Do floral odor profiles geographically vary with the degree of specificity for pollinators? Investigation in two sapromyophilous Arum species (Araceae)

We compared floral odour profiles among populations of two Arum species which show different degrees of specificity for their fly pollinators. Insects were collected from inflorescences in four populations of Arum italicum and two populations of Arum maculatum. In six Arum populations, we compared inflorescences odour profiles collected by Solid Phase Micro Extraction (SPME) and analysed by gas chromatography. We confirmed that from a pollination point of view, A. italicum is an opportunist species, as it is mainly pollinated by insects of the families Psychodidae, Chironomidae and Sciaridae, whereas A. maculatum is a specialist species, as it is 90% pollinated by Psychodidae. In all populations, Arum italicum was less attractive to pollinators than Arum maculatum. Floral odour profiles of A. italicum were not geographically structured among populations, suggesting a high gene flow or adaptation to a fluctuant guild of pollinators. On the contrary, odour profiles of A. maculatum varied between the two populations studied suggesting a lower gene flow or adaptation to different local pollinator preferences     

Stomata in some species of genus <Emphasis Type="Italic">Arum</Emphasis> from the Eastern Slavonia and Baranya region

Types and the number of stomata in the following Arum species: Arum italicum Mill., Arum maculatum var. maculatum L. and Arum maculatum var. immaculatum L., Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. in three different locations in Zablaće, Normanci and Bilje were investigated. The most prevalent stomata type at both upper and lower epidermis for each Arum species was mostly stomata type paracytic, followed by hexacytic, tetracytic or brachyparacytic as far as locality is concerned. Helicocytic type was more prevalent in Arum alpinum var. pannonicum Terpo. and Arum alpinum var. intermedium Schur. at Bilje. Other stomata types were very rare. A striking regularity in the occurrence of stomata types was not found within a single species. Some stomata types, however, were found either at adaxial or abaxial epidermis or were not present at all. The number of stomata per square mm varied from 25 to 651. A statistically significant difference in the number of stomata per square mm at upper and lower epidermis among Arum species was determined in locations Zablaće and Normanci, whereas no statistically significant differences were found in location Bilje.     

Reproduction of the West Mediterranean endemic Arum pictum (Araceae) on Corsica

Pollination in the genus Arum appears to be in general a complex deceptive pollination process. The genus Arum is composed of 28 species, all belonging to the subgenus Arum, except A. pictum, the only species of the subgenus Gymnomesium, which is basal and sister to all other Arum species. The aim of this paper is to document the pollination ecology of the insular Arum pictum, the only Arum species to flower in autumn, on the island of Corsica (France). The anthesis cycle of A. pictum occurs during the day, attracting sphaerocerid flies and staphylinid beetles early in the morning and late in the afternoon of the first day. The pollen is released from the anthers early in the morning of the second day before the departure of the insects. Its thermogenic cycle matches the anthesis cycle with an original and unique, bimodal temperature pattern of the appendix (morning and afternoon), contrary to the unimodal pattern found in all other studied Arum species. Data from reproductive success and seed sets suggest that sexual reproduction in this species is limited by pollen (e.g. attracting lured insects) rather than by resources. The biology of this Western Mediterranean species appears to be different from other Western European Arum and close to some Eastern Mediterranean species. Further studies are needed to establish whether Arum pictum represents some ancestral stage or whether its peculiar biological traits are adapted to its insular distribution.     

Antennal sensilla in five Psychodini moth flies (Diptera: Psychodidae: Psychodinae) pollinators of Arum spp. (Araceae)

