|© 2005 Chris Wagner, SBNF|
Ash-grey Indian paintbrush prefers, but is not limited to, pebble plain habitats. These habitats are surrounded by montane Jeffrey pine (Pinus jeffreyi) forest and pinyon-juniper woodlands dominated by singleleaf pinyon (P. monophylla), Sierra juniper (Juniperus occidentalis ssp. australis) and/or Utah juniper (J. osteosperma). The flora of the pebble plains consists of small cushion forming plants that are low growing, well spaced and sun tolerant [28,32,33]. Species associated with ash-grey Indian paintbrush on the pebble plains are Parish's rock cress (Arabis parishii), Bear Valley sandwort (Arenaria ursina), southern mountain buckwheat (Eriogonum kennedyi var. austromontanum), silver-haired ivesia (Ivesia argyrocoma), Big Bear Valley phlox (Phlox dolichantha), and San Bernardino bluegrass (Poa atropurpurea). Many of these species also have limited distribution on or near the San Bernardino National Forest [10,12,33].
Montane coniferous forest species associated with ash-grey Indian paintbrush include incense-cedar (Calocedrus decurrens), Jeffrey pine, ponderosa pine (P. ponderosa), Sierra juniper, white fir (Abies concolor), California black oak (Quercus kelloggii), and canyon live oak (Q. chrysolepis). Common shrub species within the montane coniferous forest are mountain big sagebrush ( A. tridentata ssp. vaseyana), manzanita (Arctostaphylos spp.), ceanothus (Ceanothus spp.), and curlleaf mountain mahogany (Cercocarpus ledifolius) [12,28,30].
Singleleaf pinyon pine mixed with Sierra juniper, Utah juniper and oaks (Quercus spp.) dominate the pinyon-juniper woodlands where ash-grey Indian paintbrush can be found. Other species commonly associated in these pinyon-juniper woodlands include basin big sagebrush ( A. tridentata ssp. tridentata), bigberry manzanita (Arctostaphylos glauca), birchleaf mountain-mahogany (Cercocarpus betuloides), desert ceanothus (Ceanothus greggii), rubber rabbitbrush (Chrysothamnus nauseosus), blackbrush (Coleogyne ramosissima), flannelbush (Fremontodendron californicum), bitterbrush (Purshia tridentata), sage (Salvia spp.) and Joshua tree (Yucca brevifolia) [12,18,37,38].
Ash-grey Indian paintbrush also inhabits dry montane meadows of southern California. Species of bentgrass (Agrostis spp), hairgrass (Deschampsia spp), muhly grass (Muhlenbergia spp.), bluegrass (Poa spp.), along with sedges (Carex spp., Scirpus spp.), and rushes (Juncus spp.) are commonly found in these meadows .Common species associated with ash-grey Indian paintbrush in the Mojavean desert scrub are creosote bush (Larrea tridentata), white bursage (Ambrosia dumosa), burrobrush (A. salsola), brittlebush (Encelia farinosa), spiny senna (Senna armata), Nevada ephedra (Ephedra nevadensis), and boxthorn (Lycium sp.) .
|© 2005 Aaron Schusteff||© 2005 Chris Wagner, SBNF|
Ash-grey Indian paintbrush is a perennial herb 2 to 6 inches (5-15 cm) in height. The plant has a taproot and can have several stems growing from the root crown. The spike-like inflorescence ranges in color from greenish yellow to a crimson red. Variations in flower size and color are due to aspect. Northerly exposures have larger more yellow flowers, whereas flowers become smaller and more reddish in color to the south. The fruit is 6 to 10 mm long. Encapsulated seeds have a net-like surface and range from 0.8 to 1.3 mm in length [10,21,22,33].
Ash-grey Indian paintbrush is a hemiparasitic plant that obtains some nutrients and water from a host plant. Host plant species parasitized by ash-grey Indian paintbrush include southern mountain buckwheat, Kennedy's buckwheat (Eriogonum kennedyi var. kennedyi) Wright's buckwheat (Eriogonum wrightii var. subscaposum), basin big sagebrush, black sagebrush (A. nova), and other Artemisia species. [10,21,22,33].RAUNKIAER  LIFE FORM:
Breeding system: Ash-grey Indian paintbrush is monoecious with bisexual flowers [10,22].
