Lying to Pinocchio: floral deception in an orchid pollinated by long-proboscid flies

Plants that lack floral rewards may nevertheless attract pollinators if their flowers sufficiently resemble those of rewarding plants. Flowers of the South African terrestrial orchid Disa nervosa are similar in floral dimensions and spectral reflectance to those of a sympatric nectar-producing irid ( Watsonia densiflora s.l. ). Observations showed that the orchid and Watsonia share the same pollinator, a long-proboscid tabanid fly Philoliche aethiopica . These flies visited inflorescences of both species during their foraging bouts and most (64%) observed or captured on Watsonia inflorescences carried pollinaria of the orchid on their proboscides. They probe an average of 6.3 flowers on Watsonia inflorescences, but just 1.9 flowers on the Disa inflorescences, a behaviour which would strongly promote cross-pollination in the self-compatible orchid. The orchid generally achieves high levels of pollination success, with approximately 50% of flowers receiving or exporting pollen at some sites. Pollination success was also high at one site that lacked Watsonia plants, suggesting that the orchid does not have an obligate dependence on Watsonia . Its pollination system may therefore be characterized as intermediate between generalized food deception and specific floral mimicry.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 271–278.     

Phytophagous insect oviposition shifts in response to probability of flower abortion owing to the presence of basal fruits

Phytophagous insects use a wide range of indicators or associated cues to avoid laying eggs in sites where offspring survival is low. For insects that lay eggs in flowers, these unsuitable sites may be created by the host plant's resource allocation to flowers. In the sequentially flowering host plant, Yucca glauca, late‐opening distal flowers are more likely to be aborted in the presence of already‐initiated basal fruits because they are strong resource sinks. If flowers are aborted, all eggs of the phytophagous insect, Tegeticula yuccasella, within the flower die. We used the phytophagous insect T. yuccasella that lays eggs in and pollinates host plant Y. glauca flowers to test the hypothesis that phytophagous insect females are less likely to invest eggs in host plant flowers if basal fruits are present because they are more likely to be aborted. We also investigated potential predictors of arrival of T. yuccasella at inflorescences at the onset of flowering. These factors may influence a phytophagous insect's decisions to select oviposition sites. We carried out a behavioral experiment using wild‐caught T. yuccasella females on manipulated inflorescences with distal flowers with basal fruits and without fruits. As potential predictors of T. yuccasella arriving at inflorescences, we used floral display size and day of onset of flowering. In support of our hypothesis, our experimental results showed that T. yuccasella was significantly less likely to oviposit in distal flowers on inflorescences with basal fruits. We also found that T. yuccasella arrival was higher at inflorescences with larger floral display size and earlier in the flowering season. These findings uncover a novel indicator of unsuitable oviposition sites—the presence of basal fruits, that phytophagous insects use to make oviposition decisions. Further, our study contributes to the growing body of evidence that shows that females prefer sites that increase the probability of survival of their offspring.     

Lack of floral nectar reduces self-pollination in a fly-pollinated orchid

One explanation for the widespread absence of floral nectar in many orchids is that it causes pollinators to visit fewer flowers on a plant, and thus reduces self-pollination. This, in turn, could increase fitness by reducing inbreeding depression in progeny and promoting pollen export. The few previous investigations of this hypothesis have all involved bee-pollinated orchids and some have given contradictory results. We studied the effects of adding artificial nectar (sucrose solution) to the spurs of a non-rewarding long-proboscid fly-pollinated orchid, Disa pulchra. Addition of nectar significantly increased the number of flowers probed by flies (2.6-fold), the time spent on a flower (5.4-fold), the number of pollinia removed per inflorescence (4.8-fold) and the proportion of removed pollen involved in self-pollination (3.5-fold). The level of self-pollination increased dramatically with the number of flowers probed by flies. Experimental self-pollination resulted in fruits with only half as many viable seeds as those arising from cross-pollination. Pollinators were more likely to fly long distances (>40 cm) when departing from non-rewarding inflorescences than when departing from rewarding ones. These findings provide support for the idea that floral deception serves to reduce pollinator-mediated self-pollination.     

