''Calochortus'' is a genus of

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 lily family. The group includes

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 ...

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 ...

and

bulb In botany, a bulb is a short underground stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs duri ...

ous species, all native to

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 ...

(primarily the

Western United States The Western United States (also called the American West, the Western States, the Far West, the Western territories, and the West) is List of regions of the United States, census regions United States Census Bureau. As American settlement i ...

).Flora of North America, Vol. 26 Page 119 ''Calochortus'' Pursh, Fl. Amer. Sept. 1: 240. 1814.
/ref> The genus ''Calochortus'' includes mariposas (or mariposa lilies) with open wedge-shaped petals, globe lilies and fairy lanterns with globe-shaped flowers, and cat's ears and star tulips with erect pointed petals. The word ''Calochortus'' is derived from Greek and means "beautiful grass".

Description

''Calochortus'' flowers have six

tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...

s. Unlike most other

Liliaceae The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fai ...

, ''Calochortus'' tepals are in two series that differ in size and color. The outer three are generally narrower and more

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'' ...

-like, while the inner three are larger, usually with bright marks at the base, and are often described as

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. The flowers are borne on a stem that arises from a bulb, generally in the spring or early summer. Flowers can be white, yellow, pink, purple, bluish, or streaked. The insides of the petals are often very 'hairy'. These hairs, along with the nectaries, are often used in distinguishing species from each other. ;Species

Distribution and habitat

The genus ''Calochortus'' includes approximately 70 species distributed from southwestern

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 ...

, through

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 ...

and

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 ...

, to northern

Guatemala Guatemala, officially the Republic of Guatemala, is a country in Central America. It is bordered to the north and west by Mexico, to the northeast by Belize, to the east by Honduras, and to the southeast by El Salvador. It is hydrologically b ...

and eastwards to

New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...

Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...

and the

Dakotas The Dakotas, also known as simply Dakota, is a collective term for the U.S. states of North Dakota and South Dakota. It has been used historically to describe the Dakota Territory, and is still used for the collective heritage, culture, geo ...

. ''Calochortus'' is the most widely dispersed genus of

on the North American Pacific Coast.Dale, Nancy; ''Flowering Plants of the Santa Monica Mountains'', Capra Press, 1986; pg. 28 Of these, 28 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 ...

. In 1998, T.B. Patterson conducted a phylogenetic analysis of the genus, dividing it into seven main clades (see Subdivision update below). The study indicated highly localized speciation, so that different floral syndromes were strongly linked to specific habitats, as follows: *Mariposas: dry grasslands, open

chaparral Chaparral ( ) is a shrubland plant plant community, community found primarily in California, southern Oregon, and northern Baja California. It is shaped by a Mediterranean climate (mild wet winters and hot dry summers) and infrequent, high-intens ...

, semideserts *Star-tulips:

wet meadow A wet meadow is a type of wetland with soils that are Solubility, saturated for part or all of the growing season which prevents the growth of trees and brush. Debate exists whether a wet meadow is a type of marsh or a completely separate type of ...

s *Cat's ears: montane woodlands *Fairy lanterns:

oak woodland An oak woodland is a plant community with a tree canopy dominated by oaks (''Quercus spp.''). In terms of canopy closure, oak woodlands are intermediate between oak savanna, which is more open, and oak forest, which is more closed. Although the c ...

s, closed forests.

Taxonomy

History

''Calochortus'' was first proposed in 1814 by Frederick Pursh to accommodate a specimen—''

C. elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' ( ...

''—received from the

Lewis and Clark expedition The Lewis and Clark Expedition, also known as the Corps of Discovery Expedition, was the United States expedition to cross the newly acquired western portion of the country after the Louisiana Purchase. The Corps of Discovery was a select gro ...

. In the 1800s, several species were added to the genus; however, much mistakes in naming conventions led to confusion and minimal knowledge gained by the end of the century. In 1940, Francis Marion Ownbey wrote a comprehensive

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 ...

on ''Calochortus'', referencing morphological evidence, geographical distribution, and his own study of cytological material. Ownbey proposed a treatment dividing ''Calochortus'' into three sections (later corroborated by J.M. Beal): # Eucalochortus #* Ten basic chromosomes and two known cases of tetraploidy #* Includes subsections Pulchelli, Eleganti, Nudi, Nitidi # Mariposa #* Basic chromosome numbers between six and nine #* Includes subsections Venusti, Macrocarpi, Nuttalliani, Gunnisoniani # Cyclobothra #* Nine basic chromosomes #* Includes subsection Weediani In 1985, F.N. Rasmussen developed a new treatment splitting ''Calochortus'' from Liliaceae, moving it into a separate family—Calochortaceae—based on chromosomal evidence, septicidal fruit, and a Polygonum type embryo sac formation. Rasmussen found that the basic chromosome numbers of Calochortus vary between seven and twenty.

Subdivision update

In the late 1990s and early 2000s, Thomas B. Patterson and Thomas J. Givnish gathered additional evidence to create a new ''Calochortus'' treatment, subdividing it into seven sections and providing reasoning behind ''Calochortus'' being separate from Liliaceae. In 1999, Patterson used cpDNA (specifically

rbcL Ribulose-1,5-bisphosphate carboxylase/oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the Photosynthesis#Light-independent reactions, light-independent (or "dark") part of photosyn ...

and ndhF sequences) isolated from frozen or silica dried leaf tissue to develop a

molecular phylogeny 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 ...

