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SPECIES:  Darlingtonia californica

Introductory

SPECIES: Darlingtonia californica
Photo used with permission of Sherry Ballard © California Academy of Sciences

AUTHORSHIP AND CITATION: 
Crane, M. F. 1990. Darlingtonia californica. In: Fire Effects Information System, [Online]. 
U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, 
Fire Sciences Laboratory (Producer). Available: 
https://www.fs.usda.gov/database/feis/plants/forb/darcal/all.html [].


Revisions: Photo added on 06 February 2017.
ABBREVIATION: DARCAL SYNONYMS: Chrysamphora californica NRCS PLANT CODE: DACA5 COMMON NAMES: California pitcherplant cobra plant California pitcher cobra lily TAXONOMY: The currently accepted scientific name of California pitcherplant is Darlingtonia californica Torr. [16]. LIFE FORM: Forb FEDERAL LEGAL STATUS: No special status OTHER STATUS: California pitcherplant is listed as threatened in the Family Lists of Candidate Endangered and Threatened Plant Species in the Continental United States (Smithsonian Institution 1975) [9]. It is also on List Four, Plants of Limited Distribution--A Watch List, of the California Native Plant Society [17].


DISTRIBUTION AND OCCURRENCE

SPECIES: Darlingtonia californica
GENERAL DISTRIBUTION: California pitcherplant is endemic to the northern Sierra Nevada and Coast Ranges of southwestern Oregon and northern California, including the Klamath, Siskiyou, Salmon, and Trinity mountains. In the Sierra Nevada, it occurs as far south as Nevada County [14,27]. Most Oregon populations are found south of Florence in Lane County, but natural populations are recorded near Hidden Lake in Lincoln County and Sand Lake in Tillamook County [35]. A transplanted population is reported near Seattle, Washington [14]. ECOSYSTEMS: FRES21 Ponderosa pine FRES23 Fir - spruce FRES27 Redwood STATES: CA OR BLM PHYSIOGRAPHIC REGIONS: 1 Northern Pacific Border 4 Sierra Mountains KUCHLER PLANT ASSOCIATIONS: K005 Mixed conifer forest K006 Redwood forest K007 Red fir forest K010 Ponderosa shrub forest SAF COVER TYPES: 207 Red fir 231 Port Orford-cedar 232 Redwood SRM (RANGELAND) COVER TYPES: NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES: Published classifications listing California pitcherplant as an indicator species or a dominant part of the vegetation in community types (cts) are presented below: Area Classification Authority CA general veg. cts Thorne 1976 CA general veg. cts Holland 1986

MANAGEMENT CONSIDERATIONS

SPECIES: Darlingtonia californica
IMPORTANCE TO LIVESTOCK AND WILDLIFE: NO-ENTRY PALATABILITY: Leaves of California pitcherplants emit a putrid odor when cut [26]. This could reduce palatability. NUTRITIONAL VALUE: NO-ENTRY COVER VALUE: NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES: NO-ENTRY OTHER USES AND VALUES: The main threat to California pitcherplant's existence in accessible habitats is human collectors. Its strange shape and carnivorous habit have made it a curiosity [21]. However, its environmental needs are so exacting that it seldom survives for long in cultivation [6,21,32]. Collecting seed, which germinates readily, is the best method for obtaining plants [21]. Several authors provide cultivation information [21,32,33]. OTHER MANAGEMENT CONSIDERATIONS: No entry

