Enhanced reproductive success revealed key strategy for persistence of devastated populations in Himalayan food‐deceptive orchid,Dactylorhiza hatagirea

Anthropogenic disturbances adversely affect populations of rare and endemic plants, resulting in reduction of their population size and performance. Among different plant groups, deceptive terrestrial orchids are vulnerable and possess greater extinction risks because of rarity in occurrence. To understand the response of food‐deceptive terrestrial orchids to disturbances, we selected Dactylorhiza hatagirea as our representative species, which is endemic to Himalaya, and studied its natural populations. This species is rare for being habitat specific, pollination limited and threatened in its natural habitats. We tested the hypothesis that disturbances lead to reduction in population size and plant performance of food‐deceptive terrestrial orchids. For assessing the impact of disturbance, two contrasting groups, heavily devastated (HD) and lightly devastated (LD), were identified on the basis of frequency and intensity of disturbance (harvesting of plant for tubers) by interviewing local people, medicinal plant extractors and shepherds. HD sites, in comparison to LD sites, were found to have smaller population sizes, but showed an increase in plant growth traits (plant height, specific leaf area, leaf N and specific shoot length). Similarly, plants at HD sites were found to have invested less in inflorescence (inflorescence size, inflorescence length, inflorescence length fraction and flowers per length), but despite that showed higher reproductive success. This was a clear indication of enhanced performance of its populations driven by disturbances. Our findings suggested that food‐deceptive species in small populations tend to reduce the probability of population extinction and have the capability to recover rapidly if conserved in time.     

Orchid pollination by sexual deception: pollinator perspectives

The extraordinary taxonomic and morphological diversity of orchids is accompanied by a remarkable range of pollinators and pollination systems. Sexually deceptive orchids are adapted to attract specific male insects that are fooled into attempting to mate with orchid flowers and inadvertently acting as pollinators. This review summarises current knowledge, explores new hypotheses in the literature, and introduces some new approaches to understanding sexual deception from the perspective of the duped pollinator. Four main topics are addressed: (1) global patterns in sexual deception, (2) pollinator identities, mating systems and behaviours, (3) pollinator perception of orchid deceptive signals, and (4) the evolutionary implications of pollinator responses to orchid deception, including potential costs imposed on pollinators by orchids. A global list of known and putative sexually deceptive orchids and their pollinators is provided and methods for incorporating pollinator perspectives into sexual deception research are provided and reviewed. At present, almost all known sexually deceptive orchid taxa are from Australia or Europe. A few sexually deceptive species and genera are reported for New Zealand and South Africa. In Central and Southern America, Asia, and the Pacific many more species are likely to be identified in the future. Despite the great diversity of sexually deceptive orchid genera in Australia, pollination rates reported in the literature are similar between Australian and European species. The typical pollinator of a sexually deceptive orchid is a male insect of a species that is polygynous, monandrous, haplodiploid, and solitary rather than social. Insect behaviours involved in the pollination of sexually deceptive orchids include pre‐copulatory gripping of flowers, brief entrapment, mating, and very rarely, ejaculation. Pollinator behaviour varies within and among pollinator species. Deception involving orchid mimicry of insect scent signals is becoming well understood for some species, but visual and tactile signals such as colour, shape, and texture remain neglected. Experimental manipulations that test for function, multi‐signal interactions, and pollinator perception of these signals are required. Furthermore, other forms of deception such as exploitation of pollinator sensory biases or mating preferences merit more comprehensive investigation. Application of molecular techniques adapted from model plants and animals is likely to deliver new insights into orchid signalling, and pollinator perception and behaviour. There is little current evidence that sexual deception drives any species‐level selection on pollinators. Pollinators do learn to avoid deceptive orchids and their locations, but this is not necessarily a response specific to orchids. Even in systems where evidence suggests that orchids do interfere with pollinator mating opportunities, considerable further research is required to determine whether this is sufficient to impose selection on pollinators or generate antagonistic coevolution or an arms race between orchids and their pollinators. Botanists, taxonomists and chemical ecologists have made remarkable progress in the study of deceptive orchid pollination. Further complementary investigations from entomology and behavioural ecology perspectives should prove fascinating and engender a more complete understanding of the evolution and maintenance of such enigmatic plant‐animal interactions.     

