Ophrys

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
About 40

The genus Ophrys is a large group of orchids from the alliance Orchis in the subtribe Orchidinae. There are many natural hybrids. The type species is Ophrys insectifera L.1753

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.

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

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 are considered the most important group of European terrestrial orchids.

During summer, all Ophrys orchids are dormant as an underground bulbous tuber, which serves 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 already start to wither.

Most Ophrys orchids are dependent on symbiotic fungi. Because of this, some species only develop small alternate leaves. Transplanting specimens, especially wild specimens, is difficult, sometimes impossible, due to this symbiosis. The shiny, basal leaves have a green or bluish color. Two to twelve flowers grow on an erect stem with basal leaves.

Orchids of the genus Ophrys use sexual deception to attract pollinators to their flowers (Schiestl 2005). 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 (Borg-Karlson 1990; Gogler et al. 2009; Stokl et al. 2009). 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 the pollinators to the flower (Schiestl 2005; Schluter et al. 2009).

The allomone produced by an orchid is specific for 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).

The Bumblebee Orchid (Ophrys bombyliflora) is a typical example. It has flowers that look and smell so much like female bumblebees that males flying nearby are irresistibly drawn in by this chemical signal, stimulating them sexually. The insect gets so excited that he starts to copulate with the flower. The firmness, the smoothness and the velvety hairs of the lip are a further incentive for the insect to enter the flower. The pollinia inadvertently stick to the head or the abdomen of the male bumblebee. On visiting another orchid of the same species, the bumblebee pollinates its sticky stigma with the pollinia. The filaments of the pollinia have, during transport, taken such a position that the waxy pollen are able to stick to the stigma. Such is the refinement of the reproduction. If the filaments hadn’t taken the new position, the pollinia could not have pollinated the new orchid.

Every Ophrys orchid has its own pollinator insect and is completely dependent on this species for its survival. Furthermore, duped males are not likely to return. They even ignore other plants of the same species. Therefore, 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.

Species

The genus has been subdivided in several sections and subsections:

Ophrys annae

• Ophrys sect. Apiferae Quentin
• Ophrys sect. Argolicae Verm. ex Quentin
  • Ophrys subsect. Atticae Quentin
• Ophrys sect. Bertoloniorum Quentin
• Ophrys sect. Mammosae Quentin
• Ophrys sect. Reinholdiae Quentin
• Ophrys sect. Tenthrediniferae Quentin
• Ophrys sect. Umbilicatae Quentin

• Ophrys aegirtica
• Ophrys alasiatica (Cyprus).
• Ophrys amanensis (E.Nelson ex Renz & Taubenheim) P. Delforge
  • Ophrys amanensis subsp. antalyensis (Kreutz & Seckel) Kreutz
  • Ophrys amanensis subsp. iceliensis (Kreutz) Kreutz
• Ophrys annae
• Ophrys antiochiana (Turkey).
• Ophrys apifera Huds. : Bee Orchid (Europe, Medit. to Caucasus).
  • Ophrys apifera var. apifera
  • Ophrys apifera var. aurita
  • Ophrys apifera var. basiliensis
  • Ophrys apifera var. botteroni
  • Ophrys apifera var. bicolor
  • Ophrys apifera var. curviflora A.Soulié 2004 (France)
  • Ophrys apifera var. friburgensis
  • Ophrys apifera var. immaculata
  • Ophrys apifera var. trollii
• Ophrys arachnitiformis Gren. & Philippi: (Europa, Asia Minor, Syria)
  • Ophrys arachnitiformis subsp. archipelagi (Gölz & H.R.Reinhard) Kreutz
  • Ophrys arachnitiformis subsp. cilentana (Devillers-Tersch. & Devillers) Kreutz

Ophrys araneola

• Ophrys araneola Rchb. : Small Spider Ophrys (Europe, Asia Minor)
  • Ophrys araneola subsp. argentaria (Devillers-Tersch. & Devillers) Kreutz
  • Ophrys araneola subsp. cretensis (H.Baumann & Künkele) Kreutz
  • Ophrys araneola subsp. illyrica (S.Hertel & K.Hertel) Kreutz
  • Ophrys araneola subsp. quadriloba (Rchb.f.) Kreutz
  • Ophrys araneola subsp. tommasinii (Vis.) Kreutz
  • Ophrys araneola subsp. virescens (Philippe ex Gren.) Kreutz
• Ophrys aranifera Huds.
  • Ophrys aranifera var. quadriloba Rchb.f.
  • Ophrys aranifera [beta]. virescens Gren
• Ophrys argentoriensis
• Ophrys argolica H.Fleischm. ex Vierh. (S. Greece to SW. Turkey).
  • Ophrys argolica subsp. biscutella (O.Danesch & E.Danesch) Kreutz
  • Ophrys argolica subsp. icariensis (M.Hirth & H.Spaeth) Kreutz
  • Ophrys argolica subsp. morisii (Martelli) Kreutz
  • Ophrys argolica subsp. pollinensis (O.Danesch & E.Danesch) Kreutz
• Ophrys akhdarensis (B.Baumann & H.Baumann) P. Delforge
• Ophrys atlantica (S. Spain, Sicilia, NW. Africa).
  • Ophrys atlantica subsp. atlantica (S. Spain, NW. Africa). Tuber geophyte
  • Ophrys atlantica subsp. hayekii (Sicilia, Tunisia). Tuber geophyte
• Ophrys attaviria D.Rückbr., U.Rückbr., Wenker & S. Wenker
  • Ophrys attaviria subsp. cesmeensis - now synonym of Ophrys cesmeensis (Kreutz) P. Delforge 2004

