Colour similarity to rewarding model plants affects pollination in a food deceptive orchid, Orchis boryi

We studied the pollination of Orchis boryi at five different locations on the Greek mainland. Orchis boryi is food deceptive and obligatorily insect pollinated. Primary pollinators were Apis mellifera and Bombus spp., which foraged on rewarding plant species nearby and visited O. boryi in between. To analyse floral colour similarity among rewarding plants and O. boryi as perceived by bees, a model of bee colour vision was employed. For each food plant an index was calculated that described the probability of a bee foraging on it to subsequently choose an orchid flower. This choice probability correlated to colour distance according to the model of bee colour vision, indicating that bees chose the deceptive orchid more frequently if they foraged on more similarly coloured species. At different sites different plant species served as models. Bees foraging on food plants from which a high choice rate to the orchid was observed visited the orchid less often after approaching it than other bees, which is likely to reflect avoidance learning. In general, the pollination syndrome appears to be a generalized form of Batesian mimicry, in which similarity to rewarding plants determines reproductive success. As expected by negative density-dependent selection, individual fruit set and pollinia export rate correlated negatively with orchid density, but were unaffected by food plant density, orchid frequency, individual variation of labellum colour, labellum size, or mouth width of the flowers.     

Effects of habitat type change on taxonomic and functional composition of orchid bees (Apidae: Euglossini) in the Brazilian Amazon

Land use change impact species richness and functional diversity (FD). In the Brazilian Amazon, we examined the impacts of oil palm plantations on orchid bee (Apidae: Euglossini) species using abundance and FD. We collected male orchid bees in oil palm plantation (PALM), legal reserves (LR), and riparian corridors (APP), and then we used morphological and life-history traits to characterize each species. We evaluated differences in bee body size by comparing intertegular span values. We tested the influence of habitat on taxonomic and functional parameters of orchid bees by applying a partial redundancy analysis (pRDA). We contrasted FD by calculating species richness, functional richness, and functional dispersion. We sampled 1176 bees from 30 species in 18 sampling days across 2015 and 2016. Males from PALM were 13.6% bigger than those in LR areas, and bees from APP showed a similar pattern compared to LR and PALM. Less than 15% of the variation in species composition was related to the distance among sampling sites, and 8% was due to habitat structure. In our pRDA, the spatial difference explained 6% of the variation in orchid bee traits, but there were no effects of habitat parameters upon FD. FD was reduced with land use change caused by oil palm plantations. Our findings support the belief that many bees are impacted by cultivated lands. Nevertheless, the functional similarity between LRs and APPs reflects common structural elements between them, although we did not find significant relationship between functional composition and habitat structure that we evaluated.     

Evidence for pollinator sharing in Mediterranean nectar-mimic orchids: absence of premating barriers?

Pollinator specificity has traditionally been considered the main reproductive isolation mechanism in orchids. Among Mediterranean orchids, however, many species attract and deceive pollinators by mimicking nectar-rewarding plants. To test the extent to which deceptive orchid species share pollinators, we collected and identified hemipollinaria-carrying insects, and used ribosomal sequences to identify the orchid species from which hemipollinaria were removed. We found that social and solitary bees, and also flies, carried hemipollinaria belonging to nine orchid species with different degrees of specialization. In particular, Anacamptis morio, Dactylorhiza romana and Orchis mascula used a large set of pollinator species, whereas others such as Orchis quadripunctata seemed to be pollinated by one pollinator species only. Out of the insects with hemipollinaria, 19% were found to carry hemipollinaria from more than one orchid species, indicating that sympatric food-deceptive orchids can share pollinators. This sharing was apparent even among orchid sister-species, thus revealing an effective overlap in pollinator sets among closely related species. These results suggest varying degrees of pollinator specificity in these orchids, and indicate that pollinator specificity cannot always act as the main isolation mechanism in food-deceptive temperate orchids.     