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Darlingtonia californica
GENERAL BOTANICAL CHARACTERISTICS: California pitcherplant is a native perennial forb. It is a carnivorous plant that traps insects in its unique leaves. With plentiful light and water in its open bog habitat, the production of nectar is relatively "cheap". Insects attracted by the nectar are used as a nutrient source [15]. Adult California pitcherplants have slowly spreading rhizomes which produce a single leaf (pitcher) and roots at each node [11,26]. The diameter of the rhizome is from 0.6 to 0.8 inch (15-20 mm), although it narrows near the apical tip [11]. Internodes are short and the pitchers appear to arise in a terminal rosette [33]. Each leaf has a sheathing base that encloses the apical bud and the base of the next leaf [11]. Individuals have been aged by counting old leaf bases attached to the rhizome [18]. Pitcher size may vary from over 39 inches (1 m) to as small as 0.4 inches (1 cm), although most pitchers are between 8 and 24 inches (2-6 dm) tall [14,27,28]. Pitcher Morphology: Germinating seeds of California pitcherplant have 3 lanceolate and nontubular cotyledons which are followed by 5 to 10 juvenile leaves from a basal rosette [11,12]. Juvenile leaves are tubular and somewhat twisted so that the opening may either be erect or horizontal. Above the opening, each of these leaves tapers into a flattened beak and the outside surface has many nectar glands [12,23]. The distinctive adult leaves have an elongated tubular portion with a keeled area the length of the tube. The tube is twisted from 90 to 180 degrees. Functionally, the twisting turns the mouth of the tube to one side making it more accessible to insects and preventing the next leaf from growing into the mouth and plugging it [11]. At the top of the tube, a hood arches over the mouth. The front edge of the hood has a curious appendage shaped like a fishtail hanging in front of the mouth. The outer surface of the fishtail appendage has many nectaries, with nectar concentrated along the rim of the fishtail and opening of the pitcher [32,33]. At maturity, groups of hood cells lose chlorophyll and become translucent windows or fenestrations. The inner walls of the hood have short, stiff, downward or backward pointing hairs. Below the nectar roll at the mouth's edge, the inner walls are smooth and waxy with long, thin hairs pointing downward near the base [32,33]. Pitchers contain water secreted by the leaf, while rainwater is excluded by the hood [23,18,28,33]. This fluid increases in volume as insects or other nitrogen containing materials are added to it [23,18,33]. Pitcher Ecology: Insects are attracted to pitchers by the color and nectar glands which cover the outside of the hood but which are more numerous on the fishtail and wing [26]. There is a heavy exudation of nectar on the nectar roll inside the mouth of the pitcher. Light from the hood fenestrations assists in attracting insects inside the hood [25]. The fenestrations also may appear to be exits and confuse some insects [26]. Insects rarely escape once they have fallen into the fluid at the pitcher base [26]. However, many if not most insect visitors ingest nectar and leave without becoming trapped [15]. This system may be mutually beneficial to both the California pitcherplants, which gain a nutrient source, and the insect visitors, which gain a nectar source [15]. The bodies of insects which drown in the pitchers decompose by bacterial action, as California pitcherplant secretes no enzymes for that purpose [23,33]. Twenty or more arthropod species may be found living in pitchers of California pitcherplants, many of which are obligate inhabitants [8,29]. Larvae of several species feed on dead insects trapped by the pitchers, thus increasing the rate of decomposition [28]. Spiders construct webs in the domes [8]. Some insects feed on pitcher tissue or lay eggs in the pitchers so that their larvae can feed on them [8]. RAUNKIAER LIFE FORM: Geophyte REGENERATION PROCESSES: Vegetative reproduction of California pitcherplant by rhizomes is more common than sexual reproduction [32]. Each leaf axil contains a single bud which can develop into a lateral rhizome branch [11]. Although most remain dormant, the buds that grow can cover a suitable location with California pitcherplants [11,32]. Each pendant flower is borne on a stalk up to 39 inches (1 m) tall [14,27]. The yellowish sepals are longer than the closed, reddish corolla, which has openings to admit insect pollinators at indentations near the ends of the petals [32,33]. The corolla shape, bell-shaped ovary, and positions of both stigma and stamens greatly increase the probability of cross-pollination [32]. Each five-chambered capsule bears a hundred or more seeds which are 0.08 to 0.1 inch (2-3 mm) long, hairy, and light reddish-brown [14,26,27]. The more rounded end of each seed has many short projections which may aid in animal dispersal and help the seed float [32,43]. Seeds mature within 10 weeks of fertilization [33]. SITE CHARACTERISTICS: Along the Oregon and northern California coast, California pitcherplant is found in sphagnum bogs, seeps, and along trickling streams [14,30]. In the Siskiyou Mountains it is found only on sites with running water [1]. In California's Klamath Ranges and the northern Sierra Nevada, California pitcherplant grows where there are slowly draining bogs formed by springs or seepage slopes and open marshy meadows [34]. It is very rarely found in bogs with standing water. Normally it is restricted to sites where its rhizomes and roots can be kept cool by cold, moving water [26,32,33]. In northwestern California, bogs with California pitcherplant as the dominant vascular species intermingle with bog forests where Port-Orford-cedar (Chamaecyparis lawsoniana), western white pine (Pinus monticola), and shore pine (Pinus contorta) dominate [17]. Other plants found with California pitcherplant include sundew (Drosera rotundifolia), northern grass-of-parnassus (Parnassia palustris), common butterwort (Pinguicula vulgaris), California coneflower (Rudbeckia californica), smallhead burnet (Sanguisorba microcephala), lance-leaved violet (Viola lanceolata occidentalis), white rushlily (Hastingsia alba), California bog-asphodel (Narthecium californicum), western tofieldia (Tofieldia glutinosa occidentalis), California ladyslipper (Cypripedium californicum), canyon bogorchid (Habenaria sparsiflora), hairy bulrush (Scirpus criniger), lilies (Lilium spp), rushes (Juncus spp.), and beaked-rushes (Rhynchospora spp.) [34]. In lower elevation bogs near the ocean, some associated species include Yosemite aster (Aster occidentalis var. yosemitanus), scarlet paintbrush (Castilleja miniata ssp. elata), Mendocino gentian (Gentiana setigera), California coneflower, Pacific reedgrass (Calamagrostis nutkaensis), smallhead burnet, largehorned butterwort (Pinguicula macroceras), and tufted hairgrass (Deschampsia caespitosa ssp. beringensis) [17]. Soils: California pitcherplant commonly grows on sphagnum or poor peat soils and gravel where the parent material is serpentine. It is often considered an indicator of serpentine [14,26,32,38,39]. It is also found on olivine grabbro in the central Siskiyou Mountains [39]. California pitcherplant occurs from sea level to 8,500 feet (2,592 m) in elevation [27,33]. SUCCESSIONAL STATUS: Bogs containing California pitcherplant can develop directly from wet sand on deflation plains along the Oregon coast [41]. Carnivorous plants are characteristic of an early stage of succession in bog habitats. Their growth helps build soil which can then be occupied by other perennials, shrubs, and eventually trees [39]. Pitcher plants in the southeastern United States do not compete well with other plants in the absence of fire or other disturbance [15]. California pitcherplant is primarily found in the bright sunshine of open bogs [10,21]. SEASONAL DEVELOPMENT: Flowers are initiated during late summer, overwinter as buds and bloom in early spring before new leaves appear [43]. Bloom can be from April to August depending on altitude [28,32]. The first pitchers produced after bloom are the tallest of the year [28]. Annual growth of the rhizome may include one or two full-sized pitchers, one that is half size or smaller, and three to five very small pitchers [29]. The smallest pitchers tend to be prostrate, with the fishtail appendage touching the ground and forming a ramp for insects [23].