, finding that ''Calochortus'' should be divided into seven major clades based on geographic location: * Bay Area * Pacific Northwest * San Diego * Great Basin- Rocky Mountains * Coast Ranges- Sierra Nevada * Southwestern California * Central Mexico Patterson also determined at the time that concerted

convergence Convergence may refer to: Arts and media Literature *''Convergence'' (book series), edited by Ruth Nanda Anshen *Convergence (comics), "Convergence" (comics), two separate story lines published by DC Comics: **A four-part crossover storyline that ...

and

phylogenetic niche conservatism The term phylogenetic niche conservatism has seen increasing use in recent years in the scientific literature, though the exact definition has been a matter of some contention. Fundamentally, phylogenetic niche conservatism refers to the tendency ...

may have confounded the idea that Calochortaceae (''Calochortus'') and Liliaceae are closely related. In 2002, Patterson and Givnish expanded on these arguments, showing that concerted convergence was demonstrated through independent evolution of characteristics such as bulbs and showy flowers and the distinct differences of these appearing as a result of survival in specific habitats. Regarding phylogenetic niche conservatism, Patterson and Givnish make the argument that this phenomenon is present in the

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...

characteristics of

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, inconspicuous flowers, berries, broad leaves, and

reticulate venation A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, fl ...

. In 2004, Patterson and Givnish made the shift to lump ''Calochortus'' within Liliaceae within their paper per the recommendations of Bremer et al. (2003) and Bremer, Chase, and Stevens (1998). Using similar DNA collection techniques to Patterson (1999), Patterson and Givnish developed a more detailed molecular phylogeny, comparing the seven recently determined sections to Ownbey's original three and finding that Ownbey's Eucalochortus section is

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 ...

, Mariposa is

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 ...

, and Cyclobothra is

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 ...

. As a result of their research, Patterson and Givnish (2004) found that the two main factors of ''Calochortus''

speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...

are: # Poor dispersal caused by heavy, passively dispersed seeds # Chromosomal evolution allowing different clades to "double up" and radiate sympatrically without hybridizing

Serpentine tolerance

Within ''Calochortus'', almost one-third of species are characterized by

ultramafic Ultramafic rocks (also referred to as ultrabasic rocks, although the terms are not wholly equivalent) are igneous and meta-igneous rocks with a very low silica content (less than 45%), generally >18% MgO, high FeO, low potassium, and are usua ...

(form

serpentine soil Serpentine soil is an uncommon soil type produced by weathered ultramafic rock such as peridotite and its metamorphic derivatives such as serpentinite. More precisely, serpentine soil contains minerals of the serpentine subgroup, especially an ...

s) habitat preferences or specific

edaphic Edaphology (from Greek , ''edaphos'' 'ground' + , '' -logia'') is concerned with the influence of soils on living beings, particularly plants. It is one of two main divisions of soil science, the other being pedology. Edaphology includes the st ...

requirements, with several being

to their environments. Thus, scientists have used serpentine tolerance in understanding evolutionary relationships within the genus. For instance, Patterson and Givnish (2004) created a serpentine tolerance

phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...

. 18 serpentine tolerant species were found (classified by occurring in whole or in part on serpentine soils) and the largest presence of tolerance was found in the Bay Area and Pacific Northwest clades—areas with unusually high numbers of serpentine rocks at the Earth's surface. In addition, Patterson and Givnish (2004) found that 11 out of 18 species displayed only two origins of serpentine tolerance in evolutionary history.

Uses

Culinary

The bulbs of many species were eaten by Native Americans. These bulbs were eaten raw or gathered in the fall and boiled, and the flower buds when young and fresh. They were eaten by the

Mormon Mormons are a religious and cultural group related to Mormonism, the principal branch of the Latter Day Saint movement started by Joseph Smith in upstate New York during the 1820s. After Smith's death in 1844, the movement split into several ...

settlers between 1853 and 1858 when famine threatened new immigrants in the

Great Salt Lake The Great Salt Lake is the largest saltwater lake in the Western Hemisphere and the eighth-largest terminal lake in the world. It lies in the northern part of the U.S. state of Utah and has a substantial impact upon the local climate, partic ...

Valley, due to crop failures. The bulbs are a starchy food source similar to a potato tuber. Some Native Americans called ''Calochortus'' "sego". They used it as food, in ceremonies and as a traditional

medicinal plant Medicinal plants, also called medicinal herbs, have been discovered and used in traditional medicine practices since prehistoric times. Plants synthesize hundreds of chemical compounds for various functions, including Plant defense against h ...

Cultivation

Some ''Calochortus'' species are cultivated as

ornamental plants Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...

by specialty nurseries and botanic gardens to sell. The bulbs are planted for their flowers, in traditional,

native plant In biogeography, a native species is indigenous to a given region or ecosystem if its presence in that region is the result of only local natural evolution (though often popularised as "with no human intervention") during history. The term is equi ...

, and

wildlife garden A wildlife garden (or habitat garden or backyard restoration) is an Biophysical environment, environment created with the purpose to serve as a sustainable haven for surrounding wildlife. Wildlife gardens contain a variety of habitats that cater t ...

s; in

rock garden A rock garden, also known as a rockery and formerly as a rockwork, is a garden, or more often a part of a garden, with a landscaping framework of rocks, stones, and gravel, with planting appropriate to this setting. Usually these are small ...

s; and in potted container gardens for those needing unwatered Summer dormancy.

Notes

References

*
Treatment from the Jepson Manual (TJM93)
* Gerritsen, Mary E and Parsons, R. ''Calochortus. Mariposa Lilies and Their Relatives''. Timber Press, 2007.
Pacific Bulb Society

External links

* *
Calflora Database: ''Calochortus''
— ''all species native to California''.
photos by Mark Egger, Flickriver search for ''Calochortus''
many photos of many species
Jepson Manual (TJM93): Key to California ''Calochortus'' speciesGenus overview, Key to North American species

Gallery of Photos. US and Mexican Calochortus Species and Natural Hybrids
{{Authority control Liliaceae genera Taxa named by Frederick Traugott Pursh