The pollination of the genus Arum (Araceae) is mainly achieved by deception, the floral odour mimicking the pollinator ovipositing site. In order to discover the sensory organs involved in this attraction, we have studied the antennae of five species of psychodine moth-flies (former Psychoda sensu lato=Psychodini), pollinators of Arum spp. The antennae of the five Psychodini reveal seven types of sensilla: multiporous tribranched sensilla basiconica (sensilla ascoidea), multiporous sensilla basiconica, multiporous sensilla coeloconica, multiporous sensilla auricillica, uniporous sensilla basiconica, aporous sensilla chaetica, aporous Böhm’s sensilla. Each species possesses three, five or six of these sensillum types. All the multiporous sensilla are probably olfactory receptors while the uniporous sensilla basiconica must possess a contact chemoreceptive function. The multiporous tribranched sensilla basiconica (s. ascoidea), present in all the species, are the best candidates for the reception of the odours given off by the ovipositing sites and the inflorescences of Arum. The multiporous sensilla basiconica and the multiporous sensilla coeloconica may be involved respectively as CO₂ receptors or thermoreceptors. Psychoda phalaenoides, which is the main pollinator of A. maculatum, is the species which possesses the largest number of antennal sensilla. The sexual dimorphism, studied only in Psycha grisescens, as concerns the number of sensilla and the absence of a sensillum type which differ according to sex, is difficult to interpret.     

Reliability of odour–genes covariance despite diet changes: a test in mound‐building mice

Odour–genes covariance (i.e. that individual odour similarities covary with genetic similarity) allows animals to assess their genetic relatedness to one another by comparing the similarities between their individual odours. The reliability of the odour–gene covariance phenomenon under environmental changes has not yet been explored. Using an innovative habituation‐generalization procedure, we tested whether the modifications of body odours induced by a change in diet interfered with the perceived similarity of individual odours of more genetically similar individuals. The effects of a change in diet, by adding aniseed flavour to the drinking water, was assessed in male mound‐building mice, a species in which odour–genes covariance has already been demonstrated. The results obtained indicate that mice perceive more similarities in the odours of sibling males than in those of nonrelatives, despite the effects of diet on body odours. We suggest that the reliability of signals arising from odour–genes covariance is likely a result of the independence between traits indicating genetic relatedness and indicator signals of biological states (e.g. diet) in body odours. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 682–688.     

The design of trapping devices in pollination traps of the genus Arum (Araceae) is related to insect type

Pollinators have long been known to select for floral traits, but the nature of this relationship has been little investigated in trap pollination systems. We investigated the trapping devices of 15 Arum spp. and compared them with the types of insects trapped. Most species shared a similar general design of trap chamber walls covered in downward‐pointing papillate cells, lacunose cells in the chamber wall and elongated sterile flowers partially blocking the exit of the trap. However, there was significant variation in all these morphological features between species. Furthermore, these differences related to the type of pollinator trapped. Most strikingly, species pollinated by midges had a slippery epidermal surface consisting of smaller papillae than in species pollinated by other insects. Midge‐pollinated species also had more elongated sterile flowers and tended to have a larger lacunose area. We conclude that pollination traps evolve in response to the type of insect trapped and that changes to the slippery surfaces of the chamber wall are an important and previously little recognized variable in the design of pollination traps. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 385–397.     

New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history

The heat‐ and odour‐producing genus Arum (Araceae) has interested scientists for centuries. This long‐term interest has allowed a deep knowledge of some complex processes, such as the physiology and dynamics of its characteristic lure‐and‐trap pollination system, to be built up. However, mainly because of its large distributional range and high degree of morphological variation, species' limits and relationships are still under discussion. Today, the genus comprises 28 species subdivided into two subgenera, two sections and six subsections. In this study, the phylogeny of the genus is inferred on the basis of four plastid regions, and the evolution of several morphological characters is investigated. Our phylogenetic hypothesis is not in agreement with the current infrageneric classification of the genus and challenges the monophyly of several species. This demonstrates the need for a new infrageneric classification based on characters reflecting the evolution of this enigmatic genus. To investigate the biogeography of Arum deeply, further spatiotemporal analyses were performed, addressing the importance of the Mediterranean basin in the diversification of Arum. Our results suggest that its centre of origin was the European–Aegean region, and that major diversification happened during the last 10 Myr. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 14–32.     

