Incomplete resupination during floral development leads to pollination failure in a slipper orchid.

In many families, plants undergo floral resupination by twisting through approximately 180° during floral development so that the flower is effectively positioned upside down. In most orchids, resupination results in the median petal (i.e., the labellum) becoming lowermost, which plays a crucial role in pollination by serving as a landing platform or as a trapping device, or both. Incomplete resupination is predicted to lead to reduced pollination, although tests of this assumption are still lacking. We investigated the effect of resupination using Phragmipedium vittatum, a rare lady's slipper orchid whose specialized labellum forms a trapping device. First, we surveyed the natural occurrence of incomplete resupination. Then we manipulated flowers into non- (≈0°), half- (≈90°), and fully resupinate (≈180°) positions to test the effect of orientation on pollen smear removal and deposition by pollinators (female hoverflies). We found that ca. 10% of flowers in the natural population were not fully resupinate, being either non- (upward, 0-60°) or half-resupinate (sideward, 60-120°). The change in orientation prevented the effectiveness of pollination by hoverflies since no pollen smear removal or deposition were found in flowers from non- and half-resupinate treatments. Although these flowers still attracted hoverflies, they were not trapped effectively. As this orchid is incapable of autonomous self-pollination, flowers that do not resupinate fail to set fruits. These results highlight the importance of correct floral orientation provided by resupination to ensure pollination in orchids and other resupinate flowers.

Flower orientation and corolla length as reproductive barriers in the pollinator-driven divergence of Erica shannonea and Erica ampullacea.

A variety of reproductive barriers can enable reproductive isolation and stable coexistence of plant species. Differing floral traits might play an important role in reproductive isolation imposed by pollinators. Such shifts in pollinator use have been hypothesized to contribute to the radiation of Erica (Ericaceae) in the Cape Floristic Region, South Africa. The sister species Erica shannonea and Erica ampullacea co-occur and overlap in flowering phenology. Both have unscented long-tubed flowers consistent with adaptations for pollination by long-proboscid flies (LPFs), but differences in flower orientation and corolla tube length are indicative of a shift in pollinator species. We conducted controlled pollination experiments and pollinator observations to determine the breeding system and pollinators of the two species. Both species are self-incompatible and require pollinator visits for seed production, suggesting that pollinators could strongly influence flower evolution. The horizontally orientated flowers of E. shannonea were found to be pollinated by Philoliche rostrata (Tabanidae), which has a long, fixed forward-pointing proboscis, while the vertically upright orientated flowers of E. ampullacea were found to be pollinated by Prosoeca westermanni (Nemestrinidae), which has a shorter proboscis that can swivel downwards. The nemestrinid fly's proboscis is too short to access the nectar in the relative long-tubed flowers of E. shannonea and the tabanid fly's proboscis cannot swivel down to access the upright flowers of E. ampullacea. Consequently, these traits are likely to act as reproductive barriers between the two Erica species and thereby might have contributed to speciation and enable stable coexistence.

A generalized bird pollination system in Schotia brachypetala (Fabaceae).

Bird pollination systems are diverse, ranging from narrow-tubed flowers pollinated by specialist nectarivores such as hummingbirds and sunbirds, to relatively open flowers pollinated by opportunistic (i.e. generalist) nectarivores. The role of opportunistic avian nectarivores as pollinators has historically been under-appreciated. A key aspect to understanding the importance of opportunistic birds as pollinators is to investigate how efficiently they transfer pollen among flowers. Here, we document the pollination and breeding systems of Schotia brachypetala, a southern African tree known as the 'weeping boer-bean' on account of its prolific production of dilute hexose-dominated nectar. The cup-shaped flowers of this tree attract a large number of bird species, including both opportunistic and specialist nectarivores. We identified floral visitors using observations and camera traps and quantified the floral traits responsible for animal attraction. We documented the breeding system, used selective pollinator exclusion to test the contribution of birds to fecundity, and performed supplemental pollination to test for pollen limitation. Single-visit pollen deposition trials were undertaken to determine the efficacy of bird pollinators. Controlled hand-pollination experiments showed that S. brachypetala is genetically self-incompatible and therefore dependent on pollinators for seed production. Supplemental hand-pollination experiments showed that natural fecundity is limited by either the amount and/or the quality of pollen on stigmas. Flowers from which birds but not insects were experimentally excluded set fewer seeds than open control flowers. Opportunistic birds deposited more pollen per visit than did specialist sunbirds. We conclude that S. brachypetala has a generalized bird pollination system that mainly involves opportunistic nectarivores.