Patterns of reproductive isolation in Mediterranean deceptive orchids

The evolution of reproductive isolation is of central interest in evolutionary biology. In plants, this is typically achieved by a combination of pre- and postpollination mechanisms that prevent, or limit, the amount of interspecific gene flow. Here, we investigated and compared two ecologically defined groups of Mediterranean orchids that differ in pollination biology and pollinator specificity: sexually deceptive orchids versus food-deceptive orchids. We used experimental crosses to assess the strength of postmating prezygotic, and postzygotic reproductive isolation, and a phylogenetic framework to determine their relative rates of evolution. We found quantitative and qualitative differences between the two groups. Food-deceptive orchids have weak premating isolation but strong postmating isolation, whereas the converse situation characterizes sexually deceptive orchids. Only postzygotic reproductive isolation among food-deceptive orchids was found to evolve in a clock-like manner. Comparison of evolutionary rates, within a common interval of genetic distance, showed that the contribution of postmating barriers was more relevant in the food-deceptive species than in the sexually deceptive species. Asymmetry in prezygotic isolation was found among food-deceptive species. Our results indicate that postmating barriers are most important for reproductive isolation in food-deceptive orchids, whereas premating barriers are most important in sexually deceptive orchids. The different rate of evolution of reproductive isolation and the relative strength of pre- and postmating barriers may have implication for speciation processes in the two orchid groups.     

Female and male fitness of a sexually deceptive orchid with a narrow distribution area: from phenotypic traits to spatial distribution patterns

• The Orchidaceae family presents one of the most extravagant pollination mechanisms: deception. While many studies on reproductive success have been performed on food‐deception orchids, less have been performed on sexually deceptive orchids. Here, we focused on Ophrys balearica P. Delforge, an endemic orchid of the Balearic Islands, to study its reproductive ecology, the spatio‐temporal variation of its reproductive success and the individual (floral display and geospatial position) and population parameters (patch size, shape and density) that affect its reproductive success.
• We performed hand‐pollination experiments, along with the recording of floral display parameters and GPS position of over 1,100 individuals from seven populations in two consecutive years. We applied, for the first time, GIS tools to analyse the effects of individual’s position within the population on the reproductive success. Reproductive success was measured both in male (removed pollinia) and female (fruit set) fitness.
• The results confirm that this species is pollinator‐dependent and mostly allogamous, but also self‐compatible. This species showed high values for the cumulative inbreeding depression index and high pollen limitation. Male fitness was almost equal to female fitness between years and populations, and reproductive success exhibited huge spatio‐temporal variation.
• Although we did not find strong correlations between floral display and reproductive success, patches with low‐plant density and individuals in the external portion of the population showed significantly higher plant fitness. These findings must be considered in conservation actions for endangered orchid species, especially considering that most orchids are strongly dependent on pollinators for their species’ fitness.

     

Pollen transfer efficiency and its effect on inflorescence size in deceptive pollination strategies

Pollination systems differ in pollen transfer efficiency, a variable that may influence the evolution of flower number. Here we apply a comparative approach to examine the link between pollen transfer efficiency and the evolution of inflorescence size in food and sexually deceptive orchids. We examined pollination performance in nine food‐deceptive, and eight sexually deceptive orchids by recording pollen removal and deposition in the field. We calculated correlations between reproductive success and flower number (as a proxy for resources allocated during reproductive process), and directional selection differentials were estimated on flower number for four species. Results indicate that sexually deceptive species experience decreased pollen loss compared to food‐deceptive species. Despite producing fewer flowers, sexually deceptive species attained levels of overall pollination success (through male and female function) similar to food‐deceptive species. Furthermore, a positive correlation between flower number and pollination success was observed in food‐deceptive species, but this correlation was not detected in sexually deceptive species. Directional selection differentials for flower number were significantly higher in food compared to sexually deceptive species. We suggest that pollination systems with more efficient pollen transfer and no correlation between pollination success and number of flowers produced, such as sexual deception, may allow the production of inflorescences with fewer flowers that permit the plant to allocate fewer resources to floral displays and, at the same time, limit transpiration. This strategy can be particularly important for ecological success in Mediterranean water‐deprived habitats, and might explain the high frequency of sexually deceptive species in these specialised ecosystems.     

