Ophrys

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Ophrys
Bee Orchid
(Ophrys apifera var. aurita)
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Monocots
Order: Asparagales
Family: Orchidaceae
Subfamily: Orchidoideae
Tribe: Orchideae
Subtribe: Orchidinae
Alliance: Orchis
Genus: Ophrys
L., 1753
Species

See text

The genus Ophrys is a large group of orchids from the alliance Orchis in the subtribe Orchidinae. These plants are remarkable in that they successfully reproduce through pseudocopulation, that is, their flowers mimic female insects to such a degree that amorous males are fooled into mating with the flowers, thereby pollinating them. There are many natural hybrids. The type species is Ophrys insectifera L.

They are referred to as the "Bee orchids" due to the flowers of some species resemblance to the furry bodies of bees and other insects. Their scientific name Ophrys is the Greek word for "eyebrow", referring to the furry edges of the lips of several species.[1]

Ophrys was first mentioned in the book "Natural History" by Pliny the Elder (23-79 AD).

Biology

They are terrestrial or ground orchids from central to South Europe, North Africa, Asia Minor, up to the Caucasus Mountains, but mostly in the Mediterranean region. They have been said to be the most important group of European terrestrial orchids.[2]

During summer, all Ophrys orchids are dormant as underground bulbous tubers, which serve as a food reserve. In late summer/autumn they develop a rosette of leaves. Also a new tuber starts to grow and matures until the following spring; the old tuber slowly dies. The next spring the flowering stem starts to grow. During flowering the leaves have already started to wither.[1]

Most Ophrys orchids are dependent on symbiotic fungi. Transplanting specimens, especially wild specimens, is difficult, sometimes impossible, due to this symbiosis unless a large amount of surrounding earth is also taken with the plant. All orchids are protected under CITES II and should not be removed or disturbed in habitat.

The shiny, basal leaves have a green or bluish color. Two to twelve flowers grow on an erect stem with basal leaves.[2] These species are successfully cultivated by specialist growers of terrestrial orchids and are reported to be difficult to grow, being sensitive to rotting and damping off diseases if not properly subjected to a cool and dry aestivation period over the summer months with no water.[1]

Pollination

Orchids of the genus Ophrys use sexual deception to attract pollinators to their flowers.[3] In sexual deception, an orchid attracts male pollinators by producing the sex pheromone of virgin female pollinators in addition to providing visual and tactile cues (Schiestl 2005; Schluter et al. 2009; Stokl et al. 2009). These signals stimulate mating behavior in the male pollinators, which then attempt copulation, called “pseudocopulation”, with the orchid labellum (Schluter et al. 2009). During pseudocopulation, pollen from the flower’s column becomes attached to some part of the pollinator, usually the head or abdomen, and the pollinator inadvertently carries and transfers this pollen to other flowers when they are once again enticed into pseudocopulation.[4][5][6] While the morphological cues such as the shape and texture of the labellum play a role especially at close range in inducing the pollinator mating behavior, the orchid’s pheromone mimic, or allomone, has been shown to play the most important role in enticing pollinators to the flower (Schiestl 2005; Schluter et al. 2009).[2]

The allomone produced by an orchid is specific to its pollinator, of which it usually only has one (Ayasse et al. 2007; Gogler et al. 2009, Schluter et al. 2009) . The allomone is a mixture of alkenes and alkanes (Schiestl and Cozzolino 2008). There are one or more active species in this mixture that account for the attraction of pollinators (Vereeken and Schiestl 2008). Pollinators and orchids use the same chemical compounds in the same absolute amounts in their pheromones and allomones, respectively (Schiestl 2008).

Every Ophrys orchid has its own pollinator insect and is completely dependent on this species for its survival. Duped males are less likely to return and may ignore other plants of the same species. Only about 10% of an Ophrys population gets pollinated. This is enough to preserve the population, since each Ophrys orchid produces about 12,000 minute seeds.[1]

Species

The number of species recognized within the genus varies very widely between authorities. Flora Europaea in 1980 and Pedersen & Faurholdt in 2007 listed about 20 species in Europe as a whole;[7][8] Buttler in 1991 increased this to 53 for slightly larger geographical area;[9] Delforge in 1995 gave a total of 130 species.[10] By contrast, a molecular phylogenetic study in 2008 suggested that there were around 10 distinguishable groups.[11] As of March 2012, the World Checklist of Selected Plant Families recognizes 34 species of Ophrys worldwide, along with their subspecies:[12]

