Do changes in floral odor cause speciation in sexually deceptive orchids?

 We investigated differences in floral odor between two sympatric, closely related sexually deceptive orchid species, Ophrys fusca and O. bilunulata, which are specifically pollinated by Andrena nigroaenea and A. flavipes, respectively. We identified biologically active compounds by gas chromatography with electroantennographic detection using antennae of the pollinator bees. Alkanes, alkenes, aldehydes, and farnesyl hexanoate released electroantennographic reactions. The relative amounts of alkanes were mostly the same between the two orchid species, whereas the relative amounts of most alkenes were significantly different. On the grounds of these findings and behavioral experiments conducted in earlier studies, we suggest that the difference in relative amounts of alkenes is responsible for the selective attraction of pollinators in the two orchids. Speciation in this group of Ophrys orchids may be brought about by changes in pattern of alkenes, which lead to attraction of a different pollinator species and therefore reproductive isolation. Received November 22, 2001; accepted February 21, 2002 Published online: November 7, 2002 Addresses of the authors: Florian P. Schiestl* and Manfred Ayasse, Department of Evolutionary Biology, Institute of Zoology, University of Vienna, Althanstrasse 14, A-1090 Vienna. *Present address: Geobotanical Institute ETH, Zollikerstrasse 107, CH-8008 Zürich. (e-mail: schiestl@geobot.umnw.ethz.ch)     

Does floral trait variability enhance reproductive success in deceptive orchids?

Unusually high intra-specific floral trait variability has often been described within deceptive orchid populations, as opposed to rewarding ones. Such variability is traditionally thought to have consequences on reproduction in this orchid group, i.e. phenotypically variable deceptive species may have a reproductive success advantage compared to those with a constant floral display. The proposed reason for this hypothetic pattern is that floral trait variability decreases pollinator avoidance learning in dealing with nectarless flowers, hence increasing their visitation rate. However, despite an intuitive and appealing hypothesis and a possible mechanism to explain it, the often-cited higher reproductive success induced by floral trait variability still remains unsupported.Here, we review the literature and consider eight studies that have experimentally or correlatively tested this hypothesis in deceptive orchids. In all these experiments, we have found no difference in average reproductive success between populations with high versus low flower trait variability, either in scent variable or colour polymorphic species. We discuss possible explanations for the lack of this pattern including the incapability of pollinators in perceiving the variability, the scarce relevance of polymorphic traits in the choice of species to forage on, or a different pollinator behaviour than the one proposed. We suggest that the high phenotypic variability is not likely to enhance deceptive orchids’ reproductive success, but is more likely to be a consequence of relaxed selection by pollinators. Nonetheless, information regarding orchid pollination strategy or pollinator cognitive abilities is often superficial, hence calling researchers for additional investigations that can contribute to a better understanding of this debated and yet unsupported hypothesis.     

Antagonistic effects of floral scent in an insect–plant interaction

In southwestern USA, the jimsonweed Datura wrightii and the nocturnal moth Manduca sexta form a pollinator–plant and herbivore–plant association. Because the floral scent is probably important in mediating this interaction, we investigated the floral volatiles that might attract M. sexta for feeding and oviposition. We found that flower volatiles increase oviposition and include small amounts of both enantiomers of linalool, a common component of the scent of hawkmoth-pollinated flowers. Because (+)-linalool is processed in a female-specific glomerulus in the primary olfactory centre of M. sexta, we hypothesized that the enantiomers of linalool differentially modulate feeding and oviposition. Using a synthetic mixture that mimics the D. wrightii floral scent, we found that the presence of linalool was not necessary to evoke feeding and that mixtures containing (+)- and/or (−)-linalool were equally effective in mediating this behaviour. By contrast, females oviposited more on plants emitting (+)-linalool (alone or in mixtures) over control plants, while plants emitting (−)-linalool (alone or in mixtures) were less preferred than control plants. Together with our previous investigations, these results show that linalool has differential effects in feeding and oviposition through two neural pathways: one that is sexually isomorphic and non-enantioselective, and another that is female-specific and enantioselective.     