A new species of Psidium L. (Myrtaceae) from southern Brazil

Rewardless kettle‐trap flowers Cypripedium yunnanense and C. flavum were watched for pollinators during 73 and 101 man‐hours, respectively, in north‐west Yunnan at 3490–3590 m a.s.l. They differ from typical Cypripedium, such as C. calceolus, in having a broad infolded flap of the lip extending all around the entrance of the pouch (instead of being restricted to the vicinity of the staminode) and in the flap not being slippery. Cypripedium yunnanense is pollinated by Lasioglossum zonulum euronotum (Halictidae), and C. flavum is pollinated by Andrena orchidea and Andrena sp. (Andrenidae, two of nine new hymenopterans discovered at the sites). The bees do not inadvertently fall into the trap by slipping (as often occurs in other slipper orchids), but enter it by crawling down the flap in full control of their movements. No natural hybrids between the two orchids are known, although they occur in close‐by or mixed stands, are co‐flowering and size compatible with regard to their pollinators, and produce fruits following manual cross‐pollination in situ. Analyses of the (non‐Cypripedium) pollen carried indicate that: (1) the two andrenids are probably oligolectic; (2) the andrenids and the halictid do not share the same flower species; and (3) the halictid is polylectic. Points (1) and (2) are probable reasons for the lack of hybrids. The other reason why Lasioglossum zonulum euronotum does not visit C. flavum despite being polylectic may be the flower's odour; cases of discrimination of closely related flowers by polylectic Lasioglossum have been reported elsewhere. Blow flies Calliphora vomitoria and Calliphora pattoni (Calliphoridae) also enter the orchids, some smearing themselves with pollen, yet they are not pollinators. They are too large to leave by the exit and die imprisoned. However, they may be accidental pollinators of the rather larger C. tibeticum present at the sites. Such accidental pollinators probably play an important role in the evolution of new pollination syndromes. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156 , 51–64.     

Responses of Snow Voles, Chionomys nivalis, Towards Conspecific Cues Reflect Social Organization during Overwintering Periods

Among microtine rodents, reaction to chemical cues from conspecifics is assumed to reflect social and spatial relationships. Generally, strong attraction of particular odours correlates with non‐aggressive behaviour and high spatial tolerance towards odour donors, whereas weak attraction correlates with greater levels of aggression and spatial segregation. In the present study, we examined whether winter odour preferences of the snow vole Chionomys nivalis, a rock‐dwelling microtine principally found at high‐mountainous regions, differ from that of other vole species, owing to their different social organization during overwintering periods. The social structure of C. nivalis over the winter period is relatively unusual among vole species in that they become nomadic and solitary. In odour choice trials under laboratory conditions, we found that both males and females avoided zones with conspecific odours of both sexes in comparison with unscented control zones or own odours. These results are consistent with the elevated levels of intraspecific aggression and spatial isolation of C. nivalis during overwintering periods. Furthermore, scent‐elicited self‐grooming increased when their own odour was offered against conspecific cues. This, in combination with an active avoidance of conspecific odours, might functionally contribute to minimize direct confrontations between solitary individuals, thereby reducing the risks of aggressive encounters during overwintering periods.     

Distribution,habitat disturbance and pollination of the endangered orchid Broughtonia cubensis (Epidendrae: Laeliinae)

The geographical distribution, population structure and pollination ecology are key aspects in the conservation and management of rare orchids. Here, we address these aspects and the main threats affecting the endangered Cuban orchid Broughtonia cubensis. This rewardless orchid is self‐compatible, but pollinator dependent. However, seed production can be negatively affected by insect‐mediated selfing. Three species of small bee (genera Ceratina and Lasioglossum) act as pollinators. As in the case of other nectarless orchids, we detected two species of plant producing large amounts of nectar in the area, the floral morphology of which closely resembles that of B. cubensis. The simultaneous flowering of these species could positively affect the reproductive success of B. cubensis. Nonetheless, the fitness of this orchid in natural conditions is low, possibly related to strong pollen limitation. To the problems arising from reduced fitness is added the fact that its historical distribution range has been greatly reduced in recent years. Throughout this study, we have detected dramatic reductions in the population sizes, in some cases as a result of human plundering, but also as a consequence of hurricanes. Based on the results of this study, we propose some guidelines to manage and conserve this orchid. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 345–357.     

