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Calochortus

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Calochortus
Sego Lily (Calochortus nuttallii)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Liliales
Family: Liliaceae
Subfamily: Calochortoideae
Genus: Calochortus
Pursh
Type species
Calochortus elegans
Synonyms[2]
  • Cyclobothra D.Don
  • Mariposa (Alph.Wood) Hoover

Calochortus /ˌkæləˈkɔːrtəs, -loʊ-/[3][4] 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).[5][6]

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".[5]

Description

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Calochortus flowers have six tepals. Unlike most other Liliaceae, Calochortus tepals are in two series that differ in size and color.[7] The outer three are generally narrower and more sepal-like, while the inner three are larger, usually with bright marks at the base,[8] and are often described as petals.[9] 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.[5]

Species[2][10][11][12][13]


Image Scientific name Subspecies Distribution
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
  • Calochortus clavatus var. avius -rare,
  • Calochortus clavatus var. gracilis
  • Calochortus clavatus ssp. clavatus
  • Calochortus clavatus ssp. pallidus
  • Calochortus clavatus ssp. recurvifolius — rare
 CA
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
  • 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 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
  • Calochortus gunnisonii var. gunnisonii
  • Calochortus gunnisonii var. perpulcher
 ID MT WY SD NE CO UT AZ NM
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
  • Calochortus kennedyi var. kennedyi
  • Calochortus kennedyi var. munzii
 CA NV AZ Sonora, Chihuahua
Calochortus leichtlinii smokey mariposa CA NV OR
Calochortus longibarbatus longbeard mariposa lily
  • Calochortus longibarbatus var. longibarbatus — (threatened by grazing)
  • Calochortus longibarbatus var. peckii
 CA OR WA
Calochortus luteus yellow mariposa lily CA
Calochortus lyallii Lyall's mariposa lily WA, 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 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
  • Calochortus palmeri var. munzii Munz's mariposa lily
  • Calochortus palmeri var. palmeri — rare
 CA
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
120px 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

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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.[7] Of these, 28 species are endemic to California.[14]

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:[15]

Taxonomy

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History

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Calochortus was first proposed in 1814 by Frederick Pursh to accommodate a specimen—C. elegans—received from the Lewis and Clark expedition. [16] 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. [16]

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[17]):

  1. Eucalochortus
    • Ten basic chromosomes and two known cases of tetraploidy
    • Includes subsections Pulchelli, Eleganti, Nudi, Nitidi
  2. Mariposa
    • Basic chromosome numbers between six and nine
    • Includes subsections Venusti, Macrocarpi, Nuttalliani, Gunnisoniani
  3. 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. [18] Rasmussen found that the basic chromosome numbers of Calochortus vary between seven and twenty.

Subdivision update

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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:[19]

  • 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. [20] 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)[21] and Bremer, Chase, and Stevens (1998).[22] 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.[23] 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

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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.[24] 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.[23] In addition, Patterson and Givnish (2004) found that 11 out of 18 species displayed only two origins of serpentine tolerance in evolutionary history.

Uses

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Culinary

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The bulbs of many species were eaten by Native Americans.[25] These bulbs were eaten raw or gathered in the fall and boiled, and the flower buds when young and fresh.[25] 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.[16]

Native Americans called Calochortus "sego". They used it as food, in ceremonies and as a traditional medicinal plant.[25]

Cultivation

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Some Calochortus species are cultivated as ornamental plants by specialty nurseries and botanic gardens to sell.[26] 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.