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

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

Evolutionary relationships among pollinators and repeated pollinator sharing in sexually deceptive orchids

The mechanism of pollinator attraction is predicted to strongly influence both plant diversification and the extent of pollinator sharing between species. Sexually deceptive orchids rely on mimicry of species‐specific sex pheromones to attract their insect pollinators. Given that sex pheromones tend to be conserved among related species, we predicted that in sexually deceptive orchids, (i) pollinator sharing is rare, (ii) closely related orchids use closely related pollinators and (iii) there is strong bias in the wasp lineages exploited by orchids. We focused on species that are pollinated by sexual deception of thynnine wasps in the distantly related genera Caladenia and Drakaea, including new field observations for 45 species of Caladenia. Specialization was extreme with most orchids using a single pollinator species. Unexpectedly, seven cases of pollinator sharing were found, including two between Caladenia and Drakaea, which exhibit strikingly different floral morphology. Phylogenetic analysis of pollinators using four nuclear sequence loci demonstrated that although orchids within major clades primarily use closely related pollinator species, up to 17% of orchids within these clades are pollinated by a member of a phylogenetically distant wasp genus. Further, compared to the total diversity of thynnine wasps within the study region, orchids show a strong bias towards exploiting certain genera. Although these patterns may arise through conservatism in the chemical classes used in sex pheromones, apparent switches between wasp clades suggest unexpected flexibility in floral semiochemical production. Alternatively, wasp sex pheromones within lineages may exhibit greater chemical diversity than currently appreciated.     

Behaviour of sexually deceived ichneumonid wasps and its implications for pollination in Cryptostylis (Orchidaceae)

Pollination via sexual deception is hypothesised to be associated with more frequent outcrossing and greater pollen dispersal distances than strategies involving food‐foraging behaviour. In this study, we investigated the behaviour and movement distances of Lissopimpla excelsa (Hymenoptera: Ichneumonidae), and their implications for the pollination of the sexually deceptive Cryptostylis ovata (Orchidaceae). Pollinator observations revealed that while L. excelsa will alight on multiple flowers within a single visit to a patch of orchids, the frequency of attempted copulation decreases with successive visits, suggesting that pollinator learning may inhibit within‐patch pollen transfer. Mark‐recapture demonstrated that 25% of wasps revisited inflorescences within a day and 50% revisited within a week. Despite the apparent site fidelity of some individuals, L. excelsa often move over large distances (maximum = 625 m), and are capable of dispersing pollen between patches. To resolve the consequences of pollination by sexual deception of ichneumonids, we compared our results with those from studies of other sexually deceptive systems. While pollination rates were comparable with other sexually deceptive orchids, L. excelsa showed high rates of column contact and moved over large distances relative to other sexually deceived pollinators. Among sexually deceptive orchids in general, the frequency of column contact was not correlated either with the frequency of pseudocopulation or with pollination rate. These results suggest that the consequences of pollination by sexual deception may vary extensively between plant taxa due to variation in floral traits, and behavioural differences between pollinator groups.     

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.     

Generalized food-deceptive orchid species flower earlier and occur at lower altitudes than rewarding ones

Aims Food-deceptive pollination, in which plants do not offer any food reward to their pollinators, is common within the Orchidaceae. As food-deceptive orchids are poorer competitors for pollinator visitation than rewarding orchids, their occurrence in a given habitat may be more constrained than that of rewarding orchids. In particular, the success of deceptive orchids strongly relies on several biotic factors such as interactions with co-flowering rewarding species and pollinators, which may vary with altitude and over time. Our study compares generalized food-deceptive (i.e. excluding sexually deceptive) and rewarding orchids to test whether (i) deceptive orchids flower earlier compared to their rewarding counterparts and whether (ii) the relative occurrence of deceptive orchids decreases with increasing altitude.Methods To compare the flowering phenology of rewarding and deceptive orchids, we analysed data compiled from the literature at the species level over the occidental Palaearctic area. Since flowering phenology can be constrained by the latitudinal distribution of the species and by their phylogenetic relationships, we accounted for these factors in our analysis. To compare the altitudinal distribution of rewarding and deceptive orchids, we used field observations made over the entire Swiss territory and over two Swiss mountain ranges.Important findings We found that deceptive orchid species start flowering earlier than rewarding orchids do, which is in accordance with the hypotheses of exploitation of naive pollinators and/or avoidance of competition with rewarding co-occurring species. Also, the relative frequency of deceptive orchids decreases with altitude, suggesting that deception may be less profitable at high compared to low altitude.     

