''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 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 in 1852 for the species ''

Ponerorchis graminifolia ''Ponerorchis graminifolia'' is a species of flowering plant in the family Orchidaceae, native to southern Korea and Japan. They are short herbaceous perennials, growing from a tuber, with small flowers in shades of pink to purple. Many varieties ...

''.

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

, making none of the three genera

as then

. ''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. 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, tribe Orchideae, 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 below.

Species

Species accepted as of March 2018: * ''

Ponerorchis alpestris ''Ponerorchis alpestris'' is a species of flowering plant in the family Orchidaceae, native to Taiwan. Taxonomy The species was first described in 1935 by Noriaki Fukuyama as ''Amitostigma alpestre''. A molecular phylogenetic study in 2014 found ...

'' (Fukuy.) X.H.Jin, Schuit. & W.T.Jin – N. & Central Taiwan * ''

Ponerorchis amplexifolia ''Ponerorchis amplexifolia'' is a species of flowering plant in the family Orchidaceae, native to south-central China (west Sichuan). Taxonomy The species was first described in 1936 by Tsin Tang and Fa Tsuan Wang as ''Amitostigma amplexifolium' ...

'' (Tang & F.T.Wang) X.H.Jin, Schuit. & W.T.Jin – China (W. Sichuan) * ''

Ponerorchis basifoliata ''Ponerorchis basifoliata'' is a species of flowering plant in the family Orchidaceae, native to south-east China (south-east Sichuan, north Yunnan). Taxonomy The species was first described in 1902 by Achille Eugène Finet, as ''Peristylus tetr ...

'' (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 * ''

'' 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

(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