Individual Odour Similarity and Discrimination in the Coruro (Spalacopus cyanus, Octodontidae)

Previous studies of subterranean, fossorial and above‐ground rodents have demonstrated that similarities in individual odours covary with genetic similarities thus supporting the theory of odour‐genes covariance (i.e. the closer the individuals are genetically, the greater the similarities between their odours). We used the habituation‐generalization paradigm, where the subject is exposed to the same odour stimulus in three consecutive habituation trials followed by two test trials in which the odour from two other individuals are presented successively. Using this test design, we showed that the socially living coruros (Spalacopus cyanus) discriminate individuals on the basis of their ano‐genital odours and also respond to odour similarities between individuals. Male and female subjects treated ano‐genital odours of two same‐sex family members and also the odours of two sibling strangers as different to each other. At the same time, they treated the odours of siblings as similar compared with the odour of an unrelated individual. No gender differences were detected. Our results contrast with those from other rodent species that did not spontaneously discriminate between individual odours of siblings from a different family than their own. The polygyneous lifestyle may provide the selective background for that difference. Additional research will be necessary to explore this hypothesis and to rule out differences due to dietary preferences and due to the type of paradigm chosen for the tests.     

Pollination by deceit and floral mimesis inThelymitra antennifera (Orchidaceae)

Thelymitra antennifera is found to be a general mimic of the search image of other co-blooming yellow or cream-flowered species including members of the following genera:Hibbertia, Goodenia and, to a lesser extent,Helichrysum, Pimelia andStackhousia. The most common pollinators areLasioglossum (Chilalictus) spec. and the introducedSyrphus damaster (hover fly) as well as a solitary wasp,Eurys spec.—T. antennifera acts by deceit, involving opticals (yellow colour and pollen imitation) as well as olfactory (sweet odour) signals. Capsule production is low and density dependent, a situation demonstrated in other species which have evolved pollination syndromes involving deceitful mimicry.     

Fly pollination in Cypripedium: a case study of sympatric C. sichuanense and C. micranthum

Most Cypripedium spp. are known to be pollinated by bees. However, myiophilous traits are found in some species, especially in sections Trigonopedia and Sinopedilum. Here we chose C. micranthum and C. sichuanense, two sympatric species endemic to Sichuan, China, to test whether these orchids are fly pollinated. Artificial pollination showed that both flowers are self‐compatible but need pollen vectors for successful reproduction. Field observation showed that C. micranthum was pollinated by fruit flies and C. sichuanense by dung flies, both novel pollinators of Cypripedium orchids. These sympatric Cypripedium spp. are also cross‐compatible, but hybrids were not found in nature. The pollination syndromes of C. sichuanense and C. micranthum fit into the complex sapromyiophily pattern. It appears that pollinator specificity is responsible for their reproductive isolation. The discovery of fly pollination in C. sichuanense and C. micranthum, which belong to the related sections Trigonopedia and Sinopedilum, suggests a shift from bee to fly pollination in the genus Cypripedium. Unlike most Cypripedium spp., the anthers of C. micranthum release discrete pollinia with narrow stalks instead of the usual amorphous pollen smears. This ‘proto‐pollinarium’ is described, probably for the first time. These pollinia are most likely an adaptation for pollination by microdiptera, so the fly can carry the contents of both chambers in the same anther. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 50–58.     

A sheep in wolf's clothing: do carrion and dung odours of flowers not only attract pollinators but also deter herbivores?

Carrion and dung odours of various flowers have traditionally been considered an adaptation for attracting the flies and beetles that pollinate them. While we accept the role of such odours in pollinator attraction, we propose that they may also have another, overlooked, anti‐herbivore defensive function. We suggest that such odours may deter mammalian herbivores, especially during the critical period of flowering. Carrion odour is a good predictor for two potential dangers to mammalian herbivores: (1) pathogenic microbes, (2) proximity of carnivores. Similarly, dung odour predicts faeces‐contaminated habitats that present high risks of parasitism. These are two new types of repulsive olfactory aposematic mimicry by plants: (1) olfactory feigning of carcass (thanatosis), a well‐known behavioural defensive strategy in animals, (2) olfactory mimicry of faeces, which also has a defensive visual parallel in animals.     