Chemical ecology and pollinator-driven speciation in sexually deceptive orchids

Sexually deceptive orchids mimic females of their pollinator species to attract male insects for pollination. Pollination by sexual deception has independently evolved in European, Australian, South African, and South American orchid taxa. Reproductive isolation is mainly based on pre-mating isolation barriers, the specific attraction of males of a single pollinator species, mostly bees, by mimicking the female species-specific sex-pheromone. However, in rare cases post-mating barriers have been found. Sexually deceptive orchids are ideal candidates for studies of sympatric speciation, because key adaptive traits such as the pollinator-attracting scent are associated with their reproductive success and with pre-mating isolation.During the last two decades several investigations studied processes of ecological speciation in sexually deceptive orchids of Europe and Australia. Using various methods like behavioural experiments, chemical, electrophysiological, and population-genetic analyses it was shown that minor changes in floral odour bouquets might be the driving force for pollinator shifts and speciation events. New pollinators act as an isolation barrier towards other sympatrically occurring species. Hybridization occurs because of similar odour bouquets of species and the overlap of flowering periods. Hybrid speciation can also lead to the displacement of species by the hybrid population, if its reproductive success is higher than that in the parental species.     

Review. Specificity in pollination and consequences for postmating reproductive isolation in deceptive Mediterranean orchids

The type of reproductive isolation prevalent in the initial stages of species divergence can affect the nature and rate of emergence of additional reproductive barriers that subsequently strengthen isolation between species. Different groups of Mediterranean deceptive orchids are characterized by different levels of pollinator specificity. Whereas food-deceptive orchid species show weak pollinator specificity, the sexually deceptive Ophrys species display a more specialized pollination strategy. Comparative analyses reveal that orchids with high pollinator specificity mostly rely on premating reproductive barriers and have very little postmating isolation. In this group, a shift to a novel pollinator achieved by modifying the odour bouquet may represent the main isolation mechanism involved in speciation. By contrast, orchids with weak premating isolation, such as generalized food-deceptive orchids, show strong evidence for intrinsic postmating reproductive barriers, particularly for late-acting postzygotic barriers such as hybrid sterility. In such species, chromosomal differences may have played a key role in species isolation, although strong postmating-prezygotic isolation has also evolved in these orchids. Molecular analyses of hybrid zones indicate that the types and strength of reproductive barriers in deceptive orchids with contrasting premating isolation mechanisms directly affect the rate and evolutionary consequences of hybridization and the nature of species differentiation.     

EFFECTS OF DISPLAY SIZE,FLOWERING PHENOLOGY,AND NECTAR AVAILABILITY ON EFFECTIVE VISITATION FREQUENCY IN COMPARETTIA FALCATA (ORCHIDACEAE)

Comparettia falcata is an epiphytic, neotropical orchid that produces nectar as a pollinator reward. In Puerto Rico, C. falcata is allogamous and pollinated by the endemic hummingbird Chlorostilbon maugaeus. Autogamous pollinations are possible, but may result in reduced fruit set. For the 1989 and 1990 flowering seasons, the probability of pollinarium removals and natural pollinations increased with individual inflorescence display size. However, the frequency of effective pollinator visits was independent of flowering phenology in both years. A positive correlation between inflorescence size and reproductive success occurred in 1990 but not in 1989. In 1990 plants produced longer spurs, a higher standing crop of nectar, and a more concentrated nectar than in 1989. There was no relation between nectar availability (= standing crop of nectar) and sugar concentration in either year. Nectar availability and sugar concentration did not vary among the first four flowers of an inflorescence in either season. Nectar availability was not a good predictor of effective visitation. Comparettia falcata has a higher natural fruit set than tropical deceptive orchids, suggesting that pollinator visitation may be enhanced by nectar reward. The small, dilute nectar volumes secreted by C. falcata may benefit the plant by increasing interplant pollinator movement and pollen dispersal.     

CONTRASTING THOUGHTS ABOUT DECEPTIVE ORCHIDS: A RESPONSE TO SOBEL AND RANDLE

Sobel and Randle (2009) challenge several methodological choices in the comparative study of the evolution of reproductive isolation in Mediterranean deceptive orchids of Scopece et al. (2007) including the species concept used and the selection of taxa, together with the perceived comparison of clades of different ages. They further criticize that pollinator information was taken from the literature and that two different methods were used to estimate pollinator specificity in food-deceptive and sexually deceptive orchids, respectively. Here we reply to these challenges.     

