- General Description
Mostly small perennial herbs. Rootstock tuberous, tubers two (sometimes three), globose or ovoid, entire, sessile or stipitate. Stem glabrous. Leaves basal, rosulate and/ or cauline, unspotted. Inflorescence lax. Flowers usually showy, insectiform. Sepals and petals free, ± spreading. Sepals larger, sometimes reflexed, usually glabrous, green, rose-pink, or whitish. Petals smaller, glabrous or hirsute. Labellum spurless, flattened, arcuate or strongly convex, entire, subentire or three-lobed, sometimes with two conical lateral protuberances at the base, or at the base of the lateral lobes, often with a distinct, deflexed, horizontal or upturned, glabrous apical appendage (which sometimes has a hair tuft at its base) or a less distinct apical mucro, velutinous above, with a variously shaped, often shiny, glabrous central area or markings (speculum), above which, in most species, is an often brightly coloured, shield-shaped area (basal field) at the base, and to either side of which may be two small swellings (basal callosities). Column having a short or rostrate, obtuse, acute or acuminate connective; rostellum minute; lateral appendages often indicated by coloured points, one on either side of the stigmatic cavity; pollinia two, attached to separate viscidia. Ovary sessile, not or slightly twisted, glabrous. (JW).
The majority of Ophrys orchids grow on calcareous soils, such as those derived from chalk and limestone which are alkaline, relatively warm and dry, and nutrient-poor (Delforge 1995). In the British Isles, O. sphegodes in southern England primarily occurs on chalk in Sussex and Kent, on the Purbeck limestones in Dorset, and on oolitic and Liassic limestones in Northamptonshire (Summerhayes 1951; Hutchings 1987). Only a few species in the genus also sometimes occur on more acidic soils, for example O. scolopax and 0. tenthredinifera and/or on more wet ones, such as O. bombyliflora and O. insectifera (Davies et al. 1988; Delforge 1995). Since orchids such as O. sphegodes and 0. insectifera can tolerate a wider range of soil pH and moisture conditions in the Mediterranean region, it has been suggested that their restriction to chalk and limestone soils in the northern parts of their ranges is advantageous in promoting their growth and survival in the cooler and damper climates of northern Europe (Summerhayes 1951). In the Mediterranean, Ophrys species grow most frequently in maquis or garrigue shrublands and unimproved grasslands and fallow fields (Delforge 1995). In Britain, Ophrys plants are typically found growing in open grasslands such as on the chalk downs of south-east England, where the turf is short (Summerhayes 1951; Lang 1989). The recent National Vegetation Classification of the British Isles cited the calcicolous grassland type as including O. fuciflora and O. sphegodes as well as other rare orchids such as Orchis simia Lam., Neotinea ustulata, and Orchis anthropophora (L.) All. (Rodwell 1992). Hutchings (1989) suggested for O. sphegodes in Sussex that such chalk grassland is a sub-optimal refuge habitat because its populations require careful habitat management for their survival, as the species can quickly be eliminated from ungrazed or fertilized sites through competition from other plants. Most Ophrys species prefer to grow in full sunlight but can tolerate partial shade and often grow in olive groves and open woods of pine, oak, and beech. Ophrys insectifera is unusual in that it frequently grows in dense woodlands in heavy shade, for example in the beechwoods of the North and South Downs in southern England (Summerhayes 1951; Lang 1989), although it does grow in more open habitats, such as among the limestone pavement of the Burren on the west coast of Ireland and in the peat bogs of the Central Plain with other marshland orchids such as Epipactis palustris and Gymnadenia conopsea (Summerhayes 1951). Man-made habitats, including disused quarries, railway embankments, and roadside verges, are other typical Ophrys habitats. Wells and Cox (1991) suggested that O. apifera behaves as a pioneer species because