Are pollination syndromes useful predictors of floral visitors in Tasmania?

Abstract Diurnal visitors to the flowers of many native plant species were identified in a wide range of Tasmanian sclerophyllous vegetation between September 1996 and April 1997. These foraging profiles were analysed to determine whether they were characteristic of various floral morphologies in predictable ways. It was found that although visitor profiles were sometimes consistent with classic pollination syndromes, these syndromes were unreliable predictors of floral visitors. Very few flowers were exclusively bird‐pollinated, and none were strictly fly‐, beetle‐, wasp‐, or butterfly‐pollinated. The majority of flowering plants were unspecialized in their morphology, and consequently hosted a diverse array of visitors. In addition, visitor profiles to congeners with similar floral morphologies, and even to conspecifics, differed between habitats. Altitude was a major factor in determining visitors, with flies being the most abundant visitors above 700 m. However, congeners in several genera of Epacridaceae, as well as the genus Correa, which differed in floral morphology also differed in visitor profiles. Tubular flowers were associated with birds, while flowers with more accessible nectar were visited by insects. The only taxa exhibiting a bee‐pollination syndrome that were largely visited by bees were the Fabaceae and Goodenia ovata Sm. Several species with purple or pink flowers were also predominantly visited by bees, but did not strictly conform to the melittophilous syndrome. In contrast, other flowers exhibiting an ostensibly mellitophilous syndrome hosted very few bees. Of these, species that occurred at high altitude were mainly visited by flies, while others received very few potential pollen vectors.     

Sources of floral scent variation: Can environment define floral scent phenotype?

Studies of floral scent generally assume that genetic adaptation due to pollinator-mediated natural selection explains a significant amount of phenotypic variance, ignoring the potential for phenotypic plasticity in this trait. In this paper, we assess this latter possibility, looking first at previous studies of floral scent variation in relation to abiotic environmental factors. We then present data from our own research that suggests among-population floral scent variation is determined, in part, by environmental conditions and thus displays phenotypic plasticity. Such an outcome has strong ramifications for the study of floral scent variation; we conclude by presenting some fundamental questions that should lead to greater insight into our understanding of the evolution of this trait, which is important to plant-animal interactions.Key words: abiotic factors, aromatics, floral scent, GxE interaction, phenotypic plasticity, pollination, terpenoids, volatilesFloral scent is thought to function as a major non-visual attractive cue for many pollinators in a large number of plant systems^1,2 and therefore most research on this plant trait has proceeded in the context of pollination ecology. Such studies have revealed the physiological and behavioral responses of pollinators to various floral volatiles (reviewed in refs. ³ and ⁴), convergent evolution of odor phenotypes attractive to specific pollinator classes (reviewed in refs. ⁵ and ⁶), reproductive isolation of plant species due to differences in pollinator attraction by scent,⁷ and instances of deception in which flowers mimic insect pheromones to effect pollination.⁸ Together, this body of evidence suggests that specific floral scent profiles can have important implications for the reproductive potential of many plant species.This pollinator-centered viewpoint has carried through to research on floral scent variation, including our most recent work on the insect-pollinated species Hesperis matronalis (Brassicaceae).⁹ Such studies usually suggest that the floral scent variation commonly found within and among individuals, populations and species (reviewed in ref. ²) is due to genetic differentiation as a result of selection by pollinators over time (reviewed in ref. ¹⁰). But an organism''s genes are only one factor determining phenotype. Both biotic (living) and abiotic (non-living) environmental conditions can profoundly affect phenotype expression, leading to significant variation. For plants, abiotic factors such as climate and soil chemistry can have particularly strong effects on phenotypes. When these environmental conditions cause changes in phenotype, we would say that a trait displays phenotypic plasticity.^11–13 A number of studies have uncovered phenotypic plasticity for many different plant traits.¹² However, while phenotypic variation in floral scent has been well-documented^1,2 and correlated with variation in biotic factors like pollinator behavior,^14–17 these studies were decidedly focused on natural selection, rather than phenotypic plasticity, as an organizational framework.However, in examining the scientific literature on floral scent, we found four studies in which the effects of naturally variable abiotic factors on floral scent profiles were examined, three of which were performed by the same research group (18^–21 (21). Moreover, these studies are decidedly not analyzed and interpreted using standard protocols for phenotypic plasticity studies.¹³

