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''Ponerorchis'' is a genus of Asian terrestrial tuberous orchids (family
Orchidaceae Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...
). It is native to temperate Eurasia, from Poland to Japan, to the north of the
Indian subcontinent The Indian subcontinent is a list of the physiographic regions of the world, physiographical region in United Nations geoscheme for Asia#Southern Asia, Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian O ...
and to northern
Indochina Mainland Southeast Asia, also known as the Indochinese Peninsula or Indochina, is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to the west an ...
.Flora of China v 25 p 92, 小红门兰属 xiao hong men lan shu, ''Ponerorchis''
/ref> They can be found in evergreen forests and meadows in temperate, mountainous regions. The species ''Ponerorchis chidorii'', endemic to Japan, is also known to grow
epiphytically An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
. In the horticultural trade, this genus is abbreviated as "Pnr". A wide range of mostly
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
s of ''Ponerorchis graminifolia'' are in cultivation, but are rarely grown outside Japan.


Description

''Ponerorchis'' species grow from an ovoid
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
. They are slender plants with one to three usually slightly fleshy leaves. The flowers are all borne on the same side of the stem. The upper
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 The term ''sepalum'' was coined b ...
and the two lateral
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s form a hood. The
column A column or pillar in architecture and structural engineering is a structural element that transmits, through compression, the weight of the structure above to other structural elements below. In other words, a column is a compression member. ...
is short and has obvious side appendages. There is a
rostellum The rostellum is a projecting part of the column in Orchidaceae flowers, and separates the male androecium from the female gynoecium, commonly preventing self-fertilisation. In many orchids, such as ''Orchis mascula'', the pollinia or pollen masse ...
separating the
pollinia A pollinium (plural 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 mil ...
from the stigma. The genus was substantially expanded in 2014, so older descriptions do not necessarily apply to the revised genus. ''Ponerorchis'' was said to be differentiated from ''Amitostigma'' and ''Neottianthe'' by the presence of a bursicle or "sac" around the viscidium (a sticky pad at the base of the pollinium). However, this feature was found to be present in some species formerly assigned to ''Amitostigma'' and to be difficult to detect in some species of ''Ponerorchis''. In the expanded genus, a bursicle is present is some species and not in others.


Taxonomy


Phylogeny and classification

The genus ''Ponerorchis'' was erected by
Heinrich Gustav Reichenbach Heinrich Gustav Reichenbach (Dresden, 3 January 1823 – Hamburg, 6 May 1889) was a botanist and the foremost German orchidologist of the 19th century. His father Heinrich Gottlieb Ludwig Reichenbach (author of ''Icones Florae Germanicae et Helve ...
in 1852 for the species '' Ponerorchis graminifolia''.
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies between 1997 and 2003 showed that ''Ponerorchis'', when broadly defined, formed a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
with the genera ''Amitostigma'', ''Neottianthe'' and ''
Hemipilia ''Hemipilia'' is a genus of plants in the family Orchidaceae. It is native to China, the Himalayas and Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia ...
''. ''Ponerorchis'', as then
circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...
, was not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. In 2003, in advance of further studies involving more species, ''Ponerorchis'' was broadened to include ''Chusua'', and an "orphan" species was transferred from ''Ponerorchis'' to become '' Hemipilia brevicalcarata''. A more detailed
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study in 2014 confirmed the strong relationship between ''Amitostigma'', ''Neottianthe'' and ''Ponerorchis'', whose species were mixed together in a single
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, making none of the three genera
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
as then
circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...
. ''Amitostigma'' and ''Neottianthe'' were subsumed into ''Ponerorchis'', making it a considerably larger genus. ''Ponerorchis'' was previously distinguished from related genera by the presence of bursicles (pouchlike receptacles) enclosing the
pollinia A pollinium (plural 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 mil ...
. The presence or absence of a bursicle was found not to be a useful diagnostic character, having arisen independently at least four times in the Orchideae. ''Ponerorchis'' is placed in the orchid subfamily
Orchidoideae The Orchidoideae, or the orchidoid orchids, are a subfamily of the orchid family (Orchidaceae) that contains around 3630 species. Species typically have a single (monandrous), fertile anther which is erect and basitonic. Description The subfa ...
, tribe
Orchideae Orchideae is a tribe of orchids in the subfamily Orchidoideae. Historically, it was divided into 2 subtribes, Orchidinae and Habenariinae. The subtribe Orchidinae alone contains about 1,800 species. However, although some phylogenetic studies hav ...
, subtribe Orchidineae. After the genera were re-circumscribed as explained above, the relationship between ''Ponerorchis'' and the rest of the subtribe is as shown in the
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
below.