Foods Implicated in U.S. Outbreaks Differ from the Types Most Commonly Consumed.

ABSTRACT: Foodborne disease outbreak investigations identify foods responsible for illnesses. However, it is not known the degree to which foods implicated in outbreaks reflect the distribution of food consumption in the U.S. population or the risk associated with their consumption. We compared the distribution of 24 categories of foods implicated in outbreaks with the distribution of foods consumed by the U.S. population. Beef, chicken, eggs, fish, herbs, mollusks, pork, sprouts, seeded vegetables, and turkey were implicated in outbreaks significantly more often than expected based on the frequency of their consumption by the general population, suggesting a higher risk of contamination or mishandling from foods in these categories than from foods in other categories. In contrast, pasteurized dairy, fruits, grains and beans, oils and sugars, and root and underground vegetables were less frequently implicated in outbreaks than their frequency of consumption by the general population, suggesting a lower health risk associated with these food categories.

Breeding systems of floral colour forms in the Drosera cistiflora species complex.

Variation in plant breeding systems has implications for pollinator-mediated selection on floral traits and the ecology of populations. Here we evaluate pollinator contribution to seed production, self-compatibility and pollen limitation in different floral colour forms of Drosera cistiflora sensu lato (Droseraceae). These insectivorous perennial plants are endemic to fynbos and renosterveld vegetation in the Cape Floristic Region of South Africa, and the species complex includes five floral colour forms (pink, purple, red, white and yellow), some of which are known to be pollinated by beetles. Controlled hand-pollination experiments were conducted in 15 populations of D. cistiflora s.l. (two to four populations per floral colour form) to test whether the colour forms vary in their degree of self-compatibility and their ability to produce seeds through autonomous self-fertilization. Yellow-flowered forms were highly self-incompatible, while other floral colour forms exhibited partial self-compatibility. Seed set resulting from autonomous selfing was very low, and pollinator dependence indices were high in all populations. Since hand cross-pollination resulted in greater seed set than open pollination in 13 of the 15 populations, we inferred that seed production is generally pollen-limited. Drosera cistiflora s.l. typically exhibits high levels of pollinator dependence and pollen limitation. This is unusual among Drosera species worldwide and suggests that pollinators are likely to mediate strong selection on attractive traits such as floral colour and size in D. cistiflora s.l. These results also suggest that the floral colour forms of D. cistiflora s.l. which are rare and threatened are likely to be vulnerable to local extinction if mutualisms were to collapse indefinitely.

Breeding systems in Cyrtanthus (Amaryllidaceae): variation in self-sterility and potential for ovule discounting.

Breeding systems of plants determine their reliance on pollinators and ability to produce seeds following self-pollination. Self-sterility, where ovules that are penetrated by self-pollen tubes that do not develop into seeds, is usually considered to represent either a system of late-acting self-incompatibility or strong early inbreeding depression. Importantly, it can lead to impaired female function through ovule or seed discounting when stigmas receive mixtures of self and cross pollen, unless cross pollen is able to reach the ovary ahead of self pollen ('prepotency'). Self-sterility associated with ovule penetration by self-pollen tubes appears to be widespread among the Amaryllidaceae. We tested for self-sterility in three Cyrtanthus species - C. contractus, C. ventricosus and C. mackenii - by means of controlled hand-pollination experiments. To determine the growth rates and frequency of ovule penetration by self- versus cross-pollen tubes, we used fluorescence microscopy to examine flowers of C. contractus harvested 24, 48 and 72 h after pollination, in conjunction with a novel method of processing these images digitally. To test the potential for ovule discounting (loss of cross-fertilisation opportunities when ovules are disabled by self-pollination), we pollinated flowers of C. contractus and C. mackenii with mixtures of self- and cross pollen. We recorded full self-sterility for C. contractus and C. ventricosus, and partial self-sterility for C. mackenii. In C. contractus, we found no differences in the growth rates of self- and cross-pollen tubes, nor in the proportions of ovules penetrated by self- and cross-pollen tubes. In this species, seed set was depressed (relative to cross-pollinated controls) when flowers received a mixture of self and cross pollen, but this was not the case for C. mackenii. These results reveal variation in breeding systems among Cyrtanthus species and highlight the potential for gender conflict in self-sterile species in which ovules are penetrated and disabled by pollen tubes from self pollen.

