Aquilegia amurensis: Difference between revisions
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{{Short description|Species of flowering plant}} |
{{Short description|Species of flowering plant}} |
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{{Speciesbox |
{{Speciesbox |
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| image = Aquilegia |
| image = Aquilegia amurensis.jpg |
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| image_caption = |
| image_caption = Preserved specimen of ''Aquilegia amurensis'' |
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| image2 = |
| image2 = |
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| image2_caption = |
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| genus = Aquilegia |
| genus = Aquilegia |
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| species = |
| species = amurensis |
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| authority = {{Abbr|Not. Syst. Herb. Hort. Bot.|Botanicheskie Materialy Gerbariia Glavnogo Botanicheskogo Sada SSSR}} |
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| authority = [[Paul Carpenter Standley|Standley]] ex. [[Edwin Blake Payson|Payson]] |
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| synonyms_ref = <ref name="POWO_272432-2">{{cite POWO |id= |
| synonyms_ref = <ref name="POWO_272432-2">{{cite POWO |id=708796-1 |title=''Aquilegia amurensis'' Kom. |access-date=December 6, 2024 }}</ref><ref name=Nold/>{{rp|64}} |
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| synonyms = ''Aquilegia |
| synonyms = ''Aquilegia flabellata'' {{small|Siebold & Zucc.}}<br/>''Aquilegia flabellata'' var. ''alpina'' {{small|{{Abbr|Izv. Imp. Akad. Nauk|Izvestiya Imperatorskoi Akademii Nauk}} Kuzen.}} |
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}} |
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'' |
'''''Aquilegia amurensis''''' is a partially accepted species of flowering plant in the genus ''[[Aquilegia]]'' (columbines) in the family [[Ranunculaceae]] that is native to the northeastern Asian. Its natural range is in the northern [[Greater Khingan]] mountain range and [[Amur River]] of China, as well as Siberian Russia, Mongolia, and possibly [[North Korea]].<ref name=Div>{{cite journal|title=Rapid Divergence Followed by Adaptation to Contrasting Ecological Niches of Two Closely Related Columbine Species ''Aquilegia japonica'' and ''A. oxysepala''|first1=Ming-Rui|last1=Li|first2=Hua-Ying|last2=Wang|first3=Ning Ding|last3=Tianyuan Lu|first4=Ye-Chao|last4=Huang|first5=Hong-Xing|last5=Xiao|first6=Bao|last6=Liu|first7=Lin-Feng|last7=Li|journal=[[Genome Biology and Evolution]]|date=March 2019|page=920|volume=11|issue=3}}</ref><ref name=Mongolia/><ref name=Nold>{{cite book|title=Columbines: Aquilegia, Paraquilegia, and Semiaquilegia|first=Robert|last=Nold|publisher=Timber Press|date=2003|location=[[Portland, Oregon|Portland]], [[Oregon|OR]]|isbn=0881925888}}</ref>{{rp|64}} Flowers of this plant have petal blades that are whitish or white-tipped, with blue-violet [[nectar spur]]s and [[sepal]]s. The plant is rarely cultivated. |
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==Description== |
==Description== |
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''Aquilegia amurensis'' grows to be about {{convert|50|cm|in}} tall. It possesses leaves in a [[Phyllotaxis|basal arrangement]] (sprouting from base of the shoot), seldom if ever with leaves on the stem. The [[Leaflet (botany)|leaflets]] are [[glabrous]] (smooth) on their topsides with soft, downy bottoms.<ref name=Nold/>{{rp|64}} The leaflets are trisected to their base. The [[petiole (botany)|petiole]]s are extremely slender and subglabrous, extending between {{convert|6|cm|in}} and {{convert|15|cm|in}} in length.<ref name=Munz/> |
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''Aquilegia chaplinei'' is a [[perennial plant]] with a height from {{convert|20|cm|in}} to {{convert|50|cm|in}},<ref name=USDA>{{cite web|url=https://www.fs.usda.gov/wildflowers/beauty/columbines/aquilegia_chaplinei.shtml|title=''Aquilegia chaplinei'', Chaplin’s columbine|work=Aquilegia Express: Yellow Columbines|publisher=[[United States Department of Agriculture]]|access-date=December 2, 2024}}</ref> averaging {{convert|40|cm|in}} tall. This is somewhat shorter than the closely related ''[[Aquilegia chrysantha]]'', which can reach up to {{convert|120|cm|in}} tall.<ref name=Nold/>{{rp|74–75}} ''A. chaplinei''{{apostrophe}}s [[Type (biology)|type locality]] at an altitude of {{convert|1650|m|ft}} suggests that it is better adapted than ''A. chrysantha'' to arid environments.<ref name=Payson/>{{rp|157}} ''A. chaplinei'' has a slender stem that is [[glabrous]] (smooth) with the exception of the [[inflorescence]].<ref name=Payson/>{{rp|156}} |