Species

Species accepted as of March 2018: * '' Ponerorchis alpestris'' (Fukuy.) X.H.Jin, Schuit. & W.T.Jin – N. & Central Taiwan * '' Ponerorchis amplexifolia'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan) * '' Ponerorchis basifoliata'' (Finet) X.H.Jin, Schuit. & W.T.Jin – China (S.E. Sichuan, N. Yunnan) * '' Ponerorchis bidupensis'' (Aver.) X.H.Jin, Schuit. & W.T.Jin – S. Vietnam * '' Ponerorchis bifoliata'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – China (N. Sichuan, S. Gansu) * '' Ponerorchis brevicalcarata'' (Finet) Soó – China (S.W. Sichuan to N.W. Yunnan) * '' Ponerorchis camptoceras'' (Rolfe ex Hemsl.) X.H.Jin, Schuit. & W.T.Jin – China (W. & S.W. Sichuan) * '' Ponerorchis capitata'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – China (Sichuan, W. Hubei) * '' Ponerorchis chidorii'' (Makino) Ohwi – Japan * '' Ponerorchis chusua'' (D.Don) Soó – Siberia to Himalaya * '' Ponerorchis compacta'' (Schltr.) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan) * '' Ponerorchis crenulata'' Soó – China (N.W. Yunnan) * '' Ponerorchis cucullata'' (L.) X.H.Jin, Schuit. & W.T.Jin – Poland to Japan and Himalaya * '' Ponerorchis dolichocentra'' (Tang, F.T.Wang & K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan) * '' Ponerorchis exilis'' (Ames & Schltr.) S.C.Chen – China (Central & N.E. Yunnan) * '' Ponerorchis faberi'' (Rolfe) X.H.Jin, Schuit. & W.T.Jin – China (Sichuan, N.W. Yunnan, N.E. Guizhou) * '' Ponerorchis farreri'' (Schltr.) X.H.Jin, Schuit. & W.T.Jin – S.E. Tibet to China (N.W. Yunnan) * '' Ponerorchis fujisanensis'' (K.Inoue) J.M.H.Shaw – N. & Central Japan * '' Ponerorchis gonggashanica'' (K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (S.W. Sichuan) * '' Ponerorchis gracilis'' (Blume) X.H.Jin, Schuit. & W.T.Jin – China to Temp. E. Asia * '' Ponerorchis graminifolia'' Rchb.f. – Korea, S. Central & S. Japan * '' Ponerorchis hemipilioides'' (Finet) Soó – China (N.W. Yunnan, Central Guizhou) * '' Ponerorchis joo-iokiana'' (Makino) Nakai – N. Korea, Japan (Central Honshu) * '' Ponerorchis keiskei'' (Finet) X.H.Jin, Schuit. & W.T.Jin – Japan (W. Honshu. Shikoku) * '' Ponerorchis kinoshitae'' (Makino) X.H.Jin, Schuit. & W.T.Jin – S. Kuril Islands to Japan (Hokkaido, N. Honshu) * '' Ponerorchis kiraishiensis'' (Hayata) Ohwi – Taiwan * '' Ponerorchis lepida'' (Rchb.f.) X.H.Jin, Schuit. & W.T.Jin – Japan (S. Kyushu) to Central Nansei-shoto * '' Ponerorchis limprichtii'' (Schltr.) Soó – Central China * '' Ponerorchis luteola'' (K.Y.Lang & S.C.Chen) X.H.Jin, Schuit. & W.T.Jin – China (N.W. Yunnan) * '' Ponerorchis monantha'' (Finet) X.H.Jin, Schuit. & W.T.Jin – S.E. Tibet to Central China * '' Ponerorchis oblonga'' (K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (N.W. Yunnan) * '' Ponerorchis omeishanica'' (Tang, F.T.Wang & K.Y.Lang) S.C.Chen – China (Sichuan: Emei Shan) * '' Ponerorchis ovata'' (K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan) * '' Ponerorchis papilionacea'' (Tang, F.T.Wang & K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (N.W. Sichuan) * '' Ponerorchis parceflora'' (Finet) X.H.Jin, Schuit. & W.T.Jin – China (N.E. Sichuan, Chongqing) * '' Ponerorchis pathakiana'' (Av.Bhattacharjee) J.M.H.Shaw – Arunachal Pradesh * '' Ponerorchis physoceras'' (Schltr.) X.H.Jin, Schuit. & W.T.Jin – China (N.W. & W. Sichuan) * '' Ponerorchis puberula'' (King & Pantl.) Verm. – E. Himalaya * '' Ponerorchis pugeensis'' (K.Y.Lang) S.C.Chen, P.J.Cribb & S.W.Gale – China (S.W. Sichuan) * '' Ponerorchis renzii'' Deva & H.B.Naithani – W. Himalaya * '' Ponerorchis secundiflora'' (Kraenzl.) X.H.Jin, Schuit. & W.T.Jin – Himalaya to China (W. Sichuan, N.W. Yunnan) * '' Ponerorchis sichuanica'' (K.Y.Lang) S.C.Chen, P.J.Cribb & S.W.Gale – China (W. Sichuan) * '' Ponerorchis simplex'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan, N.W. Yunnan) * '' Ponerorchis taiwanensis'' (Fukuy.) Ohwi – Central & S. Taiwan * '' Ponerorchis takasago-montana'' (Masam.) Ohwi – Central & E. Taiwan * '' Ponerorchis tetraloba'' (Finet) X.H.Jin, Schuit. & W.T.Jin – China (S.W. Sichuan, W. Yunnan) * '' Ponerorchis thailandica'' (Seidenf. & Thaithong) X.H.Jin, Schuit. & W.T.Jin – N. Thailand * '' Ponerorchis tibetica'' (Schltr.) X.H.Jin, Schuit. & W.T.Jin – S.E. Tibet to China (N.W. Yunnan) * '' Ponerorchis tominagae'' (Hayata) H.J.Su & Jr J.Chen – Taiwan * '' Ponerorchis trifurcata'' (Tang, F.T.Wang & K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (N.W. Yunnan) * '' Ponerorchis wenshanensis'' (W.H.Chen, Y.M.Shui & K.Y.Lang) X.H.Jin, Schuit. & W.T.Jin – China (Yunnan) * '' Ponerorchis yueana'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – S.E. Tibet to China (N.W. Yunnan)


