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Introductory

SPECIES: Claytonia perfoliata | Miner's-Lettuce
ABBREVIATION : CLAPER SYNONYMS : Montia perfoliata (Donn.) Howell [18,22,27,36] SCS PLANT CODE : CLPE MOPE2 COMMON NAMES : miner's-lettuce claspleaf miner's-lettuce Indian-lettuce TAXONOMY : The currently accepted scientific name of miner's-lettuce is Claytonia perfoliata Donn. (Portulacaceae) [38,44]. The Claytonia perfoliata complex is a polyploid group of considerable complexity, with several subspecies and many ecotypes [39,40]. The following subspecies are recognized: C. perfoliata spp. perfoliata [39] C. perfoliata ssp. mexicana (Rydb.) John M. Miller & Chambers [38] C. perfoliata ssp. viridis (A. Davidson) Fellows [39]. Varieties under the synonym Montia perfoliata are listed in several floras [19,27,35,36,38]. Miner's-lettuce hybridizes with C. parviflora, C. sibirica, and C. rubra [38,40]. LIFE FORM : Forb FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Robin F. Matthews, December 1993 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Matthews, Robin F. 1993. Claytonia perfoliata. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Claytonia perfoliata | Miner's-Lettuce
GENERAL DISTRIBUTION : Miner's-lettuce is distributed from British Columbia south to Guatemala and east to North Dakota, South Dakota, Wyoming, Utah, and Arizona [17,18,19,22,36,38]. ECOSYSTEMS : FRES20 Douglas-fir FRES21 Ponderosa pine FRES22 Western white pine FRES23 Fir - spruce FRES27 Redwood FRES28 Western hardwoods FRES29 Sagebrush FRES30 Desert shrub FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES36 Mountain grasslands FRES37 Mountain meadows FRES42 Annual grasslands STATES : AZ CA CO ID MT NV ND OR SD UT WA WY BC MEXICO ADMINISTRATIVE UNITS : BICA CHIS CRLA DEVA GLAC GRCA JODA JOTR KICA LAME LABE NOCA OLYM ORPI PINN PORE REDW SAGU SAJH SAMO SEQU TICA WHIS YOSE ZION BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 4 Sierra Mountains 5 Columbia Plateau 6 Upper Basin and Range 7 Lower Basin and Range 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains 12 Colorado Plateau 15 Black Hills Uplift 16 Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K005 Mixed conifer forest K006 Redwood forest K007 Red fir forest K009 Pine - cypress forest K010 Ponderosa shrub forest K011 Western ponderosa forest K012 Douglas-fir forest K013 Cedar - hemlock - pine forest K015 Western spruce - fir forest K016 Eastern ponderosa forest K017 Black Hills pine forest K018 Pine - Douglas-fir forest K019 Arizona pine forest K020 Spruce - fir - Douglas-fir forest K023 Juniper - pinyon woodland K024 Juniper steppe woodland K025 Alder - ash forest K026 Oregon oakwoods K028 Mosaic of K002 and K026 K029 California mixed evergreen forest K030 California oakwoods K033 Chaparral K034 Montane chaparral K035 Coastal sagebrush K037 Mountain-mahogany - oak scrub K038 Great Basin sagebrush K039 Blackbrush K041 Creosotebush K048 California steppe K051 Wheatgrass - bluegrass K055 Sagebrush steppe SAF COVER TYPES : 206 Engelmann spruce - subalpine fir 207 Red fir 210 Interior Douglas-fir 211 White fir 213 Grand fir 215 Western white pine 217 Aspen 221 Red alder 222 Black cottonwood - willow 224 Western hemlock 229 Pacific Douglas-fir 230 Douglas-fir - western hemlock 232 Redwood 233 Oregon white oak 234 Douglas-fir - tanoak - Pacific madrone 235 Cottonwood - willow 237 Interior ponderosa pine 238 Western juniper 239 Pinyon - juniper 243 Sierra Nevada mixed conifer 244 Pacific ponderosa pine - Douglas-fir 245 Pacific ponderosa pine 246 California black oak 247 Jeffrey pine 248 Knobcone pine 249 Canyon live oak 250 Blue oak - Digger pine 255 California coast live oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : In addition to the species previously listed under DISTRIBUTION AND OCCURRENCE information, miner's-lettuce is associated with bigcone Douglas-fir (Pseudotsuga macrocarpa), interior live oak (Quercus wislizenii), and Coulter pine (Pinus coulteri) [40].

