The paper attempts an arrangement of genus Ophrys into first, second and, when possible, third and fourth order divisions, with a definition of the limits of species groups and species based on consideration of features resolvable into discrete character states. Some of them have long been known, others were little used in previous analyses.

Of the stigmatic cavity and column, we take into account:

-- the shape, spherical, open at the upper anterior quarter (Ophrys lutea, Ophrys fusca, Ophrys omegaifera and their allies), or hemispheric, with a lower surface level or nearly so (most species),

-- the sides of the opening, with a set of decorations: the staminodes, that either have or lack a terminal dark spot; the labia, dark, shining pairs of crests that surround the cavity and may break into callosities that are either similarly coloured or undifferentiated from the colour of the cavity walls; the pseudo-eyes, present in nearly all species except those that are pollinated by insects in abdominal position, brilliant protuberances derived from these callosities;

-- the floor, limited to species with a hemispherical cavity, flat or undulating, greyish or invaded by the colours of either the macula or the basal field;

-- the throat, V-shaped in some species (Ophrys fusca, Ophrys lutea), sill-shaped or straight in others, in the latter case forming a gular ridge that can be sharp or soft, straight or convex.

-- the connective, which is either truncated, not covering the pollinia (Ophrys lutea, Ophrys speculum, Ophrys bombyliflora, Ophrys insectifera), or beaked (most species);

Of the labellum we consider:

-- the configuration, either orchidoïd, with flattish outspread lateral lobes, or globular, with a shoulder at the base and interdependent or fused lobes;

-- the extremity of the central lobe, indented or not, bearing an appendix or not;

-- the pilosity, its configuration, texture, length and colour;

-- the macula, not in its detailed shape, but in its positioning, its system of coloration and ornamentation and its interaction with the cavity;

Of the other perianth segments, we use:

-- the shape of the petals, strap-like, long-triangular, short-triangular, rolled laterally or transversely, auriculated or not, with straight or undulated edges;

-- the shape of the lateral sepals, asymmetrical (the lower half bulging near the base), triangular or suborbicular;

-- the position of the dorsal sepal, erect or cupped over the connective;

-- the pilosity of the petals, absent, reduced to a velvety puberulence or well developed.

The major division of the genus into two sections, Pseudophrys and Euophrys, proposed by Godfery (1928), is upheld, and redefined in terms of synapomorphies. Within Pseudophrys two constellations are distinguished, that of Ophrys fusca-Ophrys lutea-Ophrys iricolor, with a V-shaped throat, and that of Ophrys omegaifera-Ophrys atlantica, with a sill-shaped throat. Euophrys is divided into three ensembles, organized around Ophrys speculum, Ophrys insectifera and Ophrys bombyliflora-Ophrys fuciflora-Ophrys sphegodes, respectively. Shape of the labellum, details of the stigmatic cavity and organization of the labellum pilosity are used to characterize them. It is argued that the Ophrys speculum complex, with the most divergent cavity, is the sister-group of the rest of the section.

Third order divisions are attempted for the larger constellations, those of Ophrys fusca-Ophrys lutea-Ophrys iricolor, Ophrys omegaifera-Ophrys atlantica and Ophrys bombyliflora-Ophrys fuciflora-Ophrys sphegodes. That of the second is straightforward, though perhaps artificial. Those of the other two present various difficulties.

The constellation of Ophrys fusca-Ophrys lutea-Ophrys iricolor is extremely difficult to organize because of the great morphological similarity between species, in particular between some that are known to be reproductively isolated by distinct pollinators (PAULUS and GACK, 1990c). We propose a subdivision into nine groups based on the detailed topography of the labellum and of its macula and on the distribution and coloration of its pilosity:

-- The Ophrys iricolor group is characterized by a very elongated labellum with basal crests inclined obliquely towards the exterior where they present an abrupt, often concave, slope bordering a raised and elongated central plateau. The labellum pilosity extends to, or almost to, the edges. The underside of the lip is generally red or tinged with red.

-- The Ophrys fusca group is characterized by an elongated to moderately elongated lip with parallel longitudinal mammosities, often well developed, inflating the bluish or sombre nuclei of the lunulae of the macula, and located more or less near the base of the lip. The lip pilosity is uniformly coloured or with a paler tuft nestled between the lunulae of the macula. The glabrous edge of the lip is often thin and generally separated from the pilose region by a progressive transition often with an irregular contour. The underside of the lip is green or tinged with red.