Distribution

''Ponerorchis'' species are widely distributed in Eurasia, mainly in temperate regions, but extending into northern Indo-China. In Europe, they are found in
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...
and
Eastern Europe Eastern Europe is a subregion of the Europe, European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural, and socio-economic connotations. The vast majority of the region is covered by Russ ...
. In temperate Asia, they extend from
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
to the
Russian Far East The Russian Far East (russian: Дальний Восток России, r=Dal'niy Vostok Rossii, p=ˈdalʲnʲɪj vɐˈstok rɐˈsʲiɪ) is a region in Northeast Asia. It is the easternmost part of Russia and the Asian continent; and is admini ...
,
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
and
Eastern Asia East Asia is the eastern region of Asia, which is defined in both Geography, geographical and culture, ethno-cultural terms. The modern State (polity), states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. ...
. They are also found in the north of the
Indian subcontinent The Indian subcontinent is a list of the physiographic regions of the world, physiographical region in United Nations geoscheme for Asia#Southern Asia, Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian O ...
(the western
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
,
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
, the eastern Himalayas and the Assam region) and in northern Indochina (
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
and
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
).


References


External links

* *
''Ponerorchis''
{{Taxonbar, from=Q2719981, from2=Q20842, from3=Q2900972, from4=Q1319684, from5=Q2417814 Orchideae genera Flora of Poland Flora of Eastern Europe Flora of Siberia Flora of the Russian Far East Flora of China Flora of Mongolia Flora of Eastern Asia Flora of Assam (region) Flora of East Himalaya Flora of Nepal Flora of West Himalaya Flora of Myanmar Flora of Thailand Flora of Vietnam