Specificity and mutual dependency of the orchid-euglossine bee interaction

Seasonal and geographic relationships, and host pollinator specificities are examined for indications of interdependency in the orchid-euglossine bee interaction. The orchids are dependent on the bees for pollination, and their flowering seasonality corresponds well with peak activity of their pollinators. However, there is little evidence that the bees are dependent on these fragrance hosts. The orchids tap the majority of euglossine species and individuals for pollinator services during any given season, but most of those bee species that temporarily lack orchid fragrance hosts persist in the area, continually emerge from nests, and seek floral fragrance compounds. Pollinator specificity occurs in less than half of the orchids, and host specificity is rare. Geographic distributions of nearly all orchid-pollinator pairs are not mutually inclusive. Moreover, nearly a third of the local male euglossine bee species censused are not pollinators of any fragrance orchids in the area. Local alternative fragrance sources occur. The orchid-male euglossine bee interaction does not appear to represent a mutually obligatory relationship. The orchids may have exploited a preexisting behavioural phenomenon of the bees, and reciprocal evolutionary responses may not have occurred.     

Seasonal cycles,phylogenetic assembly,and functional diversity of orchid bee communities

Neotropical rainforests sustain some of the most diverse terrestrial communities on Earth. Euglossine (or orchid) bees are a diverse lineage of insect pollinators distributed throughout the American tropics, where they provide pollination services to a staggering diversity of flowering plant taxa. Elucidating the seasonal patterns of phylogenetic assembly and functional trait diversity of bee communities can shed new light into the mechanisms that govern the assembly of bee pollinator communities and the potential effects of declining bee populations. Male euglossine bees collect, store, and accumulate odoriferous compounds (perfumes) to subsequently use during courtship display. Thus, synthetic chemical baits can be used to attract and monitor euglossine bee populations. We conducted monthly censuses of orchid bees in three sites in the Magdalena valley of Colombia – a region where Central and South American biotas converge – to investigate the structure, diversity, and assembly of euglossine bee communities through time in relation to seasonal climatic cycles. In particular, we tested the hypothesis that phylogenetic community structure and functional trait diversity changed in response to seasonal rainfall fluctuations. All communities exhibited strong to moderate phylogenetic clustering throughout the year, with few pronounced bursts of phylogenetic overdispersion that coincided with the transition from wet‐to‐dry seasons. Despite the heterogeneous distribution of functional traits (e.g., body size, body mass, and proboscis length) and the observed seasonal fluctuations in phylogenetic diversity, we found that functional trait diversity, evenness, and divergence remained constant during all seasons in all communities. However, similar to the pattern observed with phylogenetic diversity, functional trait richness fluctuated markedly with rainfall in all sites. These results emphasize the importance of considering seasonal fluctuations in community assembly and provide a glimpse to the potential effects that climatic alterations may have on both pollinator communities and the ecosystem services they provide.     

