Calochortus

Calochortus is a genus of flowering plants in the lily family. The group includes herbaceous, perennial and bulbous species, all native to North America (primarily the Western United States).

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 tepals. Unlike most other Liliaceae, Calochortus tepals are in two series that differ in size and color. The outer three are generally narrower and more sepal-like, while the inner three are larger, usually with bright marks at the base, and are often described as petals. 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
 * Calochortus albus white globelily - CA, Baja California
 * Calochortus amabilis Diogenes' lantern - CA
 * Calochortus ambiguus doubting mariposa lily - UT AZ NM Sonora
 * Calochortus amoenus purple globelily - CA
 * Calochortus apiculatus pointed-tip mariposa lily - British Columbia, Alberta, WA OR ID MT WY
 * Calochortus argillosus - CA
 * Calochortus aureus golden mariposa lily - UT CO AZ NM
 * Calochortus balsensis - Oaxaca, Guerrero
 * Calochortus barbatus yellow globe lily - from Chihuahua to Oaxaca
 * Calochortus bruneaunis Bruneau mariposa lily - CA OR NV UT ID MT
 * Calochortus catalinae Santa Catalina mariposa lily — (threatened by development) CA
 * Calochortus cernuus - Morelos
 * Calochortus ciscoensis - UT
 * Calochortus clavatus club-hair mariposa lily - CA
 * Calochortus clavatus var. avius -rare,
 * Calochortus clavatus var. gracilis
 * Calochortus clavatus ssp. clavatus
 * Calochortus clavatus ssp. pallidus
 * Calochortus clavatus ssp. recurvifolius — rare
 * Calochortus coeruleus beavertail grass - CA
 * Calochortus concolor golden-bowl mariposa lily - CA, Baja California
 * Calochortus coxii Cox's mariposa lily - OR
 * Calochortus dunnii Dunn's mariposa lily — rare - CA, Baja California
 * Calochortus elegans northwestern mariposa lily; star tulip - CA OR WA ID MT
 * Calochortus elegans var. elegans elegant mariposa lily
 * Calochortus elegans var. nanus
 * Calochortus elegans var. oreophilus elegant mariposa lily
 * Calochortus elegans var. selwayensis Selway mariposa lily
 * Calochortus eurycarpus white mariposa lily - OR WA NV ID MT WY
 * Calochortus excavatus Inyo mariposa lily — (threatened by groundwater development) - CA
 * Calochortus exilis - Hidalgo
 * Calochortus fimbriatus late-blooming mariposa lily — rare - CA
 * Calochortus flexuosus winding mariposa lily - CA NV UT CO AZ NM Baja California, Sonora
 * Calochortus foliosus - Michoacán
 * Calochortus fuscus - Mexico
 * Calochortus greenei Green's mariposa lily — rare - CA OR
 * Calochortus ghiesbreghtii - Mexico, Guatemala
 * Calochortus gunnisonii Gunnison's mariposa lily - ID MT WY SD NE CO UT AZ NM
 * Calochortus gunnisonii var. gunnisonii
 * Calochortus gunnisonii var. perpulcher
 * Calochortus hartwegii - Aguascalientes, Nayarit, Jalisco
 * Calochortus howellii Howell's mariposa lily - OR
 * †Calochortus indecorus Sexton Mountain mariposa lily - OR - extinct
 * Calochortus invenustus plain mariposa lily - CA NV
 * Calochortus kennedyi desert mariposa lily - CA NV AZ Sonora, Chihuahua
 * Calochortus kennedyi var. kennedyi
 * Calochortus kennedyi var. munzii
 * Calochortus leichtlinii smokey mariposa - CA NV OR
 * Calochortus longibarbatus longbeard mariposa lily - CA OR WA
 * Calochortus longibarbatus var. longibarbatus — (threatened by grazing)
 * Calochortus longibarbatus var. peckii
 * Calochortus luteus yellow mariposa lily - CA
 * Calochortus lyallii Lyall's mariposa lily - WA, British Columbia
 * Calochortus macrocarpus sagebrush mariposa lily - CA NV OR WA ID MT, British Columbia
 * Calochortus macrocarpus var. macrocarpus
 * Calochortus macrocarpus var. maculosus Nez Perce mariposa lily
 * Calochortus marcellae - Nuevo León, Coahuila, Tamaulipas
 * Calochortus mendozae - Querétaro, San Luis Potosí
 * Calochortus minimus Sierran mariposa lily - CA
 * †Calochortus monanthus Shasta River mariposa lily — CA (presumed extinct)
 * Calochortus monophyllus yellow startulip - CA OR
 * Calochortus nigrescens - Oaxaca
 * Calochortus nitidus broad-fruit mariposa lily - WA OR ID
 * Calochortus nudus naked mariposa lily - CA OR
 * Calochortus nuttallii sego lily, (state flower of Utah) - ND SD NE MT ID CO UT NV AZ NM
 * Calochortus obispoensis San Luis mariposa lily - CA
 * Calochortus palmeri Palmer's mariposa lily - CA
 * Calochortus palmeri var. munzii Munz's mariposa lily
 * Calochortus palmeri var. palmeri — rare
 * Calochortus panamintensis Panamint Mountain mariposa lily — rare ca NV
 * Calochortus persistens Siskiyou mariposa lily — rare - CA OR
 * Calochortus plummerae Plummer's mariposa lily — rare - CA
 * Calochortus pringlei - Morelos, Puebla, Jalisco, Oaxaca
 * Calochortus pulchellus Mount Diablo globelily - CA
 * Calochortus raichei Cedars mariposa lily - CA
 * Calochortus simulans San Luis Obispo mariposa lily - CA
 * Calochortus spatulatus - Mexico
 * Calochortus splendens splendid mariposa lily - CA, Baja California
 * Calochortus striatus alkali mariposa lily - CA NV
 * Calochortus subalpinus subalpine mariposa lily, Cascade mariposa lily, cat's ear lily - WA OR
 * Calochortus superbus superb mariposa - CA
 * Calochortus syntrophus Callahan's mariposa lily - CA
 * Calochortus tiburonensis Tiburon mariposa - CA
 * Calochortus tolmiei Tolmie's star-tulip, hairy pussy ears - CA OR WA
 * Calochortus umbellatus Oakland mariposa lily - CA
 * Calochortus umpquaensis Umpqua mariposa lily - OR
 * Calochortus uniflorus mariposa 'Cupido' - CA OR
 * Calochortus venustulus - Mexico
 * Calochortus venustus butterfly mariposa, white mariposa - CA
 * Calochortus vestae Vesta's mariposa, Coast Range mariposa - CA
 * Calochortus weedii Weed's mariposa - CA, Baja Calilfornia
 * Calochortus westonii Shirley Meadow star-tulip, Weston's mariposa - CA

