''Brodiaea'' , also known by the

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 ...

cluster-lilies, is a

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are flowering plants whose seeds contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but ...

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. One school of thought places the genus in the family , while another school of thought places it in the subfamily

Brodiaeoideae Brodiaeoideae are a monocot subfamily of flowering plants in the family Asparagaceae, order Asparagales. They have been treated as a separate family, Themidaceae. They are native to Central America and western North America, from British Colum ...

of the family

Asparagaceae Asparagaceae (), known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is based on the edible garden asparagus, '' Asparagus officinalis''. This family includes both ...

.Jepson Herbarium, Jepson eFlora: ''Brodiaea'', family Themidaceae
. accessed 29 April 2016.Calflora: ''Brodiaea'', family Themidaceae
. accessed 1 May 2016. The

USDA The United States Department of Agriculture (USDA) is an United States federal executive departments, executive department of the Federal government of the United States, United States federal government that aims to meet the needs of commerc ...

Plants Database still classifies the genus ''Brodiaea'' in the family Liliaceae. ''Brodiaea'' species occur along the Pacific Coast region of

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

, from

British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...

throughout

California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...

into the Baja California Peninsula. They are especially common in

northern California Northern California (commonly shortened to NorCal) is a geocultural region that comprises the northern portion of the U.S. state of California, spanning the northernmost 48 of the state's List of counties in California, 58 counties. Northern Ca ...

.Kew World Checklist of Selected Plant Families
/ref>

Description

''Brodiaea'' species are

herbaceous 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 of ...

perennials, growing from corms. Between one and six narrow leaves are produced from the corm. The bare flowering stem ( scape) carries an umbel of flowers. Individual flowers have six blue to purple tepals, joined at the base to form a tube with free lobes at the mouth. The outer three tepal lobes are narrower than the inner three. In almost all species, inside the tepals and joined to their bases are three sterile stamens ( staminodes), resembling small

petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...

s, each opposite one of the outer tepals. Three normal

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 are also joined to the bases of the tepals and are placed opposite the inner ones. The base of the filaments of the stamens may be expanded into various shapes, such as flaps or wings. The size and shape of the staminodes and of the structures at the base of the filaments are important diagnostic characters. The compound pistil is formed of three

carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...

s forming a superior

ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...

with three locules. The style which emerges between the three stamens has a three-lobed stigma. The seeds are black.

Taxonomy

Nomenclature

The origin of the scientific name of the genus is somewhat tangled. Specimens of what is now called ''Brodiaea'' were first collected by Archibald Menzies, botanist to the Vancouver Expedition, in 1792. Menzies collected the plant from the vicinity of the Strait of Georgia, named "New Georgia" by

George Vancouver Captain (Royal Navy), Captain George Vancouver (; 22 June 1757 – 10 May 1798) was a Royal Navy officer and explorer best known for leading the Vancouver Expedition, which explored and charted North America's northwestern West Coast of the Uni ...

. The first published reference to the plant did not give it a name. This was in James Edward Smith's 1807 ''An introduction to physiological and systematical botany'', where Smith used it to argue that the tepals of liliaceous plants are

sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...

s rather than

s: The following year, early in 1808, Richard Salisbury published a description of the first ''Brodiaea'' species in '' The Paradisus Londinensis'', naming it ''Hookera coronaria'', the genus name being in honour of the illustrator William Hooker. Shortly afterwards, Smith named a moss genus '' Hookeria'', and in April 1808, he read to the

Linnean Society of London The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and Taxonomy (biology), taxonomy. It possesses several important biological specimen, manuscript a ...

a formal description of a new genus, based on the same species as Salisbury's ''Hookera coronaria'', naming the genus ''Brodiaea'' in honour of Scottish botanist James Brodie. Formal publication did not occur, however, until Smith's presentation went to print in 1810. George Boulger, writing in the ''Dictionary of National Biography'', says that Smith's actions were deliberately intended to deprive Salisbury of credit for the genus. If this was Smith's intention it was successful, since although Salisbury's genus name ''Hookera'' has priority over Smith's name ''Brodiaea'', names as similar as ''Hookera'' and ''Hookeria'' are considered to be confusing and a formal proposal to conserve the names ''Brodiaea'' and ''Hookeria'' over the name ''Hookera'' was accepted. ''Brodiaea'' is thus a "conserved name" or "''nomen conservandum''", shown by the abbreviation "''nom. cons.''" after the name in botanical sources. The type species is now '' Brodiaea coronaria'', and the original type, ''Brodiaea grandiflora'' Sm., is an illegitimate name.

Phylogeny and classification

''Brodiaea'' belongs to a group of 12 genera whose affinities were the subject of much controversy until the end of the 20th century. Salisbury treated them as a family which he named Themidaceae. Others placed this group at lower

taxonomic rank In biology, taxonomic rank (which some authors prefer to call nomenclatural rank because ranking is part of nomenclature rather than taxonomy proper, according to some definitions of these terms) is the relative or absolute level of a group of or ...

and usually included them in Liliaceae, Alliaceae, or

Amaryllidaceae The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants in the monocot order Asparagales. The family takes its name from the genus '' Amaryllis'' and is commonly known as the amaryl ...

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 ...

studies confirmed the suspicions of many that this group was misplaced, and consequently, the

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 ...