FIRE ECOLOGY

SPECIES: Darlingtonia californica
FIRE ECOLOGY OR ADAPTATIONS: California pitcherplant grows on sites with running water which seem unlikely to burn most years. However, it does grows in bright sunlight and may well be a seral species like other pitcherplants. The shallow rhizome is covered by imbricate, sheathing leaf bases [11] which may give it some protection from fire. California pitcherplant sprouts from the rhizome, and seeds may be transported from nearby unburned sites by animals [11,32]. FIRE REGIMES: 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". POSTFIRE REGENERATION STRATEGY: Rhizomatous herb, rhizome in soil Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Darlingtonia californica
IMMEDIATE FIRE EFFECT ON PLANT: Aboveground portions of California pitcherplant are probably killed by fire. Survival of the rhizome depends on fire severity. PLANT RESPONSE TO FIRE: In the Southeast, frequent, moderate fire is necessary for some pitcher plants (Sarracenia spp.) to maintain dense populations. Such fires, as often as every second year, reduce encroachment of competing plants and increase bare, wet ground for seedling establishment [9,15,19]. In Mississippi, pitcherplants in a burned bog were more abundant and had larger leaves and rhizomes than those in an unburned bog [5]. Although historical reports indicate dense populations of California pitcher plant [15], its specific response to fire has not been documented. All the older (from 146 to 350-400 years) Port-Orford-cedar in a bog forest intermingled with California pitcherplant bogs exhibit fire scars [17]. FIRE MANAGEMENT CONSIDERATIONS: NO-ENTRY

