Analyzing the Spatial Structure of Broughtonia cubensis (Orchidaceae) Populations in the Dry Forests of Guanahacabibes,Cuba

A current key issue in ecology is the role of spatial effects on population and community dynamics. In this paper, we test several hypotheses related to spatial structures and coexistence of epiphytic tropical orchid species with special emphasis on the endemic species Broughtonia cubensis. More specifically, we explored the spatial structure of orchid–host plant communities at three different levels of organization (occupied vs. nonoccupied host trees, trees with B. cubensis vs. other orchids, and reproductive vs. nonreproductive B. cubensis plants). We mapped all potential host trees and orchids at three 20 × 20 m plots and applied techniques of spatial point pattern analysis such as mark connection and mark correlation functions to evaluate departures from randomized communities. We found spatial aggregation of trees with epiphytic orchids and segregation between trees with and without epiphytic orchids, and that there was an intraspecific spatial aggregation of B. cubensis in relation to the other seven epiphytic orchid species. Furthermore, we found spatial aggregation of reproductive B. cubensis individuals and segregation between reproductive and nonreproductive individuals on their phorophytes. Thus, orchid–host plant communities show hierarchical spatial structuring with aggregation and segregation at different levels of organization. Our results point to an enhancement of local species in the coexistence of tropical epiphytic orchid communities, by reducing competition through niche differentiation.     

Spatio‐temporal Analysis of the Effects of Hurricane Ivan on Two Contrasting Epiphytic Orchid Species in Guanahacabibes,Cuba

Hurricanes represent the dominant type of disturbance in many tropical coastal forests. Here, we focus on mortality of epiphytic orchids caused by hurricane Ivan in the Guanahacabibes National Park (Cuba) and subsequent population recovery. We analyzed different aspects of hurricane damage on two contrasting epiphytic orchids, Broughtonia cubensis and Dendrophylax lindenii, as observed in three plots of coastal vegetation and in three plots of semi‐deciduous forest, respectively. First, we quantified the damage to host trees and orchids and explored if hurricane damage depended on height, size, or identity of the host tree. Second, we used mark connection and mark correlation functions to conduct a detailed analysis of small‐scale spatial patterns in hurricane damage for host trees and orchids. Finally, we analyzed the degree of recovery after Ivan during the 6 yr following the storm. Damage of B. cubensis host trees was independent of height and size, but Ivan severely affected larger and higher host trees of D. lindenii. Spatial analysis revealed non‐random structure in damage that differed between species. Broughtonia cubensis exhibited small‐scale spatial correlation in the proportion of damaged orchids, whereas D. lindenii did not. Dendrophylax lindenii showed ‘patchy’ damage patterns, correlated with height, but B. cubensis did not. The relative growth rate of B. cubensis for the 5–17 mo following Ivan was only moderately reduced and fully recovered in subsequent years, whereas that of D. lindenii was severely reduced the first year and did not fully recover thereafter. We hypothesize that differences in the host, vegetation type, and the traits of the two orchids contribute to the different responses to the hurricane.     

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.

     

Population history of the terrestrial orchid Cremastra appendiculata var. variabilis from Korea,inferred from levels and distribution of genetic diversity

Cremastra appendiculata var. variabilis is a self‐compatible, insect‐pollinated, terrestrial orchid that is a typical member of the warm‐temperate vegetation in the Korean Peninsula. Here we examine levels and partitioning of allozyme diversity (22 loci) in 12 populations of this orchid to gain insight into its genetic structure and post‐glacial colonization history in Korea. It harboured considerably higher levels of genetic variation within populations (%P = 48.1, A = 1.70 and H_e = 0.217) and lower degree of differentiation among populations (F_ST = 0.068) than those typical of allozyme‐based studies in other terrestrial orchid species. These patterns suggest that extant populations were derived from multiple source populations (i.e. from multiple glacial refugia), although further studies are needed to confirm this scenario. In addition to population history, traits such as high potential of seed dispersal, a mixed mating system and its occurrence in large and continuous populations would have contributed to the current levels and distribution of genetic diversity in Korean populations of C. appendiculata var. variabilis. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 721–732.     