Distribution and habitat
The genus Calochortus includes approximately 70 species distributed from southwestern British Columbia, through California and Mexico, to northern Guatemala and eastwards to New Mexico, Nebraska and the Dakotas. Calochortus is the most widely dispersed genus of Liliaceae on the North American Pacific Coast. Of these, 28 species are endemic to California.

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, semideserts
 * Star-tulips: wet meadows
 * Cat's ears: montane woodlands
 * Fairy lanterns: oak woodlands, closed forests.

History
Calochortus was first proposed in 1814 by Frederick Pursh to accommodate a specimen—C. elegans—received from the Lewis and Clark expedition. 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 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 ):


 * 1) Eucalochortus
 * 2) * Ten basic chromosomes and two known cases of tetraploidy
 * 3) * Includes subsections Pulchelli, Eleganti, Nudi, Nitidi
 * 4) Mariposa
 * 5) * Basic chromosome numbers between six and nine
 * 6) * Includes subsections Venusti, Macrocarpi, Nuttalliani, Gunnisoniani
 * 7) Cyclobothra
 * 8) * Nine basic chromosomes
 * 9) * 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 and ndhF sequences) isolated from frozen or silica dried leaf tissue to develop a molecular phylogeny, 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 and phylogenetic niche conservatism 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 characteristics of rhizomes, inconspicuous flowers, berries, broad leaves, and reticulate venation.

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, Mariposa is paraphyletic, and Cyclobothra is polyphyletic. As a result of their research, Patterson and Givnish (2004) found that the two main factors of Calochortus speciation are:


 * 1) Poor dispersal caused by heavy, passively dispersed seeds
 * 2) 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 (form serpentine soils) habitat preferences or specific edaphic requirements, with several being endemic 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. 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.

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 settlers between 1853 and 1858 when famine threatened new immigrants in the Great Salt Lake Valley, due to crop failures. The bulbs are a starchy food source similar to a potato tuber.

Native Americans called Calochortus "sego". They used it as food, in ceremonies and as a traditional medicinal plant.

Cultivation
Some Calochortus species are cultivated as ornamental plants by specialty nurseries and botanic gardens to sell. The bulbs are planted for their flowers, in traditional, native plant, and wildlife gardens; in rock gardens; and in potted container gardens for those needing unwatered Summer dormancy.