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The plant's flowers are suberect with blue-violet [[nectar spur]]s and [[sepal]]s. The spurs are particularly hooked and have lengths of between {{convert|10|mm|in}} and {{convert|15|mm|in}} long. ''A. amurensis''{{apostrophe}}s sepals are elliptical with lengths of between {{convert|15|mm|in}} and {{convert|25|mm|in}} long that curve to an acute point. The [[petal]] blades are oblong, spreading roughly along the [[floral axis]]. The blades range between {{convert|7|mm|in}} and {{convert|12|mm|in}} long and are whitish or white-tipped.<ref name=Nold/>{{rp|64}} The [[stamen]] are exserted (extending beyond the length of the pedals) and capped with yellow anthers that are {{convert|1|mm|in}} long.<ref name=Munz/> |
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Possessing leaves in a [[Phyllotaxis|basal arrangement]] (sprouting from base of the shoot). Its leaves extend on slender [[petiole (botany)|petiole]]s that are {{convert|7|cm|in}} to {{convert|10|cm|in}} long.<ref name=Payson/>{{rp|156}} The leaves themselves range from bi- to barely triternately compound. ''A. chaplinei'' has a fern-like appearance when not flowering.<ref name=Payson/>{{rp|156}}<ref name=Nold/>{{rp|74–75}} The leaves are [[semi-evergreen]].<ref name=Johnson>{{cite web|url=https://www.wildflower.org/plants/result.php?id_plant=AQCHC|title=Aquilegia chrysantha var. chaplinei|website=wildflower.org|publisher=[[Lady Bird Johnson Wildflower Center]]|access-date=December 2, 2024}}</ref> |
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A variety, ''A. amurensis'' var. ''albiflora'', possesses white flowers.<ref name=Nold/>{{rp|64}} When compared to ''[[Aquilegia sibirica]]'', ''A. amurensis'' emulates its leafless stems and hooked spurs but deviates with more acute sepals, pilose leaflets and pedicels, less nodding flowers, and follicles that are not glabrous.<ref name=Munz/> |
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It has pale yellow flowers. Its spurs range from {{convert|30|mm|in}} to {{convert|40|mm|in}} and can be slender, straight, or slightly spreading.<ref name=Nold/>{{rp|74}} The short spurs and [[sepal]]s under 2 centimeters long{{snd}} between {{convert|13|mm|in}} and {{convert|16|mm|in}}<ref name=Nold/>{{rp|74}}{{snd}} are the primary distinguishing features that separate ''A. chaplinei'' from ''A. chrysantha''.<ref name=Johnson/> |
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==Taxonomy== |
==Taxonomy== |
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''Aquilegia amurensis'' received its [[Binomial nomenclature|binomial]] in 1926 within ''[[wikispecies:Botanicheskie Materialy Gerbariia Glavnogo Botanicheskogo Sada S.S.S.R.|Botanicheskie Materialy Gerbariia Glavnogo Botanicheskogo Sada S.S.S.R.]]'' in the [[Soviet Union]]. The plant was first identified under the name ''Aquilegia flabellata'' var. ''alpina'' within ''[[wikispecies:Izvestiya Imperatorskoi Akademii Nauk|Izvestiya Imperatorskoi Akademii Nauk]]'' in 1915.<ref name=Nold/>{{rp|64}}<ref>{{cite POWO|id=2644772-4|title=''Aquilegia flabellata'' var. ''alpina'' Kuzen.|access-date=December 6, 2024}}</ref> |
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''A. chaplinei'' is within the ''[[Aquilegia]]'' (columbine) genus.<ref name="POWO_272432-2"/> The plant, including its [[holotype]],<ref name=Holotype>{{cite web|url=https://www.si.edu/object/aquilegia-chaplinei-standl:nmnhbotany_2076034|title=Aquilegia chaplinei Standl.|publisher=[[Smithsonian Institution]]|access-date=December 2, 2024}}</ref> was first collected by W. R. Chapline from [[Sitting Bull Falls]] in [[New Mexico]] on May 25, 1916. It received the [[binomial nomenclature|binomial]] ''Aquilegia chaplinei'' in 1918 within [[Edwin Blake Payson]]'s "The North American Species of Aquilegia", published in ''[[Contributions from the United States National Herbarium]]''.<ref name=Payson/>{{rp|156–157}}<ref name=FNA/> The holotype is now in the collection of the [[National Museum of Natural History]].<ref name=Holotype/> |
