more likely for taller plants. Pollen deposition further depended on traits affecting the physical fit of pollinators to flowers (flower size, spur length), though the exact relationships varied in time and space. Using the fitness function to assess pollinator-mediated selection through female reproductive success acting on multiple traits, we found that selection varied detectably among taxa after accounting for sampling uncertainty. Across taxa, selection on most traits was stronger on average and more variable when pollination was less reliable.

Conclusions

These results support pollination-related trait–performance–fitness relationships and thus pollinator-mediated selection on traits functionally involved in the pollination process.     

Influences of the genetic neighborhood on ramet reproductive success in a clonal desert cactus

Clonal structure in clonal plants can affect sexual reproduction. Individual ramets can decrease reproduction if their neighbors are ramets of the same genet due to inbreeding depression or self-incompatibility. We assessed ramet reproductive success in the partial self-incompatible Ferocactus robustus (Cactaceae) as a function of floral display size in focal ramets and floral display size and clonal structure of their reproductive neighborhoods. Ramets were labeled, sized in number of stems, mapped and genetically identified through RAPD markers in one population. A pollen dispersal area of 15-m radius was established for each ramet to determine the clonal diversity in the neighborhoods. Flower production and fruit set were counted on a monthly basis during one reproductive season as a surrogate of ramet fitness. We expected a decrease in individual ramet reproductive success as a function of the number of reproductive ramets of the same genet in the neighborhood. A total of 272 sampled ramets revealed 116 multilocus genotypes, showing high clonal diversity in the population (G/N = 0.43, D = 0.98). Clonal diversity of neighborhoods ranged from 0.06 to 1 and fruit set varied from 0 to 76.9%. Individual ramet reproductive success was influenced by (1) mate availability, (2) floral display size of a genet within the reproductive neighborhood, and (3) the proportion of distinguishable genotypes. Floral display size of genets and ramets coupled with the genetic diversity within the reproductive neighborhood determines the low sexual reproduction in F. robustus.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Variation in pollinator abundance and selection on fragrance phenotypes in an epiphytic orchid

Extraordinary floral variation is common among some orchids that employ deception to attract pollinators. This variation may be maintained by frequency-dependent selection where rare phenotypes are preferred. Over a 2-yr period, 1993-1994, we monitored the reproductive success of Tolumnia variegata, an obligately outcrossing epiphytic orchid, at three localities in Puerto Rico that differed in pollinator service. Plants varied in floral morphology and fragrance characteristics. Artificial arrays of varying frequencies of scentless and fragrant phenotypes were established to test for frequency-dependent selection. Where pollinators were rare (Cambalache, range of census average = 0-0.2 bees/h), 0.9-1.2% of the flowers were effectively visited (pollinarium removals and pollinations). At Tortuguero where 0.4-1.1 bees/h were observed, 4-9.2% of the flowers were visited. At Pi;atnones where bees were the most abundant (1.4-5.2 bees/h), 20.9-25.0% of the flowers were visited. A significant portion of the variance in all measures of reproductive success (male, female, and combined) was explained by differences among populations, which we attribute mostly to variation in pollinator abundance. Neither the fragrance phenotype nor its frequency had a significant effect on success as revealed by a split-plot ANOVA. There was a significant interaction between population and phenotypic frequencies in all our measures of reproductive success, but only for the 1994 flowering season. Thus, variation in floral fragrance phenotypes is not likely maintained by frequency-dependent selection. High levels of variation remain unexplained.     

Spatially inconsistent direct and indirect effects of herbivory on floral traits and pollination success in a tropical shrub

Investigations on plant–animal interactions have traditionally focused on single interactions at a time (e.g. herbivory, pollination), yet plant fitness is generally influenced in complex ways by several interactions operating concurrently, and very little is known on the degree of spatial consistency of the direct and indirect effects that link different interactions. This paper evaluates experimentally whether direct and indirect effects of herbivory on male and female flower size and pollination success of the monoecious tropical shrub Cnidoscolus aconitifolius (Euphorbiaceae) remain consistent at three distant regions in Yucatan (southeastern Mexico). Plants were subjected to different levels of leaf defoliation, and treatment effects on floral traits (corolla area, corolla tube length, pollen production), and male and female components of pollination success (percent pollen removal, number of pollen tubes) were subsequently measured to evaluate the indirect effect of herbivory on plant reproductive success via pollination. Defoliation had significant direct effects on floral traits, but its indirect effects differed between the male and female components of pollination success. Moreover, the relationships between defoliation, floral traits and male and female pollination success varied spatially (i.e. between regions), although they were frequently more spatially‐consistent for male success than for female success. Results from this study stress the importance of explicitly testing for spatial variation in direct and indirect effects arising from plant‐animal interactions.