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Caladium bicolor (Araceae) and Cyclocephala celata (Coleoptera, Dynastinae): a well-established pollination system in the Northern Atlantic rainforest of Pernambuco, Brazil.

Flowering, pollination ecology, and floral thermogenesis of Caladium bicolor were studied in the Atlantic Rainforest of Pernambuco, NE Brazil. Inflorescences of this species are adapted to the characteristic pollination syndrome performed by Cyclocephalini beetles. They bear nutritious rewards inside well-developed floral chambers and exhibit a thermogenic cycle which is synchronized to the activity period of visiting beetles. Heating intervals of the spadix were observed during consecutive evenings corresponding to the beginning of the female and male phases of anthesis. Highest temperatures were recorded during the longer-lasting female phase. An intense sweet odour was volatized on both evenings. Beetles of a single species, Cyclocephala celata, were attracted to odoriferous inflorescences of C. bicolor and are reported for the first time as Araceae visitors. All the inflorescences visited by C. celata developed into infructescences, whereas unvisited inflorescences showed no fruit development. Findings of previous studies in the Amazon basin of Surinam indicated that Cyclocephala rustica is a likely pollinator of C. bicolor. This leads to the assumption that locally abundant Cyclocephalini species are involved in the pollination of this species.

Animals↗

Flower Structure and Reproductive Biology in Aspidosperma quebracho-blanco (Apocynaceae), a Tree Pollinated by Deceit.

Flower structure, attractants, breeding system, and visitors were studied in Aspidosperma quebracho-blanco, a South American mass-flowering tree with simple moth-pollinated flowers and without floral reward. Flowers are protandrous, having an initial male phase for 2 d. The pendulous and horizontal flower position and protandry prevent self-pollination. Although an inconspicuous annular floral nectary is located at the ovary base, nectar is not secreted. Nonstructural, extrafloral nectaries have been found on the external surface of the calyx and corolla, the first report of the kind for the family. Ants, wasps, flies, and beetles collect the extrafloral exudate composed of fructose, glucose, and sucrose. Experiments indicate that natural selfing (autogamy) and agamospermy are unlikely to occur (0% success), but artificial selfing indicates self-compatibility (70% success). A very low natural fruit to flower ratio (0.03) was obtained. Floral features and nocturnal opening indicate that these flowers would be pollinated by moths that nocturnally visit the species (Noctuidae, Pyraloidea) finding no floral reward. We suggest a nonmodel deception mechanism: providing the same cues as most moth flowers, this tree seems to be dependent upon sympatric rewarding plants that share the same pollinators.

Journal Article↗

Beetle pollination of Philodendron solimoesense (Araceae) in French Guiana.

The pollination of Philodendron solimoesense (subgenus Meconostigma) was studied in four populations of French Guiana. Flowering is asynchronous within each population during July, and the flowering cycle is a 2-d process. Numerous insects visit Philodendron inflorescences, but the main pollinator seems to be Cyclocephala colasi (Scarabaeidae, Dynastinae). The pollination process displays aspects typical of beetle pollination: the production of heat and of a strong odor, the presence of a food reward (stigmatic secretion and sterile male flowers), and the presence of a copulation chamber. Flower heat production is important (ca. 11 degrees C above the ambient air) and may help to volatilize the fragrance. Attraction and choice-test experiments showed that C. colasi is not likely to depend on chemical information (such as pheromone) to localize conspecifics but may rely instead on stimuli produced by the inflorescences in order to meet mating partners.

Journal Article↗

Herbivory reduces the strength of pollinator-mediated selection in the Mediterranean herb Erysimum mediohispanicum: consequences for plant specialization.

In this study, I tested whether selection occurring on several morphological and floral traits in Erysimum mediohispanicum (Cruciferae) is modified by the effects of herbivores. Six plots were established in 1997 in the Sierra Nevada, Spain; three were fenced to exclude native ungulates, and the remaining were open to ungulates. I determined pollinator and ungulate preferences for plant traits and their effect on plant fecundity. Then I compared the selection regimes between plants excluded from herbivores and plants open to them. When ungulates were absent, I found significant selection on flower number, reproductive stalk height, basal diameter of the stalks, petal length, and inner diameter of the flowers. When ungulates were present, selection on floral traits completely disappeared, and selection strength on flower number and morphological traits decreased. This effect was due to the ungulate preference for larger plants and the phenotypic correlations between plant size and floral traits. Results suggest that pollinator-mediated selection can be disrupted by conflicting effects of plant enemies acting during or subsequent to pollination. An accurate picture of the pollinator role as selective pressure requires the consideration of the entire life cycle of the plant as well as the ecological scenario in which the interactions occur.

