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Cypripedium references

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  • David Inouye
    Antonelli, A., C. J. Dahlberg, et al. (2009). Pollination of the Lady s slipper orchid ( Cypripedium calceolus ) in Scandinavia - taxonomic and conservational
    Message 1 of 1 , Jan 12, 2013
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      Antonelli, A., C. J. Dahlberg, et al. (2009). "Pollination of the Lady's slipper orchid (Cypripedium calceolus) in Scandinavia - taxonomic and conservational aspects." Nordic Journal of Botany 27(4): 266-273.
               The Lady's slipper orchid Cypripedium calceolus L. is considered one of the most beautiful orchids of Europe. Consequently, the species has suffered from over-collecting and is now critically endangered in many countries. Although pollination success is suspected to influence the long-term survival of Cypripedium calceolus, relatively little is known about the identity of its pollinators in mainland Sweden - a region that comprises the largest European populations. In order to identify which species pollinate eight representative populations in mainland Sweden, we observed and sampled visitors to flowers using a standardized protocol. Specimens were identified and any pollen smear found on their body was examined for the presence of Cypripedium pollen. Nine species were recognized as effective pollen vectors (Andrena cineraria, A. carantonica, A. haemorrohoa, A. helvola, A. nigroaenea, A. praecox, Colletes cunicularius, Lasioglossum fratellum and L. fulvicorne), four of them for the first time in Scandinavia. This is the first time that a species of Colletes is reported to carry pollen of Cypripedium in this region. All but one specimens were females. Our results suggest a taxonomically heterogeneous pollinator fauna for Cypripedium calceolus and are discussed in light of the management of this species.

      B H., H. Sun, et al. (2005). "Pollination of a slippery lady slipper orchid in south-west China: Cypripedium guttatum (Orchidaceae)." Botanical Journal of the Linnean Society 148: 251-264.
              
      B H., H. Sun, et al. (2008). "Pollination of wild lady slipper orchids Cypripedium yunnanense and C. flavum (Orchidaceae) in south-west China: why are there no hybrids?" Botanical Journal of the Linnean Society 156(1): 51-64.
               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.

      Barkman, T. J., J. H. Beaman, et al. (1997). "Floral fragrance variation in Cypripedium: Implications for evolutionary and ecological studies." Phytochemistry 44(5): 875-882.
              
      Bergstr G., G. Birgersson, et al. (1992). "Floral fragrance disparity between three taxa of Lady's slipper Cypripedium calceolus (Orchidaceae)." Phytochemistry 31(7): 2315-2319.
               Compared 1 European subspecies and 2 North American varieties. They had distinctly different fragrances composed mainly of fatty acid derivatives, isoprenoids, and phenyl derivatives, respectively. Sexual reproduction is dependent on the deception of pollinating solitary bees, and the three taxon-specific chemical compositions most probably reflect functional evolution in relation to different bee faunas.

      Case, M. A. and Z. R. Bradford (2009). "Enhancing the trap of lady's slippers: a new technique for discovering pollinators yields new data from Cypripedium parviflorum (Orchidaceae)." Botanical Journal of the Linnean Society 160(1): 1-10.
               Approximately one-third of orchid species offer no reward to their floral visitors and instead trick them into pollination. Typically, these deceptive systems have low visitation and fruiting rates because pollinators can learn to avoid non-rewarding species. Consequently, pollination ecology studies in these species often require long hours in the field to witness relatively few floral visitations relative to rewarding plants. Cypripedium parviflorum is a food-deceptive orchid with a pouch-like trap that temporarily imprisons pollinators. To escape, pollinators exert pressure on the stigma which facilitates pollination and widens the escape holes located near each anther. This study reports the use of a ribbon and clip to block the escape passageway of this species in order to retain and observe visiting insects. The device was tested in a large population and was shown to increase significantly the probability of observing floral visitors by nearly three-fold. Ten species of hymenopteran visitors in the families Andrenidae, Apidae, Halictidae and Megachilidae were observed, with two female Adrena tridens and one male Adrena perplexa successfully removing pollen. Insect visitation to the orchids occurred during the first half of the flowering period and was significantly associated with warm, clear days.2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160, 1201310.

      Davis, R. W. (1986). "The pollination biology of Cypripedium acaule (Orchidaceae)." Rhodora 88: 445-450.
              
      Dodson, C. H. (1966). "Studies in orchid pollination. Cypripedium, Phragmipedium and allied genera." American Orchid Society Bulletin 35: 125-128.
               Have a copy of page 1 in Diptera folder.