Pollination: Ash-grey Indian paintbrush is insect and bird pollinated [17,24,33]. On the pebble plains in the San Bernardino Mountains, ash-grey Indian paintbrush pollen transfer occurred at distances of less than 10 feet (4 m) (Freas, K. 1988, unpublished report cited in ). A pollinator exclusion experiment performed on a related species, golden paintbrush (Castilleja levisecta), found that fruit set was 5 times greater for inflorescences visited by pollinators compared to inflorescences not visited [39,40].
Seed production: No information is available on this topic.
Seed dispersal: In 1 study, seed dispersal for ash-grey Indian paintbrush was limited to 16 feet (5 meters) outside the edge of the pebble plain habitat (Freas, K. 1988, unpublished report cited in ). Ash-grey Indian paintbrush seed dispersal by animals is undocumented.
Seed banking: Information regarding seed banking for ash-grey Indian paintbrush is lacking. Limited evidence for golden paintbrush suggests that its seed longevity may be 2 years [6,39]. Research is needed on seed bank longevity of ash-grey Indian paintbrush.
Germination: There are no germination studies for ash-grey Indian paintbrush. Field and laboratory studies for golden paintbrush found higher germination rates for 1st-year seeds than 2nd-year seeds and no germination occurred in the 3rd year [39,40]. Cold stratification of golden paintbrush seeds was required for 6 weeks to obtain 80% germination (St. Hilaire, K. 1987, unpublished report cited in ). Research on seed germination for ash-grey Indian paintbrush is needed.
Seedling establishment/growth: Ash-grey Indian paintbrush produces haustoria that obtain nutrients from the host plant. Haustoria formation and host contact is usually made shortly after ash-grey Indian paintbrush germination. The radicle penetrates the soil, forming a ring of hair-like structures and then branches . The haustoria are produced by small branching roots from the lateral (main) root. Since ash-grey Indian paintbrush is only a partial root parasite, a host species is not necessary for plant survival. Many species in the genus Castilleja completed their life cycle without a host when grown in a greenhouse environment .SITE CHARACTERISTICS:
Fire regimes: Pebble plain habitats, where ash-grey Indian paintbrush commonly occurs, are open rocky areas surrounded by forests of Jeffrey pine or pinyon-juniper woodlands. Pebble plains typically support little biomass and function as a natural fuel break that prevents or slows fire spread. Fire severities vary depending on the condition of surrounding vegetation. Presence of nonnative grasses, including red brome (Bromus rubens) and cheatgrass (B. tectorum), may increase fire spread through the pebble plains and increase fire severity [32,33].
Pinyon-juniper woodlands: Pinyon-juniper woodland communities in the San Bernardino Mountains experienced long-interval stand-replacement fires with estimated fire intervals of several hundred years both before and during the fire exclusion era [20,28,38]. Postfire succession is slow in singleleaf pinyon pine communities, with initial colonization by Great Basin big sagebrush scrub and desert shrub species. Fires can move quickly through pinyon-juniper woodlands, resulting in a mosaic of small scattered burned patches within uniform old-growth stands . Thin bark and lack of self pruning makes singleleaf pinyon very susceptible to intense fire . The sharp ecotone between pinyon-juniper and mixed-conifer communities may be due to the inability of singleleaf pinyon to withstand the higher frequency and severity typical of fires in mixed-conifer communities . In 1999 several large fires occurred in the San Bernardino Mountains pinyon-juniper woodlands, parts of which reburned shortly thereafter. Currently the spread of nonnative cheatgrass in the burned areas has increased fuels and fuel continuity, which may decrease the fire return interval in these habitats .
Coniferous Pine Forest: Presettlement fire regimes for mixed pine forests were short-interval (5-30 years), low-severity, nonlethal surface fires that consumed litter, shrubs, seedlings, and immature trees, leaving large trees to thrive in open stands. Fire exclusion has eliminated low-severity understory burns, resulting in dense stands of conifers and increased dead and live fuel accumulation. Current conditions cause fires to burn more severely, which increases tree mortality [1,28]. In the current fire regime, species composition has shifted from Jeffrey pine, ponderosa pine, and California black oak dominance to incense-cedar and white fir dominance .