High levels of fecundity in small and isolated populations of a self-compatible Aloe pollinated by opportunistic birds and bees.

Plant species that are effective colonisers of transient habitats are expected to have a capacity for uniparental reproduction and show flexibility in pollination systems. Such traits may enable populations to be established from a small number of founding individuals without these populations succumbing to reductions in fecundity arising from pollinator limitation. We tested these predictions for Aloe thraskii (Xanthorrhoeaceae), a succulent treelet that colonises shifting coastal dunes and has both bird and bee pollinators. We performed hand-pollination experiments, and selectively excluded bird visitors to determine differences in pollinator effectiveness. We measured pollinator visitation rates and fecundity in populations varying in their size, density and isolation distance. Controlled hand-pollinations revealed that unlike most other Aloe species, A. thraskii is self-compatible and thus capable of uniparental reproduction. The species does however depend on pollinators and is visited by various bird species as well as by bees. Fruit and seed set are not affected by selective exclusion of birds, thus indicating that bees are effective pollinators. Bird visitation rates increased with increasing plant height and population size, while bee visitation rates increased with increasing population size and density. We found that seed set per flower was lower in large populations than in small populations. These results suggest that establishment of populations of A. thraskii from a small number of individuals is unlikely to be limited by the fecundity of individual plants.

Geographic variation in cone volatiles and pollinators in the thermogenic African cycad Encephalartos ghellinckii Lem.

Heat and odour production can have profound effects on pollination in cycads. It is therefore expected that these traits would co-vary geographically with pollinator assemblages. Such intraspecific variation, may lead to the evolution of pollination ecotypes, which can be an early stage of pollinator-mediated speciation. We measured cone temperatures using miniature temperature data loggers and examined the composition of cone volatile odours using headspace sampling and analysis with gas chromatography-mass spectrometry in four populations spanning the range of the African cycad Encephalartos ghellinckii. Pollinator assemblages were also investigated in three populations. Male and female cones were thermogenic at pollen shed and receptive stages, respectively, but patterns of thermogenesis did not vary among populations. Scent emissions from cones in populations in the Drakensberg Mountains were characterised by cis-ß-ocimene, ß-myrcene and (3E)-1,3-octadiene, while camphene and α-pinene were characteristic of scent emissions from cones in populations closer to the coast. These differences in volatile blends corresponded with differences in insect assemblages. These results confirm intraspecific variation in volatile emissions of E. ghellinckii and support the predictions that intraspecific variation in volatile emissions will be associated with shifts in pollinator assemblages. While further work needs to be done to test for local adaptation in this system, this preliminary evidence is consistent with the formation of pollination ecotypes in the E. ghellinckii species complex.

Pollination and breeding system of the enigmatic South African parasitic plant Mystropetalon thomii (Mystropetalaceae): rodents welcome, but not needed.