Experimental and phylogenetic evidence for floral mimicry in a guild of fly-pollinated plants

Mimicry, as an adaptive explanation for the resemblance between organisms, is not always readily distinguishable from, inter alia , coincidence, shared ancestry, or convergent evolution. We tested the hypothesis that two rare South African orchid taxa Brownleea galpinii ssp. major (nectar-producing) and Disa cephalotes ssp. cephalotes (non-rewarding) are mimics of the nectar-producing flowers of a relatively common species, Scabiosa columbaria (Dipsacaceae), with which they always occur sympatrically. Flowers of the orchids were apparently unscented and had similar dimensions and almost identical spectral reflectance to the flowers of Scabiosa . The orchids were pollinated exclusively by long-proboscid flies (Tabanidae and Nemestrinidae) that feed mainly on nectar in Scabiosa flowers. Choice experiments showed that these flies did not discriminate between the orchids and Scabiosa when alighting on their flat-topped inflorescences. However, flies were not attracted to related orchids dissimilar to Scabiosa , or to inflorescences of B. galpinii that had been artificially reconstructed in the shape of a spike, rather than a flat-topped capitulum. A phylogenetic analysis showed traits that give the orchids a resemblance to Scabiosa , such as a flat-topped inflorescence and cream floral colouration with dark spots and short spurs, to be mostly apomorphic features, and therefore likely to be relatively recent adaptations for mimicry. We caution that the term mimic should not be applied to species whose resemblance to another species is due entirely to plesiomorphic traits that, in all likelihood, evolved prior to the ecological association.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 289–304.     

Causes of spatial patterns of fruit set in waratah: Temporal vs. spatial interactions between flowers on an inflorescence

Spatial patterns of fruit set within inflorescences may be controlled by pollination, nutrient allocation, or inflorescence architecture. Generally, flowers that have spatial and/or temporal precedence are more likely to set fruits. We sought to separate these factors by comparing patterns of fruit set on inflorescences of two species of Telopea (Proteaceae); one that flowers from the tip to the base of the rachis, the other from base to tip. In both species, most fruits were set at the top of the inflorescence (the last flowers to open for T. speciosissima) and this was extreme for T. mongaensis, where the top flowers open first. Fruit set was not generally limited by inadequate pollination for either T. mongaensis or T. speciosissima, as hand pollinations did not increase fruit set and many abscised flowers contained pollen tubes. In T. speciosissima, we tested whether removal of developing topmost fruits would ‘release’ those that had initiated but not yet aborted lower down. There was no significant effect. Plant hormones can increase the degree to which a developing fruit is a sink for nutrients, so we applied cytokinin to the developing lower fruits on some inflorescences. There was no significant effect of the hormone treatment. We conclude that temporal precedence may contribute to the skewed pattern of fruit set in T. mongaensis, because there was an extreme concentration of fruit set on the distal part of the inflorescences, but it cannot explain this pattern of fruit set in T. speciosissima, where the distal flowers are the last to open. Some other process must therefore constrain fruit set to the topmost flowers in an inflorescence. While cytokinin application had no significant effect, the power of this experiment was low and we consider that the hypothesis of hormonal control is worth further exploration.     