      Edens-Meier, R., M. Arduser, et al. (2011). "Pollination ecology of Cypripedium reginae Walter (Orchidaceae): Size matters." Telopea 13(1-2): 327-340.
               As in most pollinator-limited orchids lacking edible rewards, a population of C. reginae in
      southern Missouri showed a low conversion ratio of flowers into fruits (0.04623) over two
      seasons. There was no relationship between the length of the secondary flowering stem, the
      number of foliage leaves on the same stem and the number of flowers (one or two) produced
      at the terminus of the stem. However, the size mattered based on the physical dimensions of
      pollinia-carrying insects vs. parameters of floral architecture. While a diverse range of floral
      visitors (Coleoptera, Diptera and Hymenoptera) to C. reginae were observed over three seasons,
      only six medium-sized bees (Anthophora, Apis and Megachile spp.) carried segments of massulate
      pollinia after three seasons of observation and collection. Pollinia were always deposited dorsally
      on the thorax. These bees had a mean width of 4.44 mm and depth of 3.41 mm whereas the
      rear exit length and width of the orchid measured 6.53 mm and 3.41 mm. respectively. In
      contrast, the more numerous but smaller bees (2.66 mm width and 2.16 mm depth) in the
      genera Augochlorella, Augochlora, Ceratina, Lasioglossum spp. etc., exited the flower via the same
      rear orifices without pressing against the dehiscent anthers. Larger bees (gynes of Bombus spp.)
      measuring 9.06 mm in width and 6.25 mm in depth, were too large to escape via the rear exits so
      they left the flower via the large, dorsal entrance (through which they first entered the labellum)
      never contacting either anther. As in the small-flowered C. plectrochilum, the larger flowered
      C. reginae receives many floral visitors but selects for pollinia-vectors of a discrete body size.

      Gregg, K. B. (2004). "Recovery of showy lady's slippers (Cypripedium reginae Walter) from moderate and severe herbivory by white-tailed deer (Odocoileus virginianus Zimmerman)." Natural Areas Journal 24(3): 232-241.
              
      Kery, M. and K. B. Gregg (2004). "Demographic analysis of dormancy and survival in the terrestrial orchid Cypripedium reginae." Journal of Ecology 92(4): 686-695.
              
      Li, P., Y. Luo, et al. (2008). "Pollination of Cypripedium plectrochilum (Orchidaceae) by Lasioglossum spp. (Halictidae): The roles of generalist attractants versus restructive floral architecture." Plant Biology 10: 220-230.
               The pollination of Cypripedium plectrochilum Franch. was studied in the Huanglong Nature Reserve, Sichuan, China. Although large bees (Bombus, Apis), small bees (Ceratina, Lasioglossum), ants (Formica sp.), true flies (Diptera) and a butterfly were all found to visit the flowers, only small bees, including three Lasioglossum spp. (L. viridiclaucum, L. sichuanense and L. sp.; Halictidae) and one Ceratina sp., carried the flowerpollen and contacted  the receptive stigma. Measurements of floral architecture showed that interior floral dimensions best fit the exterior dimensions of Lasioglossum spp., leading to the consistent deposition and stigmatic reception of dorsally-placed, pollen smears. The floral fragrance was dominated by one ketone, 3-methyl-Decen-2-one. The conversion rate of flowers into capsules in open (insect) pollinated flowers at the site was more than 38%. We conclude  that, while pigmentation patterns and floral fragrance attracted a wide variety of insect foragers, canalization of interior floral dimensions ultimately determined the spectrum of potential pollinators in this generalist, food-mimic flower. A review of the literature showed that the specialised mode of pollination-by-deceit in C. plectrochilum, limiting pollinators to a narrow and closely related guild of is typical for other members of this genus.

      Li, P., Y.-B. Luo, et al. (2008). "Pollination of the lady's slipper Cypripedium henryi Rolfe (Orchidaceae)." Botanical Journal of the Linnean Society 156(4): 491-499.
               The pollination ecology of Cypripedium henryi Rolfe, a slipper orchid endemic to west China, was investigated, and its floral shape, size, colour, and scent were analysed. Examination of the breeding system suggests that the flowers are self-compatible, but need pollen vectors for successful reproduction. The flower is rewardless; over 15 insects belonging to Araneida, Hymenoptera, Diptera, Lepidoptera, and Coleoptera were recorded as flower visitors, but most only alighted or rested on the flower. In the total 32 h of observations over 2 years, female Lasioglossum bees were found to be the most frequent visitors and the only pollinators. They showed a high visitation frequency and, surprisingly, re-visited the same flowers frequently. Cypripedium henryi probably attracts pollinators visiting the flowers through general food deception (odour components, colour, false nectar guides), as well as special structures (slippery labellum, slippery staminode). Although three Lasioglossum species visited the flowers, only L. sauterum Fan et Ebmer was found with pollen. Lasioglossum flavohirtum Ebmer was large and climbed out from the entrance. Morphologically, L. sichuanense Fan et Ebmer could be considered as a potential pollinator, but the collected specimens were found to have no pollen of C. henryi on their bodies. It was speculated that the particular floral scent of C. henryi discouraged the entrance of L. sichuanense bees. Lasioglossum sauterum was matched morphologically to the flower, but not all of the visitations resulted in effective pollinations, as some flowers of C. henryi were frequently re-visited and the pollen mass had been taken away by bees on previous visitations. c 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156, 491-499.