Unrelated plants adapted to particular pollinator types tend to exhibit convergent evolution in floral traits. However, inferences about likely pollinators from 'pollination syndromes' can be problematic due to trait overlap among some syndromes and unusual floral architecture in some lineages. An example is the rare South African parasitic plant Mystropetalon thomii (Mystropetalaceae), which has highly unusual brush-like inflorescences that exhibit features of both bird and rodent pollination syndromes. We used camera traps to record flower visitors, quantified floral spectral reflectance and nectar and scent production, experimentally determined self-compatibility and breeding system, and studied pollen dispersal using fluorescent dyes. The dark-red inflorescences are usually monoecious, with female flowers maturing before male flowers, but some inflorescences are purely female (gynoecious). Inflorescences were visited intensively by several rodent species that carried large pollen loads, while visits by birds were extremely rare. Rodents prefer male- over female-phase inflorescences, likely because of the male flowers' higher nectar and scent production. The floral scent contains several compounds known to attract rodents. Despite the obvious pollen transfer by rodents, we found that flowers on both monoecious and gynoecious inflorescences readily set seed in the absence of rodents and even when all flower visitors are excluded. Our findings suggest that seed production occurs at least partially through apomixis and that M. thomii is not ecologically dependent on its rodent pollinators. Our study adds another species and family to the growing list of rodent-pollinated plants, thus contributing to our understanding of the floral traits associated with pollination by non-flying mammals.

Effects of distance from models on the fitness of floral mimics.

Rewardless plants can attract pollinators by mimicking floral traits of rewarding heterospecific plants. This should result in the pollination success of floral mimics being dependent on the relative abundance of their models, as pollinator abundance and conditioning on model signals should be higher in the vicinity of the models. However, the attraction of pollinators to signals of the models may be partially innate, such that spatial isolation of mimics from model species may not strongly affect pollination success of mimics. We tested whether pollination rates and fruit set of the rewardless orchid Disa pulchra were influenced by proximity and abundance of its rewarding model species, Watsonia lepida. Pollination success of the orchid increased with proximity to the model species, while fruit set of the orchid increased with local abundance of the model species. Orchids that were experimentally translocated outside the model population experienced reduced pollinaria removal and increased pollinator-mediated self-pollination. These results confirm predictions that the pollination success of floral mimics should be dependent on the proximity and abundance of model taxa, and thus highlight the importance of ecological facilitation among species involved in mimicry systems.

Is the timing of scent emission correlated with insect visitor activity and pollination in long-spurred Satyrium species?

Plants are expected to emit floral scent when their pollinators are most active. In the case of long-tubed flowers specialised for pollination by crepuscular or nocturnal moths, scent emissions would be expected to peak during dawn. Although this classic idea has existed for decades, it has rarely been tested quantitatively. We investigated the timing of flower visitation, pollination and floral scent emissions in six long-spurred Satyrium species (Orchidaceae). We observed multiple evening visits by pollinaria-bearing moths on flowers of all study species, but rarely any diurnal visits. The assemblages of moth pollinators differed among Satyrium species, even those that co-flowered, and the lengths of moth tongues and floral nectar spurs were strongly correlated, suggesting that the available moth pollinator fauna is partitioned by floral traits. Pollinarium removal occurred more frequently during the night than during the day in four of the six species. Scent emission, however, was only significantly higher at dusk than midday in two species. Analysis of floral volatiles using gas chromatography coupled with mass spectrometry yielded 168 scent compounds, of which 112 were species-specific. The scent blends emitted by each species occupy discrete clusters in two-dimensional phenotype space, based on multivariate analysis. We conclude that these long-spurred Satyrium species are ecologically specialised for moth pollination, yet the timing of their scent emission is not closely correlated with moth pollination activity. Scent composition was also more variable than expected from a group of closely related plants sharing the same pollinator functional group. These findings reveal a need for greater understanding of mechanisms of scent production and their constraints, as well as the underlying reasons for divergent scent chemistry among closely related plants.

Evolution of apomixis loci in Pilosella and Hieracium (Asteraceae) inferred from the conservation of apomixis-linked markers in natural and experimental populations.