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According to a 2024 [[phylogenetic]] study by Chinese researchers Huaying Wang and Wei Zhang, ''A. amurensis'' diverged from the now geographically disjunct ''[[Aquilegia flabellata|Aquilegia japonica]]''{{refn|group=note|Commonly known as ''Aquilegia flabellata''.<ref name=Nold/>{{rp|43}}}} approximately 22,970 years ago, corresponding with the [[Last Glacial Maximum]]. However, this data might be marred with interbreeding spurred by [[Last Glacial Maximum refugia|refugia]] populations lacking conspecific mates or other events. Wang and Zhang found that [[chloroplast DNA]] suggests ''A. amurensis'' is genetically more closely related to ''[[Aquilegia parviflora|Aquilegia parviflora]]''{{snd}} with which it shares a [[clade]]{{snd}} and the North American columbine clade than with ''A. japonica''. However, a 2013 study constructed a [[phylogenetic tree]] that suggested that the inverse was true; Wang and Zhang theorized that this was the result of the 2013 study utilizing a different variant of ''A. japonica''.<ref name=Biased>{{cite journal|title=Biased Gene Introgression and Adaptation in the Face of Chloroplast Capture in ''Aquilegia amurensis''|first1=Huaying|last1=Wang|first2=Wei|last2=Zhang|first3=Yanan|last3=Yu|first4=Xiaoxue|last4=Fang|first5=Tengjiao|last5=Zhang|first6=Luyuan|last6=Xu|first7=Lei|last7=Gong|first8=Hongxing|last8=Xiao|journal=Systematic Biology|doi=10.1093/sysbio/syae039|date=November 2024|volume=73|number=6|page=886–900}}</ref> |
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In 1985, Emily J. Lott proposed reclassifying the plant as ''Aquilegia chrysantha'' var. ''chaplinei'' in the journal ''[[Phytologia]]''. Lott's proposal came out of her study of plants in the [[Chihuahuan Desert]], stemming from her 1979 unpublished master's thesis on ''Aquilegia'' in the [[Trans-Pecos]] region of Texas.<ref>{{cite journal|url=https://archive.org/details/biostor-208336|title=New combinations in Chihuahuan Desert ''Aquilegia'' (Ranunculaceae)|first=Emily J.|last=Lott|journal=[[Phytologia]]|page=488|volume=58}}</ref> The name proposed by Lott was not broadly accepted outside of Texas,<ref name=Nold/>{{rp|74}} where it is used by the [[Lady Bird Johnson Wildflower Center]] of the [[University of Texas at Austin]].<ref name=Johnson/> |
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Despite similarities in the shape of the shoots and leaves to ''[[Aquilegia barykinae]]'', another species in the same region, it is not likely to be a close relative.<ref>{{cite journal |last1=Erst |first1=Andrey S. |last2=Karakulov |first2=Anatoliy V. |last3=Luferov |first3=Alexander N. |date=October 2014 |title=Aquilegia barykinae (Ranunculaceae), a new species from the Far East of Russia |journal=Систематические заметки по материалам Гербария им. П.Н. Крылова Томского государственного университета |volume=110 |pages=3–8 |url=https://www.researchgate.net/publication/277608458_Aquilegia_barykinae_Ranunculaceae_a_new_species_from_the_Far_East_of_Russia0 |access-date=December 7, 2024}}</ref> |
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==Distribution== |
==Distribution== |
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The plant is native to the northern [[Greater Khingan]] mountain range and [[Amur River]] of China, as well as Siberian Russia, Mongolia, and possibly [[North Korea]].<ref name=Div/><ref name=Mongolia>{{cite journal|url=https://phytokeys.pensoft.net/article/79702/|title=Flora of Mongolia: annotated checklist of native vascular plants|journal=[[PhytoKeys]]|first1=Shukherdorj|last1=Baasanmunkh|first2=Magsar|last2=Urgamal|first3=Batlai|last3=Oyuntsetseg|first4=Alexander P.|last4=Sukhorukov|first5=Zagarjav|last5=Tsegmed|first6=Dong Chan|last6=Son|first7=Andrey|last7=Erst|first8=Khurelpurev|last8=Oyundelger|first9=Alexey A.|last9=Kechaykin|first10=Joscelyn|last10=Norris|first11=Petr|last11=Kosachev|first12=Jin-Shuang|last12=Ma|first13=Kae Sun|last13=Chang|first14=Hyeok Jae|last14=Choi|date=March 14, 2022|volume=192|page=63–169}}</ref><ref name=Nold/>{{rp|64}} A 2017 paper in ''[[Phytotaxa]]'' claimed to be the first to record ''A. amurensis'' as present in China.<ref>{{cite journal|title=Two new species and four new records of ''Aquilegia'' (Ranunculaceae) from China|journal=[[Phytotaxa]]|date=2017|volume=316|number=2|pages=121–137|first1=Andrey S.|last1=Erst|first2=Wang|last2=Wei|first3=Sheng-Xiang|last3=Yu|doi=10.11646/phytotaxa.316.2.2}}</ref> Wang and Zhang mapped the range of ''A. amurensis'' extending through northern China and eastern Siberian Russia.<ref name=Biased/> |
