Ophrys Apifera
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''Ophrys apifera'', known in Europe as the bee orchid, is a
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
of the genus ''
Ophrys The genus ''Ophrys'' is a large group of orchids from the alliance Orchis in the subtribe Orchidinae. They are widespread across much of Europe, North Africa, Caucasus, the Canary Islands, and the Middle East as far east as Turkmenistan. These ...
'', in the family of
Orchidaceae Orchids are plants that belong to the family (biology), family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan distribution, cosmopolitan plants that ...
. It serves as an example of sexually deceptive pollination and floral mimicry, a highly selective and highly evolved plant–pollinator relationship.


Description

''Ophrys apifera'' grows to a height of . This hardy orchid develops small rosettes of
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
in autumn that continue to grow slowly during winter. Basal leaves are ovate or oblong-lanceolate, and upper leaves and bracts are ovate-lanceolate and sheathing. Leaves exhibit parallel venation. The plant blooms from mid-April in continental Europe, but in the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
it flowers June to July. A flower spike is produced, composed from one to twelve flowers. Three large, purple
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s surround the base of the flower, which can easily be mistaken for
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s. The true petals lie just above the sepals as two short, pubescent green structures protruding laterally from a central column. A third, modified petal, the labellum, sits at the bottom of the column as a landing pad for pollinators. The labellum is trilobed, with two pronounced humps on the hairy lateral lobes and a hairy median lobe having a pattern that mimics the abdomen of a
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
.Ray, H. & Wagner, V. (2015). “Orchid pollination biology”. Environmental Horticulture Department, UF/IFAS Extension (ENH1260). https://edis.ifas.ufl.edu/pdffiles/EP/EP52100.pdf The pattern of labellum colouration is quite variable. The central column is an adaptation unique to orchids, in which the stamens and pistil fused together to form one central rib. The anther cap can be found at the top, dorsal side of the column, while the stigma is hidden below on the ventral side. Two
pollinia A pollinium (: pollinia) is a coherent mass of pollen grains in a plant that are the product of only one anther, but are transferred, during pollination, as a single unit. This is regularly seen in plants such as orchids and many species of milkwee ...
hang from the column over the labellum.


Distribution

''Ophrys apifera'' is widespread across central and southern Europe, as well as North Africa and the Middle East. Its range stretches from
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
,
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
and
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
east to
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
and the
Caucasus The Caucasus () or Caucasia (), is a region spanning Eastern Europe and Western Asia. It is situated between the Black Sea and the Caspian Sea, comprising parts of Southern Russia, Georgia, Armenia, and Azerbaijan. The Caucasus Mountains, i ...
. It is quite common in the Mediterranean region eastwards to the
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
, (Codes) but is less common in its northern range being uncommon or local in
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
and Ireland. In the United Kingdom, it has a distinct southeastern preference, being more common in England. Recently it has been found in the southwest of England in Butleigh near
Glastonbury Glastonbury ( , ) is a town and civil parish in Somerset, England, situated at a dry point on the low-lying Somerset Levels, south of Bristol. The town had a population of 8,932 in the 2011 census. Glastonbury is less than across the River ...
in
Somerset Somerset ( , ), Archaism, archaically Somersetshire ( , , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by the Bristol Channel, Gloucestershire, and Bristol to the north, Wiltshire to the east ...
and Dorchester, Dorset; whereas it is only to be found in coastal regions of
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
as well as the Hodbarrow Nature Reserve in
Millom Millom is a town and civil parish on the north shore of the estuary of the River Duddon in southernmost Cumberland, Cumbria, England. It is situated just outside the Lake District National Park, about north of Barrow-in-Furness ( by road) and ...
, Cumbria, and some parts of
Northern Ireland Northern Ireland ( ; ) is a Countries of the United Kingdom, part of the United Kingdom in the north-east of the island of Ireland. It has been #Descriptions, variously described as a country, province or region. Northern Ireland shares Repub ...
. It is relatively common in the northeast of England and in recent years large numbers have appeared in the grass verges surrounding the Metro Centre in
Gateshead Gateshead () is a town in the Gateshead Metropolitan Borough of Tyne and Wear, England. It is on the River Tyne's southern bank. The town's attractions include the twenty metre tall Angel of the North sculpture on the town's southern outskirts, ...
. In
Scotland Scotland is a Countries of the United Kingdom, country that is part of the United Kingdom. It contains nearly one-third of the United Kingdom's land area, consisting of the northern part of the island of Great Britain and more than 790 adjac ...
, it was thought to be extinct, but was rediscovered in
Ayrshire Ayrshire (, ) is a Counties of Scotland, historic county and registration county, in south-west Scotland, located on the shores of the Firth of Clyde. The lieutenancy areas of Scotland, lieutenancy area of Ayrshire and Arran covers the entirety ...
in 2003. In some countries the plants have protected status. They are unusual in that in some years they appear in great numbers, then sometimes only reappear after an absence of many years. The genus ''Ophrys'' is the most species-rich (i.e. diverse) genus of orchids in Europe and the Mediterranean with over 200 species, according to ''Orchids of Britain and Europe'' by Pierre Delforge.