Ophrys balearica

• Ophrys aurelia - now synonym of Ophrys bertolonii subsp. aurelia (P.Delforge, Devillers-Tersch. & Devillers) Kreutz
• Ophrys ausonia Devillers, Devillers-Tersch. & P. Delforge (Italy)
• Ophrys aveyronensis (J.J.Wood) H. Baumann & Künkele
• Ophrys aymoninii
• Ophrys bertolonii Moretti: Bertoloni's Bee Orchid (W. & C. Medit.)
  • Ophrys bertolonii subsp. aurelia (P.Delforge, Devillers-Tersch. & Devillers) Kreutz
  • Ophrys bertolonii subsp. balearica (Baleares).
  • Ophrys bertolonii subsp. bertolonii (W. & C. Medit.)
  • Ophrys bertolonii subsp. drumana (P.Delforge) Kreutz
• Ophrys bilunulata (now synonym of Ophrys fusca subsp. bilunulata (Risso) Aldasoro & L.Sáez 2005)
• Ophrys bornmuelleri Schulze
  • Ophrys bornmuelleri var. ziyaretiana (Kreutz & Ruedi Peter) P. Delforge
  • Ophrys bornmuelleri subsp. ziyaretiana (Kreutz & Ruedi Peter) Kreutz
• Ophrys bombyliflora : Bumblebee Orchid (Canary Is., Medit.)
• Ophrys calypsus M. Hirth & H. Spaeth - now synonym of Ophrys heldreichii subsp. calypsus (M.Hirth & H.Spaeth) Kreutz 2004
  • Ophrys calypsus var. pseudoapulica (P.Delforge) P. Delforge
  • Ophrys calypsus var. scolopaxoides (P.Delforge) P. Delforge
• Ophrys candica (E.Nelson ex Soó) H. Baumann & Künkele
  • Ophrys candica subsp. calliantha (Bartolo & Pulv.) Kreutz
  • Ophrys candica subsp. lacaena (P.Delforge) Kreutz
  • Ophrys candica subsp. lyciensis (H.F.Paulus, Gügel, D.Rückbr. & U.Rückbr.) Kreutz
• Ophrys catalaunica
• Ophrys cerastes Devillers & Devillers-Tersch (Greece)

Bumblebee Orchid (Ophrys bombyliflora)

• • Ophrys cerastes var. minuscula (G.Thiele & W.Thiele) Devillers & Devillers-Tersch
• Ophrys cesmeensis (Kreutz) P. Delforge
• Ophrys ceto Devillers, Devillers-Tersch. & P. Delforge (Greece)
• Ophrys ciliata Biv.
  • Ophrys ciliata subsp. lusitanica
  • Ophrys ciliata var. orientalis (Paulus) Kreutz
  • Ophrys ciliata subsp. regis-ferdinandii
• Ophrys cilicica (S. & SE. Turkey to N. Syria and Iran).
• Ophrys conradiae
• Ophrys cornuta Stev. ex M.Bieb.
  • Ophrys cornuta f. crassicornis Renz - now synonym of Ophrys crassicornis (Renz) Devillers-Tersch. & Devillers
• Ophrys cornutula (East Aegean Is.) - now a synonym of Ophrys oestrifera subsp. cornutula (Paulus) Kreutz 2004
• Ophrys crabronifera (C. & E. Italy, Croatia).
• Ophrys crassicornis (Renz) Devillers-Tersch. & Devillers
• Ophrys cretica (S. Greece, S. Aegean Is.)
• Ophrys dianica (Spain).
• Ophrys dodekanensis (East Aegean Is.) - now a synonym of Ophrys oestrifera subsp. dodekanensis (H.Kretzschmar & Kreutz) Kreutz 2004
• Ophrys drumana - now synonym of Ophrys bertolonii subsp. drumana (P.Delforge) Kreutz
• Ophrys eptapigiensis (East Aegean Is.)
• Ophrys eleonorae - now synonym of Ophrys iricolor subsp. eleonorae (Devillers-Tersch. & Devillers) Kreutz 2004
• Ophrys exaltata (NE. Spain to Italy).
  • Ophrys exaltata subsp. tyrrhena (Italy).
• Ophrys ferrum-equinum (Albania to SW. & S. Turkey).
  • Ophrys ferrum-equinum var. anafiensis (Greece) .
  • Ophrys ferrum-equinum var. ferrum-equinum (Albania to SW. & S. Turkey).
  • Ophrys ferrum-equinum var. gottfriediana (Greece).
  • Ophrys ferrum-equinum var. minor (Greece)
• Ophrys flavomarginata (Cyprus, Syria to Israel).
• Ophrys fleischmannii (E. Medit.)
• Ophrys forestieri
• Ophrys fusca Link: Dark Bee Orchid (Medit.)
  • Ophrys fusca ssp. bilunulata (Risso) Aldasoro & L.Sáez (2005)
  • Ophrys fusca ssp. blitopertha (Aegean Is. to SW. Turkey)

Ophrys fusca

• • Ophrys fusca ssp. cinereophila (S. Greece to N. Syria).
  • Ophrys fusca ssp. funerea
  • Ophrys fusca ssp. fusca (Medit.)
  • Ophrys fusca ssp. minima (France)
  • Ophrys fusca ssp. vasconica (SW. Europe).
• Ophrys gackiae P. Delforge (Sicilia)
• Ophrys garganica E. Nelson ex O. Danesch & E. Danesch - now a synonym of Ophrys passionis var. garganica (E.Nelson ex O. Danesch & E.Danesch) P. Delforge
  • Ophrys garganica subsp. majellensis (Helga Daiss & Herm.Daiss) Kreutz
  • Ophrys garganica subsp. sipontensis (R.Lorenz & Gembardt) Kreutz
• Ophrys halia (East Aegean Is.)
• Ophrys hebes (Kalopissis) E. Willing & B. Willing
  • Ophrys hebes var. negadensis (G.Thiele & W.Thiele) Kreutz
• Ophrys heldreichii Schltr.
  • Ophrys heldreichii subsp. calypsus (M.Hirth & H.Spaeth) Kreutz
  • Ophrys heldreichii subsp. pharia (Devillers & Devillers-Tersch.) Kreutz
  • Ophrys heldreichii subsp. pseudoapulica (P.Delforge) Kreutz
• Ophrys helios (Crete) - now synonym of Ophrys holosericea subsp. helios (Kreutz) Kreutz
• Ophrys herae M. Hirth & H. Spaeth
  • Ophrys herae subsp. janrenzii (M.Hirth) M. Hirth 2005
• Ophrys holoserica (Burm.f.) Greuter (syn. of Ophrys apifera Huds.) (W. & C. Europe to Medit.) ^[1]
  • Ophrys holosericea subsp. aeoli (P.Delforge) Kreutz
  • Ophrys holosericea subsp. aramaeorum (P.Delforge) Kreutz
  • Ophrys holoserica ssp. biancae (SE. Sicilia).
  • Ophrys holoserica ssp. bornmuelleri (E. Medit. to N. Iraq).
  • Ophrys holoserica ssp. candica (SE. Italy to SW. Turkey).
  • Ophrys holoserica ssp. chestermanii (SW. Sardegna).
  • Ophrys holosericea subsp. dinarica (Kranjcev & P.Delforge) Kreutz
  • Ophrys holoserica ssp. elatior (France to S. Germany).
  • Ophrys holosericea subsp. episcopalis (Poir.) Kreutz
  • Ophrys fuciflora ssp. gracilis