it is one of the earliest colonizers of bare or disturbed ground. It also grows on stabilized or semi-stabilized sand-dunes (Summerhayes 1951), and several other species often grow near the coast (e.g. O. sphegodes in England). In continental Europe, Ophrys species can grow at an elevation of 2000 m, although most species grow at mid-elevations of 800-1400 m (50% of the 126 Ophrys species listed by Delforge (1995) grew at mid-elevations compared with 32% at a range of 200- 700 m). Ophrys species flower in the spring in the Mediterranean, usually from March to May. In northern Europe, flowering in the summer is more common, as with O. insectifera (in May/June) and O. apifera (in June/July) (Delforge 1995). Within these flowering periods, individual Ophrys flowers are relatively long-lived (Summerhayes 1951) and can remain open for several weeks if they have not been pollinated (Neiland and Wilcock 1995). As the flowers reach anthesis, the basal rosette normally starts to senesce. These leaves emerged above ground in the previous autumn and remained green throughout the winter ('winter-green' orchids). Following complete senescence of the aerial stem at the end of the spring or summer, the plant remains underground as a tuber for several weeks or months, an adaptation that enables Ophrys species to avoid desiccation during the dry summer of the Mediterranean. Tubers and roots of Ophrys species are replaced every year. Observations made of O. apifera in England (Wells and Cox 1989) and O. sphegodes in Belgium (Delforge 1995) showed that new tubers begin to develop in November and can be seen as small white protuberances on the stem above the old tuber. During the winter, growth of the new tuber is slow, but in the spring it begins to increase in size rapidly. By the time the plant flowers, the new tuber may be as large or even larger than the old one, which shrivels as its food reserves are utilized in the formation of the flowering stem and new tuber. After flowering and senescence of the aerial parts, the old tuber has been completely re-absorbed and the new tuber becomes upright. By the early autumn, a new shoot has developed at its top, from which the new vegetative or flowering stem will develop. The proportion of plants that flower in a population in any one year varies considerably between Ophrys species. In two long-term demographic studies in southern England, the average frequency of plants flowering in O. apifera was 27.4% and in 0. sphegodes 83.7% (Hutchings 1987; Wells and Cox 1989). The probability of Ophrys plants flowering is influenced by characteristics such as tuber size and leaf number. For example, Wells and Cox (1989) found a highly significant difference in all plant characters measured in a study of O. apifera between plants going to flower and those going to remain vegetative. Plants that were going to flower in June had more leaves, a larger leaf area, more roots, larger tubers, and a greater total dry weight at four sampling times (November, March, May, and June) than plants that remained vegetative. Leaf characters (total rosette leaf area, leaf number, or length of longest leaf) were found to be particularly useful in predicting the probability of flowering in O. apifera. All plants that flowered had at least two leaves in mid -May; in plants with more than two leaves the probability of flowering increased with increasing rosette size. The number of leaves in a rosette depends not only on the age of the plant and therefore the resources available from the tuber but also on the seasonal emergence time. Wells and Cox (1989) found that plants of O. apifera that appeared above ground first tended to be those that eventually had the largest number of leaves, whereas plants that emerged later had fewer and smaller leaves. Weather conditions such as low rainfall may delay emergence time in the autumn (Rasmussen 1995). Drought can also prevent inflorescence formation (Wells and Cox 1989) or cause premature floral senescence (Neiland 1994), thereby reducing the likelihood of successful pollination and seed set in Ophrys species.
