''Mertensia'' is a genus of flowering plants in the family Boraginaceae. They are perennial herbaceous plants with blue or sometimes white flowers that open from pink-tinged buds. Such a change in flower color is common in Boraginaceae and is caused by an increase of pH in the flower tissue. ''Mertensia'' is one of several plants that are commonly called "bluebell". In spite of their

common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...

, the flowers are usually salverform (trumpet-shaped) rather than campanulate (bell-shaped). ''Mertensia'' is native to most of North America and to a large part of Asia from western China to northeastern Russia.Mare Nazaire, Xiao-Quan Wang, and Larry Hufford. 2014. "Geographic origins and patterns of radiation of ''Mertensia'' (Boraginaceae)". ''American Journal of Botany'' 101(1):104-118. . Its

center of diversity A center of origin is a geographical area where a group of organisms, either domesticated or wild, first developed its distinctive properties. They are also considered centers of diversity. Centers of origin were first identified in 1924 by Ni ...

is in the Rocky Mountains. ''Mertensia'' is mostly restricted to alpine, subalpine, and montane habitats.Mare Nazaira and Larry Hufford. 2014. "Phylogenetic Systematics of the Genus ''Mertensia'' (Boraginaceae)". ''Systematic Botany'' 39(1):268-303. . Notable exceptions are ''Mertensia maritima'', a maritime plant of Arctic and

subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...

coastlines, and ''Mertensia virginica'', which is found from the Appalachian Mountains west to Minnesota, Iowa, and Missouri. Most of the species are endemic to very small areas of the Rocky Mountains. ''Mertensia virginica'' has the largest flowers in the genus and is commonly cultivated. It is sparingly

naturalized Naturalization (or naturalisation) is the legal act or process by which a non-citizen of a country may acquire citizenship or nationality of that country. It may be done automatically by a statute, i.e., without any effort on the part of the in ...

in Europe. About 12 other species are known in cultivation. Anthony Huxley, Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. (set). The Inuit ate the

rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

s of ''Mertensia maritima''.David J. Mabberley. 2008. ''Mabberley's Plant-Book'' third edition (2008). Cambridge University Press: UK. (see ''External links'' below). Many of the species of ''Mertensia'' are hard to distinguish and some are possibly

cryptic Cryptic may refer to: In science: * Cryptic species complex, a group of species that are very difficult to distinguish from one another * Crypsis, the ability of animals to blend in to avoid observation * Cryptic era, earliest period of the Earth ...

. Around 150 species names have been published in ''Mertensia''.''Mertensia'' in Tropicos (See ''External links'' below). Most authors have recognized about 45 species, but in 2014, the authors of a

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 recommended the acceptance of at least 62.

Species

Author citations are from Tropicos or the International Plant Names Index.''Mertensia'' in International Plant Names Index. (see ''External links'' below).

Classification

''Mertensia'' is a member of the tribe Cynoglosseae.James I. Cohen. 2014. "A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of character evolution". ''Cladistics'' 30(2):139-169. Its closest relative is the monotypic Eurasian genus ''

Asperugo ''Asperugo procumbens'', known as madwort or German madwort, is the single species in the monotypic plant genus ''Asperugo''. This plant is native to Europe but has been introduced elsewhere, such as the northern half of North America. Referenc ...

''. These two are probably

close to Close may refer to: Music * ''Close'' (Kim Wilde album), 1988 * ''Close'' (Marvin Sapp album), 2017 * ''Close'' (Sean Bonniwell album), 1969 * "Close" (Sub Focus song), 2014 * "Close" (Nick Jonas song), 2016 * "Close" (Rae Sremmurd song), 201 ...

Anoplocaryum ''Anoplocaryum'' is a genus of flowering plants belonging to the family Boraginaceae. Its native range is from Siberia to Mongolia and includes Northwestern Himalaya The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separat ...

'', a genus of Central Asia and Siberia. The relationships of ''Anoplocaryum'' have never been investigated by cladistic analysis of

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

Taxonomy

The type species for ''Mertensia'' is ''M. virginica''.''Mertensia'' In: Index Nominum Genericorum. In: Regnum Vegetabile (see ''External links'' below). ''Mertensia'' is divided into two

sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...

: Stenhammaria and Mertensia. Section Stenhammaria consists of the

circumboreal The Circumboreal Region in phytogeography is a floristic region within the Holarctic Kingdom in Eurasia and North America, as delineated by such geobotanists as Josias Braun-Blanquet and Armen Takhtajan. It is the largest floristic region in t ...