Habitat and ecology

''Ophrys apifera'' generally grows on semi-dry
turf Sod is the upper layer of turf that is harvested for transplanting. Turf consists of a variable thickness of a soil medium that supports a community of turfgrasses. In British and Australian English, sod is more commonly known as ''turf'', ...
, in
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
, on limestone, calcareous dunes or in open areas in woodland. It prefers well-drained calcareous soils, low in nutrients, in bright light or dim light.PhD Thesis of the University of Liverpool – Diversity and Roles of Mycorrhizal Fungi in the Bee Orchid Ophrys Apifera
/ref> It is a major colonizer of sites disturbed by human activity, such as old quarries, roadside verges and airfields. ''O. apifera'' is one of the most likely European orchid species to establish itself within towns and cities. In order to extract sufficient nutrients from the substrata it grows in, ''Ophrys apifera'' relies upon a symbiotic relationship with
mycorrhizal fungi A mycorrhiza (; , mycorrhiza, or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...
in the genus ''
Tulasnella ''Tulasnella'' is a genus of effused (patch-forming) fungi in the order Cantharellales. Basidiocarps (fruit bodies), when visible, are typically smooth, ceraceous (waxy) to subgelatinous, frequently lilaceous to violet-grey, and formed on the und ...
'', and possibly other genera. Bee orchids are threatened by mowing during flowering, or before the seed has been released. However, they often also disappear from sites that become overgrown with shrubs and/or trees, as the orchids fail to compete with these large plants for light. For these reasons, bee orchids are often found on the edge of mown areas, beside paths or within areas that are mown very infrequently. The Sussex Wildlife Trust recommends mowing at the end of July and removing the cuttings to benefit bee orchids. In prehistory, the species presumably relied upon seasonal grazing pressure, or was limited to early succession habitats and permanent grassland.


Reproduction

''Ophrys apifera'' has been considered to preferentially practice
self-pollination Self-pollination is a form of pollination in which pollen arrives at the stigma of a flower (in flowering plants) or at the ovule (in gymnosperms) of the same plant. The term cross-pollination is used for the opposite case, where pollen from ...
. The flowers are almost exclusively self-pollinating in the northern ranges of the plant's distribution, however pollination by the
solitary bee Solitary is the state of being alone or in solitude. The term may refer to: * Solitary (album), ''Solitary'' (album), 2008 album by Don Dokken * Solitary (2020 film), ''Solitary'' (2020 film), a British sci-fi thriller film * Solitary (upcoming fi ...
''
Eucera longicornis ''Eucera longicornis'' is a species of bee in the family Apidae, subfamily Apinae, and tribe Eucerini, the long-horned bees. Description The head, thorax and abdomen of this species are black, the thorax being clad in pale brown, downy hairs on ...
'' occurs in the Mediterranean region, where ''Ophrys apifera'' is more common. ''E. longicornis'' males have been observed attempting to copulate with the flowers, which emit
allomone An allomone (from Ancient Greek ' "other" and pheromone) is a type of semiochemical produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the rec ...
s that mimic the scent of the female bee. These allomones are also known to attract bee species '' Tetralonia cressa'' and '' Eucera pulveraceae''. In addition to chemosensory mimicry, the labellum of the flower acts as a visual decoy that the male bee confuses for a female. It is believed that male bees preferentially select orchids with the most bee-like labellum and attempt copulation, at which point the pollinia stick to the bee during the
pseudocopulation Pseudocopulation is a behavior similar to Copulation (zoology), copulation that serves a reproductive function for one or both participants but does not involve actual sexual union between the individuals. It is most generally applied to a pollin ...
. This achieves pollen transfer and, potentially, pollination. ''Ophrys'' like other orchids are dependent on
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
at some point during their life cycle, but especially for
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ...
, which may take months or even years underground. Orchid roots contain
orchid mycorrhiza Orchid mycorrhizae are endomycorrhizal fungi which develop symbiotic relationships with the roots and seeds of plants of the family Orchidaceae. Nearly all orchids are myco-heterotrophic at some point in their life cycle. Orchid mycorrhizae are ...
, coils of fungal hyphae inside orchid root cells.