Adaptation, Physiological↗

The consequences of direct versus indirect species interactions to selection on traits: pollination and nectar robbing in Ipomopsis aggregata.

Organisms experience a complex suite of species interactions. Although the ecological consequences of direct versus indirect species interactions have received attention, their evolutionary implications are not well understood. I examined selection on floral traits through direct versus indirect pathways of species interactions using the plant Ipomopsis aggregata and its pollinators and nectar robber. Using path analysis and structural equation modeling, I tested competing hypotheses comparing the relative importance of direct (pollinator-mediated) versus indirect (robber-mediated) interactions to trait selection through female plant function in 2 years. The hypothesis that provided the best fit to the observed data included robbing and pollination, suggesting that both interactors are important in driving selection on some traits; however, the direction and intensity of selection through robbing versus pollination varied between years. I then increased my scope of inference by assessing traits and species interactions across more years. I found that the potential for temporal variation in the direction and intensity of selection was pronounced. Taken together, results suggest that assessing the broader context in which organisms evolve, including both direct and indirect interactions and across multiple years, can provide increased mechanistic understanding of the diversity of ways that animals shape floral and plant evolution.

Animals↗

Wettable and unsinkable: the hydrodynamics of saccate pollen grains in relation to the pollination mechanism in the two New Zealand species of Prumnopitys Phil. (Podocarpaceae).

The pollination mechanism of most genera of the Podocarpaceae involves inverted ovules, a pollination drop and bisaccate pollen grains. Saccate grains have sometimes been referred to as 'non-wettable' due to their buoyant properties, while non-saccate pollen grains have been described as 'wettable'. The hydrodynamic properties of saccate pollen grains of seven podocarp species in five genera, Dacrydium Sol. ex G. Forst., Dacrycarpus (Endl.) de Laub., Manoao Molloy, Podocarpus L'Hér. ex Pers. and Prumnopitys Phil. have been tested in water, together with saccate and non-saccate pollen of four other conifer genera, Cedrus Trew (Pinaceae), Cephalotaxus Siebold & Zucc. ex Endl. (Cephalotaxaceae), Cupressus L. (Cupressaceae) and Phyllocladus Rich. ex Mirb. (Phyllocladaceae), and spores of three fern species and one lycopod species. All four spore types studied were non-wettable, whereas the bisaccate and trisaccate pollen types, like all other conifer pollen types, were wettable, enabling the grains to cross the surface tension barrier of water. Once past this barrier, grain behaviour was governed by presence or absence of sacci. Non-saccate and vestigially saccate grains sank, whereas saccate grains behaved like air bubbles, floating up to the highest point. In addition, the grains were observed to float in water with sacci uppermost, consistent with the suggestion that distally placed sacci serve to orientate the germinal furrow of the pollen grain towards the nucellus of an inverted ovule. Observations of pollen grains in the pollen chambers of naturally pollinated Prumnopitvs ovules confirmed this. The combination of buoyancy and wettability in saccate pollen has implications for the efficiency of the typical podocarp pollination mechanism.

Biomechanical Phenomena↗

Floral and vegetative morphometrics of five Pleurothallis (Orchidaceae) species: correlation with taxonomy, phylogeny, genetic variability and pollination systems.

Morphometric analyses of vegetative and floral characters were conducted in 21 populations of five Pleurothallis (Orchidaceae) species occurring in Brazilian 'campo rupestre' vegetation. A phylogenetic analysis of this species group was also carried out using nuclear ribosomal DNA internal transcribed spacers (ITS1 and ITS2). Results of the ordination and cluster analyses agree with species' delimitation revealed by taxonomic and allozyme studies. The groups formed in ordination analysis correspond to the pollinator groups determined in a previous pollination study. Relationships among the species in the cluster analysis using only vegetative characters are similar to those found in a previous allozyme study, but those indicated by cluster analysis using only floral characters differ. These results support the hypothesis that floral similarities are due to convergence driven by similar pollination mechanisms, and therefore floral traits may not be good indicators of phylogenetic relationships in this group. The results of the phylogenetic analysis support this conclusion to some extent. There is no correlation between genetic (allozyme) and morphological variability in the populations nor in the way this variability is distributed among conspecific populations. We describe a new subspecies of Pleurothallis ochreata based on differences in vegetative and chemical characters as well as geographic distribution. Absence of differentiation in floral characters, attraction of the same pollinator species, interfertility and genetic similarity support the argument for subspecific rather than specific status.