Fruit-set in Ophrys species is often low; for example 7.6% and 21.3% of flowers produced seed capsules in populations of O. vernixia Brot. and O. bombyliflora, respectively, in Portugal (Neiland 1994). An average of 8.7% was found among populations of O. insectifera studied in England, Germany, and Denmark (Neiland and Wilcock 1998). Ophrys apifera is exceptional in that most flowers set fruit throughout its range, but this high level of reproductive success is achieved by self-pollination. Ophrys capsules are usually large and may contain 10 000-15 000 seeds (Salisbury 1942; Neiland 1994), which is more than many other European orchid genera (Neiland and Wilcock 1995). Some germination of seeds has been observed in the field in spring (Fabre 1856; Fuchs and Ziegenspeck 1927, cited in Rasmussen 1999). It has often proved difficult to germinate seeds asymbiotically (Veyet 1969; Rasmussen 1995), although Moller (1985) successfully germinated and grew seedlings of O. holoserica from seed. Some fungal endophytes (Tulasnella sp.) have been found to be effective in symbiosis with Ophrys (Clements et al. 1986), and others have been isolated from the roots and identified as Eupulorhiza spp. (Andersen 1990, cited in Rasmussen 1995). Observations in the field indicate that following protocorm growth, formation of a tuber precedes leaf development (Rasmussen 1995). With O. apifera the first leaf is usually produced in the second year after germination, but the plant may remain in a vegetative state for an additional one to six years before flowering takes place (Wells 1981 ). Ophrys sphegodes, growing at the edge of its European range in southern England, is unusual in that time to flowering is short, with over 70% of plants flowering in the first year that they emerge above ground (Hutchings 1989).
Some Ophrys species have been reported to be monocarpic and show irregular patterns of appearance in particular localities (Summerhayes 1951). Recent demographic studies, however, demonstrated that Ophrys plants often have a period of dormancy during their lives, but that they can flower (and fruit) for several years in succession (Hutchings 1989; Wells and Cox 1991). In a study of O. sphegodes in Sussex in southern England, approximately 50% of the population was estimated to be dormant in any one year, and individuals could remain dormant for up to two years before re-emerging above ground (Hutchings 1987). Some orchids in the population had life spans of at least 10 years, but most were short-lived (half-lives of cohorts ranged from 1.5-2.3 years). Therefore, although the species is capable of producing fruit more than once, individual plants in the population are effectively monocarpic because they may flower only once before dying. By contrast, in a study of O. apifera in Cambridgeshire, the average half-life for the population was 6.6 years, and it was estimated that 64% of the population was more man eight years old (Wells and Cox 1981) . Regular seed production is likely to be particularly important for the maintenance of populations, especially those with short half-lives or at the edge of the range where environmental conditions may be sub-optimal, because vegetative propagation through multiplication of the tubers is insignificant in Ophrys. For example, Hutchings (1987) estimated mat <5% of O. sphegodes plants propagated vegetatively per annum. Given this dependence on seed input for population maintenance, conservation management practices that aim to reduce me mortality of flowering plants and encourage a high level of recruitment of new plants are likely to be most successful in conserving rare or threatened Ophrys populations (Hutchings 1989; Waite and Hutchings 1991).
Ophrys includes many species with localized distributions that are rare and threatened, including some listed as Red Data Book plants. A few species have been the focus of conservation programmes. For example, in Britain, O. sphegodes has been included in a wardening scheme to prevent it being collected from the wild (Farrell and Fitzgerald 1989). Favourable grazing management regimes have been successfully adopted at some locations to encourage seed recruitment and increase population sizes (Hutchings 1989; Waite and Hutchings 1991 ). However, across the range of me genus there are potential threats to me survival of Ophrys species. In a recent survey for the IUCN (IUCN Orchid Specialist Group 1996), Delforge cited overgrazing, coastal and inland tourist development, agriculture, urbanization, land drainage, garnering of tubers for salep, forestry, and general destruction of the habitat as serious threats and also assesses me risks of extinction as medium to high for many of me most threatened species, especially where populations are small. (RN).
A genus of approximately 126 species (Delforge 1995) distributed throughout Europe from the Canary Isles to the Caspian Sea, from southern Scandinavia south to North Africa, east to western Asia (NE Iran), attaining their greatest diversity in the Mediterranean Basin. The number of species listed by Delforge and others is undoubtedly inflated by the recognition of minor variants at specific rank. (JW).