''M. maritima'' and 11 of the 12 species from Asia. The Central Asian species, ''M. dschagastanica'', is assigned to section Stenhammaria, but it is morphologically unusual and might constitute a third section of ''Mertensia''. It has never been sampled for DNA. Section Mertensia consists of all of the North American species plus ''M. rivularis'', a denizen of the Russian half of the Beringian floristic region. The North American species ''M. pilosa'' has been reported from Sakhalin and the Kuril Islands.Mikhail G. Popov. 1953. "Boraginaceae". pages 67-691 and 701-716. In: Boris K. Schischkin (editor). ''Flora of the USSR'', volume 19 - Tubiflorae. Izdatel'stvo Akademii Nauk SSSR: Moskva, Leningrad,, Russia. (published by the Academy of Sciences of the USSR).Rose Lavoott (translator). 1974. "Boraginaceae". pages 73-508 and 516-531. In: English translation of: Boris K. Schischkin (editor). ''Flora of the USSR'', volume 19 - Tubiflorae. Keter Press: Jerusalem, Israel. For: Israel Program for Scientific Translations. It is the only species of ''Mertensia'' that is native to both Asia and North America. It resembles ''M. rivularis'' and might be closer to that species than to the other species of North America. It also resembles ''M. platyphylla'' and some authors have placed it in synonymy under ''Mertensia platyphylla'' variety ''platyphylla''. It has not yet been sampled in a molecular phylogenetic study. With the possible exception of ''M. pilosa'', North American ''Mertensia'' is a

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 ...

group consisting of three

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, ...

s that are known informally as the Pacific Northwest clade, the Southern Rocky Mountain clade, and the Central Rocky Mountain clade. These groups are named for the region where most of their species occur, but each includes species from well outside of that region.

History

In 1753, in his landmark ''

Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...

'', Swedish botanist Carl Linnaeus placed five species in the genus '' Pulmonaria''.Carolus Linnaeus (Carl von Linné). 1753. ''Species Plantarum'', 1st edition, 1:135-136. Holmiae: Impensis Laurentii Salvii (Lars Salvius). (A facsimile with an introduction by William T. Stearn was published by the Ray Society in 1957). (See ''External links'' below).

Albrecht Wilhelm Roth Albrecht Wilhelm Roth (6 January 1757 – 16 October 1834) was a physician and botanist born in Dötlingen, Germany. He studied medicine at the Universities of Halle and Erlangen, where he received his doctorate in 1778. After graduation, he pra ...

, in 1797, separated what are now ''M. virginica'', ''M. maritima'', and ''M. sibirica'' from ''Pulmonaria'' to form the genus ''Mertensia'', based on their smaller and differently structured calyx, their different anther position, and the presence of

nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...

s on the inner surface of the corolla.Albrecht Wilhelm Roth. 1797. "Mertensia". ''Catalecta Botanica'' 1:34-35. (See ''External links'' below). Roth described one species as ''Mertensia pulmonarioides'', apparently unaware that Linnaeus had already described it as ''Pulmonaria virginica''.James S. Pringle. 2004. "Nomenclature of the Virginia-bluebell, ''Mertensia virginica'' (Boraginaceae)". SIDA, contributions to botany 21(2):771-775.(see ''External links'' below) He thus created a superfluous

synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...

that has been a source of confusion ever since. ''Mertensia'' was named after the German

botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...

Franz Carl Mertens Franz Carl Mertens (3 April 1764 – 19 June 1831) was a German botanist who was a native of Bielefeld. He specialized in the field of phycology. Mertens studied theology and languages at the University of Halle, and after graduation taught ...

.Umberto Quattrocchi. 2000. ''CRC World Dictionary of Plant Names'', volume III. CRC Press: Baton Rouge, New York, London, Washington DC. (vol. III). (see ''External links'' below). In the time since ''Mertensia'' was erected in 1797, it has been the subject of six major revisions. These, in chronological order, were done by George Don ( George Don, the younger (G. Don), not George Don, the elder (Don)),George Don (G.Don). 1838. "A general history of the dichlamydeous plants comprising complete descriptions of the different orders". volume 4 (Corolliflorae): ''Mertensia''. pages 318-320, 372. Gilbert and Rivington: London, UK. (see ''External links'' below).

Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...

,Asa Gray. 1886. ''Synoptical Flora of North America'', volume 2, part 1 (Gamopetalae after Compositae): "Boraginaceae": pages 177-207. ''Mertensia'': pages 179, 199-201. Ivison, Blakeman, Taylor, and Company: New York, NY, USA. (see ''External links'' below). James Francis Macbride,James Francis Macbride. 1916. "The true mertensias of western North America". ''Contributions from the Gray Herbarium. New Series'' 48:1-20. (see ''External links'' below). Per Axel Rydberg,Per Axel Rydberg. 1922. ''Flora of the Rocky Mountains and Adjacent Plains'', 2nd edition. ''Mertensia''. pages 730-736. reprinted in 1954 by Hafner Publishing Company: New York, NY, USA. (see ''External links'' below). Louis Otho Williams,Louis Otho Williams. 1937. "A monograph of the genus ''Mertensia'' in North America". ''Annals of the Missouri Botanical Garden'' 24 (1):17-159. (see ''External links'' below). and

Mikhail Grigorevich Popov Mikhail Grigorevich Popov (russian: Михаил Григорьевич Попов) (5(17) April 1893 – 18 December 1955) was a Soviet Union, Soviet botanist. He is known for developing a theory on the role of hybridization in plant evolution, a ...