Brazil↗

Preferences of pollinators and herbivores in gynodioecious Geranium sylvaticum.

BACKGROUND AND AIMS: For the maintenance of gynodioecy (i.e. the coexistence of female and hermaphroditic plants), females need to compensate for the lack of pollen production through higher seed production or better progeny quality compared to hermaphrodites. In Geranium sylvaticum, females produce more seeds per flower than hermaphrodites. This difference in seed production might be modified by biological interactions with pollinators and herbivores that may favour one sex and thus affect the maintenance of gynodioecy. METHODS: Sexual dimorphism in flower size and flowering phenology, and in attractiveness to pollinators, pre-dispersal seed predators and floral herbivores were examined in natural populations of G. sylvaticum. KEY RESULTS: Pollinators preferred hermaphrodites 25 % more often than females in two of the three study populations, and floral herbivores attacked hermaphrodites 15 % more often than females in two of the six study populations. These preferences might be explained by the larger flower size of hermaphrodites. In contrast, seed predators did not prefer either sex. CONCLUSIONS: The data suggest that pollinator preference does not benefit females, whereas the higher floral herbivory of hermaphrodites might enhance the maintenance of females in G. sylvaticum. Thus, although the data support the view that ecological factors may contribute to the maintenance of gynodioecy, they also suggest that these contributions may vary across populations and that they may function in opposite directions.

Adaptation, Physiological↗

Flowering phenology and wind-pollination efficacy of heterodichogamous Juglans mandshurica (Juglandaceae).

BACKGROUND AND AIMS: Heterodichogamy differs from normal dichogamy, in that it involves two mating types (protogyny and protandry) that occur at a 1 : 1 ratio in a population. Flowering phases of the two mating types are synchronized and reciprocal, which was considered to ensure between-type outcrossing. This study aims to quantify the flowering pattern and pollination efficacy in Juglans mandshurica, a wind-pollinated heterodichogamous tree. METHODS: The pattern of flowering phenology was monitored within individual trees and pollen traps were used to measure air-borne pollen loads during the spring in 2003 and 2004. Pollen longevity was determined by staining technique. Also a pollen supplementation experiment was performed in 2004 to assess pollen limitation of fruit production. KEY RESULTS: There was no overlap between sexual functions within individual trees. Flowering periods of the two mating types were reciprocal and synchronous in both 2003 and 2004. Air-borne pollen loads were large, and protogynous and protandrous individuals each produced a high pollination peak, consistent with the two blooming periods. Maximum pollen longevity was about 4 h for protandrous individuals, and 3 h for protogynous individuals. Pollen supplementation did not increase fruit production in either protogynous or protandrous individuals. CONCLUSIONS: Heterodichogamous flowering in Juglans mandshurica effectively avoids selfing, promotes between-type outcrossing, and leads to efficient pollination in a natural population.

Breeding↗

Nectar, floral morphology and pollination syndrome in Loasaceae subfam. Loasoideae (Cornales).

BACKGROUND AND AIMS: Loasaceae subfam. Loasoideae are mostly distributed in South America (sea level to over 4500 m) with a wide range of animals documented as pollinators. The aim was to investigate correlations between nectar parameters, flower morphology, pollination syndrome and phylogeny. METHODS: Nectar was collected from 29 species from seven genera in the subfamily. Concentration and volumes were measured and the amount of sugar calculated. Correlations of nectar data were plotted on a ternary graph and nectar characteristics compared with flower visitors, floral morphology and phylogenetic data. KEY RESULTS: Sugar concentrations are generally higher than reported for most plant families in the literature. The species investigated can be roughly grouped as follows. Group I: plants with approx. 1.5(-3.5) microL nectar with (40-)60-80% sugar and 0.19-2 mg sugar flower-1; with small, white, star-shaped corollas, pollinated by short-tongued bees. Groups II, III and IV: plants with mostly orange, balloon-, saucer-, bowl- or bell-shaped corollas. Group II: plants with approx. 9-14 microL nectar with 40-60% sugar and 4-10 mg sugar flower-1; mostly visited by long-tongued bees and/or hummingbirds. Group III: plants with 40-100 microL nectar with 30-40% sugar and 14-36 mg sugar flower-1, mostly visited by hummingbirds. Group IV: geoflorous plants with 80-90 microL with 10-15% sugar and 8.5-12 mg sugar flower-1, presumably visited by small mammals. Groups II and III include species visited by bees and/or hummingbirds. CONCLUSIONS: Pollinator switches from short-tongued bees via long-tongued bees to hummingbirds appear to have taken place repeatedly in the genera Nasa, Loasa and Caiophora. Changes in nectar amount and concentration appear to evolve rapidly with little phylogenetic constraint.