Table 1

A survey of previous studies examining changes in floral scent phenotype due to abiotic factors

Relationships between species’ floral traits and pollinator visitation in a temperate grassland

Knowledge about plant–plant interactions for pollinator service at the plant community level is still scarce, although such interactions may be important to seed production and hence the population dynamics of individual plant species and the species compositions of communities. An important step towards a better understanding of pollination interactions at the community level is to assess if the variation in floral traits among plant species explain the variation in flower visitation frequency among those species. We investigated the relative importance of various floral traits for the visitation frequency of all insects, and bumblebees and flies separately, to plant species by measuring the visitation frequency to all insect-pollinated species in a community during an entire flowering season. Visitation frequency was identified to be strongly positive related to the visual display area and the date of peak flowering of plant species. Categorical variables, such as flower form and symmetry, were important to the visitation frequency of flies only. We constructed floral similarity measures based on the species’ floral traits and found that the floral similarity for all species’ traits combined and the continuous traits separately were positively related to individual visitation frequency. On the other hand, plant species with similar categorical floral traits did not have similar visitation frequencies. In conclusion, our results show that continuous traits, such as flower size and/or density, are more important for the variation in visitation frequency among plant species than thought earlier. Furthermore, differences in visitation frequency among pollinator groups give a poor support to the expectations derived from the classical pollination syndromes.     

Life history, floral development, and mating system in Clarkia xantiana (Onagraceae): do floral and whole-plant rates of development evolve independently?

Autogamously self-fertilizing taxa have evolved from outcrossing progenitors at least 12 times in the annual wildflower genus, Clarkia (Onagraceae). In C. xantiana, individuals of the selfing subspecies (ssp. parviflora) flower at an earlier age, produce successive flowers more rapidly, and produce flowers that complete their development more rapidly than their outcrossing counterparts (ssp. xantiana). Two hypotheses have been proposed to explain the joint evolution of these whole-plant and individual floral traits. The accelerated life cycle hypothesis proposes that selection favoring a short life cycle in environments with short growing seasons (such as those typically occupied by parviflora) has independently favored genotypes with early reproduction, synchronous flower production, and rapidly developing, self-fertilizing flowers. The correlated response to selection hypothesis similarly proposes that selection in environments with short growing seasons favors early reproduction, but that rapid floral development and increased selfing evolve as correlated responses to selection due to genetic linkage (or pleiotropy) affecting both whole-plant and floral development. We conducted a greenhouse experiment using maternal families from two field populations of each subspecies to examine covariation between floral and whole-plant traits within and among populations to seek support for either of these hypotheses. Our results are consistent with the accelerated life cycle hypothesis but not with the correlated response to selection hypothesis.     

Is floral morphology a good predictor of floral visitors to Antirrhineae (snapdragons and relatives)?