Pollination mutualism between Alocasia macrorrhizos (Araceae) and two taxonomically undescribed Colocasiomyia species (Diptera: Drosophilidae) in Sabah, Borneo

Two taxonomically undescribed Colocasiomyia species were discovered from inflorescences of Alocasia macrorrhizos in Kota Kinabalu City, Sabah, Borneo, Malaysia. The aims of this study were to investigate the reproductive ecology of the flies and the plant, ascertain the importance of the flies as pollinators and examine the intimate association between flowering events and life history of the flies. We conducted sampling, observations and field pollination experiments. The flies were attracted by the odour of female‐phase inflorescences in the early morning on the first day of anthesis. They fed, mated and oviposited in the inflorescences for 1 day. On the second day, the flies, covered with pollen grains, left the male‐phase inflorescences for the next female‐phase inflorescences. The immature forms of both fly species hatched, developed and pupated within the infructescences without damaging the fruits, and developed adults emerged when the mature infructescences dehisced. The flowering events and fly behaviours were well synchronized. In field pollination experiments, inflorescences bagged with a fine mesh (insect exclusion) produced almost no fruits, whereas those bagged with a coarse mesh (bee exclusion) produced as many fruits as the open‐pollinated controls. These results indicate that these flies are the most efficient and specialised pollinators for their host, A. macrorrhizos. These flies, in return, depend on A. macrorrhizos for food and habitat through most of their life cycle. This study provides a deeper insight into the less recognised, highly intimate pollination mutualism between Araceae plants and Colocasiomyia flies.     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

A gradual reduction of female structures in the pistillate flowers ofRicinus communis L. (castor-bean) with chlorflurenol (morphactin)

Chlorflurenol (morphactin) EMD 7301 W (60, 120 and 240 mg I^?1) when applied at five to seven leaf and 10–12 leaf stage induced hermaphrodite flowers in the inflorescences ofRicinus communis. The hermaphrodite flowers were mostly apical in position unlike the apical female flower in the control. With an increase in the time gap between sowing and emergence of the inflorescence there was a reduction of the female structures in the hermaphrodite flowers. The last formed inflorescences had an apical male flower in place of a female. The largest number of inflorescences with an apical male or hermaphrodite flower were produced with 60 mg 1^?1 applied at five to seven leaf stage. The hermaphrodite flowers failed to set fruits.     

A REEXAMINATION OF THE POLLEN-DONATION HYPOTHESIS IN AN EXPERIMENTAL POPULATION OF ASCLEPIAS EXALTATA

The evolution of large floral displays in hermaphroditic flowering plants has been attributed to natural selection acting to enhance male, rather than female, reproductive success. Proponents of the “pollen-donation hypothesis” have assumed that maternal resources, rather than levels of effective pollination, limit fruit set. We investigated the pollen-donation hypothesis in an experimental population of poke milkweed, Asclepias exaltata, where effective pollination did not limit fruit set. Specifically, we examined the effects of flower number per plant, and flower number per umbel on male reproductive success (number of fruits sired) and female reproductive success (number of fruits matured). In 1990, a paternity analysis was performed on fruits collected from 53 plants whose inflorescences were not manipulated. Flower number per plant was significantly correlated with male success, but not with plant gender. Flower number per plant was also significantly correlated with female success, but umbel number and stem number per plant together explained more than half (58%) the variation in female success. The percentage of fruit set was not significantly correlated with flower number per plant. Plants with large floral displays did not disproportionately increase in male reproductive success, relative to female success, as predicted by the pollen-donation hypothesis. In 1991, the effect of flower number per umbel on male and female reproductive success was investigated. Flower number per umbel was manipulated on four umbels per plant by removing flowers to leave 6, 12, or 18 flowers in each umbel. Plants with the largest umbels effectively pollinated twice as many flowers on other plants, but produced only 1.35 times as many fruits as plants with 6 and 12 flowers per umbel. Relative maleness of plants with large umbels was nearly twice that of small and medium umbels. Although these observations are consistent with the pollen-donation hypothesis at the level of umbels, they are problematic, because much of the variation in flower number per umbel exists within, rather than among, plants in natural populations. Thus, plants consist of both reproductively male (large) and female (small) inflorescences, which act to increase total reproductive success. It is therefore inappropriate to explain the evolution of large floral displays in milkweeds solely in terms of potential male reproductive success.     