Desert Scrub: Historically, fires in desert scrublands were very infrequent. They were ignited by lightning and/or Native Americans. Discontinuous sparse fuels produced fires of low to moderate severity. Nonnative grasses and increased urban development have increased fire frequency [25,28]. Creosote bush (Larrea tridentata), a dominant species in the Mojavean desert scrub of southern California, is not fire resilient due to its limited sprouting ability and possible displacement by other species [3,13,25].
The following table provides fire return intervals for plant communities and ecosystems where ash-grey Indian paintbrush is important. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
|Community or Ecosystem||Dominant Species||Fire Return Interval Range (years)|
|western juniper||Juniperus occidentalis||20-70|
|creosote bush||Larrea tridentata||<35 to <100|
|pinyon-juniper||Pinus-Juniperus spp.||< 35 |
|Jeffrey pine||Pinus jeffreyi||5-30|
|Pacific ponderosa pine*||Pinus ponderosa var. ponderosa||1-47 |
Golden paintbrush, a federally threatened species found in Washington state and British Columbia, was monitored on 3 sites at the Washington Rocky Prairie Natural Area Preserve following a wildland fire in 1987 and 2 controlled burns in 1989 and 1994. Population numbers following the burns increased in all 3 studies, and postfire mortality was low. Postfire survivorship for new establishing plants on the burn sites was higher than for unburned plants .FIRE MANAGEMENT CONSIDERATIONS:
1. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. 
2. 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. 
3. Brown, David E.; Minnich, Richard A. 1986. Fire and changes in creosote bush scrub of the western Sonoran Desert, California. The American Midland Naturalist. 116(2): 411-422. 
4. California Department of Fish and Game, Wildlife and Habitat Data Analysis Branch. 2006. Castilleja cinerea: Full report with sources for selected occurrence. In: California Natural Diversity Database (Government version--March 30, 2006). California Department of Fish and Game (Producer). Available: http://www.dfg.ca.gov/whdab/html/cnddb.html [2006, May 31]. 
5. Center for Plant Conservation. . Plant profile: Castilleja levisecta, [Online]. In: CPC (Center for Plant Conservation ) national collection of endangered plants. St. Louis, MO: Missouri Botanical Garden (Producer). Available: http://www.centerforplantconservation.org/ASP/ CPC_ViewProfile.asp?CPCNum=824 [2006, March 27]. 
6. Dunwiddie, Peter W.; Davenport, Roberta; Speaks, Pene. 2002. Effects of burning on Castilleja levisecta at Rocky Prairie Natural Area Preserve, Washington: a summary of three long-term studies, [Online]. In: ConserveOnline. The Nature Conservancy (Producer). Available: http://conserveonline.org/docs/2002/01/RPMscFnl.doc [2006, March 27]. 
7. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. 
8. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. 
9. Heckard, L. R. 1962. Root parasitism in Castilleja. Botanical Gazette. 124(1): 21-29. 
10. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. 
11. Hickman, James C. 2006. Descriptions and keys from The Jepson Manual: higher plants of California (including updated information from The Jepson Desert Manual and other sources), [Online]. In: Jepson Flora Project: Jepson online interchange for California floristics. Berkeley, CA: University of California, University and Jepson Herbaria (Producers). Available: http://ucueps.berkeley.edu/jepman.html [2006, March 27]. 
12. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. 
13. Humphrey, Robert R. 1974. Fire in the deserts and desert grassland of North America. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and ecosystems. New York: Academic Press: 365-400. 
14. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with: The Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. 
15. Keeley, Jon E.; Zedler, Paul H. 1998. Evolution of life histories in Pinus. In: Richardson, David M., ed. Ecology and biogeography of Pinus. Cambridge, UK: The Press Syndicate of the University of Cambridge: 219-250. 
16. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. 
17. Kuijt, Job. 1969. The biology of parasitic flowering plants. Berkeley, CA: University of California Press. 246 p. 
18. Laudenslayer, William F., Jr.; Boggs, Jerry R. 1988. Pinyon-juniper vegetation. In: Mayer, Kenneth E.; Laudenslayer, William F., Jr., eds. A guide to wildlife habitats of California. Sacramento, CA: California Department of Fish and Game: 62-65. 