• The association between plants and flower visitors has been historically proposed as a main factor driving the evolutionary change of both flower and pollinator phenotypes. The considerable diversity in floral morphology within the tribe Antirrhineae has been traditionally related to pollinator types. We used empirical data on the flower visitors from 59 Antirrhineae taxa from the literature and our own field surveys, which provide an opportunity to test whether flower phenotypes are reliable predictors of visitors and pollinator niches.
• The degree of adjustment between eight key floral traits and actual visitors was explored by testing the predictive value of inferred pollinator syndromes (i.e. suites of floral traits that characterise groups of plant species related to pollination). Actual visitors and inferred pollinator niches (categorisation of visitors’ association using a modularity algorithm) were also explored using Linear Discriminant Analysis (LDA).
• The bee pollinator niche is correctly classified for flowers with dull corolla colour, without nectar guides, as the most important predictor. Both predictive value and statistical classification prove useful in classifying Antirrhineae taxa and the bee pollinator niche, mostly as a consequence of the high proportion of genera and taxa with occluded corollas primarily visited by bees. Our predictive approach rendered a high Positive Predictive Value (PPV) of floral traits in the diagnosis of visitors/pollinator niches. In particular, a high PPV was found for bees as both visitors and forming pollinator niches. In addition, LDA showed that four pollinator niches are well defined based on floral traits.
• The large number of species visited by bees irrespective of pollinator syndromes leads us to hypothesise their generalist pollinator role, despite the phenotypically specialised flowers of Antirrhineae.

     

Do floral herbivores respond to variation in flower characteristics in Gelsemium sempervirens (Loganiaceae), a distylous vine?

We tested the hypothesis that traits traditionally thought to function in pollination can also influence attack by floral herbivores. Because distylous species produce two different flower types, we used Gelsemium sempervirens, a distylous vine of southeastern USA, to examine the frequency and pattern of floral herbivory in relation to floral characteristics. Flowers of the short-styled morph had larger corollas but showed no difference in the volume or concentration of nectar produced. Over the 2 yr of this study, 20-30% of plants suffered floral herbivory. The pattern of damage was morph-specific. Long-styled flowers were more likely to have damaged pistils, while stamens were more often damaged in short-styled flowers. In this distylous species, exserted flower organs were more likely to be eaten. Such differential herbivory based on the position of floral organs within a flower may result in reduced fitness via either male (short-styled morph) or female function (long-styled morph).     

Pollination syndromes in a specialised plant‐pollinator interaction: does floral morphology predict pollinators in Calceolaria?

Pollination syndromes are defined as suites of floral traits evolved in response to selection imposed by a particular group of pollinators (e.g., butterflies, hummingbirds, bats). Although numerous studies demonstrated their occurrence in plants pollinated by radically different pollinators, it is less known whether it is possible to identify them within species pollinated by one functional pollinator group. In such a framework, we expect floral traits to evolve also in response to pollinator subgroups (e.g., species, genera) within that unique functional group. On this, specialised pollination systems represent appropriate case studies to test such expectations. Calceolaria is a highly diversified plant genus pollinated by oil‐collecting bees in genera Centris and Chalepogenus. Variation in floral traits in Calceolaria has recently been suggested to reflect adaptations to pollinator types. However, to date no study has explicitly tested that observation. In this paper, we quantitatively test that hypothesis by evaluating the presence of pollination syndromes within the specialised pollination system formed by several Calceolaria and their insect pollinators. To do so, we use multivariate approaches and explore the structural matching between the morphology of 10 Calceolaria taxa and that of their principal pollinators. Our results identify morphological matching between floral traits related to access to the reward and insect traits involved in oil collection, confirming the presence of pollinator syndromes in Calceolaria. From a general perspective, our findings indicate that the pollination syndrome concept can be also extended to the intra‐pollinator group level.     

Within-population genetic variation in the distylous Primula veris: Does floral morph anisoplethy matter in fragmented habitats?

We studied genetic variation and structure in 24 populations of Primula veris, a distylous self-incompatible perennial herb that in Flanders (northern Belgium) occurs in fragmented habitats of the intensively used agricultural landscape. Distyly, which favours cross-fertilization, is characterized in this species by two genetically determined floral morph types (‘pin’ or ‘thrum’). Using 30 allozyme loci, we investigated within-population and within-morph genetic variation and structure in relation to distyly (stochastic morph bias and pin frequency), demography (population size, plant density), and evidence of recent bottlenecks. No significant relationship was found between genetic patterns and stochastic morph bias. However, F_IS values were lower when pin frequency increases in the population. This may be explained by the partial self-compatibility of the pin morph, which, in conditions of pollination disruption, as observed for P. veris in fragmented habitats, may favour intra-pin outcrosses and increase mating opportunities in the population. A lower within-population structure can be expected because of a higher density of compatible mates and more random matings. Pin and thrum individuals showed similar levels of genetic variation. Three populations revealed a significant genetic differentiation between pin and thrum morphs (F_ST0.049), as a result of recent bottlenecks, spatial segregation of the morphs and/or disturbance of demographic processes. From our findings, we may hypothesize future changes in population mating processes in fragmented habitats, which might lead to pin excesses. However, due to slowing down life-history traits (long-lived perenniality and clonality), and demographic stochasticity randomly changing morph frequency, this may take time before being expressed. Evaluating the impact of morph frequencies on population dynamics is therefore essential for implementing conservation strategies of heterostylous plant species.     