VALUE AND USE

SPECIES: Claytonia perfoliata | Miner's-Lettuce
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Miner's-lettuce is preferred by cattle in blue oak (Quercus douglasii) savannas in California [5]. It is also grazed by pocket gophers [6]. Mourning doves, California quail, and other seed-eating birds consume the fruits [24,41]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : The nutritional composition of miner's-lettuce has been determined to be 37.1 percent protein, 42.5 percent total carbohydrate, and 12.4 percent crude fiber. The calcium:phosphorus ratio is 0.66:1.0 [37]. COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : NO-ENTRY OTHER USES AND VALUES : The blossoms, leaves, and stems of miner's-lettuce may be eaten by humans at any time during the growing season. They are eaten raw or cooked, and are a good source of vitamin C [11,37]. Historically, miner's-lettuce was used as a salad plant and potherb by white settlers and Native Americans [19]. It was also used to avert or cure scurvy [37]. MANAGEMENT CONSIDERATIONS : In California, density and overall yield of miner's-lettuce is greater in bracken fern communities than in surrounding grasslands [14,15]. This may be due to increased moisture availability in winter and early spring, when bracken fern is dormant [14]. Miner's-lettuce is a host to the beet western yellows virus, which is spread by aphids [43]. Purslane sawfly larvae, which consume the seeds, afford some biological control over miner's lettuce [42,43].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Claytonia perfoliata | Miner's-Lettuce
GENERAL BOTANICAL CHARACTERISTICS : Miner's-lettuce is a native winter or spring annual. It is branched from the base with stems growing up to 14 inches (35 cm) tall. Leaves are mostly basal, simple, and 2.4 to 8.0 inches (6-20 cm) long, including the stalk. Miner's-lettuce has two stem leaves that fuse to form a disc just below the flower stalk. The elongate stalk bears numerous small flowers. Fruits are tiny, three-valved capsules containing one to three seeds. Roots are fibrous [11,22,27,36]. RAUNKIAER LIFE FORM : Therophyte REGENERATION PROCESSES : Miner's-lettuce reproduces by seed [24,33,34]. Selfing is the most common method of pollination, but insect pollination also occurs. Seeds are dispersed by explosive dehiscence. They are capable of immediate germination [39]. SITE CHARACTERISTICS : Miner's-lettuce usually occurs on moist or vernally moist sites [10,14,18,19,22]. Miller [39] reported it from a variety of substrates including river silt, sand, gravel, road tar, loam, rock crevices, talus, and scree. He also found it on burned sites. Some polyploids occur on specialized, distinctive sites. The Columbia River Gorge octoploid, for example, occurs only on north-facing basalt talus slopes or cliff faces. Other polyploids are more plastic in site requirements [40]. In California, miner's-lettuce is most common below 6,500 feet (2,000 m) [38]; in Arizona it grows at elevations of 2,500 to 7,500 feet (750-2,270 m) [19]; in Utah it grows at elevations of 2,600 to 10,890 feet (800-3,300 m) [36]. SUCCESSIONAL STATUS : Facultative Seral Species Miner's-lettuce occurs in all seral stages. It often colonizes disturbed sites, particularly following fire [22,24]. Miner's lettuce is also found on virgin fields dominated by bluebunch wheatgrass (Pseudoroegneria spicata) and Sandberg bluegrass (Poa secunda) in southeastern Washington [8]. However, miner's-lettuce is shade tolerant [22,26,27] and is more prominent under a canopy than in openings in oak savanna, western white pine (Pinus monticola), and antelope bitterbrush (Purshia tridentata) communities [3,23,26]. SEASONAL DEVELOPMENT : Time from germination to flowering varied from 33 to 90 days in a Columbia River Gorge population [39]. Miner's-lettuce flowers from February to May in Arizona and California [19,27]. In Utah, it flowers from June to July [1].

FIRE ECOLOGY

SPECIES: Claytonia perfoliata | Miner's-Lettuce
FIRE ECOLOGY OR ADAPTATIONS : Miner's-lettuce has long-lived seeds that are stored in the soil [33] and germinate following fire [34]. It is a prolific seeder [24]; mass flowering in the years immediately following a fire recharges the seedbank [33]. Miner's-lettuce can develop high cover on exposed soil in full sun [31]. POSTFIRE REGENERATION STRATEGY : Ground residual colonizer (on-site, initial community) Secondary colonizer - on-site seed