Phytochemistry

Floral parts display the presence of
quercetin Quercetin is a plant flavonol from the flavonoid group of polyphenols. It is found in many fruits, vegetables, leaves, seeds, and grains; capers, red onions, and kale are common foods containing appreciable amounts of it. It has a bitter flavor ...
and
kaempferol Kaempferol (3,4′,5,7-tetrahydroxyflavone) is a natural flavonol, a type of flavonoid, found in a variety of plants and plant-derived foods including kale, beans, tea, spinach, and broccoli. It is also found in propolis extracts. Kaempferol i ...
glycosides, which are supposed to be acylated, as well as cinnamic acid derivatives. The pink outer tepals show the presence of anthocyanins.


Culture

The name ''Ophrys'' derives from the Greek word ', meaning "eyebrow." It is believed that women in ancient Rome used to darken their eyebrows with the orchid plant, thus giving it its name.Addam, K., Takkoush, J., Bou-hamdan, M., & Itani, J. (2015). Five established orchids Ophrys apifera var. Chlorantha, Aurita, Purpurea, Purpurea. F. Alba. and Flavescens (Orchidaceae) in Lebanon as part of the native flora. Journal of Ecology and Environmental Sciences, 6(2), 163–169. ISSN: 0976-9900 & E-ISSN: 0976-9919 The Latin
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
' means "bee-bearing" or "bee-bringing"WORDS Latin-to-English Dictionary by William Whittaker, AbleMedia Classics Technology Center
accessed 2014-11-13
and refers to the bee-shaped labellum of the orchid. Its root tubers were ground into a powder called
salep Salep, also spelled sahlep salepi or sahlab,; ; ; ; ; ; ; ; ; ; is a flour made from the tubers of the orchid genus ''Orchis'' (including species '' Orchis mascula'' and ''Orchis militaris''). These tubers contain a nutritious, starchy polysacc ...
in the Ottoman Empire, which was believed to have nutritional, medicinal, and aphrodisiac values.Kasparek, M. & Grimm, U. (1999). European trade in Turkish salep with special reference to Germany. Economic Botony, 53(4), 396–406. The product's popularity spread to England in the 17th and 18th century and then to a few other European countries. Salep is still used today in medicines and traditional dishes in Turkey, which was formerly the region of the Ottoman Empire, and throughout the Middle East. It is also a popular import in the Netherlands, Cyprus, and Germany for its supposed medicinal properties. ''Ophrys apifera'' is the
county flower In 2002 Plantlife conducted a "County Flowers" public survey to assign flowers to each of the counties of the United Kingdom and the Isle of Man. The results of this campaign designated a single plant species to a "county or metropolitan area" in ...
of
Bedfordshire Bedfordshire (; abbreviated ''Beds'') is a Ceremonial County, ceremonial county in the East of England. It is bordered by Northamptonshire to the north, Cambridgeshire to the north-east, Hertfordshire to the south and the south-east, and Buckin ...
. This species has appeared on postage stamps in many countries including
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
,
Cambodia Cambodia, officially the Kingdom of Cambodia, is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. It is bordered by Thailand to the northwest, Laos to the north, and Vietnam to the east, and has a coastline ...
,
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
,
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
,
Liechtenstein Liechtenstein (, ; ; ), officially the Principality of Liechtenstein ( ), is a Landlocked country#Doubly landlocked, doubly landlocked Swiss Standard German, German-speaking microstate in the Central European Alps, between Austria in the east ...
,
Luxembourg Luxembourg, officially the Grand Duchy of Luxembourg, is a landlocked country in Western Europe. It is bordered by Belgium to the west and north, Germany to the east, and France on the south. Its capital and most populous city, Luxembour ...
,
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
, the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
and the
USSR The Union of Soviet Socialist Republics. (USSR), commonly known as the Soviet Union, was a List of former transcontinental countries#Since 1700, transcontinental country that spanned much of Eurasia from 1922 until Dissolution of the Soviet ...
. In 2009 the bee orchid appeared in a series of stamps printed in the United Kingdom to celebrate the 150th anniversary of the publication of
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
's
Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life'')The book's full original title was ''On the Origin of Species by M ...
.