Ophrys apifera (Bee Orchid)
  • Ophrys apifera Huds. – Bee Orchid
  • Ophrys argolica H.Fleischm. – Argolian Bee-orchid
    • Ophrys argolica subsp. aegaea (Kalteisen & H.R.Reinhard) H.A.Pedersen & Faurh.
    • Ophrys argolica subsp. argolica
    • Ophrys argolica subsp. biscutella (O.Danesch & E.Danesch) Kreutz
    • Ophrys argolica subsp. crabronifera (Sebast. & Mauri) Faurh.
    • Ophrys argolica subsp. elegans (Renz) E.Nelson
    • Ophrys argolica subsp. lesbis (Gölz & H.R.Reinhard) H.A.Pedersen & Faurh.
    • Ophrys argolica subsp. lucis (Kalteisen & H.R.Reinhard) H.A.Pedersen & Faurh.
  • Ophrys atlantica Munby – Atlantic Bee-orchid
  • Ophrys bertolonii Moretti – Bertoloni's Bee-orchid
Ophrys bomybliflora (Bumblebee Orchid)
  • Ophrys bombyliflora Link – Bumblebee Orchid
  • Ophrys cilicica Schltr. – Cilician Bee-orchid
  • Ophrys ferrum-equinum Desf. – Horseshoe Bee-orchid
    • Ophrys ferrum-equinum subsp. ferrum-equinum
    • Ophrys ferrum-equinum subsp. gottfriediana (Renz) E.Nelson
  • Ophrys fuciflora (F.W.Schmidt) Moench – Late Spider-orchid
    • Ophrys fuciflora subsp. andria (P.Delforge) Faurh.
    • Ophrys fuciflora subsp. apulica O.Danesch & E.Danesch
    • Ophrys fuciflora subsp. biancae (Tod.) Faurh.
    • Ophrys fuciflora subsp. bornmuelleri (M.Schulze) B.Willing & E.Willing
    • Ophrys fuciflora subsp. candica E.Nelson ex Soó
    • Ophrys fuciflora subsp. chestermanii (J.J.Wood) H.Blatt & W.Wirth
    • Ophrys fuciflora subsp. elatior (Paulus) R.Engel & Quentin
    • Ophrys fuciflora subsp. fuciflora
    • Ophrys fuciflora subsp. grandiflora (H.Fleischm. & Soó) Faurh.
    • Ophrys fuciflora subsp. lacaitae (Lojac.) Soó
    • Ophrys fuciflora subsp. oxyrrhynchos (Tod.) Soó
    • Ophrys fuciflora subsp. pallidiconi Faurh.
    • Ophrys fuciflora subsp. parvimaculata O.Danesch & E.Danesch
Ophrys fusca (Sombre Bee-orchid)
  • Ophrys fusca Link – Sombre Bee-orchid, Dark Bee-orchid
    • Ophrys fusca subsp. blitopertha (Paulus & Gack) Faurh. & H.A.Pedersen
    • Ophrys fusca subsp. cinereophila (Paulus & Gack) Faurh.
    • Ophrys fusca subsp. fusca
    • Ophrys fusca subsp. iricolor (Desf.) K.Richt.
    • Ophrys fusca subsp. pallida (Raf.) E.G.Camus in E.G.Camus & A.A.Camus
  • Ophrys insectifera L. – Fly Orchid
    • Ophrys insectifera nothosubsp. tytecaeana P.Delforge
    • Ophrys insectifera subsp. aymoninii Breistr.
    • Ophrys insectifera subsp. insectifera
  • Ophrys isaura Renz & Taubenheim
  • Ophrys kojurensis Gölz
  • Ophrys konyana Kreutz & Ruedi Peter
  • Ophrys kopetdagensis K.P.Popov & Neshat.
  • Ophrys kotschyi H.Fleischm. & Soó – Cyprus Bee-orchid, Kotschy's Bee-orchid
    • Ophrys kotschyi subsp. ariadnae (Paulus) Faurh.
    • Ophrys kotschyi subsp. cretica (Soó) H.Sund.
    • Ophrys kotschyi subsp. kotschyi
  • Ophrys kreutzii W.Hahn
  • Ophrys lepida S.Moingeon & J.-M.Moingeon
  • Ophrys lunulata Parl.
Ophrys lutea (Yellow Bee-orchid)
  • Ophrys lutea Cav. – Yellow Bee-orchid
    • Ophrys lutea subsp. aspea (Devillers-Tersch. & Devillers) Faurh.
    • Ophrys lutea subsp. galilaea (H.Fleischm. & Bornm.) Soó
    • Ophrys lutea subsp. lutea