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The species is endemic to the [[Guadalupe Mountains]] of [[West Texas]] and southeastern [[New Mexico]] in the [[West South Central states|West South Central United States]].<ref name="POWO_272432-2"/> The Guadalupe Mountains are an extremely arid environment, and ''A. chaplinei'' is found where the ground is moist such as along streams, canyons, and at the base of rocks.<ref name=USDA/><ref name=Nold/>{{rp|74}} It is also native to the New Mexican [[Sacramento Mountains (New Mexico)|Sacramento Mountains]].<ref name=Johnson/> |
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Botanist [[Philip A. Munz]] described the [[type locality (biology)|type locality]] of the species as "R. Lagar, near Radde, [[Amur Oblast|Amur]], Siberia".<ref name=Munz/><ref name=Nold/>{{rp|64}} [[Roy Lancaster]] included an image of ''A. amurensis'' in his ''Travels in China: A Plantman's Paradise''. The plant pictured was found the [[Changbai Mountains]] on the Chinese side of the nation's border with North Korea. The range of ''A. amurensis'' possibly extends over this border.<ref name=Nold/>{{rp|64}}{{refn|group=note|Nold noted that the image in Lancaster's book did not precisely correspond with the description provided by Munz and given by ''Flora of the U.S.S.R.''<ref name=Nold/>{{rp|64}}}} Munz noted that the Soviet botanist [[Vladimir Leontyevich Komarov]] had reported ''A. amurensis'' in northern Korea from the [[Yalu River]].<ref name=Munz/> |
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''A. chaplinei'' is considered a rare flower.<ref name=Johnson/> The species is of conservation concern.<ref name=FNA>{{cite book|chapter-url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500094|chapter=Aquilegia chaplinei|title=[[Flora of North America]]|volume=3}}</ref> |
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==Cultivation== |
==Cultivation== |
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In his 1946 ''Aquilegia: The Cultivated and Wild Columbines'', Munz said that he was aware of seeds being offered for sale by a Japanese company. He added that he was not aware of any other cultivation of the species.<ref name=Munz>{{cite book|title=Aquilegia: The Cultivated and Wild Columbines|series=Gentes Herbarum|volume=VII|location=[[Ithaca, New York|Ithaca]], [[New York (state)|NY]]|author-link=Philip A. Munz|first=Philip A.|last=Munz|date=March 25, 1946|publisher=[[New York State College of Agriculture and Life Sciences at Cornell University|The Bailey Hortorium of the New York State College of Agriculture at Cornell University]]|url=https://archive.org/details/sim_gentes-herbarum_1946-03-25_7_1/page/68/mode/2up|via=Archive.org|pages=68–69}}</ref> {{As of|2003}}, American botanist Robert Nold reported that the plant was "rarely, if ever, encountered in cultivation".<ref name=Nold/>{{rp|64}} |
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{{As of|2004}}, both seeds and plants were sometimes available, particularly from native plant nurseries in the region of ''A. chaplinei''{{apostrophe}}s native range.<ref name=Nold/>{{rp|74}} |
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==Notes== |
==Notes== |
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Category:Aquilegia| |
[[Category:Aquilegia|amurensis]] |
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Category:Flora of |
[[Category:Flora of Siberia]] |
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[[Category:Flora of China]] |
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[[Category:Flora of Mongolia]] |
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[[Category:Flora of North Korea]] |
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[[Category:Plants described in 1926]] |
Latest revision as of 18:54, 12 December 2024
Aquilegia amurensis | |
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Preserved specimen of Aquilegia amurensis | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Order: | Ranunculales |
Family: | Ranunculaceae |
Genus: | Aquilegia |
Species: | A. amurensis
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Binomial name | |
Aquilegia amurensis Not. Syst. Herb. Hort. Bot.