The Hieracium and Pilosella (Lactuceae, Asteraceae) genera of closely related hawkweeds contain species with two different modes of gametophytic apomixis (asexual seed formation). Both genera contain polyploid species, and in wild populations, sexual and apomictic species co-exist. Apomixis is known to co-exist with sexuality in apomictic Pilosella individuals, however, apomictic Hieracium have been regarded as obligate apomicts. Here, a developmental analysis of apomixis within 16 Hieracium species revealed meiosis and megaspore tetrad formation in 1 to 7% of ovules, for the first time indicating residual sexuality in this genus. Molecular markers linked to the two independent, dominant loci LOSS OF APOMEIOSIS (LOA) and LOSS OF PARTHENOGENESIS (LOP) controlling apomixis in Pilosella piloselloides subsp. praealta were screened across 20 phenotyped Hieracium individuals from natural populations, and 65 phenotyped Pilosella individuals from natural and experimental cross populations, to examine their conservation, inheritance and association with reproductive modes. All of the tested LOA and LOP-linked markers were absent in the 20 Hieracium samples irrespective of their reproductive mode. Within Pilosella, LOA and LOP-linked markers were essentially absent within the sexual plants, although they were not conserved in all apomictic individuals. Both loci appeared to be inherited independently, and evidence for additional genetic factors influencing quantitative expression of LOA and LOP was obtained. Collectively, these data suggest independent evolution of apomixis in Hieracium and Pilosella and are discussed with respect to current knowledge of the evolution of apomixis.

Male interference with pollination efficiency in a hermaphroditic orchid.

Hermaphroditism can lead to both intra- and intersexual conflict between male and female gender functions. However, the effect that such gender conflicts have on pollination efficiency has seldom been investigated. By artificially reducing the number of available male gametes on an individual, we quantified whether male interference with pollination efficiency occurs in the self-compatible, moth-pollinated orchid Satyrium longicauda. We partially emasculated S. longicauda inflorescences and compared pollination success and fecundity in these plants to intact controls. Pollen in both groups of plants was colour-labelled so that its dispersal by pollinators could be tracked directly in the field. Intact flowers on partially emasculated inflorescences exported more pollen and received more cross-pollen and less self-pollen than those on intact inflorescences. Proportion of fruit set per plant was similar between the two treatments; however, fruits on partially emasculated plants had proportionally more viable seeds than those on intact controls. These results provide empirical evidence that male interference with pollination efficiency can occur in a hermaphroditic plant and that such interference can compromise fecundity. The most likely mechanism for such male interference is competition for placement on the proboscis of hawkmoth pollinators. Consequently, male competition for siring success may influence the evolution of sexual systems in hermaphroditic pollinator-dependent plants.

Generalised pollination systems for three invasive milkweeds in Australia.

Because most plants require pollinator visits for seed production, the ability of an introduced plant species to establish pollinator relationships in a new ecosystem may have a central role in determining its success or failure as an invader. We investigated the pollination ecology of three milkweed species - Asclepias curassavica, Gomphocarpus fruticosus and G. physocarpus - in their invaded range in southeast Queensland, Australia. The complex floral morphology of milkweeds has often been interpreted as a general trend towards specialised pollination requirements. Based on this interpretation, invasion by milkweeds contradicts the expectation than plant species with specialised pollination systems are less likely to become invasive that those with more generalised pollination requirements. However, observations of flower visitors in natural populations of the three study species revealed that their pollination systems are essentially specialised at the taxonomic level of the order, but generalised at the species level. Specifically, pollinators of the two Gomphocarpus species included various species of Hymenoptera (particularly vespid wasps), while pollinators of A. curassavica were primarily Lepidoptera (particularly nymphalid butterflies). Pollinators of all three species are rewarded with copious amounts of highly concentrated nectar. It is likely that successful invasion by these three milkweed species is attributable, at least in part, to their generalised pollinator requirements. The results of this study are discussed in terms of how data from the native range may be useful in predicting pollination success of species in a new environment.

Floral scent in bird- and beetle-pollinated Protea species (Proteaceae): chemistry, emission rates and function.