      Li, P., R. Pemberton, et al. (2012). "Fly pollination in Cypripedium: a case study of sympatric C. and C.." Botanical Journal of the Linnean Society 170(1): 50-58.
               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.and C. 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.was pollinated by fruit flies and C.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.and C.fit into the complex sapromyiophily pattern. It appears that pollinator specificity is responsible for their reproductive isolation. The discovery of fly pollination in C.and C. 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.release discrete pollinia with narrow stalks instead of the usual amorphous pollen smears. This 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.

      Li, P., R. Pemberton, et al. (2012). "Fly pollination in Cypripedium: a case study of sympatric C. and C.." Botanical Journal of the Linnean Society 170(1): 50-58.
               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.and C. 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.was pollinated by fruit flies and C.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.and C.fit into the complex sapromyiophily pattern. It appears that pollinator specificity is responsible for their reproductive isolation. The discovery of fly pollination in C.and C. 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.release discrete pollinia with narrow stalks instead of the usual amorphous pollen smears. This 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.

      Light, M. H. S. and M. MacConaill (1998). "Factors affecting germinable seed yield in Cypripedium calceolus var. pubescens (Willd.) Correll and Epipactis helleborine (L.) Crantz (Orchidaceae)." Botanical Journal of the Linnean Society 126: 3-26.
               We comment on pollinator movement patterns.

      Lipow, S. R., P. Bernhardt, et al. (2002). "Comparative rates of pollination and fruit set in widely separated populations of a rare orchid (Cypripedium fasciculatum)." International Journal of Plant Sciences 163(5): 775-782.
              
      O'Connell, L. M. and M. O. Johnston (1998). "Male and female pollination success in a deceptive orchid, a selection study." Ecology 79(4): 1246-1260.
               "If there is selection for earlier opening flowers [had higher fitness], why do pink lady's slippers not evolve to open earlier?  This species is one of the first flowers to open in the spring.  A late frost can destroy buds, flowers, and early-forming capsules (Luer 1975)."  Goes on to talk about constraints on phenology in other species too.

      Primack, R. and E. Stacy (1998). "Cost of reproduction in the pink lady's slipper orchid (Cypripedium acaule, Orchidaceae): an eleven-year experimental study of three populations." American Journal of Botany 85(12): 1672-1679.
              
      Primack, R. B. and P. Hall (1990). "Costs of reproduction in the pink lady's slipper orchid: A four-year experimental study." American Naturalist 136: 638-656.
              
      Primack, R. B., S. L. Miao, et al. (1994). "Costs of reproduction in the pink lady's slipper orchid (Cypripedium acaule): defoliation, increased fruit production, and fire." American Journal of Botany 81(9): 1083-1090.
              
      Ren, Z.-X., D.-Z. Li, et al. (2011). "Flowers of Cypripedium fargesii (Orchidaceae) fool flat-footed flies (Platypezidae) by faking fungus-infected foliage." Proceedings of the National Academy of Sciences.
               Charles Darwin was fascinated by the orchidinteractions, but he did not realize that many orchid species are pollinated by deceit. Cypripedium, a model lineage of nonrewarding orchid flowers, is pollinated primarily by bees. Here we present both an example of floral mimesis of fungus-infected foliage in orchids and an example of flat-footed flies (Agathomyia sp.; Platypezidae) as pollen vectors for angiosperms. Cypripedium fargesii is a nectarless, terrestrial, endangered orchid from southwestern China that requires cross-pollination to produce the maximum number of viable embryos. All insects caught entering or leaving the labellum sac were Agathomyia sp. carrying conidia of Cladosporium sp. on their mouthparts and legs, suggesting mycophagy. Blackish hairy spots on the upper surface of foliage may imitate black mold spots, serving as short-term visual lures. Some odor molecules also associated with Cladosporium cultures were isolated in the floral scent. Mimesis of fungus-infected foliage probably represents an overlooked but important option in angiosperm diversification, because there are three to five more Cypripedium spp. in southwestern China with the same mode of floral presentation and black-spotted hairy leaves.

