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Published date of profile: Jan-2006.
Citation: Mifsud S., (Jan-2006) on MaltaWildPlants.com
Johann Heinrich Friedrich Link, Germany, 1767-1851
= "Eyebrow", referring to the furry edges of the lips of several species. (Greek);
= Dark Brown (Latin).
No Main Synonyms
There are several recently-split species names of this variable plant. The ones in Malta are so far thought to belong to the Ophrys fusca/Ophrys iricolor complex group. -
Plant description and characters
Soon to be updated...
Localities in Malta:
Locally frequent such as Dingli Cliffs, Fawwara, Wardija (eg: San Martin area), Mellieha (eg: Red Tower area), Qrendi, Wied Hoxt and wied Babu. It was recently found by Stephen Mifsud in Gozo.
Protection in Malta:
Not Protected by Law (LN200/2011 or LN311/2006)
Red List 1989:
Not listed in the Red Data Book of the Maltese Islands
Unlikely to be poisonous.
The plant is seen in small communities or scattered isolates in soil carpets on garigue. They do not usually grow among dense vegetation. Their light green leaves are somehow more conspicuous than the dark brown flowers in their habitat.
The Sombre Bee Orchid is a small erect plant with a basal rosette of 4 to 6 leaves and a flowering stem that can raise up to 35cm. The leaves are broad lanceolate and sometimes slightly twisted. The base of the upper leaves tend to sheath around the flowering stem. Leaves have a smooth outline and show parallel venation.
The flower, as most orchids is complex and decorated. There are 3 outer segments, about 10mm long, that are perpendicular to each other (T-shaped) and considered as the sepals. They are broadly ovate and the upper one (median sepal) is curved inwards like a hood, while the lateral ones are less curved in. The corolla consist of 3 segments or petals of which the lateral ones are small, oblong, green and rather inconspicuous, while the lower segment is large, colourful and lip-shaped; referred to as the labellum. It has a velvet-like texture.
The labellum has 3 lobes, 2 lateral and small, and a central one which is large and often has a notch at its base. The outer border and most of the lower part is dark brown/maroon. At the central and inner part there is a rather pale violet-blue patch, often shaped as a 'W'. The edge of the labellum is curved down.
There is a degree of variety between flowers of different plants. This takes place in the pattern and hue of the pale violet-blue patch, the notch of the labellum's central lobe and the size of the flower. Some flowers show reddish-brown markings on the bluish patch. The flower does not possess a spur and is not particularly scented, though some state that it has the scent of phermones of female bees to attract male bees and assist in pollination.
The stamens are 2 special structures called pollinia. They are located at the roof of the flower's mouth, just under the hooded median sepal and have a sub-spherical sticky base at the end called viscidium. They produce a sticky yellow pollen. The stigma is also very small (barely visible) and located in a cavity at the floor of the flower's mouth (inner-most part of the labellum).
The flower has an inferior ovary that is directly attached to the flowering stem. It develops into an oval/elongated fruit capsule that produces and stores a large number of tiny seeds (powder form).
Information, uses and other details
Nativity and distribution
According to reference [WWW-03] the Genus Ophrys is native to the Mediterranean region. Ophrys fusca occur in the occidental and central Mediterranean basins, namely in Spain, France, Tunisia, Algeria and also in Italy and Sardinia. It is very common in Portugal, where it was first described in 1800. [WWW-143]
The plant has the following medicinal properties according to reference: [WWW-66]
|| A medication (in the form of an oil or salve etc.) that soothes inflamed or injured skin. [WWW-32]
||Of or pertaining to nutrition; having the quality of nourishing; nutritious; nutrimental; alimental [WWW-32]
Salep is very nutritive and demulcent. It has been used as a diet of special value for children and convalescents, being boiled with water, flavoured and prepared in the same way as arrowroot. Rich in mucilage, it forms a soothing and demulcent jelly that is used in the treatment of irritations of the gastro-intestinal canal. One part of salep to fifty parts of water is sufficient to make a jelly. The tuber, from which salep is prepared, should be harvested as the plant dies down after flowering and setting seed. 
