''Mertensia'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

s in the

borage Borage ( or ; ''Borago officinalis''), also known as starflower, is an annual herb in the flowering plant family Boraginaceae native to the Mediterranean region. Although the plant contains small amounts of pyrrolizidine alkaloids, some parts ar ...

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Boraginaceae. They are

perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...

s with blue or sometimes white flowers that open from pink-tinged buds. Such a change in flower color is common in

Boraginaceae Boraginaceae, the Borago, borage or forget-me-not family, includes about 2,000 species of shrubs, trees, and herbs in 146 to 154 genus, genera with a worldwide distribution. The APG IV system from 2016 classifies the Boraginaceae as single famil ...

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 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 con ...

, the flowers are usually salverform (trumpet-shaped) rather than

campanulate This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary ...

(bell-shaped). ''Mertensia'' is

native Native may refer to: People * '' Jus sanguinis'', nationality by blood * '' Jus soli'', nationality by location of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Nat ...

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 The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...

. ''Mertensia'' is mostly restricted to alpine, subalpine, and

montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...

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 The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...

and

subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...

coastlines, and ''Mertensia virginica'', which is found from the

Appalachian Mountains The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...

west to Minnesota, Iowa, and Missouri. Most of the

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

are

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

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-national of a country acquires the nationality of that country after birth. The definition of naturalization by the International Organization for Migration of the ...

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 Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...

ate the

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

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 In law, to distinguish a case means a court decides the holding or legal reasoning of a precedent case that will not apply due to materially different facts between the two cases. Two formal constraints constrain the later court: the expresse ...

and some are possibly cryptic. Around 150

species name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...

s have been

published Publishing is the activities of making information, literature, music, software, and other content, physical or digital, available to the public for sale or free of charge. Traditionally, the term publishing refers to the creation and distribu ...

in ''Mertensia''.''Mertensia'' in

Tropicos Tropicos is an online botanical database containing taxonomic information on plants, mainly from the Neotropical realm ( Central, and South America South America is a continent entirely in the Western Hemisphere and mostly in the Sou ...

(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

According to

Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...

there are 52 accepted species in the genus.

Classification

''Mertensia'' is a member of the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

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 In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...

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 Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

analysis of

DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...

Taxonomy

The

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

for ''Mertensia'' is ''M. virginica''.''Mertensia'' In: Index Nominum Genericorum. In:

Regnum Vegetabile The International Association for Plant Taxonomy (IAPT) is an organization established to promote an understanding of plant biodiversity, facilitate international communication of research between botanists, and oversee matters of uniformity and ...

(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 ''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 Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

. 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 Sakhalin ( rus, Сахали́н, p=səxɐˈlʲin) is an island in Northeast Asia. Its north coast lies off the southeastern coast of Khabarovsk Krai in Russia, while its southern tip lies north of the Japanese island of Hokkaido. An islan ...

and the

Kuril Islands The Kuril Islands or Kurile Islands are a volcanic archipelago administered as part of Sakhalin Oblast in the Russian Far East. The islands stretch approximately northeast from Hokkaido in Japan to Kamchatka Peninsula in Russia, separating the ...

.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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

group consisting of three

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

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 genus, genera. It is the first work to consistently apply binomial nomenclature ...

'', Swedish botanist

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

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 University of Halle, Halle and University of Erlangen, Erlangen, where he received his ...

, 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 The International Plant Names Index (IPNI) describes itself as "a database of the names and associated basic bibliographical details of seed plants, ferns and lycophytes." Coverage of plant names is best at the rank of species and genus. It inclu ...

. (see ''External links'' below). based on their smaller and differently structured calyx, their different anther position, and the presence of

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

gland A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...

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 precisely 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 a ...

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

botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...

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 botany, botanist of the 19th century. His ''Darwiniana'' (1876) was considered an important explanation of how religion and science were not necessaril ...

,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 (19 May 1892 16 June 1976) was an American botanist who devoted most of his professional life to the study of the flora of Peru. Early life and education Born on 19 May 1891 in Rock Valley, Iowa, MacBride graduated from th ...

,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 (July 6, 1860 – July 25, 1931) was a Swedish-born, American botanist who was the first curator of the New York Botanical Garden Herbarium. Biography Per Axel Rydberg was born in Odh, Västergötland, Sweden and emigrated to ...

,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 () (5(17) April, 1893 – 18 December, 1955) was a Soviet botanist. He is known for developing a theory on the role of hybridization in plant evolution, and studying the flora of the Soviet Union and Eastern Europe. Ep ...

. 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 Lichenology is the branch of mycology that studies the lichens, symbiotic organisms made up of an intimate symbiotic association of a microscopic alga (or a cyanobacterium) with a Hypha, filamentous fungus. Lichens are chiefly characterized by th ...

. Gray defined the section Stenhammaria as consisting only of the

littoral The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely i ...

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 generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...

of ''Mertensia'', many species were discovered by

botanical expedition Botany, also called plant science, is the branch of natural science and biology studying plants, especially their anatomy, taxonomy, and ecology. A botanist or plant scientist is a scientist who specialises in this field. "Plant" and "botany ...

s 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 In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

analysis of DNA data has shown that many of the species are

polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...

, 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 A rank is a position in a hierarchy. It can be formally recognized—for example, cardinal, chief executive officer, general, professor—or unofficial. People Formal ranks * Academic rank * Corporate title * Diplomatic rank * Hierarchy ...

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

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

. 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 (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

s (> 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 The following terms are used to describe leaf plant morphology, morphology in the description and taxonomy (biology), taxonomy of plants. Leaves may be simple (that is, the leaf blade or 'lamina' is undivided) or compound (that is, the leaf blade ...

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

nutlet A nut is a fruit consisting of a hard or tough nutshell protecting a kernel which is usually edible. In general usage and in a culinary sense, many dry seeds are called nuts, but in a botanical context, "nut" implies that the shell does not open ...

s and

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

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

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

s of ''Mertensia''. A

molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleot ...

analysis has estimated that ''Mertensia'' diverged from ''Asperugo'' in the late

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

or early

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

. ''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 continental 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 ...

s, ''Mertensia'' spread southward and to lower elevation during periods of

Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate be ...

, 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 i ...

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 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...

land bridge In biogeography, a land bridge is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and colonize new lands. A land bridge can be created by marine regression, in which sea le ...

to North America. Most of the groups that originated in Asia are more diverse 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'')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 franciscana1.jpg, Franciscan bluebells ('' Mertensia franciscana'') Mertensia lanceolata NPS-1.jpg, Prairie bluebells ('' Mertensia lanceolata'') 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'') 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 a worldwide consortium of natural history, botanical, research, and national libraries working ...

): :
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
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''Mertensia''BoraginaceaeBoraginaleslamiidsasteridsEmbryophytaStreptophytinaStreptophytaViridiplantaeEukaryotaTaxonomyUniProt

''Mertensia''BoraginaceaeBoraginaleslamiidsasteridsEmbryophytaStreptophytinaStreptophytaViridiplantaeEukaryotaTaxonomy BrowserTaxonomy DatabaseTaxonomyNCBI
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''Mertensia''BoraginaceaeBoraginalesMagnoliopsidaTracheophytaPlantsGlobal Species

''Mertensia''BoraginaceaeAngiospermsBrowseThe Plant List

''Mertensia''

USDA Plants Profile
* * {{Taxonbar, from=Q765547 Boraginaceae genera Flora of Northern America