FIRE EFFECTS

SPECIES: Claytonia perfoliata | Miner's-Lettuce
IMMEDIATE FIRE EFFECT ON PLANT : Miner's-lettuce is probably killed by fire. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Miner's-lettuce was present in the first growing season after the stand-destroying Marble-Cone wildfire in the Santa Lucia Range of California in August 1977. Peak cover was reached in postfire year 2 and declined by postfire year 3. Percent frequency of miner's-lettuce on two study sites that had been dominated by Coulter pine follows [16]: Site 1978 1979 1980 ______________________________________________ Chews Ridge site 1 9 36 8 Chews Ridge site 2 7 48 2 Miner's-lettuce is common in recently burned chaparral [20]. A year after a fire in chaparral in the Sierra Nevada foothills, miner's-lettuce had high seed production on moist north-east slopes. Postfire cover quickly exceeded prefire levels [24]. Miner's-lettuce was also present the year following a severe fire in a chaparral riparian zone in the Los Padres National Forest, California, but its frequency was reduced by postfire year 2 [9]. Miner's-lettuce is also common after fire in more northern portions of its range. It was present in the first growing season after a fall wildfire in ponderosa pine (Pinus ponderosa) stands in the Selway-Bitterroot Wilderness, Idaho, and had increased in frequency by postfire year 3 [25]. In burned ponderosa pine shelterwood cut units in Idaho, miner's-lettuce was present in postfire year 1 on sites burned with dry fuels, but was not present on sites burned with moist fuels. It also was not present in the preburn vegetation or in unburned control plots [30]. Miner's-lettuce was present in the first growing season following the stand-destroying Pattee Canyon wildfire in a Douglas-fir (Pseudotsuga menziesii)/ninebark (Physocarpus malvaceus) habitat type in west-central Montana [7]. It was still present in the herbaceous layer 10 years later [34]. Basin big sagebrush (Artemisia tridentata spp. tridentata)-Idaho fescue (Festuca idahoensis)-bluebunch wheatgrass communities at the John Day Fossil Beds National Monument in eastern Oregon were burned in the spring and fall. Although not in the preburn vegetation, miner's-lettuce was present in trace amounts (less than 2% frequency) the summer after the fall prescribed fire. It was not present after the spring fire or in control plots [29]. Miner's-lettuce establishes after fire in disturbed and climax grasslands in southeastern Washington [8]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Rapid growth of miner's-lettuce after fire in chaparral in the Sierra Nevada foothills contributes to an increased food supply for flocking bird species such as mourning dove and western meadowlark [24].