See also

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Notes

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  1. ^ Tropicos, Calochortus Pursh
  2. ^ a b Kew World Checklist of Selected Plant Families
  3. ^ "Calochortus". Dictionary.com Unabridged (Online). n.d. Retrieved 2016-01-22.
  4. ^ Sunset Western Garden Book, 1995:606–607
  5. ^ a b c Flora of North America, Vol. 26 Page 119 Calochortus Pursh, Fl. Amer. Sept. 1: 240. 1814.
  6. ^ Pursh, Frederick Traugott. 1813. Flora Americae Septentrionalis; or, A systematic arrangement and description of the plants of North America. Containing, besides what have been described by preceding authors, many new and rare species, collected during twelve years travels and residence in that country 1: 240 in English and Latin
  7. ^ a b Dale, Nancy; Flowering Plants of the Santa Monica Mountains, Capra Press, 1986; pg. 28
  8. ^ Mathew, Brian (1987). The Smaller Bulbs. London: B.T. Batsford. p. 22. ISBN 978-0-7134-4922-8.
  9. ^ Taylor, Ronald J. (1994) [1992]. Sagebrush Country: A Wildflower Sanctuary (rev. ed.). Missoula, MT: Mountain Press Pub. Co. p. 74. ISBN 0-87842-280-3. OCLC 25708726.
  10. ^ Gerritsen, Mary E and Parsons, R. Calochortus. Mariposa Lilies and Their Relatives. Timber Press, 2007.
  11. ^ Biota of North America Program 2034 county distribution maps
  12. ^ Espejo Serena, A. & López-Ferrari, A.R. (1994). Las Monocotiledóneas Mexicanas una Sinopsis Florística 1(3): 1-74. Consejo Nacional de la Flora de México, México D.F.
  13. ^ Gerritsen, M.E. & Parsons, R. (2007). Calochortus: Mariposa lilies & their relatives: 1-232. Timber press, Inc. Portland, U.S.A.
  14. ^ USDA Natural Resource Conservation Service, Plant Profile for Calochortus Pursh; Data contributed by John K. Kartesz and USDA-NRCS National Plant Data Center
  15. ^ P. L. Fiedler & R. K. Zebell, Flora of North America; 18. Calochortus Pursh, Fl. Amer. Sept. 1: 240. 1814.
  16. ^ a b c Ownbey, Marion (1940). "A Monograph of the Genus Calochortus". Annals of the Missouri Botanical Garden. 27 (4): 371–560. doi:10.2307/2394384. JSTOR 2394384.
  17. ^ Beal, J. M.; Ownbey, Marion (1943). "Cytological Studies in Relation to the Classification of the Genus Calochortus. III". Botanical Gazette. 104 (4): 553–562. doi:10.1086/335169. ISSN 0006-8071.
  18. ^ Rasmussen, F. N. (1985), Dahlgren, Rolf M. T.; Clifford, H. Trevor; Yeo, Peter F. (eds.), "Superorder Liliiflorae", The Families of the Monocotyledons: Structure, Evolution, and Taxonomy, Berlin, Heidelberg: Springer, pp. 107–274, doi:10.1007/978-3-642-61663-1_11, ISBN 978-3-642-61663-1, retrieved 2023-12-27
  19. ^ Patterson, TB. 1999. Phylogeny, biogeography, and evolutionary trends in the core Liliales and Calochortus (Calochortaceae): Insights from DNA sequenced data (Ph.D.). United States -- Wisconsin: The University of Wisconsin - Madison.
  20. ^ Patterson, Thomas B.; Givnish, Thomas J. (2002). "PHYLOGENY, CONCERTED CONVERGENCE, AND PHYLOGENETIC NICHE CONSERVATISM IN THE CORE LILIALES: INSIGHTS FROM rbcL AND ndhF SEQUENCE DATA". Evolution. 56 (2): 233–252. doi:10.1111/j.0014-3820.2002.tb01334.x. ISSN 0014-3820. PMID 11926492.
  21. ^ Bremer, B; Bremer, K; Chase, MW; Reveal, JL; Soltis, DE; Soltis, PS; Stevens, PF; Anderberg, AA; Fay, MF; Goldblatt, P; Judd, WS; Kallersjo, M; Karehed, J; Kron, KA; Lundberg, J (2003). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II". Botanical Journal of the Linnean Society. 141 (4): 399–436. doi:10.1046/j.1095-8339.2003.t01-1-00158.x. ISSN 0024-4074.
  22. ^ Bremer, K; Chase, MW; Stevens, PF (1998). "An Ordinal Classification for the Families of Flowering Plants". Annals of the Missouri Botanical Garden. 85 (4): 531. doi:10.2307/2992015. JSTOR 2992015.
  23. ^ a b Patterson, Thomas B.; Givnish, Thomas J. (2004). "Geographic cohesion, chromosomal evolution, parallel adaptive radiations, and consequent floral adaptations in Calochortus (Calochortaceae): evidence from a cpDNA phylogeny". New Phytologist. 161 (1): 253–264. doi:10.1046/j.1469-8137.2003.00951.x. ISSN 0028-646X.
  24. ^ Fiedler, Peggy Lee (1985). "Heavy Metal Accumulation and the Nature of Edaphic Endemism in the Genus Calochortus (Liliaceae)". American Journal of Botany. 72 (11): 1712–1718. doi:10.2307/2443728. JSTOR 2443728.
  25. ^ a b c "University of Michigan at Dearborn, Native American Ethnobotany: Calochortus". Archived from the original on 2013-12-04. Retrieved 2015-04-17.
  26. ^ "Telos Rare Bulbs Nursery database: Calochortus". Archived from the original on 2020-04-06. Retrieved 2015-04-17.

References

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