Environmental determinants of genetic diversity in Caragana microphylla (Fabaceae) in northern China

Non‐rewarding orchids rely on various ruses to attract their pollinators. One of the most common is for them to resemble flowers sought by insects as food sources. This can range from generalized food deception to the mimicry of specific sympatric food plants. We investigated the basis of pollinator deception in the European food‐deceptive orchid Traunsteinera globosa, which has unusually compact flowerheads resembling those of sympatric rewarding species of Knautia and Scabiosa (Dipsacaceae), and Valeriana (Caprifoliaceae). Visual signals of T. globosa are similar in both fly and bee vision models to those of the sympatric food plants used in the choice experiments, but scent signals are divergent. Field experiments conducted in Austria and the Czech Republic showed that both naive and experienced (with respect to visitation of T. globosa) insect species approached the orchids at the same rate as food plants, but direct contact with orchid flowers was taxon specific. Flies were most easily duped into probing the orchid, and, in doing so, frequently received and deposited pollinaria, whereas most bees and butterflies avoided landing on orchid flowers. We conclude that T. globosa is a mimic of a guild of fly‐pollinated plants, but the ecological dependence of the orchid on its models remains to be fully tested. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 269–294.     

Orchid seed removal by ants in Neotropical ant‐gardens

• Most plants that inhabit ant‐gardens (AGs) are cultivated by the ants. Some orchids occur in AGs; however, it is not known whether their seeds are dispersed by AG ants because most orchid seeds are tiny and dispersed by wind.
• We performed in situ seed removal experiments, in which we simultaneously provided Azteca gnava ants with seeds of three AG orchid species and three other AG epiphyte species (Bromeliaceae, Cactaceae and Gesneriaceae), as well as the non‐AG orchid Catasetum integerrimum.
• The seeds most removed were those of the bromeliad Aechmea tillandsioides and the gesneriad Codonanthe uleana, while seeds of AG orchids Coryanthes picturata, Epidendrum flexuosum and Epidendrum pachyrachis were less removed. The non‐AG orchid was not removed. Removal values were positively correlated with the frequency of the AG epiphytes in the AGs, and seeds of AG orchids were larger than those of non‐AG orchids, which should favour myrmecochory.
• Our data show that Azt. gnava ants discriminate and preferentially remove seeds of the AG epiphytes. We report for the first time the removal of AG orchid seeds by AG ants in Neotropical AGs.

     

Opportunistic pollinator shifts among sexually deceptive orchids indicated by a phylogeny of pollinating and non-pollinating thynnine wasps (Tiphiidae)

The thynnine wasp genus Neozeleboria Rohwer is the main pollinating group of the sexually deceptive Australian orchid genus, Chiloglottis R.Br. In a highly specialized interaction, Chiloglottis species attract males from a single or very few Neozeleboria species through the chemical mimicry of the female wasp's sex pheromone. An earlier study examining the historical association among Chiloglottis and Neozeleboria using DNA sequence data found matching phylogenetic patterns suggestive of cospeciation between orchids and pollinators. However, patterns of constraint in Neozeleboria emergence phenology and sex pheromones suggested that the close association among orchid and wasp clades may be due to pollinator switching among closely related wasp taxa that have similar traits. In this study, we further examine the association by incorporating a morphological phylogenetic analysis of non‐pollinating as well as pollinating Neozeleboria. The morphological analysis is then compared with DNA sequence data from one nuclear and one mitochondrial gene for an increased sample of outgroup genera. The combined molecular data set finds a monophyletic Neozeleboria, although support for this was not strong in the individual data sets. A high congruence between molecular and morphological analyses was found among higher groupings of Neozeleboria. Neozeleboria species that pollinate Chiloglottis species are not found as a monophyletic group but, rather, are scattered throughout a phylogeny comprising pollinators and non‐pollinators. Under the cospeciation model, the presence of related Neozeleboria non‐pollinators carries the unlikely implication that the association between plant and pollinator has been repeatedly lost. Instead, we favour the alternative ‘preferential pollinator switching’ model that accounts for the specialization among orchid and wasp lineages in terms of similarities in traits among related Neozeleboria. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 381–395.     

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.     