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Synonyms[1][2]: 64 | |
Aquilegia flabellata Siebold & Zucc. |
Aquilegia amurensis is a partially accepted species of flowering plant in the genus Aquilegia (columbines) in the family Ranunculaceae that is native to the northeastern Asian. Its natural range is in the northern Greater Khingan mountain range and Amur River of China, as well as Siberian Russia, Mongolia, and possibly North Korea.[3][4][2]: 64 Flowers of this plant have petal blades that are whitish or white-tipped, with blue-violet nectar spurs and sepals. The plant is rarely cultivated.
Description
[edit]Aquilegia amurensis grows to be about 50 centimetres (20 in) tall. It possesses leaves in a basal arrangement (sprouting from base of the shoot), seldom if ever with leaves on the stem. The leaflets are glabrous (smooth) on their topsides with soft, downy bottoms.[2]: 64 The leaflets are trisected to their base. The petioles are extremely slender and subglabrous, extending between 6 centimetres (2.4 in) and 15 centimetres (5.9 in) in length.[5]
The plant's flowers are suberect with blue-violet nectar spurs and sepals. The spurs are particularly hooked and have lengths of between 10 millimetres (0.39 in) and 15 millimetres (0.59 in) long. A. amurensis's sepals are elliptical with lengths of between 15 millimetres (0.59 in) and 25 millimetres (0.98 in) long that curve to an acute point. The petal blades are oblong, spreading roughly along the floral axis. The blades range between 7 millimetres (0.28 in) and 12 millimetres (0.47 in) long and are whitish or white-tipped.[2]: 64 The stamen are exserted (extending beyond the length of the pedals) and capped with yellow anthers that are 1 millimetre (0.039 in) long.[5]
A variety, A. amurensis var. albiflora, possesses white flowers.[2]: 64 When compared to Aquilegia sibirica, A. amurensis emulates its leafless stems and hooked spurs but deviates with more acute sepals, pilose leaflets and pedicels, less nodding flowers, and follicles that are not glabrous.[5]
Taxonomy
[edit]Aquilegia amurensis received its binomial in 1926 within Botanicheskie Materialy Gerbariia Glavnogo Botanicheskogo Sada S.S.S.R. in the Soviet Union. The plant was first identified under the name Aquilegia flabellata var. alpina within Izvestiya Imperatorskoi Akademii Nauk in 1915.[2]: 64 [6]
According to a 2024 phylogenetic study by Chinese researchers Huaying Wang and Wei Zhang, A. amurensis diverged from the now geographically disjunct Aquilegia japonica[note 1] approximately 22,970 years ago, corresponding with the Last Glacial Maximum. However, this data might be marred with interbreeding spurred by refugia populations lacking conspecific mates or other events. Wang and Zhang found that chloroplast DNA suggests A. amurensis is genetically more closely related to Aquilegia parviflora – with which it shares a clade – and the North American columbine clade than with A. japonica. However, a 2013 study constructed a phylogenetic tree that suggested that the inverse was true; Wang and Zhang theorized that this was the result of the 2013 study utilizing a different variant of A. japonica.[7]
Despite similarities in the shape of the shoots and leaves to Aquilegia barykinae, another species in the same region, it is not likely to be a close relative.[8]
Distribution