Conservation


Ecological concerns

There are local extinctions of ''Ophrys spp''. and potentially ''O. apifera'' in Turkey and Iran due to the excessive harvesting of wild orchids to produce salep products. Since ''O. apifera'' relies heavily on its symbiotic relationship with ''Tulasnella'' fungi for survival, it is vulnerable to any chemicals, particularly fungicides, and threats which might reduce the fungi's prevalence.


Species status

Protected species status in Northern Ireland. Endangered (EN) species status in Turkey, Ukraine, and Carpathian Mountains.Osiadacz, B. & Kreciala, M. (2014). Ophrys apifera Huds. (Orchidaceae), a new orchid species to the flora of Poland. Biodiversity Research and Conservation, 36, 11–16. Sandal Erzurumlu, G., Tuluhan Yilmaz, K., & Sogut, Z. (2017). A regional scale evaluation of conservation status of orchid species recorded in the eastern Mediterranean region of Turkey. Turkish Journal of Agriculture– Food Science and Technology, 5(9), 996–1001. Critically Endangered (CR) species status in Czech Republic and Slovakia.


Laws/regulations

The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) named ''O. apifera'' an endangered species that is potentially an object of trade, setting strict regulations on trade of salep products. Turkey, Iran, Germany, the Netherlands, and Cyprus are all Parties to CITES and, in theory, must follow those regulations. Turkey has banned the export of true salep (salep made from an orchid root).


Taxonomy

The specific evolutionary relationships between members of this genus are still poorly understood, and the best method for classifying orchid species and their variants is highly debated. Some researchers propose classifying species based on their pollination tactics, while others propose classifying them based on the type of chemical emitted from the flower. A widely used and generally accepted method of classification distinguishes orchid species based on flower morphology. The taxonomy of ''O. apifera'' based on this classification scheme is as follows: Kingdom Plantae, Phylum Tracheophyta, Subphylum Angiospermae, Class Liliopsida, Order Asparagales, Family Orchidaceae, Subfamily Orchidoideae, Genus ''Ophyrus'', Species ''O. apifera''.


Varieties

* ''Ophrys apifera'' var. ''apifera'' * ''Ophrys apifera'' var. ''aurita'' * ''Ophrys apifera'' var. ''bicolor'' * ''Ophrys apifera'' var. ''botteronii'' * ''Ophrys apifera'' var. ''friburgensis'' * ''Ophrys apifera'' var. ''immaculata'' * ''Ophrys apifera'' var. ''trollii'' * ''Ophrys apifera'' var. ''fulvofusca'' * ''Ophrys apifera'' var. ''belgarum'' * ''Ophrys apifera'' var. ''jurana'' * ''Ophrys apifera'' var. ''atrofuscus''


References


External links

* *
Biolib

Ophrys apifera

Ophrys apifera , Plants of the World Online


{{Taxonbar, from=Q159291 apifera Orchids of Europe Flora of North Africa Flora of the Caucasus Flora of Western Asia Plants described in 1762 Taxa named by William Hudson (botanist)