Long-term ecology of euglossine orchid-bees (Apidae: Euglossini) in Panama

Summary Abundance patterns during 6–7 years and orchid visitation were determined for 51 species of the 57 local euglossine bees. Male bees were counted at 3 chemical attractants presented in the same manner each month. Sites were separated by 75 km but included wet Atlantic forest at 500 m elevation, moist forest at 180 m near Barro Colorado Island, and cloud forest at 900 m near the Pacific ocean. 1. From 15 to 30 euglossine species of 4 genera were active in each month and site; monthly species number and general bee abundance were positively correlated. Many species had 3 annual abundance peaks (range 1–4) and were active throughout the year, but peak annual abundances rarely occurred during late wet or early dry seasons. In contrast, Eufriesea generally were present as adults only 1–2 months in a year. 2. Euglossine populations were exceptionally stable. Species at each site were more stable than any known insect population, and stability and abundance were positively associated. However, year-to-year population stability and the degree of seasonality were not correlated. Among the three sites, the more diverse (species rich) bee assemblages displayed lower stability; these were the wetter and upland sites. 3. The most abundant bees visited more orchid species. Eg. and El. each visited and average of 4 orchid species (range 0–13); Ex. and Ef. visited 0–3. Stable populations did not visit more or fewer orchid species than did unstable populations. 4. Less than 68% of species at each site visited orchid flowers; less than a few dozen of the 100–800 bees counted in a day carried orchid pollinaria. Over 20% of the euglossine species never were seen with pollinaria at any site and probably seldom visit orchids in central Panama. 5. Most bee species visited 1 or no fragrance orchids in a given habitat. Orchids tended to utilize common pollinators that seldom included more than 1 species, and they utilized stable or unstable, seasonal or aseasonal bees. However, the most stable and abundant bee, Eg. imperialis, rarely pollinated orchids; fewer than 10 of ca. 20000 bees carried pollinaria. 6. Orchids may interact primarily with discrete seasonal bee population peaks-probably the emerging adults. Although specialized orchid preferences are implicated for species that visit few or no local orchids but pollinate other species and carry pollinaria in other areas, euglossine bees do not need orchids to survive or reproduce.     

Solitary invasive orchid bee outperforms co-occurring native bees to promote fruit set of an invasive Solanum

Our understanding of the effects of introduced invasive pollinators on plants has been exclusively drawn from studies on introduced social bees. One might expect, however, that the impacts of introduced solitary bees, with much lower population densities and fewer foragers, would be small. Yet little is known about the potential effects of naturalized solitary bees on the environment. We took advantage of the recent naturalization of an orchid bee, Euglossa viridissima, in southern Florida to study the effects of this solitary bee on reproduction of Solanum torvum, an invasive shrub. Flowers of S. torvum require specialized buzz pollination. Through timed floral visitor watches and two pollination treatments (control and pollen supplementation) at three forest edge and three open area sites, we found that the fruit set of S. torvum was pollen limited at the open sites where the native bees dominate, but was not pollen limited at the forest sites where the invasive orchid bees dominate. The orchid bee’s pollination efficiency was nearly double that of the native halictid bees, and was also slightly higher than that of the native carpenter bee. Experiments using small and large mesh cages (to deny or allow E. viridissima access, respectively) at one forest site indicated that when the orchid bee was excluded, the flowers set one-quarter as many fruit as when the bee was allowed access. The orchid bee was the most important pollinator of the weed at the forest sites, which could pose additional challenges to the management of this weed in the fragmented, endangered tropical hardwood forests in the region. This specialized invasive mutualism may promote populations of both the orchid bee and this noxious weed. Invasive solitary bees, particularly species that are specialized pollinators, appear to have more importance than has previously been recognized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Macroevolution of perfume signalling in orchid bees

Theory predicts that both stabilising selection and diversifying selection jointly contribute to the evolution of sexual signalling traits by (1) maintaining the integrity of communication signals within species and (2) promoting the diversification of traits among lineages. However, for many important signalling traits, little is known about whether these dynamics translate into predictable macroevolutionary signatures. Here, we test for macroevolutionary patterns consistent with sexual signalling theory in the perfume signals of neotropical orchid bees, a group well studied for their chemical sexual communication. Our results revealed both high species‐specificity and elevated rates of evolution in perfume signals compared to nonsignalling traits. Perfume complexity was correlated with the number of congeners in a species’ range, suggesting that perfume evolution may be tied to the remarkably high number of orchid bee species coexisting together in some neotropical communities. Finally, sister‐pair comparisons were consistent with both rapid divergence at speciation and character displacement upon secondary contact. Together, our results provide new insight into the macroevolution of sexual signalling in insects.     

Are body size and volatile blends honest signals in orchid bees?