The cooked root is a source of 'salep', a fine white to yellowish-white powder that is obtained by drying the tuber and grinding it into a powder  . Salep is said to be very nutritious and is made into a drink or added to other cereals and used in bread etc  . One ounce of salep is said to be enough to sustain a person for a day [100, 115] . The salep can also be made into a drink  .
Plants can be grown in a lawn, but the lawn must not be cut until the plants have set seed  . Plants are best grown in the shade  . Orchids are, in general, shallow-rooting plants of well-drained low-fertility soils. Their symbiotic relationship with a fungus in the soil allows them to obtain sufficient nutrients and be able to compete successfully with other plants. They are very sensitive to the addition of fertilizers or fungicides since these can harm the symbiotic fungus and thus kill the orchid  . This symbiotic relationship makes them very difficult to cultivate, though they will sometimes appear uninvited in a garden and will then thrive. Transplanting can damage the relationship and plants might also thrive for a few years and then disappear, suggesting that they might be short-lived perennials  . The flowers resemble a female insect and also emit a scent similar to female pheromones, they are pollinated by a male insect of that species attempting to copulate with the flower  . Tubers should be planted out whilst they are dormant, this is probably best done in the autumn  . They should be planted at least 5cm below soil level  .
Sow seed at the soil surface, preferably as soon as it is ripe, in the greenhouse and do not allow the compost to dry out. The seed of this species is extremely simple, it has a minute embryo surrounded by a single layer of protective cells. It contains very little food reserves and depends upon a symbiotic relationship with a species of soil-dwelling fungus. The fungal hyphae invade the seed and enter the cells of the embryo. The orchid soon begins to digest the fungal tissue and this acts as a food supply for the plant until it is able to obtain nutrients from decaying material in the soil  .
It is best to use some of the soil that is growing around established plants in order to introduce the fungus, or to sow the seed around a plant of the same species and allow the seedlings to grow on until they are large enough to move. This species only rarely forms new offsets and so division is seldom feasible, the following methods can be tried, however  .
Division of the tubers as the flowers fade  . This species produces a new tuber towards the end of its growing season. If this is removed from the plant as its flowers are fading, the shock to the plant can stimulate new tubers to be formed. The tuber should be treated as being dormant, whilst the remaining plant should be encouraged to continue in growth in order to give it time to produce new tubers  .
Division can also be carried out when the plant has a fully developed rosette of leaves but before it comes into flower  . The entire new growth is removed from the old tuber from which it has arisen and is potted up, the cut being made towards the bottom of the stem but leaving one or two roots still attached to the old tuber. This can often be done without digging up the plant. The old tuber should develop one or two new growths, whilst the new rosette should continue in growth and flower normally  .
The flower is pollinated by a solitary male bee of the Genus Andrena. [RC] In Malta the species found is specifically Andrena carbonaria (Black Mining Bee)  . The male pollinating bee does not enter the plant head first but goes in the other way round, that is, abdomen first, in an attempt to copulate with the plant. This abdominal pollination by pseudocopulation is a characteristic of O. fusca and O. lutea only. In fact these two species (or species complexes) are sometimes collectively referred to as the Pseudophrys group. O. lutea is also found in the Maltese islands where it is rather rare. [RC]
List of synonyms
Below is a list of synonyms of Ophrys fusca as taken from reference [WWW-141] . Note that some synonym names have been introduced recently (1998-2002) indicating that some have split the Ophrys fusca complex into individual species or subspecies. [SM]
- Arachnites fusca Tod. 1893;
