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Mertensia
	Mertensia virginica (type species)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Boraginales
Family:	Boraginaceae
Subfamily:	Boraginoideae
Genus:	Mertensia; Roth 1797
Type species
	Mertensia virginica; (L.) Persoon ex Link
Species
	See text

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.^{[citation needed]} 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.^[1] Its center of diversity is in the Rocky Mountains. Mertensia is mostly restricted to alpine, subalpine, and montane habitats.^[2] Notable exceptions are Mertensia maritima, a maritime plant of Arctic and subarctic coastlines, and Mertensia virginica, which is found from the Appalachian Mountains west to Minnesota, Iowa, and Missouri.^[1] 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 in Europe.^{[citation needed]} About 12 other species are known in cultivation.^[3] The Inuit ate the rhizomes of Mertensia maritima.^[4]

Many of the species of Mertensia are hard to distinguish and some are possibly cryptic.^[2] Around 150 species names have been published in Mertensia.^[5] Most authors have recognized about 45 species,^[4] but in 2014, the authors of a molecular phylogenetic study recommended the acceptance of at least 62.^[1]

Species

Author citations are from Tropicos^[5] or the International Plant Names Index.^[6]

section Stenhammaria

Mertensia davurica (Sims) G.Don 1837
Mertensia kamczatica (Turcz.) DC. 1846
Mertensia maritima (L.) Gray 1821 – oysterplant, sea bluebell
Mertensia meyeriana J.F.Macbr. 1916
Mertensia pterocarpa (Turcz.) Tatew. & Ohwi 1933
Mertensia sibirica (L.) G.Don 1837
Mertensia simplicissima (Ledeb.) G.Don 1837
Mertensia sinica Kamelin 2009
Mertensia stylosa (Fisch.) DC. 1846
Mertensia tarbagataica B.Fedtsch. 1915

section Mertensia (incomplete)

Mertensia alpina (Torrey) G.Don 1838 – alpine bluebells
Mertensia arizonica Greene 1897 – aspen bluebells
Mertensia bella Piper 1918 – beautiful bluebells
Mertensia brevistyla Piper 1918 – short-styled bluebells
Mertensia ciliata G.Don 1837 – mountain bell
Mertensia drummondii (Lehmann) G.Don 1838
Mertensia franciscana A.Heller 1899 – Franciscan bluebells
Mertensia lanceolata DC. 1846 – prairie bluebells
Mertensia longiflora Greene 1898 – long bluebell
Mertensia macdougalii A.Heller 1899
Mertensia oblongifolia (Nuttall) G.Don 1838 – oblongleaf bluebells
Mertensia paniculata (Aiton) G.Don 1837 – tall bluebell or northern bluebell
Mertensia pilosa (Chamisso) G.Don 1838
Mertensia platyphylla A.Heller 1899
Mertensia rivularis DC. 1846
Mertensia subcordata Greene 1899
Mertensia virginica (L.) Persoon ex Link 1829 – Virginia bluebells

Mertensia dschagastanica Regel 1880 is probably a species of Stephanocaryum (S. dschagastanicum (Regel) O.D.Nikif. 2014).

Classification

Mertensia is a member of the tribe Cynoglosseae.^[7] Its closest relative is the monotypic 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 cladistic analysis of DNA sequences.

Taxonomy

The type species for Mertensia is M. virginica.^[8] 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.^[2]

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.^[9]^[10] 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.^{[citation needed]} It has not yet been sampled in a molecular phylogenetic study.

With the possible exception of M. pilosa, North American Mertensia is a monophyletic group consisting of three clades that are known informally as the Pacific Northwest clade, the Southern Rocky Mountain clade, and the Central Rocky Mountain clade.^[1] 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, Swedish botanist Carl Linnaeus placed five species in the genus Pulmonaria.^[11] Albrecht Wilhelm Roth, in 1797, separated what are now M. virginica, M. maritima, and M. sibirica from Pulmonaria to form the genus Mertensia,^[6] 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.^[12] Roth described one species as Mertensia pulmonarioides, apparently unaware that Linnaeus had already described it as Pulmonaria virginica.^[13] He thus created a superfluous synonym that has been a source of confusion ever since.