Is floral divergence sufficient to maintain species boundaries upon secondary contact in Mediterranean food-deceptive orchids?

Analyzing the processes that determine whether species boundaries are maintained on secondary contact may shed light on the early phase of speciation. In Anacamptis morio and Anacamptis longicornu, two Mediterranean orchid sister-species, we used molecular and morphological analyses, together with estimates of pollination success and experimental crosses, to assess whether floral isolation can shelter the species' genomes from genetic admixture on secondary contact. We found substantial genetic and morphological homogenization in sympatric populations in combination with an apparent lack of postmating isolation. We further detected asymmetric introgression in the sympatric populations and an imbalance in cytotype representation, which may be due either to a difference in flowering phenology or else be a consequence of cytonuclear incompatibilities. Estimates of genetic clines for markers across sympatric zones revealed markers that significantly deviated from neutral expectations. We observed a significant correlation between spur length and reproductive success in sympatric populations, which may suggest that directional selection is the main cause of morphological differentiation in this species pair. Our results suggest that allopatric divergence has not led to the evolution of sufficient reproductive isolation to prevent genomic admixture on secondary contact in this orchid species pair.     

Plant–floral visitor network structure in a smallholder <Emphasis Type="Italic">Cucurbitaceae</Emphasis> agricultural system in the tropics: implications for the extinction of main floral visitors

Animal pollination is responsible for the majority of the human food supply. Understanding pollination dynamics in agricultural systems is thus essential to help maintain this ecosystem service in the face of human disturbances. Surprisingly, our understanding of plant–pollinator interactions in widely distributed smallholder agricultural systems is still limited. Knowledge of pollination dynamics in these agricultural systems is necessary to fully assess how human disturbances may affect pollination services worldwide. In this study, we describe the structure of a plant–floral visitor network in a smallholder Cucurbitaceae agricultural system. We further identify the main floral visitors of these crops and tested their importance by simulating how their extinction affected network structure and robustness. The observed network was highly connected and generalized but it was neither nested nor compartmentalized. Our results suggest that the structure of agricultural plant–pollinator networks could be inherently different from those in natural communities. These differences in network structure may reflect differences in spatial distribution of floral resources between agricultural and natural systems. We identified Augochlora nigrocyanea and Peponapis limitaris as the two most frequent floral visitors. However, removal of these species did not affect network structure or its robustness, suggesting high levels of interaction rewiring. To our knowledge, this is one of the first studies to describe the structure of a plant–floral visitor network in diverse agricultural systems in the tropics. We emphasize the need for more studies that evaluate network structure in agricultural systems if we want to fully elucidate the impact of human disturbances on this key ecosystem service.     