Speciation in sexually deceptive orchids: pollinator-driven selection maintains discrete odour phenotypes in hybridizing species

Ophrys orchids mimic the female sex pheromones of their pollinator species to attract males for pollination. Reproductive isolation in Ophrys is based on the selective attraction of only a single pollinator species. A change of floral odour can result in the attraction of a new pollinator species that acts as an isolation barrier towards other sympatrically occurring Ophrys species. Ophrys lupercalis, Ophrys bilunulata, and Ophrys fabrella grow sympatrically and bloom consecutively on Majorca and are pollinated by three species of Andrena. We investigated variation of phenotypic and genotypic flower traits, aiming to study the role of the floral odour for reproductive isolation and speciation. Using chemical and electrophysiology (gas chromatography coupled with an electroantennographic detector) methods, we show that the three Ophrys species use the same odour compounds for pollinator attraction, but in different proportions. A comparison of the floral odour bouquets in a multivariate analysis revealed a clear grouping of plants from the same species, although with an overlap between species. A comparison of the same plants using molecular markers gave a contrasting result. Although O. lupercalis and O. fabrella were genetically well separated, plants of O. bilunulata did not form a distinct group but were similar to either O. lupercalis or O. fabrella. Our data indicate gene flow and hybridization to occur between O. bilunulata and O. lupercalis as well as between O. bilunulata and O. fabrella. All plants of O. bilunulata, despite having different genotypes, showed a very similar floral odour. This reflects a strong selective pressure by the pollinating males. The overlap of genotypes of O. bilunulata and O. fabrella supports our hypothesis that O. fabrella diverged from O. bilunulata by scent variation and the attraction of a new pollinator species, Andrena fabrella. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 439–451.     

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

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

     

Odour–genes covariance within a natural population of subterranean Spalax galili blind mole rats

Previous research, using habituation techniques with multiple rodent species as subjects, has demonstrated (from kin to across species) the greater perceptual similarity in the qualities of individual odours of more closely genetically related individuals ('odour–genes covariance', abbreviated 'OGC'). This predictable relationship between individual genotypes and individual odours has only been assessed in rodents in which the genetic similarities and dissimilarities are known either through laboratory breeding or by selecting odour donors from different populations or species. To study OGC within a natural population of rodents, we genotyped Spalax galili blind mole rats using ten microsatellite markers, and conducted pairwise genetic distance comparisons to estimate genetic similarity and to calculate the relatedness coefficient between pairs of individuals. We then used habituation-generalization techniques to assess whether perceived odour similarities covaried with the genetic similarities we had identified. Indeed, animals treated odours from genetically closer donors as more similar than odours from less genetically similar donors. The results suggest an accurate and subtle ability to resolve genetically determined odour distinctions among familially unrelated animals within a population. Graded differential responses to conspecific odours based on degrees of similarity between other individuals' odours and one's own have been demonstrated in kinship and species discriminations and preferences. The evidence presented here provides the basis for hypothesizing a similar process that could promote optimal outbreeding and inclusive fitness within populations of conspecifics. The evolutionary implications of these findings are discussed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 483–490.     

Effect of belowground herbivory on parasitoid associative learning of plant odours

Root herbivores can influence both the performance and the behaviour of parasitoids of aboveground insect herbivores through changes in aboveground plant quality and in the composition of the plant's odour blend. Here we show that root herbivory by Delia radicum larvae did not influence the innate preferences for plant odours of the two closely related parasitoid species Cotesia glomerata and C. rubecula, but did affect their learned preferences, and did so in an opposite direction. While C. glomerata learned to prefer the odour of plants with intact roots, C. rubecula learned to prefer the odour of root‐infested plants. The learned preference of C. glomerata for the odour of plants with intact roots matches our previously published result of its better performance when developing in P. brassicae hosts feeding on this plant type. In contrast, the relatively stronger learned preference of C. rubecula for the odour of root‐infested plants cannot be merely explained by its performance, as the results of our present study indicate that D. radicum root herbivory did not influence the performance of C. rubecula nor of its host P. rapae. Our results stress the importance of assessing the influence of root herbivores on both innate and learned responses of parasitoids to plant odours.     