The effect of inflorescence display size and flower position on pollination success in two deceptive and one rewarding orchid

• Inflorescence display size and flower position on the inflorescence play important roles in plant reproduction, in the formation of fruits and are primarily linked to pollinator behaviour. We used three orchids to determine how visitation rates and choice of pollinator depend on number and position of the flowers along the inflorescence.
• We measured reproductive success in (1) natural conditions, (2) hand-pollination experiments and (3) an experimental design, by modifying composition of inflorescences in populations of two deceptive orchids, Orchis anthropophora and O. italica, and one rewarding orchid, Anacamptis coriophora subsp. fragrans.
• There were no differences in natural fruit production in relation to flower position on the inflorescence (i.e. upper versus lower part), suggesting no preference of pollinators for different parts of the inflorescence. Hand-pollination experiments highlighted low pollen limitation in A. coriophora subsp. fragrans but high limitation in O. italica and O. anthropophora. Reproductive success of deceptive orchids in experimental plots decreased significantly when flowers on the upper half of the inflorescence were removed leading to reduced floral display, while reproductive success of the nectariferous species did not differ significantly.
• Our data highlight that in the examined orchids there is no clear relationship between fruit formation and flower position along inflorescences. Thus we can affirm that, for orchids, the entire inflorescence plays a dominant role in insect attraction but the part of the flower spike does not influence the choice of the insect. This implies that all flowers have the same possibility of receiving visits from pollinators, and therefore each flower has the same opportunity to set fruit.

     

Are deception-pollinated species more variable than those offering a reward?

In most pollination systems, animals transfer pollen among plants of a given species. Pollinator visitations do not come without cost, so plants usually offer a reward. However, the flowers of some plant species, mostly orchids, lack rewards and deceive animals into visiting their flowers. Deceptive species are thought to have high levels of variation in traits associated with advertisement and pollinator attraction, which have been attributed to genetic drift, or disruptive selection due to pollinator behavior. Rewarding species are assumed to have less variation due to stabilizing selection. We compared variability in floral morphology and fragrance composition between deceptive and rewarding species. Because both suites of traits are often linked with floral advertisement and pollinator attraction, we expected variation to be greater in species with deceptive pollination systems than in those offering rewards. We obtained floral morphology metrics for 20 deceptive species and 41 rewarding species native or naturalized in Puerto Rico, Venezuela, and Ecuador. Floral fragrances were sampled from eight deceptive species and four rewarding species. We found that the amplitude of variation in floral morphology and fragrance composition covaries significantly. Comparison of coefficients of variation for morphology indicated that, overall, deceptive species show significantly higher variation than rewarding species, and this pattern was also found among just orchids or just nonorchids. There were no statistical differences in morphological variation between orchids and nonorchids within a functional pollination group. Fragrance variation, measured by Jaccard distance, tended to be greater for deceptive species than for rewarding species. Although overlap in measures of variation occurs between the two groups, the data support the hypothesis that populations of deception-pollinated species are more variable than rewarding species in traits associated with pollinator attraction.     

Modeling the two‐locus architecture of divergent pollinator adaptation: how variation in SAD paralogs affects fitness and evolutionary divergence in sexually deceptive orchids

Divergent selection by pollinators can bring about strong reproductive isolation via changes at few genes of large effect. This has recently been demonstrated in sexually deceptive orchids, where studies (1) quantified the strength of reproductive isolation in the field; (2) identified genes that appear to be causal for reproductive isolation; and (3) demonstrated selection by analysis of natural variation in gene sequence and expression. In a group of closely related Ophrys orchids, specific floral scent components, namely n‐alkenes, are the key floral traits that control specific pollinator attraction by chemical mimicry of insect sex pheromones. The genetic basis of species‐specific differences in alkene production mainly lies in two biosynthetic genes encoding stearoyl–acyl carrier protein desaturases (SAD) that are associated with floral scent variation and reproductive isolation between closely related species, and evolve under pollinator‐mediated selection. However, the implications of this genetic architecture of key floral traits on the evolutionary processes of pollinator adaptation and speciation in this plant group remain unclear. Here, we expand on these recent findings to model scenarios of adaptive evolutionary change at SAD2 and SAD5, their effects on plant fitness (i.e., offspring number), and the dynamics of speciation. Our model suggests that the two‐locus architecture of reproductive isolation allows for rapid sympatric speciation by pollinator shift; however, the likelihood of such pollinator‐mediated speciation is asymmetric between the two orchid species O. sphegodes and O. exaltata due to different fitness effects of their predominant SAD2 and SAD5 alleles. Our study not only provides insight into pollinator adaptation and speciation mechanisms of sexually deceptive orchids but also demonstrates the power of applying a modeling approach to the study of pollinator‐driven ecological speciation.     