The consequences of rewardlessness in orchids: reward-supplementation experiments with Anacamptis morio (Orchidaceae)

Pollinators are expected to respond to low reward availability in an inflorescence by visiting fewer flowers before departure, thus potentially causing reduced visitation, but also reduced geitonogamous selfing. I tested this hypothesis using Anacamptis morio, an orchid that does not reward its pollinators. Supplementation of inflorescences with artificial nectar did not result in an increase in fruit set on supplemented inflorescences compared to control inflorescences and tended to reduce pollinia removal. Supplementation resulted in reduced fruit quality, but there was no evidence that this was as a result of inbreeding depression. Behavioral experiments showed that pollinating bumble bees, as predicted, visited more flowers on supplemented inflorescences. Bumble bees also deposited more self-pollen on supplemented inflorescences, but this was marginally significant. Bumble bee queens removed significantly more pollinia from control inflorescences, while Bombus terrestris and B. lucorum workers did not. I conclude that while pollinators behaved as predicted, there was weak evidence that pollinia removal, pollen deposition, and fruit set followed the predictions of the hypothesis. I argue that this was probably because some pollinators were more efficient at removing and depositing pollen on control inflorescences, while others were not.     

Variation in fruit- and seed set among and within inflorescences of Melampyrum roseum var. japonicum (Scrophulariaceae)

Abstract Variations in fruit set and seed set among and within inflorescences of the annual herb, Melampyrum roseum var. japonicum, were studied. Under natural conditions, although the mean fruit set was slightly different among inflorescences, the mean seedset was not significantly different among inflorescences within the plants. In constrast, within the inflorescences, the flowers located at a lower position of the inflorescence and which opened earlier showed higher fruit set than those at a higher position and which opened later. However, the seed set of matured capsules were not significantly different from each other, regardless of the position of flowers within the inflorescences. Patterns of the fruit- and seed set under open pollination indicated that variation in seed reproduction of M. roseum is due to variation in fruit production. The results of clipping experiments of flowers revealed that there was no functional limitation in seed production among flowers located at various positions within the inflorescence. It seemed that the variation in the fruit set within the inflorescences of M. roseum was not attributable to 'architectural effects'. Reduction of the number of flowers within the inflorescences resulted in an increase of fruit set and seed weight, indicating that the flowers in an inflorescence compete for resources. This phenomenon supports the 'resource competition hypothesis', and variation in fruit set within the inflorescence is attributable to competition among flowers within the inflorescence for limited resources. Consequently, it was concluded that, under natural conditions, the early blooming flowers located at lower positions of the inflorescences obtain more resources and produce more fruits than the late blooming flowers located at higher positions in M. roseum .     

Expression of IAA-Lysine Synthetase Gene Caused Variations of Flower Develop ment in Transgenic Tobacco

The indoleacetic acid-lysine synthetase (iaaL) gene of Pseudornonas syringae subsp. savastanoi was fused to a rice Rchl0 promoter and introduced into tobacco plants. The expression pattern of this chimeric Rchl0-iaaL gene in the transgenic plants was studied, and the endogenous levels of indoleacetic acid (IAA) were assayed with the high-performance liquid chromatography (HPLC) method. The results showed that the Rchl0 promoter could direct the high level expression of the iaaL gene in young inflorescences, fully bloomed flowers, and immature fruits, resulted in a decrease of the IAA levels by 32%–82%. The transgenic plants displayed decreased apical dominance, enlongated stamens, abnormal inflorescences and fruits. The transgenie plant pYL614-T14, in which the endogenous IAA levels in inflorescences were reduced by 82 %, displayed extremely abnormal inflorescence with only two flowers and a much higher potential of vegetative growth than that of the controls after flowering. These Rchl 0-iaaL gene transgenic tobacco plants would pro- vide valuable materials for further studies of physiological functions of IAA in reproductive development.     

COMPETITION-DEPENDENT ABSCISSION OF SELF-POLLINATED FLOWERS OF PHORMIUM TENAX (AGAVACEAE): A SECOND ACTION OF SELF-INCOMPATIBILITY AT THE WHOLE FLOWER LEVEL?

