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Mertensia

( Boraginaceae Genera )

Species
Classification
Taxonomy
History
Evolution
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Mertensia Mode Species Taxonomy

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Mertensia is a genus of in the borage family Boraginaceae. They are Perennial plant 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 changes in soil pH. Mertensia is one of several plants that are commonly called " bluebell". In spite of their common name, 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 is in the Rocky Mountains. Mertensia is mostly restricted to Alpine climate, subalpine, and montane ecology 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 Oceanic climate plant of Arctic and subarctic coastlines, and Mertensia virginica, which is found from the Appalachian Mountains west to Minnesota, Iowa, and Missouri. Most of the species are Endemism to very small areas of the Rocky Mountains.

Mertensia virginica has the largest flowers in the genus and is commonly Gardening. It is sparingly naturalized 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 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 Species complex. Around 150 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 study recommended the acceptance of at least 62.

Species

According to Plants of the World Online there are 52 accepted species in the genus.

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 Sister group is the Monotypic taxon Eurasian genus Asperugo. These two are probably close to Anoplocaryum, a genus of Central Asia and Siberia. The relationships of Anoplocaryum have never been investigated by Cladistics analysis of DNA sequences.

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: Stenhammaria and Mertensia. Section Stenhammaria consists of the circumboreal M. maritima and 11 of the 12 species from Asia.

The Central Asian species, M. dschagastanica, is assigned to section Stenhammaria, but it is Plant morphology 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 Monophyly group consisting of three that are known informally as the Pacific Northwest clade, the Southern Rocky Mountain clade, and the Central Rocky Mountain clade. These groups are Neologism 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, 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, in 1797, separated what are now M. virginica, M. maritima, and M. sibirica from Pulmonaria to form the genus Mertensia,< Mertensia in International Plant Names Index. (see External links below). based on their smaller and differently structured calyx, their different anther position, and the presence of nectar glands 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 that has been a source of confusion ever since.

Mertensia was named after the German Botany, Franz Carl Mertens.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. 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.

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, who is best known for his work in lichenology. Gray defined the section Stenhammaria as consisting only of the Littoral zone 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 of Mertensia, many species were Plant collecting 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 Treatise 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. Phylogenetics analysis of DNA data has shown that many of the species are Polyphyly, 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 Taxonomic 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 Plant taxonomy of Mertensia, some of the Taxon were based on shared Symplesiomorphy Phenotypic trait rather than the Apomorphy character states that show true phylogenetic relationships. In Mertensia, as elsewhere, such groups have often proved to be Paraphyly. More recently, molecular phylogenetics has greatly clarified the ancestral and derived character states in Mertensia. Some of the traits Evolution later have appeared independently as many as seven times.

Ancestral states in Mertensia include short plant height (< 40 cm), long (> 1.5 mm), filaments inserted higher in the Petal, Sepal divided at least ²⁄₃ of the way to the base, and acute to Leaf shape calyx lobe apices. 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 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 of Mertensia. A molecular clock analysis has estimated that Mertensia Speciation from Asperugo in the late Oligocene or early Miocene. Asperugo and Mertensia do not closely resemble each other morphologically. Being mostly plants of subarctic climates, Mertensia spread southward and to lower elevation during periods of Pleistocene Glacial period, then retreated northward and to higher elevation during . Mertensia originated in Asia and Seed dispersal over the land bridge to North America. Most of the groups that originated in Asia are more Biodiversity 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.