Animals↗

Honey bee (Hymenoptera: Apidae) distribution and potential for supplementary pollination in commercial tomato greenhouses during winter.

This study examined the use of honey bees, Apis mellifera L., to supplement bumble bee, Bombus spp., pollination in commercial tomato, Lycopersicon esculentum Miller, greenhouses in Western Canada. Honey bee colonies were brought into greenhouses already containing bumble bees and left for 1 wk to acclimatize. The following week, counts of honey and bumble bees foraging and flying throughout the greenhouse were conducted three times per day, and tomato flowers open during honey bee pollination were marked for later fruit harvest. The same counts and flower-marking also were done before and after the presence of honey bees to determine the background level of bumble bee pollination. Overall, tomato size was not affected by the addition of honey bees, but in one greenhouse significantly larger tomatoes were produced with honey bees present compared with bumble bees alone. In that greenhouse, honey bee foraging was greater than in the other greenhouses. Honey bees generally foraged within 100 m of their colony in all greenhouses. Our study invites further research to examine the use of honey bees with reduced levels of bumble bees, or as sole pollinators of greenhouse tomatoes. We also make specific recommendations for how honey bees can best be managed in greenhouses.

Agriculture↗

Transcriptional regulation of three EIN3-like genes of carnation (Dianthus caryophyllus L. cv. Improved White Sim) during flower development and upon wounding, pollination, and ethylene exposure.

Using a combination of approaches, three EIN3-like (EIL) genes DC-EIL1/2 (AY728191), DC-EIL3 (AY728192), and DC-EIL4 (AY728193) were isolated from carnation (Dianthus caryophyllus) petals. DC-EIL1/2 deduced amino acid sequence shares 98% identity with the previously cloned and characterized carnation DC-EIL1 (AF261654), 62% identity with DC-EIL3, and 60% identity with DC-EIL4. DC-EIL3 deduced amino acid sequence shares 100% identity with a previously cloned carnation gene fragment, Dc106 (CF259543), 61% identity with Dianthus caryophyllus DC-EIL1 (AF261654), and 59% identity with DC-EIL4. DC-EIL4 shared 60% identity with DC-EIL1 (AF261654). Expression analyses performed on vegetative and flower tissues (petals, ovaries, and styles) during growth and development and senescence (natural and ethylene-induced) indicated that the mRNA accumulation of the DC-EIL family of genes in carnation is regulated developmentally and by ethylene. DC-EIL3 mRNA showed significant accumulation upon ethylene exposure, during flower development, and upon pollination in petals and styles. Interestingly, decreasing levels of DC-EIL3 mRNA were found in wounded leaves and ovaries of senescing flowers whenever ethylene levels increased. Flowers treated with sucrose showed a 2 d delay in the accumulation of DC-EIL3 transcripts when compared with control flowers. These observations suggest an important role for DC-EIL3 during growth and development. Changes in DC-EIL1/2 and DC-EIL4 mRNA levels during flower development, and upon ethylene exposure and pollination were very similar. mRNA levels of the DC-EILs in styles of pollinated flowers showed a positive correlation with ethylene production after pollination. The cloning and characterization of the EIN3-like genes in the present study showed their transcriptional regulation not previously observed for EILs.

Dianthus↗

Use of diploid male frequency data as an indicator of pollinator decline.

Pollination deficits in agricultural and natural systems are suggestive of large reductions in pollinator populations. However, actual declines are difficult to demonstrate using census data. Here, we show census data to be misleading because many abundant pollinators exhibit high levels of production of sterile diploid males usually found only in small inbred hymenopteran populations; Euglossa imperialis exhibits high levels of diploid male production induced by low effective population sizes (Ne approximately 15), despite being the most abundant orchid bee in lowland tropical forests in Panama. We caution that although some pollinators appear abundant on the basis of census data, their long-term persistence may be highly tenuous based on genetic evidence. We propose the use of diploid male frequency data as a metric for assessing the sustainability of bee populations.

Alleles↗

Mauritian coloured nectar no longer a mystery: a visual signal for lizard pollinators.