The emission of floral scent from Lilium ‘siberia’ in response to light intensity and temperature

Floral scent is an important part of volatile compounds emitted from plants, and is influenced by many environmental factors. In this study, the floral scent emitted from Lilium ‘siberia’, a common breed of lily, was collected by dynamic headspace at different levels of light intensity (0, 100, 300, 600, 1,000, and 1,500 μmol m^?2 s^?1) and temperature (10, 20, 30, and 40 °C). Using the automated thermal desorption-gas chromatography/mass spectrometry (ATD-GC/MS) technique, the components and release amounts were subsequently identified to investigate the influence of light and temperature on the emission of floral scent. The results revealed that the numbers and release amounts of floral scent components were significantly influenced by light intensity and temperature, showing the similar pattern: first increasing and then decreasing. After light intensity treatment, the maximum numbers and release amounts mainly appeared at 600 and 1,000 μmol m^?2 s^?1. For temperature treatment, 30 °C resulted in the highest numbers and release amounts of the floral scent components. At different levels of light intensity and temperature, terpenoid compounds showed the highest numbers and release amounts among the component categories. α-Ocimene and linalool were the two terpenoid compounds with the highest release amounts, and accounted for the highest proportion. The results obtained provide evidence that both light intensity and temperature trigger the emission of floral scent. The particular response mechanisms must be investigated in future research.     

Early Cretaceous floral structures and in situ tricolpate‐striate pollen: New early eudicots from Portugal

Five new taxa with affinities to extant lineages that diverged at an early stage from the main line of eudicot evolution are established from the Early Cretaceous (late Aptian or early Albian) Vale de Agua locality, Portugal. Staminate flowers of Lusistemon striatus and pistillate flowers of Lusicarpus planatus are unisexual without rudiments of the opposite gender. They are linked by the association of an unusual pollen type found in situ in the stamens and adhering to the stigmatic surface. The staminate flower, Lusistemon striatus, is composed of six stamens subtended by small perianth parts. The arrangement of the stamens is difficult to ascertain, but their variable sizes suggests a spiral arrangement. Pollen found in situ is tricolpate and striate with densely‐spaced, sparsely diverging and anastomosing muri that are aligned more or less parallel to the polar axis. The muri have a conspicuous supratectal ornamentation of fine transverse ridges. The granular infratectal layer forms an indistinct internal reticulum. The foot layer is thin. Pollen is closely similar to dispersed grains from the Aptian of Egypt described as STRIOTRI‐SEGMUR. It also resembles pollen of the dispersed pollen genus Rutihesperipites, as well as some dispersed pollen assigned to Striatopollis. Pistillate flowers of Lusicarpus planatus consist of a bicarpellate, syncarpous gynoecium borne on a short stalk. The styles are bent outwards and expose the double‐crested stigmatic regions on their ventral sides. The only organ preserved besides the gynoecium is a lateral scale‐like organ at the base of the stalk. Pollen of the same type found in Lusistemon striatus occurs on the stigmatic surface of the carpels. Comparisons with extant taxa demonstrate that Lusistemon and Lusicarpus share many characters with early diverging groups of eudicots, in particular Buxaceae. In addition to the Lusistemon‐Lusicarpus flowers, the Vale de Agua samples also contain three other pistillate reproductive structures that may be related to early diverging lineages of eudicots. Silucarpus camptostylus has a bicarpellate and syncarpous gynoecium with two styles; Valecarpus petiolatus and Aguacarpus hirsutus have tricarpellate gynoecia that are distinguished from each other in the shape and extension of the stigma as well as other details.     

How does insect visitation trigger floral colour change?

Abstract.  1. Visitation by the key pollinator, Bombus terrestris , was implicated in inducible flower colour change in Lupinus pilosus . Behaviour at the flower and rate of visitation by these bumble bees had specific effects; exclusion of this flower visitor led to retarded onset, and reduced rate, of colour change.
2. The foraging behaviour of B. terrestris was influenced by floral colour change in L. pilosus . Choice of pre-change flowers was greater than random in relation to the proportion of colour phases available within the plant population.
3. Levels of floral manipulation that mimicked the flower handling characteristics of visiting bumble bees confirmed that triggering of the pollen release mechanism is necessary for the instigation of colour change.
4. Moreover, this study suggests that, in L. pilosus , an aspect of pollination (pollen deposition by bees and/or subsequent pollen tube growth within the style) is linked with colour change and may act as the trigger for such change.     

Can floral field margins improve pollination and seed production in coriander Coriandrum sativum L. (Apiaceae)?

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