-- The Ophrys funerea group is characterized by an elongated lip without prominent relief in the nuclei of the lunulae of the macula or with long, weakly developed, longitudinal elevations. The lip pilosity comprises a paler, well-marked, fairly wide belt underlining the distal edge, and often the external edges, of the macula and contrasting with the dark lobes. The macula is generally divided into two by a narrow, perfectly rectilinear central relief, often carrying fairly long hairs. The glabrous edge of the lip is often narrow and generally separated from the pilose region by a progressive transition with an often irregular contour. The underside of the lip is green or tinged with red.

-- The Ophrys obaesa group is characterized by an elongated to moderately elongated lip adorned with long, sinuous areas of longitudinal relief in the nuclei of the lunulae of the macula and a strongly marked longitudinal groove prolonging the V-shaped throat. The extremity of the central lobe is globular, separated from the lateral lobes by a depression coinciding with the distal part of the macula that nearly reaches the sinus. The lip pilosity includes a paler, well-marked, fairly wide belt subtending the distal edge and often the external edges of the macula. The macula is divided by a narrow central band of fairly long hairs, white along the groove, brown-purple distally. The glabrous edge of the lip is often thin and generally separated from the pilose part by a progressive transition of irregular contour. The underside of the lip is greenish-yellow.

-- The Ophrys migoutiana group is characterized by an elongated lip nearly free of longitudinal relief. The lateral petals are exceptionally long and wide, particularly near the extremity. The lip pilosity is bicolored as in the two preceding groups, but the darker part is here confined to a narrow external wreath that includes only the tips of the lobes, the paler part surrounding the macula now invading the whole lip. The macula is glabrous, without bisecting relief or central line of hairs. The glabrous edge of the lip is very wide, abruptly separated from the pilose area, resulting in a sharp line that traces an even contour. The underside of the lip is green, rarely tinged red.

-- The Ophrys attaviria group is characterized by an elongated lip nearly free of longitudinal relief. The lip pilosity is sometimes nearly uniformly coloured but more often darkening progressively from the border of the macula towards the edges of the lobes, leaving paler, relatively irregular, zones. The macula is glabrous, without bisecting line or central pilosity. The glabrous edge of the lip is sometimes wide, separated from the pilose part by a demarkation line less clear-cut than in the preceding group. Each side of the base of the lip is joined to the exterior of the stigmatic cavity by a protruding dark flange. The underside of the lip is green.

-- The Ophrys "blithopertha-fusca" group is characterized by an elongated lip nearly without longitudinal relief. The pilosity of the lip is extremely short, bicolored. The macula is glabrous, with a well-marked bisecting relief. The glabrous edge of the lip often wide, separated from the pilose part by a relatively irregular demarcation line. The underside of the lip is green.

-- The Ophrys lutea group is characterized by the lip very wide, more than in all the other groups, the sides forming the least acute angle with the axis, generally greater than 45° . Well-marked mammosities inflate the blue or dark nuclei of the lunulae of the macula. The lip pilosity comprises an internal wreath of brown hairs surrounding the macula and an external wreath of yellow hairs. The macula is often divided in two by a narrow central relief, frequently carrying fairly long hairs similar to those of the external areas of the lip. The glabrous edge of the lip is very wide and generally separated from the pilose part by a progressive transition area with an irregular contour. The underside of the lip is yellowish green.

-- The Ophrys subfusca group is characterized by the lip wider than in all groups except the preceding one, the sides forming with the axis an angle of the order of 45° . Well-marked mammosities inflate the blue or dark nuclei of the lunulae of the macula. The lip pilosity comprises an internal wreath of brown hairs encircling the macula and an external wreath of yellow hairs. The macula is generally undivided, very rarely bisected by a narrow central relief. The glabrous edge of the lip is often very narrow and generally separated from the pilose part by a progressive transition zone with an irregular contour. The underside of the lip is yellowish green.