REFERENCES

SPECIES: Claytonia perfoliata | Miner's-Lettuce
REFERENCES : 1. Allman, Verl Phillips. 1953. A preliminary study of the vegetation in an exclosoure in the chaparral of the Wasatch Mountains, Utah. Utah Academy Proceedings. 30: 63-78. [9096] 2. Bainbridge, David A. 1990. Soil solarization for restorationists. Restoration & Management Notes. 8(2): 96-98. [14160] 3. Bartolome, James W. 1987. California annual grassland and oak savannah. Rangelands. 9(3): 122-125. [2861] 4. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434] 5. Borchert, Mark I.; Davis, Frank W.; Michaelson, Joel; Oyler, Lynn Dee. 1989. Intractions of factors affectting seedling recruitment of blue oak (Quercus douglasii) in California. Ecology. 70(2): 389-404. [6626] 6. Burton, Douglas H.; Black, Hugh C. 1978. Feeding habits of Mazama pocket gophers in south-central Oregon. Journal of Wildlife Management. 42(2): 383-390. [15818] 7. Crane, M. F.; Habeck, James R.; Fischer, William C. 1983. Early postfire revegetation in a western Montana Douglas-fir forest. Res. Pap. INT-319. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 29 p. plus chart. [710] 8. Daubenmire, Rexford F. 1975. Plant succession on abandoned fields, and fire influences, in a steppe area in southeastern Washington. Northwest Science. 49(1): 36-48. [745] 9. Davis, Frank W.; Keller, Edward A.; Parikh, Anuja; Florsheim, Joan. 1989. Recovery of the chaparral riparian zone after wildfire. In: Protection, management, and restoration for the 1990's: Proceedings of the California riparian systems conference; 1988 September 22-24; Davis, CA. Gen. Tech. Rep. PSW-110. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 194-203. [13883] 10. Dorn, Robert D. 1984. Vascular plants of Montana. Cheyenne, WY: Mountain West Publishing. 276 p. [819] 11. Elias, Thomas S.; Dykeman, Peter A. 1982. Field guide to North American edible wild plants. New York: Outdoor Life Books. 286 p. [21104] 12. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 13. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 14. Gliessman, S. R. 1976. Allelopathy in a broad spectrum of environments as illustrated by bracken. Botanical Journal of the Linnean Society. 73: 95-104. [9135] 15. Gliessman, S. R.; Muller, C. H. 1978. The allelopathic mechanisms of dominance in bracken (Pteridium aquilinum) in southern California. Journal of Chemical Ecology. 4(3): 337-362. [9973] 16. Griffin, James R. 1982. Pine seedlings, native ground cover, and Lolium multiflorum on the Marble-Cone burn, Santa Lucia Range, California. Madrono. 29(3): 177-188. [4935] 17. Harrington, H. D. 1964. Manual of the plants of Colorado. 2d ed. Chicago: The Swallow Press Inc. 666 p. [6851] 18. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. [1168] 19. Kearney, Thomas H.; Peebles, Robert H.; Howell, John Thomas; McClintock, Elizabeth. 1960. Arizona flora. 2d ed. Berkeley, CA: University of California Press. 1085 p. [6563] 20. Keeley, Jon E.; Keeley, Sterling C. 1986. Chaparral and wildfires. Fremontia. 14(3): 18-21. [18365] 21. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 22. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. [13798] 23. Larsen, J. A. 1922. Effect of removal of the virgin white pine stand upon the physical factors of site. Ecology. 3(4): 302-305. [12935] 24. Lawrence, George E. 1966. Ecology of vertebrate animals in relation to chaparral fire in the Sierra Nevada foothills. Ecology. 47(2): 278-291. [147] 25. Merrill, Evelyn H.; Mayland, Henry F.; Peek, James M. 1980. Effects of a fall wildfire on herbacious vegetation on xeric sites in the Selway-Bitterroot Wilderness, Idaho. Journal of Range Management. 33(5): 363-367. [1642] 26. Monsen, Stephen B.; Shaw, Nancy L. 1983. Seeding antelope bitterbrush with grasses on south-central Idaho rangelands--a 39-year response. In: Tiedemann, Arthur R.; Johnson, Kendall L., compilers. Proceedings--research and management of bitterbrush and cliffrose in western North America; 1982 April 13-15; Salt Lake City, UT. Gen. Tech. Rep. INT-152. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forestand Range Experiment Station: 126-136. [1684] 27. Munz, Philip A. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. [6155] 28. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 29. Sapsis, David S. 1990. Ecological effects of spring and fall prescribed burning on basin big sagebrush/Idaho fescue--bluebunch wheatgrass communities. Corvallis, OR: Oregon State University. 105 p. Thesis. [16579] 30. Simmerman, Dennis G.; Arno, Stephen F.; Harrington, Michael G.; Graham, Russell T. 1991. A comparison of dry and moist fuel underburns in ponderosa pine shelterwood units in Idaho. In: Andrews, Patricia L.; Potts, Donald F., eds. Proceedings, 11th annual conference on fire and forest meteorology; 1991 April 16-19; Missoula, MT. SAF Publication 91-04. Bethesda, MD: Society of American Foresters: 387-397. [16186] 31. Steele, Robert; Geier-Hayes, Kathleen. 1992. The grand fir/mountain maple habitat type in central Idaho: succession and management. Gen. Tech. Rep. INT-284. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 90 p. [17791] 32. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 7 p. [20090] 33. Stickney, Peter F. 1993. Effects of fire on upland forests in the Northern Rocky Mountains. Unpublished paper on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT: 3 p. [21627] 34. Toth, Barbara L. 1991. Factors affecting conifer regeneration and community structure after a wildfire in western Montana. Corvallis, OR: Oregon State University. 124 p. Thesis. [14425] 35. U.S. Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. SCS-TP-159. Washington, DC. 416 p. [11573] 36. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. [2944] 37. Schelstraete, Marc; Kennedy, Barbara M. 1980. Composition of miner's lettuce (Montia perfoliata). Journal of the American Dietetic Association. 77(1): 21-25. [22163] 38. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992] 39. Miller, J. M. 1976. Variation on populations of Claytonia perfoliata (Portulacaceae) . Systematic Botany. 1(1): 20-34. [22190] 40. Miller, John M. 1978. Phenotypic variation, distribution and relationships of diploid and tetr tetraploid populations of the Claytonia perfoliata complex (Portulacace. Systematic Botany. 3(3): 322-341. [18036] 41. Biswell, H. H.; Taber, R. D.; Hedrick, D. W.; Schultz, A. M. 1952. Management of chamise brushlands for game in the north coast region of California. California Fish and Game. 38(4): 453-484. [13673] 42. Gorske, S. F.; Hopen, H. J. 1978. Case of the purslane sawfly. American Vegetatble Growers. 2b(8): 14-15. [22191] 43. Smith, H. G.; Stevens, M.; Hallsworth, P. B. 1991. The use of monoclonal antibodies to detect beet mild yellowing virus and beet western yellows virus in aphids. Annals of Applied Biology. 119: 295-302. [22266] 44. Swanson, John R. 1966. A synopsis of relationships in Montioideae (Portulacaceae). Brittonia. 18: 229-241. [22265]

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