- Ophrys africana G.Foelsche & W.Foelsche 2001;
- Ophrys arnoldii P.Delforge 1999;
- Ophrys blitopertha Paulus & Gack 1998;
- Ophrys cinereophila Paulus & Gack 1998;
- Ophrys creberrima Paulus 1998;
- Ophrys cressa Paulus 1998;
- Ophrys creticola Paulus 1998;
- Ophrys ficuzzana H.Baumann & K�nkele 1986;
- Ophrys funcrea Viviani 1824;
- Ophrys fusca subsp. akhdarensis B.Baumann & H.Baumann 2001;
- Ophrys fusca subsp. blitopertha (Paulus & Gack) Faurh. & H.A.Pedersen 2002;
- Ophrys fusca subsp. creberrima (Paulus) H.Kretzschmar 2002;
- Ophrys fusca subsp. cressa (Paulus) H.Kretzschmar 2002;
- Ophrys fusca subsp. creticol (Paulus) H.Kretzschmar 2002;
- Ophrys fusca subsp. thriptiensis (Paulus) H.Kretzschmar 2002;
- Ophrys fusca var. maculata Balayer 1986;
- Ophrys fusca var. rubescens Balayer 1986;
- Ophrys marmorata G.Foelsche & W.Foelsche 1998;
- Ophrys myodes Lapeyr. 1813;
- Ophrys nicotiae Zodda 1900;
- Ophrys obaesa Lojac. 1909;
- Ophrys obscura Beck 1879;
- Ophrys oestrifera Rchb. 1830;
- Ophrys parosica P.Delforge 1995;
- Ophrys pectus Mutel 1835;
- Ophrys peraiolae G.Foelsche & al. 2000;
- Ophrys peraiolae var. rubra G.Foelsche & al. 2000;
- Ophrys phaseliana D.R�ckbr. & U.R�ckbr. 1996;
- Ophrys sphegodes subsp. moesziana So� 1927;
- Ophrys sulcata Devillers-Tersch. & Devillers 1994;
- Ophrys tetraloniae W.P.Teschner 1987;
- Ophrys thriptiensis Paulus 1998;
- Ophrys tricolor Desf. ex Nyman 1882;
- Ophrys truncata Dulac 1867;
- Ophrys zonata Devillers-Tersch. & Devillers 1994
Ophrys fusca- / iricolor- complex in Malta
The plants of the general species name 'Ophrys fusca' found in Malta are believed to be of the Ophrys fusca- / iricolor- complex aggregate, which means some are part of the A1: Ophrys iricolor complex and some of the A2: Ophrys fusca complex group.
The A1 Ophrys iricolor group is made up of 5 species, 3 of them are of oriental distribution, the forth one in Corsica, Sardinia and Tunisia, and the fifth one endemic to Tunisia. Some are quite narrow endemics. [WWW-143] 2 of these are believed to be found in Malta. 
A1-1. O. astypalaeica - (Not reported in Malta, endemic to the island of Astypalea in the Cyclades)
A1-2. O. iricolor - (Unlikely to be present in Malta. Rather similar to O. eleonorae)
A1-3. O. mesaritica - (Believed to be endemic to Crete, but now its presence is also confirmed in Malta)
A1-4. O. vallesiana -(Endemic to Tunisia, some reports say it is found in Malta)
A1-5. O. eleonorae - ( Present in Malta. Can be easily confused with O. iricolor)
Species of the Ophrys fusca complex.
This complex or aggregate is made up of 26 species which are closely related to each other and previously they where referred to as Ophrys fusca s.l. The main charactaristic of this group is that of having an acute angle of about 28-45 degrees between the margin of the labellum and the longitudinal axis. Also, they do not have plataeus at the rim near the mouth (as in the Iricolor complex). Species with wider angles are placed in the sub-fusca group. The Ophrys fusca complex is mainly of occidental distribution, with species endemic to Crete, Malta-Sicily, Corsicas and Tunisia. [WWW-147]
- Ophrys arnoldii P. Delforge
- Ophrys attaviria D. R�ckbrodt & Wenker
- Ophrys bilunulata Risso (Distribution is still poorly known, may be found in Malta)
- Ophrys caesiella P. Delforge (The species was originally discovered on Malta but also found later in the south of Sicily)
- Ophrys creberrima H. F. Paulus
- Ophrys cressa H. F. Paulus (This species appears to be endemic to the east of Minos' island.)
- Ophrys creticola H. F. Paulus (This quite rare species appears to be endemic to some Cretan mountains)
- Ophrys delforgei P. Devillers & J. Devillers - Terschuren (This is the "O. fusca" form with the smallest flowers in continental France)
- Ophrys eptapigiensis H.F. Paulus
- Ophrys forestieri (Reichenbach Fil.) Lojacono
- Ophrys funerea Viviani
- Ophrys fusca Link (First described in Portugal, but most probably grows in many places of the western Mediterranean basin. Found in Malta)
- Ophrys hespera J. Devillers-Terschuren & P. Devillers
- Ophrys leucadica Renz
- Ophrys lucifera J. Devillers - Terschuren & P. Devillers (It grows in Italy, Tuscany, Sicily and maybe in other places of the central Mediterranean basin. Its distribution area, like that of O. fusca is not yet well understood.)