Recurrent polymorphic mating type variation in Madagascan Bulbophyllum species (Orchidaceae) exemplifies a high incidence of auto‐pollination in tropical orchids

The transition from outcrossing to self‐fertilization is one of the most common evolutionary changes in angiosperms. The orchid family exemplifies this evolutionary trend but, because of a general lack of large‐scale surveys on auto‐pollination in orchid taxa, the incidence and modes of auto‐pollination among (sub)tropical orchids remain poorly known. In the present study, we assessed the frequency and mode of auto‐pollination within and among species of a largely monophyletic group of Madagascan Bulbophyllum. The capacity for autonomous fruit set was investigated by bagging experiments in the greenhouse and the field, complemented with detailed floral micromorphological studies of the gynostemium. Our survey comprises 393 accessions, representing at least 78 species, and thus approximately 37% of the species diversity of the genus in the Madagascan region. Our studies revealed that mating type is directly related to gynostemium structure, most often involving the presence or absence of a physical barrier termed ‘rostellum’. As a novel and unexpected finding, we identified eight species of a single lineage of Madagascan Bulbophyllum (termed ‘clade C’), in which auto‐pollinating morphs (selfers), either lacking a rostellum or (rarely) possessing a stigmatic rostellum, co‐exist with their pollinator‐dependent conspecifics (outcrossers). We hypothesize that auto‐pollination via rostellum abortion has a simple genetic basis, and probably evolved rapidly and recurrently by subtle changes in the timing of rostellum development (heterochrony). Thus, species of clade C may have an intrinsic genetic and developmental lability toward auto‐pollination, allowing rapid evolutionary response under environmental, perhaps human‐disturbed conditions favouring reproductive assurance. Overall, these findings should stimulate further research on the incidence, evolution, and maintenance of mating type variation in tropical orchids, as well as how they adapt(ed) to changing environmental conditions. © 2014 The Authors. Botanical Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 242–258.     

Pollen limitation and fruiting failure related to canopy closure in Calypso bulbosa (Orchidaceae), a northern food‐deceptive orchid with a single flower

Natural fruit set is constrained by pollen limitation and fruiting failure, and pollen limitation is expected to be especially severe in deceptive orchids. We performed hand cross‐pollinations in ten populations of a food‐deceptive orchid, Calypso bulbosa, under sparse and dense canopies in three non‐consecutive years. We explored the relationships between natural fruit set, pollen limitation and fruiting failure. Mean natural fruit set over the years was 60%, which is exceptionally high for a deceptive orchid. On average, hand cross‐pollination increased fruit set by 23%. Among open‐pollinated plants that did not set a fruit, 55.5% were estimated to be pollen limited and 44.5% to be limited by fruiting failure, i.e. inability to set a fruit after pollination. In species with high natural fruit set, hand cross‐pollination experiments may not always detect statistically significant pollen limitation. In our case, pollen limitation tended to become significant when the natural fruit set dropped below 60%. Canopy cover had a significant effect on fruiting failure, which was more severe under a dense canopy. Although our results demonstrate pollen limitation in many cases, they also highlight the fact that food deception can be a very effective pollination strategy. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 744–750.     

Genetic diversity of nectar‐rewarding Platanthera chlorantha and nectarless Cephalanthera rubra

We examined the genetic diversity of two orchid species, the nectar‐rewarding Platanthera chlorantha and the nectarless Cephalanthera rubra, in north‐eastern Poland. We found lower differences in genetic diversity between the species than we expected. The level of genetic variation at species level was lower in C. rubra (P_POL = 14%, A = 1.14 and H_E = 0.060) than in P. chlorantha (P_POL = 25.5%, A = 1.35 and H_E = 0.078). In the majority of populations of both species a high proportion of unique genotypes was noted. The overall F_ST values for all populations were moderate and similar for both species (P. chlorantha: 0.251, P < 0.001; C. rubra: 0.267, P < 0.001). No relationship was found between genetic and geographical distances in either species (P > 0.05, Mantel test). We discuss the breeding systems, small population size and population subdivision as the most important factors affecting the genetic diversity of this species. We suggest that conservation programmes should be initiated to maintain or even increase the fitness and genetic variation of populations of both species. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 751–763.     