    • Ophrys lutea subsp. melena Renz
  • Ophrys lycia Renz & Taubenheim
  • Ophrys oblita Kreutz
  • Ophrys omegaifera H.Fleischm. – Omega Bee-orchid
    • Ophrys omegaifera subsp. dyris (Maire) Del Prete
    • Ophrys omegaifera subsp. fleischmannii (Hayek) Del Prete
    • Ophrys omegaifera subsp. hayekii (H.Fleischm. & Soó) Kreutz
    • Ophrys omegaifera subsp. israelitica (H.Baumann & Künkele) G.Morschek & K.Morschek
    • Ophrys omegaifera subsp. omegaifera
  • Ophrys pseudomammosa Renz
Ophrys reinholdii (Reinhold's Bee-orchid)
  • Ophrys reinholdii Spruner ex Fleischm. – Reinhold's Bee-orchid
    • Ophrys reinholdii subsp. antiochiana (H.Baumann & Künkele) H.Baumann & R.Lorenz
    • Ophrys reinholdii subsp. reinholdii
    • Ophrys reinholdii subsp. straussii (H.Fleischm.) E.Nelson
  • Ophrys schulzei Bornm. & Fleischm.
  • Ophrys scolopax Cav. – Woodcock Bee-orchid
    • Ophrys scolopax subsp. apiformis (Desf.) Maire & Weiller
    • Ophrys scolopax subsp. conradiae (Melki & Deschâtres) H.Baumann & al.
    • Ophrys scolopax subsp. cornuta (Steven) E.G.Camus
    • Ophrys scolopax subsp. heldreichii (Schltr.) E.Nelson
    • Ophrys scolopax subsp. rhodia (H.Baumann & Künkele) H.A.Pedersen & Faurh.
    • Ophrys scolopax subsp. scolopax
Ophrys speculum (Mirror Bee Orchid)
  • Ophrys speculum Link – Mirror Bee-orchid, Mirror Orchid
    • Ophrys speculum subsp. lusitanica O.Danesch & E.Danesch
    • Ophrys speculum subsp. regis-ferdinandii (Acht. & Kellerer ex Renz) Soó
    • Ophrys speculum subsp. speculum
  • Ophrys sphegodes Mill. – Early Spider-orchid
    • Ophrys sphegodes nothosubsp. jeanpertii (E.G.Camus) Del Prete & Conte
    • Ophrys sphegodes subsp. aesculapii (Renz) Soó ex J.J.Wood
    • Ophrys sphegodes subsp. araneola (Rchb.) M.Laínz
    • Ophrys sphegodes subsp. atrata (Rchb.f.) A.Bolòs
    • Ophrys sphegodes subsp. aveyronensis J.J.Wood
    • Ophrys sphegodes subsp. catalcana Kreutz
    • Ophrys sphegodes subsp. cretensis H.Baumann & Künkele
    • Ophrys sphegodes subsp. epirotica (Renz) Gölz & H.R.Reinhard
    • Ophrys sphegodes subsp. gortynia H.Baumann & Künkele
    • Ophrys sphegodes subsp. helenae (Renz) Soó & D.M.Moore
    • Ophrys sphegodes subsp. mammosa (Desf.) Soó ex E.Nelson
    • Ophrys sphegodes subsp. passionis (Sennen) Sanz & Nuet
    • Ophrys sphegodes subsp. sipontensis (R.Lorenz & Gembardt) H.A.Pedersen & Faurh.
    • Ophrys sphegodes subsp. sphegodes
    • Ophrys sphegodes subsp. spruneri (Nyman) E.Nelson
Ophrys tenthredinifera
  • Ophrys tenthredinifera Willd.
  • Ophrys tremoris Gämperle & Gölz
  • Ophrys ulupinara W.Hahn
  • Ophrys umbilicata Desf.
    • Ophrys umbilicata subsp. beerii Shifman
    • Ophrys umbilicata subsp. bucephala (Gölz & H.R.Reinhard) Biel
    • Ophrys umbilicata subsp. calycadniensis Perschke
    • Ophrys umbilicata subsp. flavomarginata (Renz) Faurh.
    • Ophrys umbilicata subsp. lapethica (Gölz & H.R.Reinhard) Faurh.
    • Ophrys umbilicata subsp. latakiana (M.Schönfelder & H.Schönfelder) Faurh. & H.A.Pedersen
    • Ophrys umbilicata subsp. umbilicata
  • Ophrys urteae Paulus
  • Ophrys zagrica Gölz
Ophrys sulcata - MHNT