19. McLean, Alastair. 1968. Fire resistance of forest species as influenced by root systems. Journal of Range Management. 22(2): 120-122. 
20. Minnich, Richard A. 1999. Vegetation, fire regimes, and forest dynamics. In: Miller, P. R.; McBride, J. R., eds. Oxidant air pollution impacts in the montane forests of southern California: a case study of the San Bernardino Mountains. Ecological Studies: Analysis and Synthesis. Vol. 134. New York: Springer-Verlag: 44-80. 
21. Munz, Philip A. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. 
22. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. 
23. NatureServe. 2005. Comprehensive species report--Castilleja cinerea, [Online]. In: NatureServe Explorer: an online encyclopedia of life [Web application], (Version 4.6). Arlington, VA: NatureServe (Producer). Available: http://www.natureserve.org [2005, November 21]. 
24. O'Brien, Mary H. 1980. The pollination biology of a pavement plain: pollinator visitation patterns. Oecologia. 47: 213-218. 
25. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. 
26. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
27. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. 
28. Stephenson, John R.; Calcarone, Gena M. 1999. Southern California mountains and foothills assessment. Gen. Tech. Rep. PSW-GTR-172. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 402 p. 
29. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
30. Thorne, Robert F. 1977. Montane and subalpine forests of the Transverse and Peninsular ranges. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 537-557. 
31. Tibor, David P., ed. 2001. California Native Plant Society's inventory of rare and endangered plants of California. 6th edition. Sacramento, CA: California Native Plant Society. 387 p. 
32. U.S. Department of Agriculture, Forest Service, Pacific Southwest Region. 2005. Habitat status reports--Pebble plain. In: Reading Room. In: Revised Land Management Plans and Final Environmental Impact Statement--Angeles National Forest, Cleveland National Forest, Los Padres National Forest, San Bernardino National Forest [CD ROM]. R5-MB-086-CD. Vallejo, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Region (Producer). Available: San Diego, CA: U.S. Department of Agriculture, Forest Service, Cleveland National Forest [2006, March 27]. 
33. U.S. Department of Agriculture, Forest Service, Pacific Southwest Region. 2005. Species accounts--plants: Castilleja cinerea. In: Reading Room. In: Revised Land Management Plans and Final Environmental Impact Statement--Angeles National Forest, Cleveland National Forest, Los Padres National Forest, San Bernardino National Forest [CD ROM]. R5-MB-086-CD. Vallejo, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Region (Producer). Available: San Diego, CA: U.S. Department of Agriculture, Forest Service, Cleveland National Forest [2006, March 27]. 
34. U.S. Department of Agriculture, Forest Service, San Bernardino National Forest. 2002. Effects of the 1999 Willow Fire on threatened, endangered, sensitive, and watch list plant species and their habitats on the San Bernardino National Forest. Final accomplishment report--monitoring funded under the National Fire Plan in 2001. San Bernardino, CA: San Bernardino National Forest. 41 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 
35. U.S. Department of Agriculture, Natural Resources Conservation Service. 2006. PLANTS database (2006), [Online]. Available: http://plants.usda.gov/. 
36. U.S. Department of the Interior, Fish and Wildlife Service. 2016. Endangered Species Program, [Online]. Available: http://www.fws.gov/endangered/. 
37. Vasek, Frank C.; Thorne, Robert F. 1977. Transmontane coniferous vegetation. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 797-832. 
38. Wangler, Michael J.; Minnich, Richard A. 1996. Fire and succession in pinyon-juniper woodlands of the San Bernardino Mountains, California. Madrono. 43(4): 493-514. 
39. Wentworth, Jane B. 1994. The demography and population dynamics of Castilleja levisecta, an endangered perennial. Seattle, WA: University of Washington. 53 p. Dissertation. 
40. Wentworth, Jane B. 2002. Demography and population dynamics of Castilleja levisecta, [Online]. In: Botanical Electronic News (BEN). BEN Archives Issue No. 290. Norman, OK: University of Oklahoma, Department of Botany and Microbiology (Producer). Available: http://www.ou.edu/cas/botany-micro/ben/ben290.html [2006, March 27].