Ophrys holoserica

• • Ophrys holosericea subsp. gresivaudanica (O.Gerbaud) Kreutz
  • Ophrys holosericea subsp. halia (Paulus) Kreutz
  • Ophrys holoserica nothosubsp. halicarnassia (Turkey).
  • Ophrys holosericea subsp. helios (Kreutz) Kreutz
  • Ophrys holoserica ssp. holoserica (W. & C. Europe to Medit.)
  • Ophrys holosericea subsp. holubyana (András.) Kreutz
  • Ophrys holosericea subsp. homeri (M.Hirth & H.Spaeth) Kreutz
  • Ophrys holoserica ssp. lacaitae (Sicilia to S. Italy).
  • Ophrys holosericea subsp. linearis (Moggr.) Kreutz
  • Ophrys holosericea subsp. lorenae (E.De Martino & Centur.) Kreutz
  • Ophrys holoserica ssp. maxima (Crete to SW. & S. Turkey, Israel).
  • Ophrys holoserica ssp. oxyrrhynchos (Sardegna to Sicilia, Italy).
  • Ophrys holosericea (subsp. posidonia (P.Delforge) Kreutz
  • Ophrys holosericea subsp. serotina (Rolli ex Paulus) Kreutz
  • Ophrys holosericea subsp. tetraloniae (W.P.Teschner) Kreutz
  • Ophrys holosericea subsp. untchjii (M.Schulze) Kreutz
• Ophrys iceliensis (Turkey) - now synonym of Ophrys amanensis subsp. iceliensis (Kreutz) Kreutz 2004
• Ophrys illyrica (Croatia) - now synonym of Ophrys araneola subsp. illyrica (S.Hertel

& K.Hertel) Kreutz 2004

• Ophrys incantata Devillers & Devillers-Tersch (Croatia)
• Ophrys incubacea Bianca ex Tod. : Spider Orchid (S. Europe).
  • Ophrys incubacea subsp. brutia (P.Delforge) Kreutz
  • Ophrys incubacea subsp. castri-caesaris Looken (2005) (France)
  • Ophrys incubacea subsp. garganica
• Ophrys insectifera : Fly Orchid (Europe).
  • Ophrys insectifera subsp. insectifera (Europe).
  • Ophrys insectifera f. luteomarginata L. Lewis 2006 (England)

Fly Orchid (Ophrys insectifera)

• Ophrys iricolor Desf. (Medit.)
  • Ophrys iricolor subsp. astypalaeica (P.Delforge) Kreutz
  • Ophrys iricolor subsp. eleonorae (Devillers-Tersch. & Devillers) Kreutz
  • Ophrys iricolor subsp. iricolor (E. Medit.)
  • Ophrys iricolor subsp. lojaconoi (P.Delforge) Kreutz
  • Ophrys iricolor subsp. maxima (Corse, Sardegna, Tunisia) T
  • Ophrys iricolor subsp. mesaritica (Paulus, C. Alibertis & A.Alibertis) Kreutz
• Ophrys isaura (S. Turkey).
• Ophrys israelitica (Aegean Is. to Jordan).
• Ophrys kopetdagensis (Turkmenistan).
• Ophrys kotschyi (Cyprus).
• Ophrys kojurensis Gölz 2006 (Iran)
• Ophrys lacaena P. Delforge (Greece)
• Ophrys latakiana (Syria) - now a synonym of Ophrys oestrifera subsp. latakiana (M.Schönfelder & H.Schönfelder) Kreutz 2004
• Ophrys laurensis Geniez & Melki - now a synonym of Ophrys lutea subsp. laurensis (Geniez & Melki) Kreutz 2004
• Ophrys liburnica Devillers & Devillers-Tersch (Croatia)
• Ophrys lindia (East Aegean Is.)
• Ophrys lunulata (Sicilia to S. Italy).
• Ophrys lupercalis
• Ophrys lutea Cav. : Yellow Ophrys (Medit.)
  • Ophrys lutea subsp. archimedea (P.Delforge & M.Walravens) Kreutz
  • Ophrys lutea subsp. battandieri (E.G.Camus) Kreutz
  • Ophrys lutea subsp. laurensis (Geniez & Melki) Kreutz
  • Ophrys lutea ssp. lutea (W. & C. Medit.)
  • Ophrys lutea subsp. numida (Devillers-Tersch. & Devillers) Kreutz
  • Ophrys lutea ssp. pseudospeculum (Medit.)