References

  1. ↑ 1.0 1.1 1.2 1.3 Illustrated Encyclopedia of Orchids. ISBN 0-88192-267-6. 
  2. ↑ 2.0 2.1 2.2 Robert L. Dressler. The Orchids, Natural History and Classification. ISBN 0-674-87526-5. 
  3. ↑ Schiestl, F. P. (2005). "On the success of a swindle: pollination by deception in orchids". Naturwissenschaften 92: 255–264. 
  4. ↑ Borg-Karlson A.-K. 1990 Chemical and ethological studies of pollination in the genus Ophrys (Orchidaceae). Phytochemistry 29: 1359-1387
  5. ↑ Gögler, J.; Stökl, J.; Sramkova, A.; Twele, R.; Francke, W.; Cozzolino, S.; Cortis, P.; Scrugli, A.; Ayasse, M. (2009). "Ménage à Trois — Two Endemic Species of Deceptive Orchids and One Pollinator Species". Evolution 63 (9): 2222–2234. doi:10.1111/j.1558-5646.2009.00712.x. 
  6. ↑ Stokl, J., P. M. Schluter, T. F. Stuessy, H. F. Paulus, R. Fraberger, D. Erdmann, C. Schulz et al. 2009. Speciation in sexually deceptive orchids: pollinator-driven selection maintains discrete odour phenotypes in hybridizing species. Biological Journal of the Linnean Society 98:439-451.
  7. ↑ Tutin, T.G.; Heywood, V.H.; Burges, N.A.; Moore, D.M.; Valentine, D.H.; Walters, S.M. & Webb, D.A., eds. (1980), Flora Europaea, Vol. 5, Cambridge University Press, ISBN 978-0-521-20108-7 
  8. ↑ Pedersen, H.Æ. & Faurholdt, N. (2007), Ophrys : The Bee Orchids of Europe, Richmond, Surrey, UK: Kew Publishing, ISBN 978-1-84246-152-5 
  9. ↑ Buttler, Karl Peter (1991), Field Guide to Orchids of Britain and Europe, Swindon: Crowood, ISBN 978-1-85223-591-8 
  10. ↑ Delforge, Pierre (1995), Orchids of Britain and Europe, transl. Christine Grey-Wilson, London: HarperCollins, ISBN 978-0-00-220024-0 
  11. ↑ Devey, D.S.; Bateman, R.M.; Fay, M.F. & Hawkins, J.A. (2008), "Friends or Relatives? Phylogenetics and Species Delimitation in the Controversial European Orchid Genus Ophrys", Annals of Botany 101: 385–402 
  12. ↑ Search for "Ophrys", World Checklist of Selected Plant Families, Royal Botanic Gardens, Kew, retrieved 2012-03-29 
  • Ayasse, M. 2007. Chemical mimicry in sexually deceptive orchids of the genus Ophrys. Phyton- Annales Rei Botanicae 46:221-223.
  • Schiestl, F. P., and S. Cozzolino. 2008. Evolution of sexual mimicry in the orchid subtribe orchidinae: the role of preadaptations in the attraction of male bees as pollinators. Bmc Evolutionary Biology 8.
  • Schluter, P. M., P. M. Ruas, G. Kohl, C. F. Ruas, T. F. Stuessy, and H. F. Paulus. 2009. Genetic patterns and pollination in Ophrys iricolor and O-mesaritica (Orchidaceae): sympatric evolution by pollinator shift. Botanical Journal of the Linnean Society 159:583-598.
  • Vereecken, N. J., and F. P. Schiestl. 2008. The evolution of imperfect floral mimicry. Proceedings of the National Academy of Sciences of the United States of America 105:7484-7488.

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