[edit]The plant is native to the northern Greater Khingan mountain range and Amur River of China, as well as Siberian Russia, Mongolia, and possibly North Korea.[3][4][2]: 64 A 2017 paper in Phytotaxa claimed to be the first to record A. amurensis as present in China.[9] Wang and Zhang mapped the range of A. amurensis extending through northern China and eastern Siberian Russia.[7]
Botanist Philip A. Munz described the type locality of the species as "R. Lagar, near Radde, Amur, Siberia".[5][2]: 64 Roy Lancaster included an image of A. amurensis in his Travels in China: A Plantman's Paradise. The plant pictured was found the Changbai Mountains on the Chinese side of the nation's border with North Korea. The range of A. amurensis possibly extends over this border.[2]: 64 [note 2] Munz noted that the Soviet botanist Vladimir Leontyevich Komarov had reported A. amurensis in northern Korea from the Yalu River.[5]
Cultivation
[edit]In his 1946 Aquilegia: The Cultivated and Wild Columbines, Munz said that he was aware of seeds being offered for sale by a Japanese company. He added that he was not aware of any other cultivation of the species.[5] As of 2003[update], American botanist Robert Nold reported that the plant was "rarely, if ever, encountered in cultivation".[2]: 64
Notes
[edit]References
[edit]- ^ "Aquilegia amurensis Kom". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved December 6, 2024.
- ^ a b c d e f g h i j k l Nold, Robert (2003). Columbines: Aquilegia, Paraquilegia, and Semiaquilegia. Portland, OR: Timber Press. ISBN 0881925888.
- ^ a b Li, Ming-Rui; Wang, Hua-Ying; Tianyuan Lu, Ning Ding; Huang, Ye-Chao; Xiao, Hong-Xing; Liu, Bao; Li, Lin-Feng (March 2019). "Rapid Divergence Followed by Adaptation to Contrasting Ecological Niches of Two Closely Related Columbine Species Aquilegia japonica and A. oxysepala". Genome Biology and Evolution. 11 (3): 920.
- ^ a b Baasanmunkh, Shukherdorj; Urgamal, Magsar; Oyuntsetseg, Batlai; Sukhorukov, Alexander P.; Tsegmed, Zagarjav; Son, Dong Chan; Erst, Andrey; Oyundelger, Khurelpurev; Kechaykin, Alexey A.; Norris, Joscelyn; Kosachev, Petr; Ma, Jin-Shuang; Chang, Kae Sun; Choi, Hyeok Jae (March 14, 2022). "Flora of Mongolia: annotated checklist of native vascular plants". PhytoKeys. 192: 63–169.
- ^ a b c d e f Munz, Philip A. (March 25, 1946). Aquilegia: The Cultivated and Wild Columbines. Gentes Herbarum. Vol. VII. Ithaca, NY: The Bailey Hortorium of the New York State College of Agriculture at Cornell University. pp. 68–69 – via Archive.org.
- ^ "Aquilegia flabellata var. alpina Kuzen". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved December 6, 2024.
- ^ a b Wang, Huaying; Zhang, Wei; Yu, Yanan; Fang, Xiaoxue; Zhang, Tengjiao; Xu, Luyuan; Gong, Lei; Xiao, Hongxing (November 2024). "Biased Gene Introgression and Adaptation in the Face of Chloroplast Capture in Aquilegia amurensis". Systematic Biology. 73 (6): 886–900. doi:10.1093/sysbio/syae039.
- ^ Erst, Andrey S.; Karakulov, Anatoliy V.; Luferov, Alexander N. (October 2014). "Aquilegia barykinae (Ranunculaceae), a new species from the Far East of Russia". Систематические заметки по материалам Гербария им. П.Н. Крылова Томского государственного университета. 110: 3–8. Retrieved December 7, 2024.
- ^ Erst, Andrey S.; Wei, Wang; Yu, Sheng-Xiang (2017). "Two new species and four new records of Aquilegia (Ranunculaceae) from China". Phytotaxa. 316 (2): 121–137. doi:10.11646/phytotaxa.316.2.2.