Yellow Ophrys (Ophrys lutea)

• Ophrys lycia (SW. Turkey)
• Ophrys lyciensis (Turkey) - now synonym of Ophrys candica subsp. lyciensis (H.F.Paulus, Gügel, D.Rückbr. & U.Rückbr.) Kreutz 2004
• Ophrys magniflora
• Ophrys mammosa Desf. : Bosom Orchid (SE. Europe to S. Turkmenistan).
  • Ophrys mammosa subsp. leucophthalma (Devillers-Tersch. & Devillers) Kreutz
  • Ophrys mammosa subsp. macedonica (H.Fleischm. ex Soó) Kreutz
• Ophrys marmorata
• Ophrys maxima (Crete).
• Ophrys medea Devillers & Devillers-Tersch (Croatia)
• Ophrys morio Paulus & Kreutz (Cyprus) - now a synonym of Ophrys transhyrcana subsp. morio (Paulus & Kreutz) Kreutz 2004
• Ophrys morisii (Martelli) G. Keller & Soó (S. Corse, Sardegna).
• Ophrys negadensis (Greece) - now synonym of Ophrys hebes var. negadensis (G.Thiele & W.Thiele) Kreutz 2004
• Ophrys oestrifera M.Bieb.
  • Ophrys oestrifera subsp. akcakarae Kreutz 2006 (Turkey)
  • Ophrys oestrifera subsp. bicornis (Sadler) Kreutz
  • Ophrys oestrifera subsp. cornutula (Paulus) Kreutz
  • Ophrys oestrifera subsp. dodekanensis (H.Kretzschmar & Kreutz) Kreutz
  • Ophrys oestrifera subsp. hygrophila (Gügel, Kreutz
  • Ophrys oestrifera subsp. karadenizensis (M. Schönfelder & H. Schönfelder) Kreutz
  • Ophrys oestrifera subsp. latakiana (M. Schönfelder & H.Schönfelder) Kreutz
  • Ophrys oestrifera subsp. leptomera (P. Delforge) Kreutz
  • Ophrys oestrifera subsp. minutula (Gölz & H.R. Reinhard) Kreutz
  • Ophrys oestrifera subsp. rhodostephane (Devillers & Devillers-Tersch.) Kreutz
  • Ophrys oestrifera subsp. schlechteriana (Soó) Kreutz
  • Ophrys oestrifera subsp. zinsmeisteri (A.Fuchs) Kreutz
• Ophrys omegaifera H.Fleischm. (Medit.)
  • Ophrys omegaifera subsp. algarvensis (D. Tyteca, Benito & M. Walravens) Kreutz
  • Ophrys omegaifera ssp. dyris (S. Iberian Pen, Baleares, Morocco).
  • Ophrys omegaifera subsp. hayekii (H.Fleischm. & Soó) Kreutz
  • Ophrys omegaifera ssp. omegaifera (C. & E. Medit.)
  • Ophrys omegaifera subsp. sitiaca (Paulus, C. Alibertis & A. Alibertis) Kreutz
  • Ophrys omegaifera subsp. vasconica (O.Danesch & E.Danesch) Kreutz

Ophrys morisii

• Ophrys ortuabis (Sardegna).
• Ophrys oxyrrhynchos Tod
  • Ophrys oxyrrhynchossubsp. biancae
  • Ophrys oxyrrhynchossubsp. calliantha
• Ophrys pallida (Sicilia, Malta, Algeria, Tunisia).
• Ophrys panattensis (Sardegna). Probably a hybrid between O. argolica and O. lunulata
• Ophrys parvula (East Aegean Is.).
• Ophrys passionis Sennen
  • Ophrys passionis var. garganica (E. Nelson ex O. Danesch & E. Danesch) P. Delforge
• Ophrys perpusilla Devillers-Tersch. & Devillers (Greece)
• Ophrys persephonae (E. Aegean Is.)
• Ophrys pharia Devillers & Devillers-Tersch (Croatia)
• Ophrys picta - now synonym of Ophrys scolopax subsp. picta (Link) Kreutz
• Ophrys pollinensis (Italy). Probably a hybrid between O. argolica and O. holosericea
• Ophrys praecox (S. Corse to NW. Sardegna)
• Ophrys promontorii (C. & S. Italy).
• Ophrys provincialis
• Ophrys pseudapifera
• Ophrys pseudoatrata S. Hertel & H. Presser (Italy)
• Ophrys pseudobertolonii (W. & C. Medit.)
  • Ophrys pseudobertolonii ssp. bertoloniiformis (W. & C. Medit.) Tuber geophyte
  • Ophrys pseudobertolonii ssp. pseudobertolonii (N. Italy) Tuber geophyte

Mirror Bee Orchid (Ophrys speculum)

• Ophrys pseudomammosa (Greece to N. Turkey).
• Ophrys reinholdii (Balkan Pen. to NW. Iran)
  • Ophrys reinholdii ssp. reinholdii (Balkan Pen. to SW. Turkey) Tuber geophyte
  • Ophrys reinholdii ssp. straussii (S. & SE. Turkey to NW. Iran) Tuber geophyte
• Ophrys rhodostephane Devillers & Devillers-Tersch (Croatia) - now a synonym of ** Ophrys oestrifera subsp. rhodostephane (Devillers & Devillers-Tersch.) Kreutz 2004
• Ophrys sabulosa Paulus & Gack ex P. Delforge (Sicilia)
• Ophrys santonica
• Ophrys schulzei (CS. & SE. Turkey to Lebanon and Iran)
• Ophrys scolopax Cav. : Woodcock Orchid (Hungary, Medit. to Caucasus).
  • Ophrys scolopax ssp. cornuta : Horned Orchid (Hungary to Turkey).
  • Ophrys scolopax ssp. heldreichii (Greece to SW. Turkey, Cyprus)
  • Ophrys scolopax subsp. picta (Link) Kreutz
  • Ophrys scolopax ssp. scolopax (Medit. to Caucasus)
  • Ophrys scolopax subsp. vetula (Risso) Kreutz
• Ophrys sepioides Devillers & Devillers-Tersch (Greece)
• Ophrys sicula : Lesser Yellow Bee Orchid (S. & E. Medit.)
• Ophrys speculum : Mirror Bee Orchid (Medit.)
• Ophrys sphegodes Mill. : Early Spider Orchid (W. & S. Europe to Turkey).
  • Ophrys sphegodes ssp. aesculapii (Albania to S. Turkey).
  • Ophrys sphegodes subsp. argensonensis (J.-C.Guérin & A.Merlet) Kreutz
  • Ophrys sphegodes subsp. grammica (B.Willing & E.Willing) Kreutz
  • Ophrys sphegodes subsp. herae (M.Hirth & H.Spaeth) Kreutz