- Ophrys lupercalis J. Devillers-Terschuren & P. Devillers (This is a very early flowering plant in the South of France. It (or similar species) appears to be found in Malta)
- Ophrys marmorata G. Foelsche & W. Foelsche
- Ophrys obaesa Lojacono
- Ophrys pallid a Rafinesque
- Ophrys parvula H. F. Paulus
- Ophrys peraiolae Foelsche W., Foelsche G., Gerbaud O. & Gerbaud M. (This species appears to be endemic to a very small region close to the cove of Peraiola on the northeastern coast of Corsica)
- Ophrys perpusilla J. Devillers - Terschuren & P. Devillers
- Ophrys sabulosa H. F. Paulus & Gack ex P. Delforge
- Ophrys sulcata J. Devillers-Terschuren & P. Devillers
- Ophrys thriptiensis H. F. Paulus
- Ophrys zonata J. Devillers - Terschuren & P. Devillers
The Ophrys fusca species that are 'officially' reported to be found in Malta
In one report  about the Ophrys fusca related species found in the Maltese islands written in 2001 by 4 botanists ( G.Bartolo, S.Pulvirenti, E.Lanfranco, D.Stevens), the following 5 species are given:
Ophrys fusca Link,
O. caesiella P. Delforge,
O. parosica, P.Delforge,
O. pectus Mutel,
O. mesaritica P. Delforge (Iricolor group)
N.B. Ophrys pectus is a hybrid of O. mesaritica (A1: Ophrys iricolor complex group) x Ophrys bilunulata (A2: Ophrys fusca complex group) [WWW-142]
Finally, according to the Belgian botanist Delforge, who dedicated much work on the Ophrys group and visited Malta during the end of February, indicated that the following species are found:
O. mesaritica P. Delforge
O. lupercalis J. Devillers-Terschuren & P. Devillers
O. pectus Mutel
O. fusca Link (possibly)
But during his stay in Malta, Delforge did not encounter O. parosica (fusca complex), or O. eleonorae from the Iricolor complex. .
The Ophrys fusca- / lutea- complex in Sicily.
There is a big number of Pseudophrys species in Sicily, i.e. abdominal pollinated Ophrys fusca s.l. and lutea s.l., of which most can be distinguished easily. However, one should look at the whole population, and be careful picking out of individually deviating stems, since there are individual 'outliers' in size or shape of the bloom in every bigger Ophrys fusca s.l. population. In Sicily there are also many sites where several species are occurring simultaneously or successively. Not yet all Sicilian Pseudophrys have been described as species, so some have still to be named provisionally according to the form of the labellum, the pollinating bee, or an important site. [WWW-142]
Ophrys mesaritica is an early flowering plant - initially believed to be endemic to Crete, the place where it was found by Kretzschmar & al. (2002: 194). [WWW-148] However Pierre Delforge confirmed its occurrence also in Malta. [368, WWW-148] . The key identification features of this Ophrys are:
1) It is part of the Ophrys iricolor complex, distinguished by having characteristic elevated ridges at the stigmatic cavity (inner end of the lip).
2) Labellum between 11-16mm long.
3) The earliest Ophrys to flower, found with flowers from mid-December to February.
Ophrys eleonorae starts flowering in March and is a fairly large plant with broad leaves and tall scape. The key identification features of this Ophrys are:
1) It is part of the Ophrys iricolor complex, distinguished by having characteristic elevated ridges at the stigmatic cavity (inner end of the lip).
2) Labellum between 17-23mm long.
3) Red coloration at the underside of the labellum which does not reach the edges.
Ophrys caesiella starts flowering in mid February and is a smallish plant with broad leaves and rather short scape (8-12cm c.). The key identification features of this Ophrys are:
1) It is part of the Ophrys fusca complex, distinguished by having the angle between the outer margin of labellum and longitudinal axis less from 45 degrees. Furthermore, it lacks characteristic elevated ridges at the stigmatic cavity (inner end of the lip).