. Asa Gray divided ''Mertensia'' into two sections: Stenhammaria and Mertensia.Asa Gray. 1874. "Notes on Borraginaceae" (sic). ''Proceedings of the American Academy of Arts and Sciences'' 10(New Series volume 2):48-62. (see ''External links'' below). The section Stenhammaria was named for the Swedish naturalist and clergyman

Christian Stenhammar Christian Stenhammar (1783-1866) was a Swedish naturalist interested in lichens and an entomologist who specialised in Diptera.His collection is held by Uppsala University.He was a clergyman. Works *''Försök till Gruppering och Revision af de ...

, who is best known for his work in lichenology. Gray defined the section Stenhammaria as consisting only of the littoral species ''M. maritima'', but in 2014, it was expanded to comprise 12 species. In 1886, Asa Gray described seven species in ''Mertensia''. After Gray completed his

monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...

of ''Mertensia'', many species were discovered by botanical expeditions in the western United States. Many of the species recognized by Macbride and Rydberg were later placed in synonymy by Louis O. Williams in his treatment of North American ''Mertensia'' in which he accepted only 24 species, far fewer than Macbride and Rydberg. ''M. pilosa'' was not mentioned anywhere in the monograph by Williams, not even as a synonym. Popov (1953) recognized the same 24 North American species as Williams as well as 14 species from Asia, including ''M. pilosa''. Phylogenetic analysis of DNA data has shown that many of the species are polyphyletic, but no comprehensive revision of the Asian or of the North American species has been attempted since 1953. In 1967, one of the sections delineated by Popov was raised to the rank of genus as '' Pseudomertensia''.Harald Udo von Riedl. 1967. ''Pseudomertensia''. pages 58-63. In: "Boraginaceae". pages 1-281. In: Karl Heinz Rechinger (editor). ''Flora Iranica'', monograph 48. Akademische Druck- und Verlagsanstalt: Graz, Austria. This reclassification has been confirmed by molecular phylogenies which place ''Pseudomertensia'' closer to '' Myosotis'' than any of the genera that have been sampled so far.

Evolution

In the earlier infrageneric classifications of ''Mertensia'', some of the groups were based on shared "primitive" characters rather than the

derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...

character states that show true phylogenetic relationships. In ''Mertensia'', as elsewhere, such groups have often proved to be

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

. More recently, molecular phylogenetics has greatly clarified the ancestral and derived character states in ''Mertensia''. Some of the traits evolving later have appeared independently as many as seven times. Ancestral states in ''Mertensia'' include short plant height (< 40 cm), long

stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

s (> 1.5 mm), filaments inserted higher in the

corolla Corolla may refer to: *Corolla (botany), the petals of a flower, considered as a unit *Toyota Corolla, an automobile model name * Corolla (headgear), an ancient headdress in the form of a circlet or crown * ''Corolla'' (gastropod), a genus of moll ...

, calyces divided at least ²⁄₃ of the way to the base, and acute to acuminate calyx lobe

apices The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics), a teenaged super villainess in the Marvel Universe * Ape-X, a super-intelligent ape in the Squadron Supreme universe *Apex ...

. Their derived alternatives are greater plant height (> 40 cm), short stamens (< 1.5 mm), filaments attached lower in the corolla, and calyces divided less than halfway to the base, and obtuse calyx lobe apices. The nutlets and

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

of ''Mertensia'' are nearly uniform and consequently, are not of much taxonomic value.Tomoko Fukuda and Hiroshi Ikeda. 2012. "Palynological analysis and taxonomic position of the genus ''Mertensia'' (Boraginaceae)". ''Botany'' 90(8):722-730. . There are no known fossils of ''Mertensia''. A molecular clock analysis has estimated that ''Mertensia'' diverged from ''Asperugo'' in the late

Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...

or early Miocene. ''Asperugo'' and ''Mertensia'' do not closely resemble each other morphologically. Being mostly plants of

subarctic climate The subarctic climate (also called subpolar climate, or boreal climate) is a climate with long, cold (often very cold) winters, and short, warm to cool summers. It is found on large landmasses, often away from the moderating effects of an ocean, ge ...