Evolutionary shifts between pollination systems are often accompanied by modifications of floral traits, including olfactory cues. We investigated the implications of a shift from passerine bird to beetle pollination in Protea for floral scent chemistry, and also explored the functional significance of Protea scent for pollinator attraction. Using headspace sampling and gas chromatography-mass spectrometry, we found distinct differences in the emission rates and chemical composition of floral scents between eight bird- and four beetle-pollinated species. The amount of scent emitted from inflorescences of beetle-pollinated species was, on average, about 10-fold greater than that of bird-pollinated species. Floral scent of bird-pollinated species consists mainly of small amounts of "green-leaf volatiles" and benzenoid compounds, including benzaldehyde, anisole and benzyl alcohol. The floral scent of beetle-pollinated species is dominated by emissions of linalool, a wide variety of other monoterpenes and the benzenoid methyl benzoate, which imparts a fruity odour to the human nose. The number of compounds recorded in the scent of beetle-pollinated species was, on average, greater than in bird-pollinated species (45 versus 29 compounds, respectively). Choice experiments using a Y-maze showed that a primary pollinator of Protea species, the cetoniine beetle Atrichelaphinis tigrina, strongly preferred the scent of inflorescences of the beetle-pollinated Protea simplex over those of the bird-pollinated sympatric congener, Protea roupelliae. This study shows that a shift from passerine bird- to insect-pollination can be associated with marked up-regulation and compositional changes in floral scent emissions.

Floral volatiles, pollinator sharing and diversification in the fig-wasp mutualism: insights from Ficus natalensis, and its two wasp pollinators (South Africa).

Combining biogeographic, ecological, morphological, molecular and chemical data, we document departure from strict specialization in the fig-pollinating wasp mutualism. We show that the pollinating wasps Elisabethiella stuckenbergi and Elisabethiella socotrensis form a species complex of five lineages in East and Southern Africa. Up to two morphologically distinct lineages were found to co-occur locally in the southern African region. Wasps belonging to a single lineage were frequently the main regional pollinators of several Ficus species. In South Africa, two sister lineages, E. stuckenbergi and E. socotrensis, pollinate Ficus natalensis but only E. stuckenbergi also regularly pollinates Ficus burkei. The two wasp species co-occur in individual trees of F. natalensis throughout KwaZulu-Natal. Floral volatile blends emitted by F. natalensis in KwaZulu-Natal were similar to those emitted by F. burkei and different from those produced by other African Ficus species. The fig odour similarity suggests evolutionary convergence to attract particular wasp species. The observed pattern may result from selection for pollinator sharing among Ficus species. Such a process, with one wasp species regionally pollinating several hosts, but several wasp species pollinating a given Ficus species across its geographical range could play an important role in the evolutionary dynamics of the Ficus-pollinating wasp association.

Effects of pollen reward removal on fecundity in a self-incompatible hermaphrodite plant.

The pollen of hermaphrodite plants is often utilised by flower-visiting animals. While pollen production has obvious benefits for plant male fitness, its consequences for plant female fitness, especially in self-incompatible hermaphrodite species, are less certain. Pollen production could either enhance seed production though increased pollinator attraction, or reduce it if ovules are discounted by deposition of self pollen, as can occur in species with late-acting self-incompatibility. To test the effects of pollen reward provision on female fitness, we artificially emasculated flowers in two populations of the succulent Aloe maculata (Asphodelaceae), which has a late-acting self-incompatibility system, over the course of its flowering period. Flowers of this species are visited by sunbirds (for nectar) and native bees (for pollen and nectar). We measured floral visitation rates, floral rejection rates, pollen deposition on stigmas and fruit and seed set in both emasculated and non-emasculated plants. We found that flowers of emasculated plants suffered reduced visitation and increased rejection (arrival without visitation) by bees, but not by sunbirds; had fewer pollen grains deposited on stigmas and showed an overall decrease in fruit set and seed set. Rates of seed abortion were, however, greatly reduced in emasculated flowers. This study shows that pollen rewards can be important for seed set, even in self-incompatible plants, which have been assumed to rely on nectar rewards for pollinator attraction. Seed abortion was, however, increased by pollen production, a result that highlights the complexity of selection on pollen production in hermaphrodite flowers.