Within the constellation of Ophrys omegaifera, defined by a silled throat, adorned with a rectangular brush of white pilosity, the Ophrys omegaifera group forms a homogeneous ensemble. Its shared characters, macular omega, pilosity configuration, dark flanges at the base of the lip, pink or red coloration of the underside of the lip, are, however, not apomporphic at the level of the group, the very rigid arching of the lip being the only exception. The eastern group around Ophrys omegaifera and the only western representative, Ophrys dyris are divergent. The two taxa of the Ophrys atlantica group are highly distinct from each other as well as from any other group. Two species, Ophrys sitiaca and Ophrys vasconica, from opposite sides of the Mediterranean basin, have characters intermediate between the complexes of Ophrys omegaifera and Ophrys fusca-Ophrys lutea-Ophrys iricolor.

-- The constellation of Ophrys bombyliflora-Ophrys fusciflora-Ophrys sphegodes is well characterized by the rigid, globular structure of the labellum. The stigmatic cavity is very constant, hemispheric, with a flat or nearly flat floor terminated in the front by a gular crest more or less jutting outward. Pseudo-eyes are developed, most often from internal callosities. The labellum is comprised of four well defined regions, the basal field, the macula, the lobes and the appendix. The petals are hairy, except in species that have secondarily lost this pilosity, and the appropriate length for the pollinators is attained, contrary to what is found in other groups of the constellation, without significant longitudinal or transversal inrolling. In spite of the differences between species being larger then in other constellations, the limits of groups are not easy to establish. Eight are here thought to have a reasonable chance

of being monophyletic:

-- Ophrys tenthredinifera and Ophrys bombyliflora are very distinct and probably relatively distant, but are nevertheless united by very rounded, nearly orbicular, sepals, and a stigmatic cavity which is framed by labia better marked than in the rest of the constellation except Ophrys speculum. The internal labia constitute the ridge of the cavity, forming substantial internal callosities on the sides, long and sometimes contiguous in Ophrys bombyliflora, shorter, more like pseudo-eyes in Ophrys tenthredinifera. The external labia and their basal callosities are very marked in Ophrys bombyliflora, less so in Ophrys tenthredinifera. These species, alone in the constellation, have the extremity of the connective obtuse.

-- The complex of Ophrys fuciflora has elongated sepals, a pointed extremity to the connective and a stigmatic cavity without complete labia, characteristic of the constellation except for the preceding group, which it nevertheless resembles, especially Ophrys tenthredinifera, by the remaining characters. The pseudo-eyes, formed by the internal callosities, are often elongated transversally. They are at the extremities of a well marked, angular, gular crest to which they are linked by short pedicels, the ensemble making a mere suggestion of traces of the internal labia. More or less prominent vestiges of the basal callosities rise from the sides of the basal field. Staminodial points are present in most species. The basal field is often extensive, of a colour different from that of the floor of the cavity, concolorous or not with the labellum, the latter condition occasionally variable within the same species (Ophrys fuciflora). The pilosity of the labellum tends to be organized in a complete marginal or submarginal wreath reinforced by a tuft above the appendix, which is well developed. The strongly ciliated petals are often very short, or, if not, abruptly auriculated, the extremity always triangular and acute. The complex is relatively homogeneous. The transversal curving of the labellum, which has often been used to divide it, to the point of constructing separate sections, varies within the same species (Ophrys fuciflora, Ophrys apulica). A more natural partition into three groups (and one little-differentiated cluster) based on the pilosity of the labellum is proposed. They are:

-- The group of Ophrys episcopalis, mainly eastern, with a few species in Italy and the large Tyrrhenian islands, has the labellum sides with an abundant pilosity, long and silky, forming a wide, continuous marginal or submarginal wreath. The form of the labellum is somewhat variable, generally fairly globulous or prismatic, sometimes more spread, occasionally strongly inrolled.

-- The group of Ophrys fuciflora, formed mostly of central or sub-Mediterranean species, has a reduced labellum pilosity, located at the shoulders and sometimes immediately above the appendix. The external distal regions of the labellum are glabrescent or velvety. The labellum tends to be spread.

- The cluster of entities around Ophrys tetraloniae involves a number of recently detected populations of late flowering ophrys that have been attached either to Ophrys fuciflora or to Ophrys scolopax, depending on the degree of lateral inrolling of the labellum. They have in common an extremely glabrous labellum as in Ophrys fuciflora. The presence and denseness of the band of pilosity along the sides of the labellum is not constant, yet many individuals differ little from Ophrys fuciflora. The curvature of the labellum is variable. All the populations belong to this cluster, which we believe is situated phylogenetically within the group of Ophrys fuciflora, rather than to Ophrys scolopax.