s, ''Mertensia'' spread southward and to lower elevation during periods of Pleistocene glaciation, then retreated northward and to higher elevation during

interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...

s. ''Mertensia'' originated in Asia and dispersed over the

Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...

n land bridge to North America. Most of the groups that originated in Asia are more

diverse Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce * Diversity marketing, marketing communication targeting diverse customers ...

there,Sandy P. Harrison, G. Yu, H. Takahara, and Iain Colin Prentice. 2001. "Diversity of temperate plants in East Asia". ''Nature'' 413(6852):129-130.Robert E. Ricklefs, Hong Qian, and Peter S. White. 2004. "The region effect on mesoscale plant species richness between eastern Asia and eastern North America". ''Ecography'' 27(2):129-136. but ''Mertensia'' is a conspicuous exception in having most of its species in North America.

Gallery

Mertensia alpina (7977046090).jpg, Alpine bluebells ('' Mertensia alpina'') Mertensiaarizonica.jpg, Aspen Bluebells (''Mertensia arizonica'') Mertensia brevistyla TPrendusi lg.jpg, Short-styled Bluebell (''

Mertensia brevistyla ''Mertensia'' is a genus of flowering plants in the family Boraginaceae. They are perennial herbaceous plants with blue or sometimes white flowers that open from pink-tinged buds. Such a change in flower color is common in Boraginaceae and is ca ...

'')Teresa Prendusi
Short-styled Bluebell (Mertensia brevistyla)
/ref> Shot of blue mountain bluebells flowers hanging from a stem mertensia ciliata.jpg, Mountain bell (''

Mertensia ciliata ''Mertensia ciliata'' is a species of flowering plant in the borage family known by the common names tall fringed bluebells, mountain bluebells, and streamside bluebells. Distribution It is native to the western United States, in California, Nev ...

'') Mertensia franciscana1.jpg, Franciscan bluebells (''

Mertensia franciscana ''Mertensia'' is a genus of flowering plants in the Family (biology), family Boraginaceae. They are Perennial plant, perennial herbaceous plants with blue or sometimes white flowers that open from pink-tinged buds. Such a change in flower color ...

'') Mertensia lanceolata NPS-1.jpg, Prairie bluebells (''

Mertensia lanceolata ''Mertensia lanceolata'', known as prairie bluebells, narrow-leaved languid lady, lance-leaved bluebells, and lance-leaved lungwort is a species of flowering plant native to western North America. A herbaceous perennial it has blue-green leaves a ...

'') Mertensia longiflora 1718.JPG, Long bluebells ('' Mertensia longiflora'') Mertensia maritima (3701701247).jpg, Oysterplant ('' Mertensia maritima'') Mertensia oblongifolia flowers (3525613924).jpg, Oblongleaf bluebells ('' Mertensia oblongifolia'') Tall Bluebells (3816794344).jpg, Tall bluebells (''

Mertensia paniculata ''Mertensia paniculata'', also known as the tall lungwort, tall bluebells, or northern bluebells, is an herb or dwarf shrub with drooping bright-blue, bell-shaped flowers. It is native to northwestern North America and the Great Lakes. Distribu ...

'') Virginia Bluebells at Rocky River.jpg, Virginia bluebells ('' Mertensia virginica '')

References

Biodiversity Heritage Library The Biodiversity Heritage Library (BHL) is the world’s largest open access digital library for biodiversity literature and archives. BHL operates as worldwide consortiumof natural history, botanical, research, and national libraries working toge ...

): :
page 135page 136''Species Plantarum'', 1st ed. (1753)
At:View Record of title 25
At:Titles by Carl von Linné (1707-1778)AuthorsBHL

Nomenclature of the Virginia bluebellVolume 21, View BookSIDA, contributions to botanyBHL
:
page 318page 372volume 4A general history of the dichlamydeous plantsDon, George, 1798-1856authorsBHL
:
page 179page 199volume 2, part 1View BookSynoptical flora of North America:Gray, Asa, 1810-1888authorsBHL
:
Louis Otho Williams 1937volume 24Annals of the Missouri Botanical GardentitlesBHL
:
Asa Gray 1874volume 10 (New Series volume 2)Proceedings of the American Academy of Arts and SciencestitlesBHL

James Francis Macbride 1916

Per Axel Rydberg 1922Internet Archive

''Mertensia''
At:Species Records for ''Mertensia''
At:''Mertensia''
At:List of genera
At:Boraginaceae
At:List of families
At:Families and Genera in GRIN
At:Queries
At:GRIN taxonomy for plants

''Mertensia''BoraginaceaeBoraginaleslamiidsasteridsEmbryophytaStreptophytinaStreptophytaViridiplantaeEukaryotaTaxonomyUniProt

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* * {{Taxonbar, from=Q765547 Boraginaceae genera Flora of North America