Sugar preferences and digestive efficiency of the village weaver: a generalist avian pollinator of African plants.

Recent research has shown that nectar properties of flowers pollinated by generalist avian nectarivores differ markedly from those of flowers pollinated by specialist avian nectarivores. In particular, flowers pollinated by generalist avian nectarivores tend to have very dilute nectar dominated by hexose sugars. To establish whether pollinator-mediated selection can explain these traits, we tested nectar sugar preferences and digestive capabilities of the village weaver (Ploceus cucullatus), a common generalist passerine nectarivore in South Africa. When offered pairwise choices of equicaloric hexose and sucrose solutions, village weavers preferred hexose solutions at 5% and 10% sucrose equivalents (SE) but did not show significant preference for either type of sugar when higher concentrations were offered (15%, 20% and 25% SE). Birds were less efficient at absorbing sucrose than hexose sugars, as revealed by high-performance liquid chromatography (HPLC) analysis of excreta sugar content. This was true at both concentrations tested (8.22% and 25%), although apparent sucrose assimilation rates were still relatively high (89.6+/-2.9% at low concentrations and 93.6+/-1.7% at high concentrations). Transit times indicated that sucrose also passes through the digestive tract faster than hexose sugars, particularly when consumed at high concentrations. This may limit the rate at which sucrose can be hydrolyzed before absorption. These results indicate that hexose preferences in generalist avian nectarivores may help explain the low sucrose content in flowers pollinated by these birds. Moreover, the preference for hexose sugars in weavers was most evident at the low concentrations (ca. 9% sugar by mass) that are typical of nectar in flowers pollinated by generalist avian nectarivores.

A key role for floral scent in a wasp-pollination system in Eucomis (Hyacinthaceae).

BACKGROUND AND AIMS: Floral scent may play a key role as a selective attractant in plants with specialized pollination systems, particularly in cases where floral morphology does not function as a filter of flower visitors. The pollination systems of two African Eucomis species (E. autumnalis and E. comosa) were investigated and a test was made of the importance of scent and visual cues as floral attractants. METHODS AND KEY RESULTS: Visitor observations showed that E. autumnalis and E. comosa are visited primarily by pompilid wasps belonging to the genus Hemipepsis. These wasps carry considerably more Eucomis pollen and are more active on flowers than other visiting insects. Furthermore, experiments involving virgin flowers showed that these insects are capable of depositing pollen on the stigmas of E. autumnalis, and, in the case of E. comosa, pollen deposited during a single visit is sufficient to result in seed set. Experimental hand-pollinations showed that both species are genetically self-incompatible and thus reliant on pollinators for seed set. Choice experiments conducted in the field and laboratory with E. autumnalis demonstrated that pompilid wasps are attracted to flowers primarily by scent and not visual cues. Measurement of spectral reflectance by flower petals showed that flowers are cryptically coloured and are similar to the background vegetation. Analysis of headspace scent samples using coupled gas chromatography-mass spectrometry revealed that E. autumnalis and E. comosa scents are dominated by aromatic and monoterpene compounds. One hundred and four volatile compounds were identified in the floral scent of E. autumnalis and 83 in the floral scent of E. comosa, of which 57 were common to the scents of both species. CONCLUSIONS: This study showed that E. autumnalis and E. comosa are specialized for pollination by pompilid wasps in the genus Hemipepsis and achieve specialization through cryptic colouring and the use of scent as a selective floral attractant.

An analysis of stressors and co-morbid mental health problems that contribute to youth's paths to substance-specific services.

Substance-specific services are tailored to address the inappropriate use of chemicals such as alcohol and marijuana. Unfortunately, few teens ever access such need-based services. This article explores the paths to substance-specific service use in a sample of urban adolescents who are public service sector users: a direct path from substance misuse and an indirect path from general mental health service use. Results indicate that even though a significant percentage of youths frequently consume substances and display negative use-related behaviors, few actually received substance-specific services. Contributors to the substance misuse path included youths' family substance dependence and environmental stressors. Contributors to the mental health services path were comorbid depression and substance misuse.