Most floral nectars are clear as water, and the enigmatic coloured nectar in three endemic plant species in Mauritius has puzzled scientists studying it. One hypothesis about the possible ecological function of coloured nectar is that it serves as a visual signal for pollinators. Recent studies have shown that at least two of the three Mauritian plant species with coloured nectar are visited and pollinated by endemic Phelsuma geckos. We here provide experimental evidence for the visual signal hypothesis by showing that Phelsuma ornata geckos prefer coloured over clear nectar in artificial flowers. In flowering plants, coloured nectar could additionally function as an honest signal that allows pollinators to assert the presence and judge the size of a reward prior to flower visitation, and to adjust their behaviour accordingly, leading to increased pollinator efficiency. Our study provides a first step in understanding this rare and intriguing floral trait.

Animals↗

Genetic rescue of remnant tropical trees by an alien pollinator.

Habitat fragmentation is thought to lower the viability of tropical trees by disrupting their mutualisms with native pollinators. However, in this study, Dinizia excelsa (Fabaceae), a canopy-emergent tree, was found to thrive in Amazonian pastures and forest fragments even in the absence of native pollinators. Canopy observations indicated that African honeybees (Apis mellifera scutellata) were the predominant floral visitors in fragmented habitats and replaced native insects in isolated pasture trees. Trees in habitat fragments produced, on average, over three times as many seeds as trees in continuous forest, and microsatellite assays of seed arrays showed that genetic diversity was maintained across habitats. A paternity analysis further revealed gene flow over as much as 3.2 km of pasture, the most distant pollination precisely recorded for any plant species. Usually considered only as dangerous exotics, African honeybees have become important pollinators in degraded tropical forests, and may alter the genetic structure of remnant populations through frequent long-distance gene flow.

Animals↗

How an orchid harms its pollinator.

Certain orchids produce flowers that mimic the sex pheromones and appearance of female insects in order to attract males by sexual deception for the purpose of pollination. In a series of field experiments, we found that the sexually deceptive orchid, Chiloglottis trapeziformis, can have a negative impact on its wasp pollinator Neozeleboria cryptoides. Male and female wasps, however, were affected differently by the orchid's deceit because of their different roles in the mimicry system. Male wasps could not discriminate between the chemical cues of orchids and female wasps, a vital signal in long-range attraction. Males, however, learn to avoid areas containing orchids. This strategy has implications for females attempting to attract mates in areas occupied by orchids. Compared with circumstances when females were on their own, females in the presence of orchids elicited fewer male approaches and no copulation attempts. Females in a large orchid patch also elicited fewer male approaches than females in a small patch. The nature of the orchid's impact on its wasp pollinator indicates an arms race evolutionary scenario in this interaction between plant and pollinator.

Animals↗

Towards the theory of pollinator-mediated gene flow.

I present a new exposition of a model of gene flow by animal-mediated pollination between a source population and a sink population. The model's parameters describe two elements: (i) the expected portion of the source's paternity that extends to the sink population; and (ii) the dilution of this portion by within-sink pollinations. The model is termed the portion-dilution model (PDM). The PDM is a parametric restatement of the conventional view of animal-mediated pollination. In principle, it can be applied to plant species in general. I formulate a theoretical value of the portion parameter that maximizes gene flow and prescribe this as a benchmark against which to judge the performance of real systems. Existing foraging theory can be used in solving part of the PDM, but a theory for source-to-sink transitions by pollinators is currently elusive.

Animals↗

Pollination-Induced Ethylene in Carnation (Role of Pollen Tube Growth and Sexual Compatibility).

The pollen-pistil interactions that result in the stimulation of ethylene production and petal senescence in carnation (Dianthus caryophyllus L.) flowers were investigated. Pollination of White Sim flowers with Starlight pollen elicited an increase in ethylene production by styles, leading to increased petal ethylene and premature petal senescence. In contrast, pollination with 87-29G pollen led to an early increase in ethylene production, but this was not sustained, and did not lead to petal senescence. Both Starlight and 87-29G pollen germinated on White Sim stigmas and their tubes grew at similar rates, penetrating the length of the style. Crosses between Starlight and White Sim led to the production of viable seeds, whereas 87-29G pollen was infertile on White Sim flowers. Pollination of other carnations with 87-29G elicited ethylene production and petal senescence and led to the production of viable seeds. These results suggest that physical growth of pollen tubes is insufficient to elicit a sustained increase in ethylene production or to lead to the production of signals necessary for elicitation of petal ethylene production and senescence. Rather, the cell-cell recognition reactions leading to sexual compatibility in Dianthus appear to play a role in this interorgan signaling after pollination.

Journal Article↗