Ophrys tenthredinifera

• • Ophrys sphegodes subsp. massiliensis (Viglione & Véla) Kreutz
  • Ophrys sphegodes ssp. oodicheila
  • Ophrys sphegodes subsp. panormitana (Tod.) Kreutz
  • Ophrys sphegodes ssp. parnassica (Greece, Crete).
  • Ophrys sphegodes ssp. sphegodes (W. & S. Europe to Crimea).
  • Ophrys sphegodes ssp. sicula
  • Ophrys sphegodes subsp. tarquinia (P.Delforge) Kreutz
  • Ophrys sphegodes subsp. zeusii (M.Hirth) Kreutz,
• Ophrys splendida
• Ophrys spruneri (Sicilia to Israel)
  • Ophrys spruneri ssp. panormitana (Sicilia).
  • Ophrys spruneri ssp. spruneri (Greece to Israel)
• Ophrys straussii H.Fleischm. & Bornm.
  • Ophrys straussii subsp. antiochiana (H.Baumann & Künkele) Kreutz
• Ophrys sulcata
• Ophrys tenthredinifera Willd. (Medit.)
  • Ophrys tenthredinifera subsp. aprilia (Devillers & Devillers-Tersch.) Kreutz
  • Ophrys tenthredinifera subsp. grandiflora (Ten.) Kreutz
  • Ophrys tenthredinifera var. guimaraesii (D.Tyteca) Kreutz
• Ophrys transhyrcana Czerniak.
  • Ophrys transhyrcana subsp. morio (Paulus & Kreutz) Kreutz
  • Ophrys transhyrcana subsp. sintenisii (H.Fleischm. & Bornm.) Kreutz
• Ophrys turcomanica (NE. Iran).
• Ophrys umbilicata (WC. Medit. to Iran).

Ophrys umbilicata

• • Ophrys umbilicata ssp. attica (Corse to Iran).
  • Ophrys umbilicata ssp. umbilicata (E. Medit. to S. Iran).
• Ophrys vasconica
• Ophrys vernixia (Medit.)
  • Ophrys vernixia ssp. ciliata (Medit.)
  • Ophrys vernixia ssp. orientalis (Greece) - now synonym of Ophrys ciliata var. orientalis (Paulus) Kreutz 2004
  • Ophrys vernixia ssp. regis-ferdinandii (E. Aegean Is. to SW. Turkey)
  • Ophrys vernixia ssp. vernixia (WC. & S. Portugal, N. Spain).
• Ophrys virescens Philippe ex Gren
• Ophrys zonata

Natural hybrids

• Ophrys × aghemanii (O. scolopax subsp. cornuta × O. turcomanica) (Iran).
• Ophrys × albertiana (O. apifera × O. holosericea) (Europe).
  • Ophrys × albertiana nothosubsp. albertiana (Europe). Tuber geophyte
  • Ophrys × albertiana nothosubsp. morellensis (O. apifera × O. holoserica ssp. candica) (Italy) Tuber geophyte.

Ophrys apifera (left),
Ophrys holoserica (right)
from Thomé Flora von Deutschland, Österreich und der Schweiz 1885