Albania, Algeria, Austria, Baleares, Baltic States, Belarus, Belgium, Bulgaria, Canary Is., Central European Rus, Corse, Cyprus, Czechoslovakia, Denmark, East Aegean Is., East European Russia, Finland, France, Germany, Great Britain, Greece, Hungary, Iran, Iraq, Ireland, Italy, Kriti, Krym, Lebanon-Syria, Libya, Morocco, Netherlands, North Caucasus, North European Russi, Northwest European R, Norway, Palestine, Poland, Portugal, Romania, Sardegna, Sicilia, Spain, Sweden, Switzerland, Transcaucasus, Tunisia, Turkey, Turkey-in-Europe, Turkmenistan, Ukraine, Yugoslavia
- Ophrys × albertiana E.G.Camus
- Ophrys × angelarum Gennaio, M.Gargiulo, Medagli & Chetta
- Ophrys × angelii Soca
- Ophrys × apicula J.C.Schmidt ex Rchb.f.
- Ophrys apifera Huds.
- Ophrys × arachnitiformis Gren. & M.Philippe
- Ophrys × araniferiformis Dalla Torre & Sarnth.
- Ophrys argolica H.Fleischm.
- Ophrys × barbaricina M.Allard & M.P.Grasso
- Ophrys × battandieri E.G.Camus
- Ophrys × baumanniana Soó
- Ophrys bertolonii Moretti
- Ophrys bingoelensis Yild. & Kiliç
- Ophrys bombyliflora Link
- Ophrys × borgersiae P.Delforge
- Ophrys bornmuelleri M.Schulze
- Ophrys × bourlieri Maire
- Ophrys × breieri Wallenwein & Saad
- Ophrys × breuerorum P.Delforge
- Ophrys × brigittae H.Baumann
- Ophrys × brigodeana P.Delforge
- Ophrys × cailliauana P.Delforge
- Ophrys × campolati O.Danesch & E.Danesch
- Ophrys × carica H.Baumann & Künkele
- Ophrys × carpinensis O.Danesch & E.Danesch
- Ophrys × carquierannensis E.G.Camus
- Ophrys × cesinensis Medagli, D'Emerico & Ruggiero
- Ophrys × chobautii G.Keller ex B.Tyteca & D.Tyteca
- Ophrys × cicmiriana P.Delforge
- Ophrys cilicica Schltr.
- Ophrys × circaea W.Rossi & Prola
- Ophrys × circlarium Pellegrino
- Ophrys × clapensis Balayer
- Ophrys × cranbrookeana Godfery
- Ophrys cretica (Vierh.) E.Nelson
- Ophrys × cugniensis Soca
- Ophrys × daneschiana W.J.Schrenk
- Ophrys × denisiana H.Baumann, Künkele & R.Lorenz
- Ophrys × devenensis Rchb.f.
- Ophrys × diakoptensis M.Bayer
- Ophrys × domus-maria M.P.Grasso
- Ophrys × eliasii Sennen ex E.G.Camus & A.Camus
- Ophrys × emmae G.Keller ex H.Wettst.
- Ophrys × epidavrensis G.Eberle
- Ophrys × ettlingeriana P.Delforge
- Ophrys × extorris Soó
- Ophrys × feldwegiana B.Baumann & H.Baumann
- Ophrys × fernandii Rolfe
- Ophrys × ferruginea W.Rossi & Liuti
- Ophrys ferrum-equinum Desf.
- Ophrys × flavicans Vis.
- Ophrys fusca Link
- Ophrys × grafiana Soó
- Ophrys × grampinii Cortesi
- Ophrys × gumprechtii O.Danesch & E.Danesch
- Ophrys × heraultii G.Keller ex Schrenk
- Ophrys holosericea (Burm.f.) Greuter
- Ophrys × inceburumensis Kreutz
- Ophrys insectifera L.
- Ophrys × jansenii P.Delforge
- Ophrys × kalteiseniana B.Baumann & H.Baumann
- Ophrys × kelleri Godfery
- Ophrys × kohlmuellerorum Soca
- Ophrys kojurensis Gölz
- Ophrys konyana Kreutz & Ruedi Peter
- Ophrys kopetdagensis K.P.Popov & Neshat.