      Ren, Z.-X., D.-Z. Li, et al. (2011). "Flowers of Cypripedium fargesii (Orchidaceae) fool flat-footed flies (Platypezidae) by faking fungus-infected foliage." Proceedings of the National Academy of Sciences 108(18): 7478-7480.
               Charles Darwin was fascinated by the orchidinteractions, but he did not realize that many orchid species are pollinated by deceit. Cypripedium, a model lineage of nonrewarding orchid flowers, is pollinated primarily by bees. Here we present both an example of floral mimesis of fungus-infected foliage in orchids and an example of flat-footed flies (Agathomyia sp.; Platypezidae) as pollen vectors for angiosperms. Cypripedium fargesii is a nectarless, terrestrial, endangered orchid from southwestern China that requires cross-pollination to produce the maximum number of viable embryos. All insects caught entering or leaving the labellum sac were Agathomyia sp. carrying conidia of Cladosporium sp. on their mouthparts and legs, suggesting mycophagy. Blackish hairy spots on the upper surface of foliage may imitate black mold spots, serving as short-term visual lures. Some odor molecules also associated with Cladosporium cultures were isolated in the floral scent. Mimesis of fungus-infected foliage probably represents an overlooked but important option in angiosperm diversification, because there are three to five more Cypripedium spp. in southwestern China with the same mode of floral presentation and black-spotted hairy leaves.

      EVOLUTION Correction for of Cypripedium fargesii (Orchidaceae) fool flat-footed flies (Platypezidae) by faking fungus-infected foliage,by Zong-Xin Ren, De-Zhu Li, Peter Bernhardt, and Hong Wang, which appeared in issue 18, May 3, 2011, of Proc Natl Acad Sci USA (108:7478 first published April 18, 2011; 10.1073/pnas.1103384108).

      The authors note that, due to misidentification by the entomologist, the flat-footed fly (Agathomyia sp.) is dropped and reidentified as Cheilosia lucida Barkalov et Cheng (Diptera, Syrphidae) by K. K. Huo of Shaanxi University of Technology. The genus Cheilosia includes some fungal feeders (1). Due to this error, we cannot prove that flat-footed flies (Platypezidae) are pollen vectors of an angiosperm species. Nevertheless, the specimens of Cheilosia lucida carried conidia of Cladosporium, thus this misidentification does not affect the main conclusions or interpretations in this article.

      Shefferson, R. P., T. Kull, et al. (2005). "Adult whole-plant dormancy induced by stress in long-lived orchids." Ecology 86(11): 3099-3104.
              
      Shefferson, R. P., T. Kull, et al. (2012). "Linking vegetative dormancy to fitness in two long-lived herbaceous perennials." Ecosphere 3(2): art13.
               Vegetative dormancy occurs in many plant families, but its evolutionary context remains a mystery. We asked whether vegetative dormancy is an adaptive response to environmental stress and environmental stochasticity in certain long-lived plant species. We conducted an in situ experimental study in two and three populations of Cypripedium calceolus and Cephalanthera longifolia, respectively, in Estonia. Plants were defoliated, shaded, or simply observed at the beginning of the growing season in 2002 and 2003, and monitored demographically through 2008. We assessed links between fitness and vegetative dormancy using stochastic life table response experiments (SLTREs), in which the impact of treatment on the log stochastic population growth rate (a = log  via shifts in projection matrix transitions in treated plants relative to controls was used to assess the fitness impacts of treatment-induced life history responses. In Cypripedium, the observed lifespans of individuals that became vegetatively dormant in 2003/04 was significantly higher than plants that had not done so (P = 0.050). Defoliation and shading resulted in lower levels of flowering in both species. Both defoliation and shading decreased a relative to controls in Cypripedium and Cephalanthera. Defoliation- and shading-induced shifts in transitions involving vegetative dormancy were generally associated with significantly positive SLTRE contributions to  and shifts in the standard deviations of demographic rates generally contributed little to  Thus, vegetative dormancy is likely to be an adaptive response to environmental stress and stochasticity. Further work on the genetic basis to vegetative dormancy will clarify whether enough heritability may have existed in the past, or exists now, to support vegetative dormancy as an adaptation.

      Shefferson, R. P., J. Proper, et al. (2003). "Life history trade-offs in a rare orchid: the costs of flowering, dormancy, and sprouting." Ecology 84(5): 1199-1206.
              