2) Labellum very small - between 9-12mm long.
3) Labellum with a well defined yellow margin
4) No hairs at the furrow at the middle of the speculum.
Comparative structure of the labellum in Ophrys fusca and Ophrys lutea
The morphology and anatomy of the labellar epidermal cells and the way in which they are arranged are described in an attempt to locate and characterize the osmophore in O. fusca and O. lutea. The micromorphology of the labellum of these two species is similar. Four types of epidermal cells are present on the adaxial surface of the labellum. Long unicellular trichomes with straight tips cover the basal region of the labellum, whereas short unicellular trichomes with polygonal flattened bases form the reflective median speculum. The apical region of the labellum possesses a villous indumentum of long acuminate trichomes with bent or sinuate tips. Large smooth-walled, dome-shaped papillae occur on the margins and on the distal region of the abaxial surface of the labellum. These remarkable papillae have high polarity; the protoplasm at the apex of each cell contains several small vacuoles, while a prominent nucleus surrounded by numerous hypertrophied amyloplasts occurs at the opposite end of the cell. Positive reactions to Vogel's staining test and to Sudan black B enabled us to conclude that the osmophores of both species are composed of these peculiar secretory epidermal cells and by two or three subsecretory layers of parenchyma cells. 
Ophrys fusca-iricolor complex group of Malta
Ophrys fusca show a large degree of variability mainly in its labellum, but also in its flowering period and dimensions. This results in flowers which show a variable range of morphological differences (namely the labellum) from one plant to another. In some extremes these would look as quite different species.
In the recent 2 decades or so, there was an interest to split the general Ophrys fusca taxon into singular species according to their slight morphological differences. This resulted in a vast number of species names, sometimes being synonyms of each other being named by different authors.
For the sake of trying to keep things simple and organised, groups or complexes were formed where each has an aggregate of Ophrys species with related characteristics, either morphological or depending upon the pollinator insect.
Apparently, the 'Ophrys fusca' species that are found in Malta come from 2 complex groups, that are: Ophrys iricolor complex and Ophrys fusca complex - sometimes referred to as the Ophrys fusca-/iricolor- complex group. In Sicily there is prevalence of Ophrys fusca-lutea complex aggregate.
These newly split names are somehow ambiguous and maybe, one should be safer to stick to this large group of related species simply as Ophrys fusca s.l.
The differences between one species and another are very minimal in several cases. This is so true that sometimes authors do not agree between themselves, and two names given by different authors may be referring to the same plant. An in-depth study would require genetic mapping and DNA sequencing. [SM]
Pollination of orchids has always been so particular and fascinating. Each species would usually have a restricted group of pollinators and in several cases just one species. The flower mimics in its shape, size and pheromones production the female gender of its pollinator species. This will in turn attract the male and fool him as if the flower was the female partner. The pheromones (sex hormones) produced by the flower, that are nearly identical to those emitted by the female insect, play an important role in attracting the male pollinator
The male sits on the flower's labellum and tries with all his efforts to copulate. The movement of the abdomen or thorax of the pollinator would eventually touch the male reproductive organs of the flower which are specialized in structure and physiology. They are detachable organs known as pollinia. They are located in longitudinal pockets at the roof of the flower's mouth. At the base of each pollinium there is a very sticky surface called the viscidium. This sticks strongly with the insect's body while he is in his copulating action. When the insect flies away, the entire pollinium is detached from the flower and carried away by the pollinator.
If the pollinator lands on another compatible Orchid flower, again to try to copulate, the pollinium would eventually fall off and ends up in the stigmatic cavity at the floor of the flower's mouth, just at the inner end of the labellum. Inside this cavity there are the female recipient organs. The clusters of pollen of the pollinium will finally come in contact with the stigma and so fertilize the flower. Pollination is now complete. [SM]
Ophrys fusca prefer to grow in non-weedy soil patches over garigue. It is rarely found growing if there is Oxalis pes-caprae, Bellis annua or high herbaceous vegetation. On the other hand it does not mind Thymbra capitata and Erica multiflora and one can find Ophrys fusca growing beside these shrubs. Another point is that this plant usually grows in scattered communities, so if one finds a plant, it is common to find several other plants in that area. [SM]
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