Genome size variation in Orchidaceae subfamily Apostasioideae: filling the phylogenetic gap

With more than 160‐fold variation, Orchidaceae are currently the most diverse angiosperm family with respect to the amount of nuclear DNA. This study provides first genome size estimates for approximately 50% of species currently recognized in subfamily Apostasioideae, which is sister to the other four orchid subfamilies. The estimated 1C‐values range from 0.38 pg in Apostasia nuda to 5.96 pg in Neuwiedia zollingeri var. javanica, a nearly 16‐fold range. The two genera show non‐overlapping genome sizes, with those in Apostasia being distinctly smaller than those in Neuwiedia. In fact, most Apostasia spp. are at the lower end of the range of orchid C‐values. Observed discontinuities in DNA amounts in genera most probably reflect interspecific variation in ploidy. In addition to ploidy heterogeneity in N. zollingeri var. javanica, intraspecific variation in genome size (up to 17.7%) was also detected in some species; this can be plausibly related to the incidence of different geographical variants or unrecognized taxonomic heterogeneity. The AT content varied from 62.6 to 66.0%, which is in the upper range recorded for angiosperms. The genome size data obtained in this study fill a major phylogenetic gap in Orchidaceae and show that (very) small genomes prevail in subfamily Apostasioideae. © 2013 The Linnean Society of London     

Size,structure and nitrogen content of seeds of Fabaceae in relation to nodulation

Looking for clues to explain the low rate and unpredictability of the pollination of allogamous, epiphytic orchids in the context of the success of the orchid family in general, we compared the pollination of two miniature twig epiphytes in Mexico: Notylia barkeri, a ‘weedy’, rewarding species, and Erycina crista‐galli, a rare, deceptive species. We measured the effects of the spatial organization of the flowers and various parameters of visibility and height above the ground in relation to the spatial organization of the seed capsules (which retrospectively measures the activity of the pollinators). The spatial presentation of the flowers of E. crista‐galli explained only 5% of the distribution of the seed capsules. For N. barkeri, with compound flowers, the spatial presentation, considering individual flowers or inflorescences, explained 12% and 45% of the distribution of the seed capsules, respectively, and all other interactions between the variables were insignificant. Both the deceptive and reward strategies resulted in the production of large numbers of seeds despite a low pollination percentage (1–5%). Notylia barkeri produced 11.8 (2005–6) and 53.7 (2007–8) times more seeds in total than E. crista‐galli. Furthermore, unlike E. crista‐galli, N. barkeri responded to loss of individuals after high winds with an increased production of flowers per plant. We suggest that orchids have evolved to specialize in chance, and instead of maximizing pollinator attraction, they maximize the seed production resulting from every casual encounter. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 251–266.     

Where do Sardinian orchids come from: a putative African origin for the insular population of Platanthera bifolia var. kuenkelei?

Because of the complex geoclimatic dynamics of the Mediterranean basin, resulting from a combination of Tertiary geological processes and Quaternary climatic oscillations, the current species composition of the main continental islands is a mosaic resulting from relative contribution of vicariance, geodispersal and long‐distance dispersal by seed. To help distinguish among the contributions of these different dispersal modes to the orchid species richness on the island of Sardinia, we investigated the genetic structure and phylogeographic relationship of the only Platanthera bifolia var. kuenkelei population on the island. By using plastid DNA sequence variation, we show that this recently discovered population is more closely related to the Tunisian P. bifolia var. kuenkelei than to the geographically closer P. bifolia populations from peninsular Italy. Results from haplotype network construction support the hypothesis that the current distribution of P. bifolia var. kuenkelei (Sardinia and Tunisia) is most likely explained by over‐sea dispersal. However, haplotype diversity and mismatch analysis of this unique Sardinian population suggest that, if a long‐distance dispersal event did occur, it was not recent. More likely, temporarily favourable climatic conditions generated a suitable habitat on the island (also in terms of suitable mycorrhizas and pollinators) and thus a transient opportunity for seedling growth and population establishment of P. bifolia var. kuenkelei from North Africa. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 466–475.     

Floral shape mimicry and variation in sexually deceptive orchids with a shared pollinator

Mimics can have both accurate mimicry and phenotypic variation if deception operates in multiple sensory modes. Australian Tongue orchids (Cryptostylis species) attract their sole pollinator, male Lissopimpla excelsa wasps (Ichneumonidae), by accurately mimicking the scent and colour of female L. excelsa wasps. To test for shape mimicry of female wasps, both traditional and geometric morphometric comparisons were performed with allopatric Cryptostylis ovata and the often sympatric Cryptostylis erecta, Cryptostylis leptochila, and Cryptostylis subulata. Although some floral parts accurately mimicked the female wasp, the overall floral shape differed dramatically among orchid species. The function (if any) of this interspecific shape variation is unknown, although it does not cause character displacement of pollen attachment locations to reduce interspecific pollen transfer. Analyses showed that floral parts involved in pollinia transfer were similarly shaped for three of the four Cryptostylis species and all attach their pollinia to the same location on the pollinator's abdomen. Shape may interact with pollinator behaviour: in the field, pollination rates doubled when two Cryptostylis species were present, regardless of orchid abundances. Perhaps variation in shape hinders pollinator recognition and the avoidance of orchids, similar to scent and colour variation in other sexually deceptive orchid systems. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 469–481.     