The relative success of fruit from paired self- and cross-pollinations was examined in Phormium tenax when the contrasted pollinations were separated by different distances on the same and different inflorescences. We determined whether the retention of selfed fruits differed from that of crossed fruits and whether it depended on the level of competition with crossed fruit, the number of seeds per fruit, and/or the presence of earlier developing fruit. We found that the success of selfed fruits is determined by the degree of competition with crossed fruits and may be an expression of self-incompatibility. Competition-dependence of the abscission of selfed flowers has not been documented previously. It is parallel to cryptic self-incompatibility in which individual self-pollen grains are not as successful as cross-pollen when competing on the same pistil. The competition-dependent abscission of self-pollinations considered here, however, operates at the level of whole flowers. The phenomenon of competition-dependent abscission of selfed flowers in P. tenax also has implications for the measurement and interpretation of self-incompatibility in other species. Self-incompatibility is a quantitative phenomenon. The facultative success of selfing shows that the effective strength of self-incompatibility can be highly susceptible to the conditions of competition under which it is measured. The competition-dependent abscission of selfed flowers allows a high level of outcrossing to be achieved while it assures seed set when pollinations are scarce. Several other causes of intermediate selfing frequencies can also be explained by this “best-of-both-worlds” hypothesis.     

ALLOCATION OF REPRODUCTIVE RESOURCES WITHIN AND AMONG INFLORESCENCES OF LAVANDULA STOECHAS (LAMIACEAE)

Patterns of fruit set were studied in Lavandula stoechas, a Mediterranean shrub commonly occurring in southern Spain. The small, hermaphroditic flowers of this shrub are aggregated into dense, headlike inflorescences and exhibit extensive variations in fecundity. It was shown that as the number of developing fruits in the inflorescence increased, the probability of a flower setting fruit, the size of seeds, and their germinability decreased, most likely because of strong within-inflorescence resource limitation. An experiment was designed to ascertain whether increased fertility in late-opening flowers could be induced through reallocation of reproductive resources between different inflorescences. The experiment consisted of removing half of the inflorescence buds from a set of plants and comparing their fecundity with that of intact individuals. Thinning did not increase the proportion of flowers setting fruit which, in fact, was slightly lower than that of intact individuals (probably due to some reduction of floral display brought about by thinning). Although treated plants produced heavier seeds than controls, results suggest that inflorescences of L. stoechas behave as autonomous modules among which resources cannot be reallocated. Predispersal seed predation by insects accounted on average for a 31% reduction in fruit set. Predation was found to be nonrandomly distributed within inflorescences, with most damage concentrated on late fruits (i.e., those with smaller and less germinable seeds).     

Temporal patterns of seed set and decelerating fitness returns on female allocation in Zigadenus paniculatus (Liliaceae), an andromonoecious lily

I describe temporal patterns of seed production in the andromonoecious lily Zigadenus particulatus. Fruit set per flower and seed set per fruit declined through time within plants. Hand pollination experiments showed that this was not due to increasing pollen limitation. Nutrient supplementation had little effect on seed output, but leaf clipping reduced seed production, especially in late-blooming flowers, and removal of early-blooming flowers increased seed set by later flowers. Thus, the temporal pattern of seed output was due to declining availability of photosynthates. Plants with larger bulbs produced larger inflorescences, a greater proportion of hermaphrodite flowers, more fruits per hermaphrodite flower, and more seeds per fruit, but lost a greater fraction of their initial bulb mass as a consequence of fruiting. After controlling for the effects of bulb mass, plants with larger inflorescences produced a greater proportion of male flowers, and plants with more hermaphrodite flowers produced fewer fruits per hermaphrodite flower and fewer seeds per fruit. Thus, the female fitness gain curve was decelerating. The temporal decline in seed output provides a partial explanation for the parallel decline in allocation to pistils. However, a complete explanation for the pattern of gamete packaging requires an understanding of factors controlling male, as well as female, fitness.     