Secondary sexual traits may convey reliable information about males’ ability to resist pathogens and that females may prefer those traits because their genes for resistance would be passed on to their offspring. In many insect species, large males have high mating success and can canalize more resources to the immune function than smaller males. In other species, males use pheromones to identify and attract conspecific mates, and thus, they might function as an honest indicator of a male's condition. The males of orchid bees do not produce pheromones. They collect and store flower volatiles, which are mixed with the volatile blends from other sources, like fungi, sap and resins. These blends are displayed as perfumes during the courtship. In this study, we explored the relationship between inter‐individual variation in body size and blend composition with the males’ phenoloxidase (PO) content in Euglossa imperialis. PO content is a common measure of insect immune response because melanine, its derived molecule, encapsulates parasites and pathogens. Body size and blend composition were related to bees’ phenolic PO content. The inter‐individual variation in body size and tibial contents could indicate differences among males in their skills to gain access to some compounds. The females may evaluate their potential mates through these compounds because some of them are reliable indicators of the males’ capacity to resist infections and parasites.     

Evolution and diversity of floral scent chemistry in the euglossine bee-pollinated orchid genus Gongora

•Background and Aims Animal-pollinated angiosperms have evolved a variety of signalling mechanisms to attract pollinators. Floral scent is a key component of pollinator attraction, and its chemistry modulates both pollinator behaviour and the formation of plant–pollinator networks. The neotropical orchid genus Gongora exhibits specialized pollinator associations with male orchid bees (Euglossini). Male bees visit orchid flowers to collect volatile chemical compounds that they store in hind-leg pouches to use subsequently during courtship display. Hence, Gongora floral scent compounds simultaneously serve as signalling molecules and pollinator rewards. Furthermore, because floral scent acts as the predominant reproductive isolating barrier among lineages, it has been hypothesized that chemical traits are highly species specific. A comparative analysis of intra- and inter-specific variation of floral scent chemistry was conducted to investigate the evolutionary patterns across the genus.•Methods Gas chromatography–mass spectrometry (GC-MS) was used to analyse the floral scent of 78 individuals belonging to 28 different species of Gongora from two of the three major lineages sampled across the neotropical region. Multidimensional scaling and indicator value analyses were implemented to investigate the patterns of chemical diversity within and among taxonomic groups at various geographic scales. Additionally, pollinator observations were conducted on a sympatric community of Gongora orchids exhibiting distinct floral scent phenotypes.•Key Results A total of 83 floral volatiles, mainly terpenes and aromatic compounds, were detected. Many of the identified compounds are common across diverse angiosperm families (e.g. cineole, eugenol, β-ocimene, β-pinene and terpinen-4-ol), while others are relatively rare outside euglossine bee-pollinated orchid lineages. Additionally, 29 volatiles were identified that are known to attract and elicit collection behaviour in male bees. Floral scent traits were less variable within species than between species, and the analysis revealed exceptional levels of cryptic diversity. Gongora species were divided into 15 fragrance groups based on shared compounds. Fragrance groups indicate that floral scent variation is not predicted by taxonomic rank or biogeographic region.•Conclusions Gongora orchids emit a diverse array of scent molecules that are largely species specific, and closely related taxa exhibit qualitatively and quantitatively divergent chemical profiles. It is shown that within a community, Gongora scent chemotypes are correlated with near non-overlapping bee pollinator assemblies. The results lend support to the hypothesis that floral scent traits regulate the architecture of bee pollinator associations. Thus, Gongora provides unique opportunities to examine the interplay between floral traits and pollinator specialization in plant–pollinator mutualisms.     

Deriving ecological relationships from geographical correlations between host and parasitic species: an example with orchid bees

Aim  To investigate biogeographical patterns of cleptoparasitic Exaerete bee species and their orchid bee hosts.
Location  Neotropical region, from Central America to southern Brazil.
Methods  Correlations between relative frequencies of cleptoparasitic Exaerete species and their host Eulaema species were employed to investigate the geographical association between such species pairs.
Results  Our data support the current proposition that the Eulaema meriana / Eulaema flavescens complex is the main host for Exaerete frontalis . Contrary to current belief, however, Eulaema nigrita apparently is not the only and, in some regions, not the most important host for Exaerete smaragdina .
Main conclusions  Current knowledge on cleptoparasite host associations among orchid bees is based on fortuitous observations, and in some instances generalizations from such observations are not corroborated by the frequencies and distributions of the bees involved. Our data suggest that cleptoparasitic pressure, rather than other features of the forest environment, may be responsible for the low abundance of E. nigrita in the Amazonian forests.     