- Ophrys kotschyi H.Fleischm. & Soó
- Ophrys × kulpensis Kreutz
- Ophrys × laconensis Scrugli & M.P.Grasso
- Ophrys × laconica H.Baumann & Künkele
- Ophrys × liceana Renz & Taubenheim
- Ophrys × lievreae Maire
- Ophrys × luizetii E.G.Camus
- Ophrys lunulata Parl.
- Ophrys lutea Cav.
- Ophrys × macchiatii E.G.Camus
- Ophrys × maladroxiensis Scrugli, Todde & Cogoni
- Ophrys × maremmae O.Danesch & E.Danesch
- Ophrys × marganaiensis Licheri & Rodi
- Ophrys × mariae F.Fohringer
- Ophrys × marmarensis H.Baumann & Künkele
- Ophrys × methonensis H.Baumann & Künkele
- Ophrys × minuticauda Duffort
- Ophrys × montis-angeli O.Danesch & E.Danesch
- Ophrys × montis-grossi Kohlmüller
- Ophrys × moreana H.Baumann & Künkele
- Ophrys × nelsonii Contré & Delamain
- Ophrys × neocamusii Godfery
- Ophrys × neoruppertii A.Camus ex Ruppert
- Ophrys × notabilis Renz & Taubenheim
- Ophrys × nouletii E.G.Camus
- Ophrys × olbiensis E.G.Camus
- Ophrys omegaifera H.Fleischm.
- Ophrys × painiana P.Delforge
- Ophrys × paphosiana H.Baumann & Künkele
- Ophrys × pardui Giotta & Piccitto
- Ophrys × parenzani Medagli, D'Emerico & Ruggiero
- Ophrys × parvaisiana P.Delforge
- Ophrys × peltieri Maire
- Ophrys × personei Cortesi
- Ophrys × peteri Steffan
- Ophrys × pietzschii Kümpel ex Rumsey & H.J.Crouch
- Ophrys × pizzulensis Soca
- Ophrys × placotica B.Willing & E.Willing
- Ophrys × plorae C.Alibertis & A.Alibertis
- Ophrys × pseudofusca Albert & Camus
- Ophrys × pseudoquadriloba Renz
- Ophrys × quintartiana P.Delforge
- Ophrys × rainei Albert & Jahand.
- Ophrys × rasbachii G.Eberle
- Ophrys × rauschertii Kümpel
- Ophrys × raynaudii P.Delforge
- Ophrys × rechingeri Soó
- Ophrys × regis-minois Halx
- Ophrys reinholdii Spruner ex Fleischm.
- Ophrys × resurrecta O.Danesch & E.Danesch
- Ophrys × saintenoy-simoniana P.Delforge
- Ophrys × salvatoris O.Danesch & E.Danesch
- Ophrys × sansimonensis Soca
- Ophrys schulzei Bornm. & Fleischm.
- Ophrys scolopax Cav.
- Ophrys × semibombyliflora E.G.Camus
- Ophrys × sicana H.Baumann & Künkele
- Ophrys × sienaertiana P.Delforge
- Ophrys × soller M.Henkel
- Ophrys × sommieri E.G.Camus ex Cortesi
- Ophrys × sorrentinoi Lojac.
- Ophrys speculum Link
- Ophrys sphegodes Mill.
- Ophrys × spuria G.Keller ex Reinhard
- Ophrys × stefaniae G.Kretzschmar & H.Kretzschmar
- Ophrys × sulcitana Scrugli, Todde & Cogoni
- Ophrys tenthredinifera Willd.
- Ophrys × terschureniana P.Delforge
- Ophrys × torrensis H.Dekker
- Ophrys tremoris Gämperle & Gölz
- Ophrys × turiana J.E.Arnold
- Ophrys umbilicata Desf.
- Ophrys × vamvakiae Kohlmüller
- Ophrys × varvarae Faller & Kreutz
- Ophrys × vernonensis O.Danesch & E.Danesch
- Ophrys × vespertilio W.Rossi & Contorni
- Ophrys × vicina Duffort
- Ophrys × waldmanniana Soó
- Ophrys zagrica Gölz
First published in Sp. Pl.: 948 (1753)
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