      Shefferson, R. P., B. K. Sandercock, et al. (2001). "Estimating dormancy and survival of a rare herbaceous perennial using mark-recapture methods." Ecology 82(1): 145-156.
              
      Shefferson, R. P. and E. L. Simms (2007). "Costs and benefits of fruiting to future reproduction in two dormancy-prone orchids." Journal of Ecology 95(4): 865-875.
               1 Reproduction is expected to occur at a cost to survival, growth or future reproduction. However, trade-offs in long-lived, clonal herbs have proven difficult to assess, particularly when they are prone to adult dormancy.
      2 We assessed the costs of fruiting in a study of two species of lady's slipper orchid, Cypripedium candidum and C. parviflorum, growing sympatrically in a wet meadow in north-eastern Illinois, USA, from 2000 to 2005.
      3 First, we characterized flowering and fruiting in both populations. We found some differences between species, with 68.6 +- 5.7% (mean +- SE) and 43.5 +- 1.4% of sprouting plants flowering, while 33.6 +- 10.0% and 33.5 +- 8.1% of flowering plants fruited in C. candidum and C. parviflorum, respectively.
      4 Next, we tested the survival, sprouting and flowering response to current fruiting using multistate mark-recapture statistics. The best-fit model posited no cost of fruiting. However, according to a model parsimonious with the best-fit model, fruiting may have resulted in a small cost to survival visible primarily in small-sized individuals of C. parviflorum (decrease from 0.846 in non-fruiting but flowering plants to 0.824 in fruiting plants). In all cases, fruiting resulted in an increased probability of future flowering, suggesting that reproduction may have a higher priority in resource allocation than survival.
      5 Finally, we tested the effects of fruiting on future fruiting using logistic regression for two years in which fruiting was particularly high, but detected no change in the probability of fruiting after fruiting.
      6 Fruiting may increase in response to internal cues, perhaps related to nutrient uptake or storage, in addition to the obvious effects of pollination. The result may be that plants with greater access to nutrients or with greater stored reserves are more likely to flower each season. We suggest a need for further research exploring the internal mechanisms governing fruiting response in long-lived, clonal herbs.

      Sugiura, N., T. Fujie, et al. (2001). "Flowering phenology, pollination, and fruit set of Cypripedium macranthos var. rebunense, a threatened lady's slipper (Orchidaceae)." Journal of Plant Research 114(1114): 171-178.
              
      Sugiura, N., M. Goubara, et al. (2002). "Bumblebee pollination of Cypripedium macranthos var. rebunense (Orchidaceae); a possible case of floral mimicry of Pedicularis schistostegia (Orobanchaceae)." Plant Systematics and Evolution 235(1-4): 189-195.
               Nectarless flowers of Cypripedium macranthos var. rebunense are pollinated by only queen Bombus pseudobaicalensis, which also pollinates nectar-producing flowers of Pedicularis schistostegia. Our previous study (Sugiura et al. 2001) suggested that they form a floral mimicry system: (1) Flowering phenology of both plants overlapped greatly. (2) Cypripedium occurred with lower frequency relative to Pedicularis. And (3) in a mixed stand of both plants, foraging bumblebees were sometimes confused between the mimic and model flowers. The present study clarified the system with new evidence. (4) Flower colour of Cypripedium and Pedicularis would be similar within the range of a bumblebee's visual spectrum. And (5) both species largely overlapped in spatial distribution. Based on these findings, we discuss how C. macranthos var. rebunense differs in pollination mechanism from other congeneric species, especially C. calceolus.

      Vogt, C. A. (1990). "Pollination in Cypripedium reginae (Orchidaceae)." Lindleyana 5: 145-150.
              
      Zheng, G., P. Li, et al. (2011). "Mixed bumblebee and blowfly pollination of Cypripedium flavum (Orchidaceae) in Sichuan, China." Ecological Research 26(2): 453-459.
               Most Cypripedium species are specialized orchids pollinated by, in a broad sense, bees or flies. Here we present the first evidence that a slipper orchid, Cypripedium flavum, is pollinated by both bees and flies, i.e., bumblebees and blowflies. Artificial pollination experiments demonstrated that the flowers of C. flavum are self-compatible, but need pollen vectors for successful reproduction. Field observations detected 25 insects visiting the flowers, and 14 of these insects entered into the labellum of the flowers, but only female bumblebees, Bombus hypnorum, B. remotus, and the blowfly Calliphora vomitoria exited of the labellum with pollen smears of C. flavum. The floral functional morphology of C. flavum appears to be more suited to bumblebees than to blowflies. The bumblebees are more efficient pollinators of the orchid, but blowflies are more frequent visitors, so they pollinated more flowers despite being less efficient.

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