• Ophrys × alibertiana (O. spruneri × O. tenthredinifera) (Crete).
• Ophrys × ambrosii (O. fusca ssp. minima × O. vasconica) (Europe)
• Ophrys × angelensis (O. incubacea × O. promontorii) (Italy).
• Ophrys × anomala (O. maxima × O. scolopax ssp. heldreichii) (Crete)
• Ophrys × apicula (O. insectifera × O. sphegodes ssp. araneola) (Europe).
  • Ophrys × apicula nothosubsp. apicula (Europe). Tuber geophyte
  • Ophrys × apicula nothosubsp. fabrei (O. insectifera subsp. aymoninii × O. sphegodes subsp. arenosa) (Europe). Tuber geophyte
• Ophrys × argentariensis (O. crabronifera × O. incubacea) (Europe)
• Ophrys × aschersonii (O. holosericea × O. sphegodes) (Europe).
• Ophrys × azurea (O. promontorii × O. pseudobertolonii) (Italy).
• Ophrys x barbaricina (. (O.ciliata x O.morisii). (Sardinia)
• Ophrys × barlae (O. bertolonii × O. incubacea) (S. Europe).
• Ophrys × bastianii (O. magniflora × O. scolopax ssp. scolopax) (Europe)
• Ophrys × battandieri (O. fusca × O. lutea) (NW. Africa).
• Ophrys × baumanniana (O. cretica × O. sphegodes) (Crete)
• Ophrys × bayeri (O. fusca × O. omegaifera ssp. fleischmannii) (Cyprus).
• Ophrys × benoitiana (O. incubacea × O. sphegodes ssp. lunulata) (Sicily).
• Ophrys × bergonii (O. saratoi × O. scolopax ssp. scolopax) (Europe)
• Ophrys × bernardii (O. aveyronensis × O. scolopax ssp. scolopax) (Europe)
• Ophrys × bilineata (O. bertolonii × O. sphegodes) (Europe)
• Ophrys × bodegomii (O. passionis × O. tenthredinifera) (Spain).
• Ophrys × borakisiana (O. fusca × O. mammosa) (E. Aegean Is.).
• Ophrys × borgersiae (O. reinholdii × O. spruneri) (Greece).
• Ophrys × boscoquartensis (O. biscutella × O. sphegodes) (Italy).
• Ophrys × bourlieri (O. iricolor × O. lutea × O. tenthredinifera). (N. Africa).
• Ophrys × braunblanquetiana (O. fusca × O. incubacea) (Corse, Italy).
• Ophrys × broeckii (O. arachnitiformis × O. araneola) (Europe)
• Ophrys × burneriana (O. sphegodes ssp. cretensis × O. spruneri) (Crete)
• Ophrys × campolati (O. promontorii × O. tenthredinifera) (Italy.
• Ophrys × camusii (O. exaltata × O. sphegodes) (Italy).
• Ophrys × carica (O. ferrum-equinum × O. holosericea) (Turkey.
• Ophrys × carpinensis (O. biscutella × O. scolopax ssp. cornuta) (Italy).
• Ophrys × carquierannensis (O. exaltata ssp. arachnitiformis × O. fusca) (France).
• Ophrys × cascalesii (O. sphegodes ssp. araneola × O. passionis) (France).
• Ophrys × chiesesica (O. drumana × O. fuciflora) (Europe)
• Ophrys × chimaera (O. bertolonii × O. sulcata) (Europe)
• Ophrys × chobautii (O. lutea × O. vernixia ssp. ciliata) (SW. Europe).
• Ophrys × circaea (O. apifera × O. bombyliflora) (Italy).
• Ophrys × clapensis (O. bombyliflora × O. lutea) (France).
• Ophrys × composita (O. scolopax × O. tenthredinifera) (France)
• Ophrys × corcyrensis (O. ferrum-equinum × O. sphegodes) (Greece).
• Ophrys × cortesii (O. incubacea × O. araneola) (France)
• Ophrys × cosana (O. bombyliflora × O. incubacea) (France)
• Ophrys × costei (O. aveyronensis × O. passionis) (France)
• Ophrys × couloniana (O. bertolonii × O. promontorii) (Italy).
• Ophrys × cranbrookeana (O. exaltata ssp. arachnitiformis × O. scolopax) (France).
• Ophrys × cugniensis (O. bertolonii × O. lutea) (Sicilia).
• Ophrys × daneschianum (O. bertoloniiformis × O. tenthredinifera) (France)
• Ophrys × delphinensis (O. argolica × O. scolopax ssp. cornuta) (S. Greece).
• Ophrys x denisiana ((O. bertolonii subsp. explanata x O. ciliata subsp. ciliata) (Sicilia)

Ophrys x devenensis

• Ophrys × devenensis (O. holosericea × O. insectifera) (Europe).
• Ophrys × diakoptensis (O. delphinensis × O. spruneri) (Greece).
• Ophrys × domitia (O. bombyliflora × O. lutea ssp. pseudospeculum) (France)
• Ophrys × domus-maria (O. apifera var. ? apifera × O. morisii) (France)
• Ophrys × duvigneaudiana (O. araneola × O. scolopax ssp. scolopax) (France)
• Ophrys × eliasii (O. fusca × O. vernixia ssp. ciliata) (S. Europe).
  • Ophrys × eliasii nothosubsp. conimbricensis (O. fusca × O. vernixia) (Portugal) Tuber geophyte
  • Ophrys × eliasii nothosubsp. eliasii (S. Europe). Tuber geophyte
• Ophrys × emmae (O. bertolonii × O. vernixia ssp. ciliata) (S. Europe).
• Ophrys × enobarbia (O. bertolonii × O. fuciflora) (France)
• Ophrys × epidavrensis (O. argolica × O. sphegodes ssp. aesculapii) (Greece).
• Ophrys × estacensis (O. fuciflora × O. splendida) (France)
• Ophrys × ettlingeriana (O. argolica × O. umblicata ssp. attica) (Greece).
• Ophrys × extorris (O. holosericea × O. insectifera × O. sphegodes) (Austria).
• Ophrys × fabrei (O. araneola × O. aymoninii) (France)
• Ophrys × fayacensis (O. arachnitiformis × O. provincialis) (France)
• Ophrys × feldwegiana (O. ferrum-equinum × O. tenthredinifera) (Greece).
• Ophrys × fernandii (O. bombyliflora × O. vernixia ssp. ciliata) (France).
• Ophrys × ferruginea (O. fusca × O. holoserica) (Italy).
• Ophrys × flahaultii (O. apifera var. ? apifera × O. sphegodes) (France)
• Ophrys × flavicans (O. araneola × O. bertolonii) (France)
• Ophrys × gauthieri (O. fusca ssp. fusca × O. lutea ssp. lutea) (France)
• Ophrys × godferyana (O. arachnitiformis × O. sphegodes) (France)
• Ophrys × grafiana (O. bombyliflora × O. umblicata ssp. attica) (Greece)
• Ophrys × grampinii (O. sphegodes × O. tenthredinifera) (Italy to Sicilia).
• Ophrys × grinincensis (O. provincialis x O. saratoi)
• Ophrys × gumprechtii (O. bertolonii × O. holosericea ssp. parvimaculata) (Italy).
  • Ophrys × gumprechtii nothosubsp. enobarbia (O. bertolonii × O. holosericea) (Italy). Tuber geophyte
  • Ophrys × gumprechtii nothosubsp. gumprechtii (Italy). Tuber geophyte