-- The group of Ophrys scolopax, less well individualized than the first groups, composed of species dispersed throughout the Mediterranean basin and to the Caucasus, may be polyphyletic or even artificial. They can be characterized by the pilosity of the labellum, intermediate between the two other groups, forming a generally much narrower and sparser, marginal or submarginal, wreath than the others, but nevertheless complete. The species of this group have a much more constant tendency to lateral inrolling than do those of the other two groups.

-- The complex of Ophrys umbilicata is extremely similar to that of Ophrys fuciflora. It does not differ in a consistent way from all the groups included in the complex of Ophrys fuciflora except by the dorsal sepal recurved over the connective.

-- The complex of Ophrys apifera comprises species similar in aspect to the species of the Ophrys fuciflora complex, in particular, of the group of Ophrys scolopax. Ophrys apifera differs from all the other groups by the details of its stigmatic cavity. Its floor is undulated, pale yellowish, contrasting strikingly with the basal field, the central elevation greyish. The pseudo-eyes are compressed laterally, carried on elevated pedicels, straight and keeled. The longly flexuous and acuminate connective is found only in distant groups. The basal field is very long, bright red-orange, contrasting strongly with the labellum. Ophrys schulzei, of uncertain affinities, is superficially similar to Ophrys apifera, in particular by labellum shape.

-- The uniqueness of the complex of Ophrys reinholdii lies mostly in its stigmatic cavity. Labial vestiges are different from those in other Fuciflorae and Araniferae. The staminodial and temporal callosities are well marked by folds or bumps of the same colour at the edge of the cavity. The staminodial callosities are otherwise often terminated by a dark spot that greatly exceeds the staminodial points. The external labia are represented by crests that come out onto the base of the labellum and that sometimes have small shiny areas similar to pseudo-eyes. Inversely, and contrary to the other Fuciflorae, the internal labia are poorly marked and do not develop lustrous black pseudo-eyes. There are white callosities around the junction of the labia that seem to be formed by the external labia. The floor of the cavity extends fairly far beyond the lateral edges which appear relatively cut away. Large maculoïd patches are often present. The basal field is often downy.

-- The existence of a group of Ophrys argolica was proposed by GÖLZ and REINHARD (1982), a plausible hypothesis. It is difficult, however, to define this group by exclusive characters. The most obvious feature, the spectacled macula, is a detail of coloration that can appear in any of the groups within the constellation. Labellum pilosity does not differ significantly from that of the other Fuciflorae. The appendix is similar to that of the Ophrys reinholdii complex. Petal shape is probably the most unique feature. The petals appear as relatively full, elongated half-spindles, without the abruptly auriculated base or the stiff, acute distal triangle of the Ophrys fuciflora complex. The pseudo-eyes are small, wide-set and circular, formed near the junction of the labia in such a way that it is difficult to detect their derivation from either the internal labia as in the complex of Ophrys fuciflora or from the external ones as sometimes in that of Ophrys reinholdii, individual species of the group differing perhaps from each other in this detail. The floor of the cavity is largely maculoïd.

-- The complex of Ophrys sphegodes, without being necessarily phylogenetically more distant, is more sharply defined than all the preceding groups are from each other. The stigmatic cavity carries fewer labial vestiges than the Bombyliflorae or the Fuciflorae. The pseudo-eyes are clearly formed by the internal callosities but they are circular and stuck to the interior walls of the stigmatic cavity, appearing disconnected from the gular ridge, which itself is less marked. Staminodial points are absent. A large part of the cavity floor is invaded by the colour of the basal field, the maculoïd area being absent or reduced to a narrow band surrounding a pale central patch. The macula, less connected to the cavity than in the other complexes, is, when complete, often attached to the shoulders or to the exterior of the cavity and not to the labia, and it does not control the colour of the pseudo-eyes nor their contour. Labellum pilosity is usually fairly thick and arranged in a continuous marginal or submarginal wreath that is often wide. The extremity of the labellum is notched, the appendix absent or weakly developed. The petals are elongated, with undulated edges, non auriculated, glabrous or very weakly ciliated. To divide this very large complex coherently we use the colour and ornamentation of the stigmatic cavity and basal field, conditions which are constant at the species level in this complex, contrary to others, such the Ophrys fuciflora group. Five types are distinguished:

-- Ophrys sphegodes is a relatively isolated species, differing from all the others in the complex by the pale colour of the interior of the stigmatic cavity and the basal field, in strong contrast with the tone of the rest of the labellum. The exterior of the cavity is dull, tinged green, greyish or pinkish. The pseudo-eyes are large and generally a greenish-grey, quite reminiscent of the colour of the eggs of Acipenser stellatus.