The interaction between pollinator size and the bristle staminode of Penstemon digitalis (Scrophulariaceae)

Penstemon digitalis, a prairie species whose flowers possess a large bristle staminode, is visited by eight bee species of varying size. Two sets of field experiments involving staminode removal were performed to test pollination efficiency in relation to bee size. Our data indicate that bristle staminode presence and function are influenced by size-dependent selection on bee body size and associated pollen transporting attributes. The first experiment compared staminode presence and removal in open-pollinated flowers. Staminode removal significantly reduced total pollen deposition but had no effect on total pollen removal. The second experiment utilized single bee visits to assess the interaction between pollinator size and staminode presence on the rate of pollen deposition and removal. This experiment indicated that staminode removal resulted in fewer pollen grains deposited on stigmas and less pollen removed from anthers for both large and small bees (the contrary was true for medium bees). Although the number of pollen grains deposited was greatly reduced for large bees, staminode removal reduced deposition efficiency most notably for small bees by 41.3%, reducing female reproductive success. Staminode removal increased pollen removal efficiency most notably for medium bees by 22.7%, reducing male reproductive success. Mechanisms of staminode function are discussed.     

A humped latitudinal phylogenetic diversity pattern of orchid bees (Hymenoptera: Apidae: Euglossini) in western Amazonia: assessing the influence of climate and geologic history

Amazonian rainforests are among the most species‐rich terrestrial habitats on Earth. The aim of this study was to analyze phylogenetic diversity (PD) patterns of orchid bee assemblages along a latitudinal gradient of 15° from northern Peru to central Bolivia and to relate them to climatic factors and geological history. We expanded an existing phylogeny of orchid bees and analyzed the PD of 15 orchid bee assemblages along a latitudinal gradient using mean pair‐wise phylogenetic distance. The resulting pattern was correlated to climatic factors and elevation. We found a hump‐shaped pattern of PD that peaked in central Peru and decreased towards the equatorial and especially towards the southern, subtropical sites. The decrease in PD towards higher latitudes is a common pattern found in many taxa, which in our case correlates with increasing climatic seasonality. However, the decrease towards the equatorial sites is unusual and may be related to a particular historic event: the northern sites with low PD are situated in the area of the former Lake Pebas, which covered western Amazonia until 3 mya. After the lake disappeared orchid bees mainly belonging to two distantly related species groups apparently colonized the region and diversified, which led to the comparatively low observed PD. In contrast, in central Peru, no in situ radiations were detected, hence the assemblages were composed of species from diverse phylogenetic lineages. Additionally, we identified multiple phylogenetically independent radiations of regionally restricted Euglossa species along the latitudinal gradient that, according to a published, dated phylogeny, diversified roughly 3–1 mya. The hump‐shaped latitudinal pattern of PD of the orchid bees of western Amazonia thus appears to have resulted from a preponderance of early divergent lineages in central Peru and of young radiations from distantly related clades colonizing higher latitudes, possibly triggered by historic climate fluctuations and major geological events.     

Scaling of nectar foraging in orchid bees

Morphology influences the rate at which foraging bees visit nectar flowers, the quantity of nectar they must consume to fuel their activities, and, consequently, the profitability of flower species. Because feeding time is a major determinant of visitation rate, I used a biomechanical model to examine how energy intake rate (E) varies with sucrose concentration, body mass (M), and proboscis length in orchid bees (Apidae: Euglossini). Under geometric scaling, the optimal sugar concentration (Smax) should be largely independent of body size, and E proportional to M1.0. In a comparative study of 30 orchid bee species ranging from 50 to 800 mg, Smax fell between 35% and 40% w/w, but E proportional to M0.54, significantly less than model predictions. Proboscis length and radius scale geometrically with body mass, but proboscis length exhibits substantial size-independent variation, particularly in small bees. One cost of a long proboscis is a reduction in both E and Smax in accordance with the scaling model. This finding highlights a difference between the lapping mechanism used by bumblebees and the suction mechanism used by orchid bees. A field study confirms that orchid bees harvest nectars with between 34% and 42% sucrose, independent of body size.     

Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries. Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related, perennial orchid species that differ in (a) floral traits important for pollination, including flowering phenology, floral display, and spur length, and (b) dominant pollinators. If plant–pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species. We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers, and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the expected direction, with selection for earlier flowering in two populations of the early‐flowering G. conopsea s.s. and for later flowering in one population of the late‐flowering G. densiflora. No divergent selection on floral morphology was detected, and there was no significant stabilizing selection on any trait in the two species. The results suggest ongoing adaptive differentiation of flowering phenology, strengthening this premating reproductive barrier between the two species. Synthesis: This study is among the first to test whether divergent selection on floral traits contribute to the maintenance of species differences between closely related plants. Phenological isolation confers a substantial potential for reproductive isolation, and divergent selection on flowering time can thus greatly influence reproductive isolation and adaptive differentiation.     

Effects of fragments and landscape characteristics on the orchid bee richness (Apidae: Euglossini) in an urban matrix,southwestern Amazonia

Orchid bees are important pollinators in tropical forests. Although studies have already detected effects of habitat loss and forest fragmentation on bee assemblages, little is known about orchid bees in urban forest fragments. The aim of this study was to analyse the influence of forest fragments (size and edge index) and landscape features (forest cover area and built-up area around the forest fragments, connectivity and spatial distance from the urban center) on the abundance, richness and composition of orchid bees. Male bees were attracted by odoriferous baits and collected in ten forest fragments of different sizes. In total, we collected 3166 male bees belonging to 4 genera and 38 species. The increase of the built-up area and the reduction of the forest cover area around the forest fragments decreased the abundance and richness of bees. We recorded a smaller number of bees in areas closer to the urban center. We failed to find a significant relationship between abundance/richness of bees and forest fragment area, edge index, and connectivity. We observed that the faunistic dissimilarity was positively correlated with the geographic distance between forest fragments. The forest fragments that presented greater landscape dissimilarity also presented greater faunistic dissimilarity. Our results suggest that the matrix structure around the forest fragments is an important factor that influences the Euglossini bee assemblages inside these forest fragments. Based on our results, we believed that the conservation of fragments with a larger forest cover area and smaller built-up areas around them contribute to the maintenance of the diversity of orchid bees and their pollination services.     

DOES SELECTION ON FLORAL ODOR PROMOTE DIFFERENTIATION AMONG POPULATIONS AND SPECIES OF THE SEXUALLY DECEPTIVE ORCHID GENUS OPHRYS?

Sexually deceptive orchids from the genus Ophrys attract their pollinators primarily through the chemical mimicry of female hymenopteran sex pheromones, thereby deceiving males into attempted matings with the orchid labellum. Floral odor traits are crucial for the reproductive success of these pollinator-limited orchids, as well as for maintaining reproductive isolation through the attraction of specific pollinators. We tested for the signature of pollinator-mediated selection on floral odor by comparing intra and interspecific differentiation in odor compounds with that found at microsatellite markers among natural populations. Three regions from southern Italy were sampled. We found strong floral odor differentiation among allopatric populations within species, among allopatric species and among sympatric species. Population differences in odor were also reflected in significant variation in the attractivity of floral extracts to the pollinator, Colletes cunicularius. Odor compounds that are electrophysiologically active in C. cunicularius males, especially alkenes, were more strongly differentiated among conspecific populations than nonactive compounds in the floral odor. In marked contrast to these odor patterns, there was limited population or species level differentiation in microsatellites (FST range 0.005 to 0.127, mean FST 0.075). We propose that the strong odor differentiation and lack of genetic differentiation among sympatric taxa indicates selection imposed by the distinct odor preferences of different pollinating species. Within species, low FST values are suggestive of large effective population sizes and indicate that divergent selection rather than genetic drift accounts for the strong population differentiation in odor. The higher differentiation in active versus non-active odor compounds suggests that divergent selection among orchid populations may be driven by local pollinator preferences for those particular compounds critical for pollinator attraction.     