Ophrys × heraultii

• Ophrys × heraultii (O. tenthredinifera × O. vernixia subsp. ciliata) (Baleares).
• Ophrys × hoeppneri (O. bombyliflora × O. sphegodes) (Italy).
• Ophrys × hybrida (O. insectifera × O. sphegodes) (France)
• Ophrys × insidiosa (O. aegirtica × O. apifera) (France)
• Ophrys × inzengae (O. bertolonii × O. tenthredinifera) (France)
• Ophrys × jacquetii (O. araneola × O. magniflora) (France)
• Ophrys × jarigei (O. fusca ssp. minima × O. sphegodes)) (France)
• Ophrys × jeanpertii (O. araneola × O. sphegodes) (France)
• Ophrys × joannae (O. atlantica × O. iricolor) (N. Africa).
• Ophrys × kalteiseniana (O. ferrum-equinum × O. reinholdii) (Greece to Turkey).
• Ophrys × kelleri (O. exaltata subsp. arachnitiformis × O. incubacea) (France to Italy, S. Turkmenistan).
• Ophrys × kohlmuellerorum (O. scolopax ssp. scolopax × O. sulcata) (France)
• Ophrys × kreutziana (O. kotschyi × O. mammosa) (Cyprus).
• Ophrys × kulpensis (O. phrygia × O. schulzei) (Turkey)
• Ophrys × kurzeorum (O. atlantica × O. omegaifera ssp. dyris) (Spain) .
• Ophrys × laconensis (O. arachniiformis × O. tenthredinifera) (France)
• Ophrys × leguerrierae (O. lutea ssp. ? × O. araneola) (France)
• Ophrys × lievreae (O. iricolor × O. tenthredinifera) (N. Africa).
• Ophrys × lithinensis (O. omegaifera × O. sitiaca) (Crete).
• Ophrys × llenasii (O. incubacea × O. scolopax) (SW. Europe).
• Ophrys × luizetii (O. apifera × O. sphegodes ssp. araneola) (France)
• Ophrys × lumenii (O. bertolonii × O. ciliata) (France)
• Ophrys × lupiae (O. bertolonii × O. incubacea × O. tenthredinifera) (Italy).
• Ophrys × lydia (O. mammosa × O. vernixia ssp. regis-ferdinandii) (Greece to Turkey).
  • Ophrys × lydia nothosubsp. lydia (Turkey). Tuber geophyte
  • Ophrys × lydia nothosubsp. magnessa (O. mammosa × O. vernixia ssp. ciliata) (Greece). Tuber geophyte
• Ophrys × lyrata (O. bertolonii × O. incubacea) (France)
• Ophrys × macchiatii (O. sphegodes × O. vernixia ssp. ciliata) (Sardegna).
• Ophrys × maladroxiensis (O. holosericea × O. morisii) (Sardegna).
  • Ophrys × maladroxiensis nothosubsp. daissiorum (O. holosericea ssp. chestermanii × O. morisii) (Sardegna) Tuber geophyte
  • Ophrys × maladroxiensis nothosubsp. maladroxiensis (Sardegna) Tuber geophyte
• Ophrys × manfredoniae (O. incubacea × O. tenthredinifera) (Italy).
• Ophrys × maremmae (O. holosericea subsp. fuciflora × O. tenthredinifera) (Italy).
• Ophrys × marmarensis (O. holosericea × O. umbilicata) (E. Aegean Is.).
• Ophrys × mastii (O. ferrum-equinum × O. spruneri) (Greece).
• Ophrys × methonensis (O. argolica × O. tenthredinifera) (Greece).
• Ophrys × minuticauda (O. apifera × O. scolopax) (France, Sardegna).
  • Ophrys × minuticauda nothosubsp. donorensis (O. apifera × O. scolopax ssp. conradiae) (Sardegna).
  • Ophrys × minuticauda nothosubsp. minuticauda (France) Tuber geophyte
• Ophrys × mirtiae (O. sicula × O. vernixia ssp. ciliata) (Greece).
• Ophrys × monachorum (O. exaltata × O. fuciflora) (France)
• Ophrys × montis-angeli (O. biscutella × O. tenthredinifera) (Italy)
• Ophrys × montis-leonis (O. arachniformis × O. fuciflora ssp. fuciflora) (France)
• Ophrys × montserratensis (SW. Europe).
  • Ophrys montserratensis nothosubsp. montserratensis (Spain) Tuber geophyte
  • Ophrys montserratensis nothosubsp. neoruppertii (O. bertolonii × O. scolopax) (France). Tuber geophyte
• Ophrys × moreana (O. argolica × O. ferrum-equinum) (Greece).
• Ophrys × morensis (O. incubacea × O. praecox) (Sardegna).
• Ophrys × mulierum (O. mammosa × O. reinholdii) (Greece).
• Ophrys × nelsonii (O. insectifera × O. scolopax) (France).
• Ophrys × neocamusii (O. arachnitiformis × O. aurelia) (France)
• Ophrys × neorupperti (O. aurelia × O. scolopax ssp. scolopax) (France)
• Ophrys × neowalteri (O. bertolonii × O. sphegodes ssp. araneola) (France).
• Ophrys × obscura (O. fuciflora ssp. fuciflora × O. sphegodes) (France)
• Ophrys × olbiensis (O. bombyliflora × O. scolopax) (France).
• Ophrys × panattensis (O. conradiae × O. morisii) (France)
• Ophrys × pano-lefkaron (O. iricolor × O. mammosa) (E. Aegean Is. (Rodos) to Cyprus).
• Ophrys × pantaliciensis (O. incubacea × O. vernixia subsp. ciliata) (Sicilia).
• Ophrys × paphosiana (O. kotschyi × O. umbilicata) (Cyprus).
• Ophrys × peloponnesiaca (O. argolica × O. mammosa) (Greece).
• Ophrys × peltieri (O. scolopax × O. tenthredinifera) (N. Africa).
• Ophrys × personii (O. lutea × O. tentherdiniffera) (Italy).
• Ophrys × perspicua (O. holosericea × O. phrygia) (Turkey).
• Ophrys × pesseiae (O. morisii × O. panattensis) (Sardegna).
• Ophrys × philippei (O. scolopax × O. sphegodes) (S. Europe).
• Ophrys × pietzschii (O. apifera × O. insectifera) (Europe).
• Ophrys × piscinica (O. incubacea × O. lutea) (Italy).
• Ophrys × plorae (O. cretica × O. spruneri) (Crete).
• Ophrys × poisneliae (O. insectifera ssp. aymoninii × O. pseudobertolonii ssp. catalaunica) (Spain)
• Ophrys × provecta (O. bilunulata × O. omegaifera subsp. dyris) (Spain).
• Ophrys × pseudofunerea (O.funerea × O. sphegodes) (France)
• Ophrys × pseudofusca (O. fusca × O. sphegodes) (S. Europe)
• Ophrys × pseudoquadriloba (O. lutea × O. mammosa) (Greece).
• Ophrys × pseudospruneri (O. mammosa × O. spruneri) (Greece).
• Ophrys × pugliana (O. holosericea × O. oxyrrhynchos) (Italy).

