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.
The genus Calochortus includes mariposas 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 stems grow to over tall.
The flowers can be white, yellow, pink, red-orange, or purplish. They are up to wide, with 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. The insides of the petals are often very 'hairy'. These hairs, along with the nectaries, are often used in distinguishing species from each other.
The group includes herbaceous, perennial, and bulbous species.

Taxonomy

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 :
  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 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:
Patterson also determined at the time that concerted convergence and phylogenetic niche conservatism may have confounded the idea that Calochortaceae 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. and Bremer, Chase, and Stevens. Using similar DNA collection techniques to Patterson, 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 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 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 created a serpentine tolerance phylogeny. 18 serpentine tolerant species were found 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 found that 11 out of 18 species displayed only two origins of serpentine tolerance in evolutionary history.

Species

The genus contains the following species:
ImageScientific nameSubspeciesDistribution
Calochortus albus white globelilyCA, Baja California
Calochortus amabilis Diogenes' lanternCA
Calochortus ambiguus doubting mariposa lilyUT AZ NM Sonora
Calochortus amoenus purple globelilyCA
Calochortus apiculatus pointed-tip mariposa lilyBritish Columbia, Alberta, WA OR ID MT WY
Calochortus argillosusCA
Calochortus aureus golden mariposa lilyUT CO AZ NM
Calochortus balsensisOaxaca, Guerrero
Calochortus barbatus yellow globe lilyfrom Chihuahua to Oaxaca
Calochortus bruneaunis Bruneau mariposa lilyCA OR NV UT ID MT
Calochortus catalinae Santa Catalina mariposa lilyCA
Calochortus cernuusMorelos
Calochortus ciscoensisUT
Calochortus clavatus club-hair mariposa lily
  • Calochortus clavatus var. avius -rare,
  • Calochortus clavatus var. gracilis
  • Calochortus clavatus ssp. clavatus
  • Calochortus clavatus ssp. pallidus
  • Calochortus clavatus ssp. recurvifoliusrare
CA
Calochortus coeruleus beavertail grassCA
Calochortus concolor golden-bowl mariposa lilyCA, Baja California
Calochortus coxii Cox's mariposa lilyOR
Calochortus dunnii Dunn's mariposa lilyrareCA, Baja California
Calochortus elegans northwestern mariposa lily; star tulip
  • 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
  • CA OR WA ID MT
    Calochortus eurycarpus white mariposa lilyOR WA NV ID MT WY
    Calochortus excavatus Inyo mariposa lily - CA
    Calochortus exilisHidalgo
    Calochortus fimbriatus late-blooming mariposa lilyrareCA
    Calochortus flexuosus winding mariposa lilyCA NV UT CO AZ NM Baja California, Sonora
    Calochortus foliosusMichoacán
    Calochortus fuscusMexico
    Calochortus greenei Green's mariposa lily — rareCA OR
    Calochortus ghiesbreghtiiMexico, Guatemala
    Calochortus gunnisonii Gunnison's mariposa lily
  • Calochortus gunnisonii var. gunnisonii
  • Calochortus gunnisonii var. perpulcher
  • ID MT WY SD NE CO UT AZ NM
    Calochortus hartwegiiAguascalientes, Nayarit, Jalisco
    Calochortus howellii Howell's mariposa lilyOR
    Calochortus indecorus Sexton Mountain mariposa lilyOR - extinct
    Calochortus invenustus plain mariposa lilyCA NV
    Calochortus kennedyi desert mariposa lily
  • Calochortus kennedyi var. kennedyi
  • Calochortus kennedyi var. munzii
  • CA NV AZ Sonora, Chihuahua
    Calochortus leichtlinii smokey mariposaCA NV OR
    Calochortus longibarbatus longbeard mariposa lily
  • Calochortus longibarbatus var. longibarbatus
  • Calochortus longibarbatus var. peckii
  • CA OR WA
    Calochortus luteus yellow mariposa lilyCA
    Calochortus lyallii Lyall's mariposa lilyWA, British Columbia
    Calochortus macrocarpus sagebrush mariposa lily
  • Calochortus macrocarpus var. macrocarpus
  • Calochortus macrocarpus var. maculosus Nez Perce mariposa lily
  • CA NV OR WA ID MT, British Columbia
    Calochortus marcellaeNuevo León, Coahuila, Tamaulipas
    Calochortus mendozaeQuerétaro, San Luis Potosí
    Calochortus minimus Sierran mariposa lilyCA
    Calochortus monanthus Shasta River mariposa lilyCA
    Calochortus monophyllus yellow startulipCA OR
    Calochortus nigrescensOaxaca
    Calochortus nitidus broad-fruit mariposa lilyWA OR ID
    Calochortus nudus naked mariposa lilyCA OR
    Calochortus nuttallii sego lily, ND SD NE MT ID CO UT NV AZ NM
    Calochortus obispoensis San Luis mariposa lilyCA
    Calochortus palmeri Palmer's mariposa lily
  • Calochortus palmeri var. munzii Munz's mariposa lily
  • Calochortus palmeri var. palmeri — rare
  • CA
    Calochortus panamintensis Panamint Mountain mariposa lilyrareCA NV
    Calochortus persistens Siskiyou mariposa lilyrareCA OR
    Calochortus plummerae Plummer's mariposa lilyrareCA
    Calochortus pringleiMorelos, Puebla, Jalisco, Oaxaca
    Calochortus pulchellus Mount Diablo globelilyCA
    Calochortus raichei Cedars mariposa lilyCA
    Calochortus simulans San Luis Obispo mariposa lilyCA
    Calochortus spatulatusMexico
    Calochortus splendens splendid mariposa lilyCA, Baja California
    Calochortus striatus alkali mariposa lilyCA NV
    Calochortus subalpinus subalpine mariposa lily, Cascade mariposa lily, cat's ear lilyWA OR
    Calochortus superbus superb mariposaCA
    Calochortus syntrophus Callahan's mariposa lilyCA
    Calochortus tiburonensis Tiburon mariposaCA
    Calochortus tolmiei Tolmie's star-tulip, hairy pussy earsCA OR WA
    Calochortus umbellatus Oakland mariposa lilyCA
    Calochortus umpquaensis Umpqua mariposa lilyOR
    Calochortus uniflorus mariposa 'Cupido'CA OR
    Calochortus venustulusMexico
    Calochortus venustus butterfly mariposa, white mariposaCA
    Calochortus vestae Vesta's mariposa, Coast Range mariposaCA
    Calochortus weedii Weed's mariposaCA, Baja Calilfornia
    Calochortus westonii Shirley Meadow star-tulip, Weston's mariposaCA

    Etymology

    The word Calochortus is derived from Greek and means "beautiful grass".

    Distribution and habitat

    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.
    The plants grow in open areas in partial shade over a wide range of elevations. T. B. Patterson's phylogenetic analysis indicated highly localized speciation, with different floral syndromes being strongly linked to specific habitats, as follows:

    Uses

    Culinary

    The bulbs of many species were eaten by Native Americans, being eaten raw or gathered in the fall and boiled. The flower buds were eaten young and fresh. They were eaten by Mormon settlers, especially in 1848 as a famine food when their cultivated crops were attacked by crickets. 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.

    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.