-- The group of Ophrys incubacea, differs from Ophrys sphegodes by the stigmatic cavity. The interior of the cavity and the basal field are concolorous with the centra-distal part of the labellum. They are generally very dark, often blackish, sometimes dark red when the labellum itself is. The pseudo-eyes are small, black or blue, often bright blue or white exteriorly. The exterior of the cavity is often brilliant white, sometimes tinged pink or pale green but retaining the bright, shining shades. The edges of the cavity are joined to the exterior of the pseudo-eyes by short bridles of the same colour, often vivid white.

-- The species of the group of Ophrys arachnitiformis have, as in the group of Ophrys incubacea, the interior of the cavity and the basal field concolorous with the labellum. The walls of the cavity are dull, the pseudo-eyes large, grey-green, often surrounded by clear yellowish or greenish-yellow circles, that are prolonged in a joining bar across the back of the cavity. Many species are arachnitiform. A small appendix is more constantly present than in the two preceding groups.

-- Ophrys provincialis combines in a unique way the characters of the groups of Ophrys incubacea and Ophrys arachnitiformis. It, with Ophrys sphegodes, is the only species of the complex to have the cavity and the basal field not concolorous with the labellum. The cavity wall is generally brilliantly white or greenish white, connected to the pseudo-eyes by white bridles as in Ophrys passionis. The back of the cavity and the basal field are clear bright red, often visibly paler than the reddish brown labellum. The pseudo-eyes are surrounded and joined by a white double bar contrasting strongly with the back. The macula is generally strongly outlined by white and there is a small appendix.

-- Ophrys araneola is a very distinct and isolated species. Its stigmatic cavity and basal field are concolorous with the labellum, the cavity walls relatively dull as in the group of Ophrys arachnitiformis. The pseudo-eyes are black, contrasting little with the bottom of the cavity and are neither circled or connected by pale bands, but often weakly joined to the cavity wall by a greenish, comma-shaped infra-ocular mark. This design is very constant, close to that of the group of Ophrys bertolonii. The very flat labellum, very small in proportion to the other floral parts, the relatively weak pilosity of the submarginal wreath, the robust port of the plant in relation to its flowers, complete the ensemble of its unique characters. The species probably has affinities with the group of Ophrys arachnitiformis and with Ophrys sphegodes.

-- The complex of Ophrys bertolonii, very close to that of Ophrys sphegodes, differs by the more elongated labellum, always with a long and luxuriant pilosity, and by the constant presence of a well developed appendix. The stigmatic cavity, uniform within the group and similar to that of Ophrys araneola, has the interior and the basal field very dark, almost black. The pseudo-eyes are black, not contrasting with the cavity, not surrounded or linked by clear bands, but often slightly underlined by a greenish comma-shaped mark connected to the cavity wall. In most cases the walls of the cavity are cut away exposing the pseudo-eyes. The petals tend to be longer, with straighter edges than in the group of Ophrys sphegodes. The macula is characteristically limited to the central zone, horseshoe- or drop-shaped, rarely lined with white.

-- The complex of Ophrys mammosa, superficially very similar to that of Ophrys sphegodes, is, for the most part, composed of species that have habitually been associated with it within the section Araniferae. However, two very obvious characters separate these ensembles. The long pilosity of the labellum, that forms a well developed continuous wreath in Ophrys sphegodes and its allies, is very reduced in the Ophrys mammosa complex where it is limited to the shoulders. The rest of the labellum is covered with an extremely short, fine pilosity, invisible with a hand lens, that gives the surface a velvety, texture, shimmering or rippled in macrophotographs made with annular flash. The appendix, usually present, is of the same texture and colour as the rest of the labellum edge and prolongs the curvature of the extremity without, or nearly without, a hiatus. Other differences, possibly phylogenetically more significant, separate the two complexes. The macular system of Ophrys mammosa and its allies is joined with the stigmatic cavity as in the Fuciflorae. It connects distinctly with the external labial vestiges and it controls, in many species, large maculoïd patches on the floor of the cavity, even when the sides of the floor are invaded by the colour of the basal field. It often also controls the colour of the pseudo-eyes, which, in several of the species groups, are more at the exterior, in the region of junction of the labia, as in the Ophrys argolica and Ophrys reinholdii complexes. Also as in these complexes, the vestiges of the temporal, external and staminodial callosities, sometimes relatively prominent, often form folds on the sides of the cavity. Two less constant attributes support the division, the lower portion of the sepals of Ophrys mammosa and its allies are usually washed with brown or purple, as in the case of most of the eastern complexes. Their petals are a little more triangular, more auriculated at the base, more often slightly ciliated. The complex of Ophrys mammosa is limited to the eastern Mediterranean basin where it replaces that of Ophrys sphegodes, which is entirely absent.