Reproductive biology of Cyrtopodium punctatum in situ: implications for conservation of an endangered Florida orchid

Cyrtopodium punctatum (Linnaeus) Lindley is an endangered epiphytic orchid restricted in the USA to southern Florida. This species has been extensively collected from the wild since the early 1900s, and today only a few plants remain in protected areas. As part of a conservation plan, a reproductive biology study was conducted to better understand the ecology of this species in Florida. Cyrtopodium punctatum relies on a deceit pollination system using aromatic compounds to attract pollinators. Nine aromatic compounds were identified as components of the fragrance of C. punctatum inflorescences, including two compounds that are known to be Euglossine bee attractants. However, this group of bees is not native to Florida. Of the four bee species observed to visit C. punctatum flowers in the present study, carpenter bees ( Xylocopa spp.) are likely to be the main pollinators. Pollination experiments demonstrated that C. puntatum is self-compatible, but requires a pollinator and thus does not exhibit spontaneous autogamy. In addition, the rates of fruit set were significantly higher for flowers that were outcrossed (xenogamy) than for those that were self-crossed. Thus, the species has evolved a degree of incompatibility. Examples of natural pollination and fruit set were observed during the present study (2007–2008), but the rates of reproduction were modest as a consequence of the low plant numbers and possible changes in insect densities as a result of anthropogenic influences.     

Colour evolution within orchids depends on whether the pollinator is a bee or a fly

• Orchids are a classic angiosperm model for understanding biotic pollination. We studied orchid species within two species‐rich herbaceous communities that are known to have either hymenopteran or dipteran insects as the dominant pollinators, in order to understand how flower colour relates to pollinator visual systems.
• We analysed features of the floral reflectance spectra that are significant to pollinator visual systems and used models of dipteran and hymenopteran colour vision to characterise the chromatic signals used by fly‐pollinated and bee‐pollinated orchid species.
• In contrast to bee‐pollinated flowers, fly‐pollinated flowers had distinctive points of rapid reflectance change at long wavelengths and a complete absence of such spectral features at short wavelengths. Fly‐pollinated flowers also had significantly more restricted loci than bee‐pollinated flowers in colour space models of fly and bee vision alike.
• Globally, bee‐pollinated flowers are known to have distinctive, consistent colour signals. Our findings of different signals for fly pollination is consistent with pollinator‐mediated selection on orchid species that results from the distinctive features of fly visual systems.

     

Species specificity and intraspecific variation in the chemical profiles of Heliconius butterflies across a large geographic range

In many animals, mate choice is important for the maintenance of reproductive isolation between species. Traits important for mate choice and behavioral isolation are predicted to be under strong stabilizing selection within species; however, such traits can also exhibit variation at the population level driven by neutral and adaptive evolutionary processes. Here, we describe patterns of divergence among androconial and genital chemical profiles at inter‐ and intraspecific levels in mimetic Heliconius butterflies. Most variation in chemical bouquets was found between species, but there were also quantitative differences at the population level. We found a strong correlation between interspecific chemical and genetic divergence, but this correlation varied in intraspecific comparisons. We identified “indicator” compounds characteristic of particular species that included compounds already known to elicit a behavioral response, suggesting an approach for identification of candidate compounds for future behavioral studies in novel systems. Overall, the strong signal of species identity suggests a role for these compounds in species recognition, but with additional potentially neutral variation at the population level.