Mertensia was named after the German botanist, Franz Carl Mertens.^[14]

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)),^[15] Asa Gray,^[16] James Francis Macbride,^[17] Per Axel Rydberg,^[18] Louis Otho Williams,^[19] and Mikhail Grigorevich Popov.^[9]^[10]

Asa Gray divided Mertensia into two sections: Stenhammaria and Mertensia.^[20] 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 species M. maritima, but in 2014, it was expanded to comprise 12 species.^[2]

In 1886, Asa Gray described seven species in Mertensia. After Gray completed his 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.^[21] 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 character states that show true phylogenetic relationships. In Mertensia, as elsewhere, such groups have often proved to be paraphyletic. 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 stamens (> 1.5 mm), filaments inserted higher in the corolla, calyces divided at least ²⁄₃ of the way to the base, and acute to acuminate 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.^[22]

There are no known fossils of Mertensia. A molecular clock analysis has estimated that Mertensia diverged 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 glaciation, then retreated northward and to higher elevation during interglacials.^[1] Mertensia originated in Asia and dispersed over the Beringian land bridge to North America. Most of the groups that originated in Asia are more diverse there,^[23]^[24] but Mertensia is a conspicuous exception in having most of its species in North America.

Gallery

Alpine bluebells (Mertensia alpina)
Aspen Bluebells (Mertensia arizonica)^[25]
Short-styled Bluebell (Mertensia brevistyla)^[26]
Mountain bell (Mertensia ciliata)
Franciscan bluebells (Mertensia franciscana)
Prairie bluebells (Mertensia lanceolata)
Long bluebells (Mertensia longiflora)
Oysterplant (Mertensia maritima)
Oblongleaf bluebells (Mertensia oblongifolia)
Tall bluebells (Mertensia paniculata)
Virginia bluebells (Mertensia virginica )

References

^ ^a ^b ^c ^d ^e 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. doi:10.3732/ajb.1300320.
^ ^a ^b ^c ^d Mare Nazaira and Larry Hufford. 2014. "Phylogenetic Systematics of the Genus Mertensia (Boraginaceae)". Systematic Botany 39(1):268-303. doi:10.1600/036364414X678107.
^ 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. ISBN 978-0-333-47494-5 (set).
^ ^a ^b David J. Mabberley. 2008. Mabberley's Plant-Book third edition (2008). Cambridge University Press: UK. ISBN 978-0-521-82071-4 (see External links below).
^ ^a ^b Mertensia in Tropicos (See External links below).
^ ^a ^b Mertensia in International Plant Names Index. (see External links below).
^ 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. doi:10.1111/cla.12036
^ Mertensia In: Index Nominum Genericorum. In: Regnum Vegetabile (see External links below).
^ ^a ^b 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).
^ ^a ^b 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.
^ 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. 1797. "Mertensia". Catalecta Botanica 1:34-35. (See External links below).
^ James S. Pringle. 2004. "Nomenclature of the Virginia-bluebell, Mertensia virginica (Boraginaceae)". SIDA, contributions to botany 21(2):771-775.(see External links below)
^ Umberto Quattrocchi. 2000. CRC World Dictionary of Plant Names, volume III. CRC Press: Baton Rouge, New York, London, Washington DC. ISBN 978-0-8493-2673-8 (vol. III). (see External links below).
^ 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. 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. 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. 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. 1937. "A monograph of the genus Mertensia in North America". Annals of the Missouri Botanical Garden 24 (1):17-159. (see External links below).
^ 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).
^ 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.
^ Tomoko Fukuda and Hiroshi Ikeda. 2012. "Palynological analysis and taxonomic position of the genus Mertensia (Boraginaceae)". Botany 90(8):722-730. doi:10.1139/b2012-024.
^ 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.
^ Mertensia Arizonica, Aspen Bluebells
^ Teresa Prendusi, Short-styled Bluebell (Mertensia brevistyla)