The species of the Ophrys mammosa complex remain too little known to allow an entirely coherent subdivision. Details of the stigmatic cavity nevertheless permit the definition of a few groups:

-- Ophrys ferrum-equinum, Ophrys gottfriediana, Ophrys lycia, Ophrys spruneri, Ophrys mammosa, Ophrys helenae, Ophrys gortynia, Ophrys leucophthalma have a very dark basal field, concolorous with the labellum, or darker (Ophrys spruneri, Ophrys ferrum-equinum, Ophrys gottfriediana) than it. The stigmatic cavity shows all the characteristics of the complex. In particular, the maculoïd patches of the floor are very prominent. They are more striking than in any other group because of the intense, luminous colours that their reflecting parts share with the macula. The pseudo-eyes, often blue or bluish, are also habitually maculoïd. The vestigial folds of the sides of the cavity are prominent. Within the group, Ophrys ferrum-equinum and Ophrys gottfriediana occupy a singular place, with obvious affinities to the Ophrys argolica group.

-- Ophrys aesculapii and Ophrys epirotica share with the previous group the very dark basal field, and the close linkage of the macula to the cavity and the pseudo-eyes, which are white in Ophrys aesculapii, black or grey-blue in Ophrys epirotica. The highly reflecting blue patches of the species in the first group are lacking. The stigmatic cavity is less swollen by temporal folds and the labellum is flatter.

-- Ophrys sintenisii, Ophrys transhyrcana, Ophrys amanensis probably form a discrete group, united by the long and very acute extremity of the connective.

-- Ophrys grammica, Ophrys cretensis, Ophrys herae are three small- to medium-flowered species, characterized by the relatively pale, dull basal field, either concolorous with the labellum that has the same characteristics, as in Ophrys cretensis, or in spectacular contrast with it, as in Ophrys herae and Ophrys grammica. The floor of the cavity is little marked, or not marked, by maculoïd patches, the pseudo-eyes, large and greenish, are clearly internal, the other labial vestiges reduced to small external callosities.

The cladogram of Fig.13 depicts the phylogenetic hypothesis postulated here for the groups within genus Ophrys. It is based on the character-state table of Table 1, which summarizes the hypotheses made in the course of the present analysis on homologies, character polarity and transformation series.

Numerous species of the genus Ophrys are extremely vulnerable to alterations of the environment. Most vulnerable are species of limited or fragmented distribution. A preliminary evaluation of the relative fragility of species of the genus, based on a method of vulnerability indices has shown that some 60 % of Ophrys species have a very high vulnerability rating, above or equal to 0,75 times the maximum index. A more detailed analysis requires the mapping of ecological traits, such as choice of pollinator and phytosociological preference, on a sufficiently robust phylogenesis.

In the course of this study, several nomenclatural steps were necessary. Appendix 1 contains the technical details. The use of Ophrys subfusca (REICHENBACH fil.) HAUSSKNECHT, Ophrys battandieri CAMUS, Ophrys bilunulata RISSO, Ophrys mirabilis GENIEZ and MELKI, Ophrys picta LINK, Ophrys sphegodes MILLER and Ophrys arachnitiformis GRENIER and PHILIPPE is discussed. New combinations are proposed for Ophrys schlechteriana, Ophrys melitensis and Ophrys macedonica. Six species are described as new, Ophrys vallesiana, Ophrys lupercalis, Ophrys zonata, Ophrys calocaerina, Ophrys sulcata, Ophrys passionis, and Ophrys leucophthalma.

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