Effect of individual height and labellum colour on the pollination of Caladenia (syn. Arachnorchis) behrii (Orchidaceae) in the northern Adelaide region, South Australia

Caladenia behrii, a sexually deceptive orchid, is an endangered terrestrial orchid endemic to the Adelaide Hills, South Australia. We examined pollination success among individuals of different colours and heights in three populations of C. behrii in the northern Adelaide region. The labellum of C. behrii varies from cream to deep maroon. Over five years, the pollination success (pollinia deposited on stigma) of four colour groups (maroon, maroon and cream, hatched, and cream) was assessed for a total of 782 flowers. Labellum colour had no effect on the pollination success of individuals, indicating no frequency-dependent selection. It is likely that pollinators use the sexual pheromone mimic produced by the flowers as the main orientation cue. For three years the height of pollinated C. behrii individuals was recorded, along with the height of the nearest non-pollinated neighbour (distance of labellum from the ground, rather than stem length). Although a range of different heights was pollinated at population level, individual flowers that were taller than their closest neighbours were significantly more likely to be pollinated. Flower head height was not correlated with opening date and was greater for orchids growing among grass-tree leaves. Results imply that pollinators are more successful in finding tall flowers than short ones for visual and/or olfactory reasons; however, the effect of height preference is eliminated at population level.     

On the roles of colour and scent in a specialized floral mimicry system

Background and Aims

Sexually deceptive orchids achieve cross-pollination by mimicking the mating signals of female insects, generally hymenopterans. This pollination mechanism is often highly specific as it is based primarily on the mimicry of mating signals, especially the female sex pheromones of the targeted pollinator. Like many deceptive orchids, the Mediterranean species Ophrys arachnitiformis shows high levels of floral trait variation, especially in the colour of the perianth, which is either green or white/pinkinsh within populations. The adaptive significance of perianth colour polymorphism and its influence on pollinator visitation rates in sexually deceptive orchids remain obscure.

Methods

The relative importance of floral scent versus perianth colour in pollinator attraction in this orchid pollinator mimicry system was evaluated by performing floral scent analyses by gas chromatography-mass spectrometry (GC-MS) and behavioural bioassays with the pollinators under natural conditions were performed.

Key Results

The relative and absolute amounts of behaviourally active compounds are identical in the two colour morphs of O. arachnitiformis. Neither presence/absence nor the colour of the perianth (green versus white) influence attractiveness of the flowers to Colletes cunicularius males, the main pollinator of O. arachnitiformis.

Conclusion

Chemical signals alone can mediate the interactions in highly specialized mimicry systems. Floral colour polymorphism in O. arachnitiformis is not subjected to selection imposed by C. cunicularius males, and an interplay between different non-adaptive processes may be responsible for the maintenance of floral colour polymorphism both within and among populations.     

COMPARATIVE APPROACHES TO THE EVOLUTION OF REPRODUCTIVE ISOLATION: A COMMENT ON SCOPECE ET AL. 2007

Speciation can be driven by the evolution of many forms of reproductive isolation. Comparative study is a powerful approach for elucidating the relative importance of individual isolating barriers in the speciation process. A recent contribution by Scopece and colleagues provides comparative data for two groups of deceptive pollination orchids and aims to test hypotheses about which forms of isolation are most important in the two clades. The authors compare pollinator isolation and postmating isolation between the two orchid groups, and conclude that food-deceptive orchid species have less isolation by pollinator specificity than sexually deceptive species, and that postmating isolation is more important in the food-deceptive clade. Although we find this approach to be novel and potentially powerful, these conclusions are called into question by the methods used to define and select species and quantify pollinator isolation. Definition and selection of taxa were performed in a biased manner that undermines the ability to infer general patterns of speciation. Furthermore, pollinator isolation was calculated inconsistently for the two groups under study, effectively nullifying the comparison.