Themidaceae was resurrected in 1996. When the Angiosperm Phylogeny Group published the APG II system in 2003, Themidaceae was accepted as an optional family for those who wanted to

circumscribe In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...

families narrowly in the order

Asparagales Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...

. When the APG III system was published in 2009, the former Themidaceae was treated as a

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

, Brodiaeoideae, of the family

sensu lato. Some sources, such as ITIS, continue to use the

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 ...

groups of obsolete taxonomic systems. Other sources, such as the Angiosperm Phylogeny Website mostly follow the Angiosperm Phylogeny Group. Brodiaea (or brodeia) is also used as a common name to refer to three genera, ''Brodiaea'', '' Dichelostemma'', and '' Triteleia''. The latter two genera were once included as part of the genus ''Brodiaea''. The

monophyly 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 comm ...

of ''Brodiaea'' as presently defined is not entirely certain. It might be intermixed with ''Dichelostemma''.

Species

, the

World Checklist of Selected Plant Families The World Checklist of Selected Plant Families (usually abbreviated to WCSP) was an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected p ...

lists the following 17 species.Search for "Brodiaea", USDA Plants Profile for ''Brodiaea'' − "Subordinate Taxa" tab
. accessed 29 April 2016. English common names are from the ''Flora of North America''. # '' Brodiaea appendiculata'' Hoover - appendage cluster-lily - central California # '' Brodiaea californica'' Lindl. ex Lem. - California cluster-lily - northern California, southwestern Oregon # '' Brodiaea coronaria'' (Salisb.) Jeps. - harvest cluster-lily; Californian hyacinth - British Columbia, Washington, Oregon, much of California # '' Brodiaea elegans'' Hoover - elegant cluster-lily - western Oregon, most of California # '' Brodiaea filifolia'' S.Watson - threadleaf cluster-lily - southern California # '' Brodiaea insignis'' (Jeps.) Niehaus - Kaweah cluster-lily - Tulare County # '' Brodiaea jolonensis'' Eastw. - chaparral cluster-lily - southern California, northern

Baja California Baja California, officially the Free and Sovereign State of Baja California, is a state in Mexico. It is the northwesternmost of the 32 federal entities of Mexico. Before becoming a state in 1952, the area was known as the North Territory of B ...

# '' Brodiaea kinkiensis'' Niehaus - San Clemente Island cluster-lily - San Clemente Island # '' Brodiaea matsonii'' R.E.Preston - Shasta County # '' Brodiaea minor'' (Benth.) S.Watson (syn. ''Brodiaea purdyi'' Eastw.) - vernalpool cluster-lily - northern California # '' Brodiaea nana'' Hoover - northern California # '' Brodiaea orcuttii'' (Greene) Baker - Orcutt's cluster-lily - southern California, northern

# '' Brodiaea pallida'' Hoover - Chinese Camp cluster-lily - Calaveras and Tuolumne Counties # '' Brodiaea santarosae'' T.J.Chester - Santa Rosa basalt brodiaea - Riverside and

San Diego San Diego ( , ) is a city on the Pacific coast of Southern California, adjacent to the Mexico–United States border. With a population of over 1.4 million, it is the List of United States cities by population, eighth-most populous city in t ...

Counties # '' Brodiaea sierrae'' R.E.Preston -

Butte In geomorphology, a butte ( ) is an isolated hill with steep, often vertical sides and a small, relatively flat top; buttes are smaller landforms than mesas, plateaus, and table (landform), tablelands. The word ''butte'' comes from the French l ...

, Yuba and

Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...

Counties # '' Brodiaea stellaris'' S.Watson - starflower cluster-lily - Sonoma, Mendocino and Humboldt Counties # '' Brodiaea terrestris'' Kellogg - dwarf cluster-lily - southwestern Oregon and much of coastal and southern California ;formerly included Numerous other names have been coined using the name ''Brodiaea,'' referring to species now regarded as better suited to other genera ''( Androstephium Beauverdia Dandya Dichelostemma Leucocoryne Nothoscordum Tristagma Triteleia Triteleiopsis).''

Distribution and habitat

''Brodiaea'' species are confined to western North America, from

in the north, through the

West Coast of the United States The West Coast of the United States, also known as the Pacific Coast and the Western Seaboard, is the coastline along which the Western United States meets the North Pacific Ocean. The term typically refers to the Contiguous United States, contig ...

region, to northwestern

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

in the south. The majority of species 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 ...

. Many are adapted to serpentine soils or other soils with particular chemical compositions, resulting in limited distributions and several rare and endangered species. An example is '' Brodiaea pallida'', known only from two populations along the border between Tuolumne County and Calaveras County, California.

Cultivation

A number of species of ''Brodiaea'' are in cultivation. Species such as '' B. californica'' and '' B. coronaria'' are recommended for sunny positions in the garden, where they extend the flowering season of most ornamental bulbs, flowering in early summer rather than in spring. The flower heads ( umbels) of larger species can be dried for use as winter decorations. Smaller species, such as '' B. terrestris'', may be grown in a bulb frame or alpine house. pp. 19–21.

References

Bibliography

External links

*
Calflora Database: ''Brodiaea'' speciesJepson Manual eFlora (TJM2) treatment of ''Brodiaea''USDA Plants Profile for ''Brodiaea'' species (brodiaea)
{{Authority control Asparagaceae genera Flora of California