Reproductive structures of an extinct platanoid from the early Cretaceous (latest Albian) of eastern North America

Two new species of platanoid reproductive structure are described from the Bull Mountain locality in the Patapsco Formation (Potomac Group) of northeastern Maryland, USA. Pistillate inflorescences and infructescences (Platanocarpus elkneckensis sp. nov.) consist of flowers and fruits in sessile globose heads that are borne on an elongate axis. Individual pistillate flowers consist of five free carpels surrounded by prominent tepals. Staminate inflorescences, flowers and isolated stamens are assigned to Hamatia elkneckensis gen. et sp. nov. Staminate flowers are borne in a globose head with a small number of stamens (five?) per flower. Stamens consist of very short filaments, long anthers with strongly valvate dehiscence and an apically expanded connective. The connective expansion is frequently very well-developed, hook-like and extends down the ventral surface of the stamen. Anthers contain small, tricolporate, reticulate pollen. Association evidence, similarity of inflorescence structure and the occurrence of Hamatia-type pollen on flowers, carpels and fruits of Platanocarpus elkneckensis suggests that the staminate and pistillate material was produced by a single species of plant. The “Hamatia-plant” provides further evidence of pentamerous floral structure in mid-Cretaceous platanoids and documents the occurrence of unequivocal tricolporate pollen in the platanoid complex.     

Fruit-Set Induced Changes in the Sex of Flowers in Caesalpinia calycina (Leguminosae)

Abstract: The proportions of hermaphrodite to staminate flowers in inflorescences of the andromonoecious species Caesalpinb calycina vary. Analysis of fruit position along the flowering rhachis, and also of the sex of floral buds in inflorescences with fruits set in different positions, indicate that fruiting success influences the sex of flowers in distal positions along the rhachis. Other reports of andromonoecy in caesalpinoid species are examined with reference to floral sex lability and the influence of fruit set.     

Three new species of Dendropanax (Araliaceae) from Bahia, Brazil

Three new species of BrazilianDendropanax from the rain forests of Bahia state are described and illustrated.Dendropanax amorimii, endemic to the rain forests near Boa Nova, in southern Bahia, is characterized by compound umbellate inflorescences with elongated primary branches each having a proximal whorl of bracts, and by leaves with well-developed tertiary venation.Dendropanax bahiensis, endemic to the Atlantic rain forests of southern Bahia, is distinguished by its large and membranous leaves with reddish schizogenous glands on the abaxial surface and by the reduced and branched inflorescences with short-pedicellate flowers and large fruits.Dendropanax geniculatus, an endemic species from the “matas de grotão” of Serra da Chapadinha, Chapada Diamantina, is characterized by simple and usually geniculate inflorescences bearing long-pedicellate flowers and fruits.     

Endophytic fungal community succession in reproductive organs of two olive tree cultivars with contrasting anthracnose susceptibilities

The factors shaping endophytic fungal assemblages in reproductive organs have been less evaluated than in vegetative organs. Here, the temporal dynamics of fungal endophytes in inflorescences and fruits of olive tree cultivars with contrasting anthracnose susceptibilities was assessed, starting before flowers open until fruit ripening. Fungal diversity assessed by rRNA sequencing of cultivable isolates revealed a higher richness and abundance of endophytes in inflorescences than in fruits. Endophytes from Sordariomycetes were dominant in inflorescences, while in fruits they were from Dothideomycetes. The overall assemblages of inflorescences and fruits were shaped mainly by the plant organ, followed by the phenological stage and lastly, by the plant cultivar. The importance of cultivar as a shaping factor varied according to the plant organ, being only significant in inflorescence-associated endophytes. A set of fungal taxa were positively associated with a particular cultivar and phenological stage, likely contributing to the plant anthracnose resistance.