Ophrys × pulchra

• Ophrys × pulchra (Ophrys araneola × Ophrys holoserica) (France, Germany)
• Ophrys × quadriloba (O. lutea ssp. lutea × O. sphegodes) (France)
• Ophrys × quintartiana (O. ariadnae × O. ferrum-equinum) (Greece).
• Ophrys × ragusana (O. bombyliflora × O. exaltata) (France)
• Ophrys × raimbaultii (O. incubacea × O. magniflora) (France)
• Ophrys × rainei (O. bombyliflora × O. incubacea) (S. Europe).
• Ophrys × rasbachii (O. argolica × O. vernixia ssp. ciliata) (Greece).
• Ophrys × rauschertii (O. apifera × O. insectifera × O. sphegodes) (Europe).
• Ophrys × raynaudii (O. argolica × O. reinholdii) (Greece).
• Ophrys × rechingeri (O. ferrum-equinum × O. mammosa) (Greece, Crete).
• Ophrys × regis-minois (O. cretica × O. scolopax ssp. heldreichii) (Crete)
• Ophrys × rovittellii (O. exaltata × O. tenthredinifera) (France)
• Ophrys × royanensis (O. drumana × O. insectifera) (France)
• Ophrys × salvatoris (O. bertolonii × O. biscutella) (Italy).
• Ophrys × samuelii (O. drumana × O. scolopax ssp. scolopax) (France)
• Ophrys × sanconoensis (O. exaltata × O. tenthredinifera) (Sicilia).
• Ophrys × sanctae-sofiae (O. iricolor × O. vernixia ssp. ciliata) (Sardegna).
• Ophrys × sancti-leonardi (O. fusca × O. tenthredinifera). (Italy).
• Ophrys x scheriana Riech. (O.bombyliflora x O.bremifera)
• Ophrys × semibombyliflora (O. bombyliflora × O. exaltata ssp. arachniformis) (Italy).
• Ophrys × simica (O. lucis × O. tenthredinifera) (E. Aegean Is.)
• Ophrys × sivana (O. episcopalis × O. holosericea ssp. candica) (Crete to Turkey).
• Ophrys × soller (O. apifera × O. vernixia ssp. ciliata) (Baleares).
• Ophrys × sommieri (O. bombyliflora × O. tenthredinifera) (Medit.)
• Ophrys × sorrentini (O. bertolonii × O. tenthredinifera) (S. Europe).
• Ophrys × souliei Soca 2005 (Ophrys aveyronensis (J.J. Wood) H. Baumann & Künkele × Ophrys fusca subsp. fusca.).) (France)
• Ophrys × spanui (O. annae × O. tenthredinifera) (France)
• Ophrys × spuria (O. bertolonii × O. fusca) (S. Europe).
• Ophrys × stefaniae (O. ferrum-equinum × O. iricolor) (Greece).
• Ophrys × sundermanniana (O. argolica × O. spruneri) (Greece).
• Ophrys × telchinensis (O. ferrum-equinum × O. lucis) (E. Aegean Is.).
• Ophrys × terrae-laboris (O. promontorii × O. sphegodes) (Italy).
• Ophrys × todaroana (O. incubacea × O. sphegodes ?)) (France)
• Ophrys × torrensis (O. argentaria × O. × pseudoscolopax) (Italy).
• Ophrys × triadensis (O. sicula × O. tenthredinifera) (Greece).
• Ophrys × tuscanica (O. crabronifera × O. tenthredinifera) (Italy).
• Ophrys × tytecana (O. aymoninii × O. insectifera) (France)
• Ophrys × varvarae (O. cretica × O. fusca) (Crete)
• Ophrys × vernonensis (O. biscutella × O. promontorii) (Italy).
• Ophrys × vespertilio (O. apifera × O. bertolonii) (Italy).
• Ophrys × vicina (O. holosericea × O. scolopax) (France, SW. Sardegna).
  • Ophrys × vicina nothosubsp. corriasiana (O. holosericea ssp. chestermanii × O. scolopax ssp. conradiae) (SW. Sardegna) Tuber geophyte
  • Ophrys × vicina nothosubsp. vicina (France) Tuber geophyte
• Ophrys × viglaensis (O. ferum-equinum var. minor × O. pruneri) (Greece)
• Ophrys × vogatsica (O. reinholdii × O. sphegodes) (Greece).
• Ophrys × waldmanniana (O. ferrum-equinum × O. vernixia ssp. ciliata) (Greece).
• Ophrys × yvonneae (O. holosericea ssp. apulica × O. incubacea) (Italy).

References

1. ^ Kew Botanical Gardens : World Checklist of Monocotyledons

Ayasse, M. 2007. Chemical mimicry in sexually deceptive orchids of the genus Ophrys. Phyton- Annales Rei Botanicae 46:221-223.

Gogler, J., J. Stokl, A. Sramkova, R. Twele, W. Francke, S. Cozzolino, P. Cortis et al. 2009. MENAGE TROIS-TWO ENDEMIC SPECIES OF DECEPTIVE ORCHIDS AND ONE POLLINATOR SPECIES. Evolution 63:2222-2234.

Schiestl, F. P. 2005. On the success of a swindle: pollination by deception in orchids. Naturwissenschaften 92:255-264.

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.

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-drivenselection maintains discrete odour phenotypes in hybridizing species. Biological Journal of the Linnean Society 98:439-451.

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.

External links

• Extensive information and beautiful slides
• Ophrys picture database
• Orchids of Greece, slide show.