Surprisingly high orchid diversity in travertine and forest areas in the Huanglong valley, China, and implications for conservation

The presence of such a large number of terrestrial orchid species in a small area (ca. 1 km²) of the Huanglong valley in southwestern China is uncommon for this country. Studying the relationship between the distribution patterns of these orchid species and their microenvironments may help us understand this uncommon phenomenon. We established 662 1 m × 1 m plots, measured the cover of each species and found that there were 33 orchid species distributed mainly in two different habitats, i.e. travertine areas and forest. In the travertine areas, 30 orchid species were found; the six most common ones being Cypripedium bardolphianum, Cypripedium flavum, Cypripedium tibeticum, Galearis diantha, Ponerorchis chusua and Phaius delavayi. However in the forested habitat, we found 21 orchid species; the most common ones being Tipularia szechuanica and Goodyera repens. Travertine areas had a higher number of orchid species as well as higher numbers of orchid species per plot as compared to forest. Light availability seems critical to the performance and distribution of orchid species. Stream flow through the travertine area during the orchids growing season appears to be an important factor in shaping and maintaining stable microenvironments favorable to the growth and reproduction of orchids. The results presented in this study suggest that some orchid species in the travertine area might be threatened if the travertine stream flows were to change or be disrupted.     

Molecular phylogenetics of Vanda and related genera (Orchidaceae)

The genus Vanda and its affiliated taxa are a diverse group of horticulturally important species of orchids occurring mainly in South‐East Asia, for which generic limits are poorly defined. Here, we present a molecular study using sequence data from three plastid DNA regions. It is shown that Vanda s.l. forms a clade containing approximately 73 species, including the previously accepted genera Ascocentrum, Euanthe, Christensonia, Neofinetia and Trudelia, and the species Aerides flabellata. Resolution of the phylogenetic relationships of species in Vanda s.l. is relatively poor, but existing morphological classifications for Vanda are incongruent with the results produced. Some novel species relationships are revealed, and a new morphological sectional classification is proposed based on support for these groupings and corresponding morphological characters shared by taxa and their geographical distributions. The putative occurrence of multiple pollination syndromes in this group of taxa, combined with complex biogeographical history of the South‐East Asian region, is discussed in the context of these results. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 549–572.     

Does facilitating pollinator learning impede deceptive orchid attractiveness? A multi‐approach test of avoidance learning

It has often been proposed that nectarless deceptive orchid species exploit naïve pollinators in search of food before they learn to avoid their flowers, and that intraspecific floral trait polymorphism, often noted in this plant group, could prolong the time needed for learning, thus increasing orchid reproductive success. We tested the importance of avoidance learning in a European deceptive orchid, Anacamptis morio, which has been reported to have a highly variable fragrance bouquet among individuals. We used an indirect approach, i.e. we facilitated pollinators’ ability to learn to avoid A. morio by adding anisaldehyde to selected inflorescences, a scent compound that is easily perceived by the natural pollinators and produced in large quantities by the closely related, nectar producing Anacamptis coriophora, a species that shares pollinator species with A. morio. In a series of three experiments (in artificial arrays, in natural populations and in bumblebee behavioural observations), we consistently found no difference either of reproductive success of or visitation rates to scent‐added versus control inflorescences. We also found that the decrease of reproductive success over time in artificial populations of this deceptive species was not as important as expected. Together, these data suggest that pollinators do not fully learn to avoid deceptive inflorescences, and that pollinator avoidance behaviour alone may explain the lower reproductive success usually found in deceptive orchids. We discuss the possible explanations for this pattern in deceptive orchids, particularly in relation to pollinator cognition and learning abilities. Lastly, in light of our results, the potential for higher average reproductive success in deceptive orchids with high phenotypic variability driven by avoidance learning thus appears to be challenged.