The smell of danger: chemical recognition of fish predators by the invasive crayfish Procambarus clarkii

1. Since avoiding predation can compromise animal fitness, prey are expected to respond to different predator species with an intensity appropriate to the level of risk. In fresh waters, the threat of predation is typically assessed by chemical cues, in particular by odours released by either injured/disturbed conspecifics (conspecific alarm odour) or predators (predator odours). Here, we used the most widely distributed crayfish in the world, the invasive North American Procambarus clarkii, to investigate the relative effectiveness of odours emitted by fish predators compared with conspecific alarm odour. We also tested whether P. clarkii is able to discriminate between fish predators of which it has ‘experience’ (either recent, via introduction to the same water body, or old, by sharing a native range), as well as between fish predators that pose low or high risk. 2. The study was carried out on introduced populations of P. clarkii from two sites, characterised by different fish assemblages: the Malewa River (a tributary of Lake Naivasha, Kenya) and Lake Trasimeno (Italy). Laboratory experiments consisted of three sequential phases (‘water’, ‘food’ and ‘smell’ phases) and five treatments. Treatments differed in the odour presented during the smell phase, i.e. no odour (plain water) and odours from either injured conspecifics or three fish species per site. Crayfish from the Malewa River population were confronted with the odours of largemouth bass (Micropterus salmoides), common carp (Cyprinus carpio) and tilapia (Tilapia zillii) (all introduced to Lake Naivasha but absent from the Malewa River), and those from the Lake Trasimeno population with the odours of the introduced largemouth bass and carp and the native chub (Squalius cephalus). Largemouth bass is the only predator that imposes a high risk to crayfish, and it also shares its native range with P. clarkii. We analysed the time spent by crayfish feeding, in locomotion and in adopting a raised or lowered posture. A reduction in the time spent feeding and in locomotion, and an increase in the time spent in the lowered posture were considered to indicate alarm. 3. Crayfish from both populations responded with a more pronounced reduction in feeding to conspecific alarm odour rather than to predator odours. Crayfish from the Malewa River reacted with the same intensity to the odours of the three fish species tested, whereas, in Lake Trasimeno, the odour of largemouth bass was significantly more threatening than the odours of the other two species. 4. Procambarus clarkii seems to perceive a general fish odour that alerts it to possible predation risk without the need of either a direct recent experience or via sharing a common native range. However, where they coexist with fish, crayfish become able to distinguish among species, adapting the intensity of their response to the effective risk. Our results confirm the relatively high learning capacity of P. clarkii reported in previous studies and suggest the existence of mechanisms that make predator recognition particularly efficient in this extraordinarily successful invader.     

Could learning of pollen odours by honey bees (Apis mellifera) play a role in their foraging behaviour?

Abstract. The role of pollen odour cues in the foraging behaviour of honey bees (Apis mellifera L.) is poorly understood. Using classical conditioning of the proboscis extension response, in which bees learn to associate an odour with a sucrose reward, the present study tests whether odours of bee-collected pollen from the hive environment or odours of fresh pollen on the anthers of flowers could be used in pollen foraging. Honey bees efficiently learn odours from field-bean (Vicia faba) bee-collected pollen and oilseed-rape (Brassica napus) bee-collected pollen, hand-collected pollen, anthers and whole flowers, demonstrating that honey bees can learn pollen odours associatively in biologically realistic concentrations. Honey bees learn pollen odours of oilseed rape better than field bean and, although they generalize these two odours, they easily distinguish between them in discrimination tests, suggesting that pollen odours may be used in species recognition/discrimination. There is little evidence that honey bees can recognize whole flowers based on previous experience of bee-collected pollen odour. However, they generalize the odours of oilseed-rape anthers and whole flowers, suggesting that anther pollen in situ may play a more prominent role than bee-collected pollen in foraging behaviour.