References for species: Darlingtonia californica


1. Atzet, Thomas; Wheeler, David L. 1984. Preliminary plant associations of the Siskiyou Mountain Province. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 278 p. [9351]
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. [434]
3. Daubenmire, Rexford. 1978. Plant geography--with special reference to North America. Physiological Ecology. New York: Academic Press. 338 p. [8949]
4. Dittberner, Phillip L.; Olson, Michael R. 1983. The plant information network (PIN) data base: Colorado, Montana, North Dakota, Utah, and Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 786 p. [806]
5. Eleuterius, L. N.; Jones, S. B., Jr. 1969. A floristic and ecological study of pitcher plant bogs in south Mississippi. Rhodora. 71: 29-34. [12333]
6. Everett, Percy C. 1957. A summary of the culture of California plants at the Rancho Santa Ana Botanic Garden 1927-1950. Claremont, CA: The Rancho Santa Ana Botanic Garden. 223 p. [7191]
7. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
8. Fashing, N. J. 1981. Arthropod associates of the cobra lily (Darlingtonia californica). Virginia Journal of Science. 23(3): 92. Abstract. [12304]
9. Folkerts, George W. 1977. Endangered and threatened carnivorous plants of North America. In: Prance, G. T.; Elias, T. S. ed, eds. Extinction is forever. Threatened and endangered species of plants in the Americas and their significance today and in t; 1976 May 11-13; New York. [Place of publication unknown]. [Publisher unknown]. 301-313. [12388]
10. Fowlie, J. A. 1982. Notes on the habitat and ecological relationships of Cypripedium californicum A. Gray and Darlingtonia californica. Orchid Digest. 46(5): 165-170. [12300]
11. Franck, Daniel H. 1975. Early histogenesis of the adult leaves of Darlingtonia californica (Sarraceniaceae) and its bearing on the nature of epiascidiate foliar a. American Journal of Botany. 62(2): 116-132. [12485]
12. Franck, Daniel H. 1976. Comparative morphology and early leaf histogenesis of adult and juvenile leaves of Darlingtonia californica and their bearing on the concept of. Botanical Gazette. 137(1): 20-34. [12486]
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. Hitchcock, C. Leo; Cronquist, Arthur. 1964. Vascular plants of the Pacific Northwest. Part 2: Salicaceae to Saxifragaceae. Seattle, WA: University of Washington Press. 597 p. [1166]
15. Joel, Daniel M. 1988. Mimicry and mutalism in carnivorous pitcher plants (Sarraceniaceae, Nepenthaceae, Cephalotaceae, Bromdiaceae). Biological Journal of the Linnean Society. 35(2): 185-197. [12303]
16. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954]
17. Keeler-Wolf, Todd. 1986. An ecological survey of the proposed Stone Corral - Josephine Peridotite Research Natural Area (L. E. Horton - Darlingtonia Bog Research Nat. Area) on the Six Rivers National Forest, Del Norte County, California. Purchase order # 40-9AD6-5-907. Unpublished report on file at: U.S. Department of Agriculture, Forest Service, Intermountain Fire Sciences Laboratory, Missoula, MT. 69 p. [12307]
18. Knight, Walter; Knight, Irja; Howell, John Thomas. 1970. A vegetation survey of the Butterfly Botanical Area, California. Wasmann Journal of Biology. 28: 1-246. [12306]
19. Komarek, E. V. 1981. History of prescribed fire and controlled burning in wildlife management in the South. In: Wood, Gene W., ed. Prescribed fire and wildlife in southern forests: Proceedings of a symposium; 1981 April 6-8; Myrtle Beach, SC. Georgetown, SC: Clemson University, Belle W. Baruch Forest Science Institute: 1-14. [14802]
20. Jones, F. M. 1921. Pitcher plants and their moths. Natural History. 21: 296-316. [12301]
21. Kruckeberg, A. R. 1982. Gardening with native plants of the Pacific Northwest. Seattle: University of Washington Press. 252 p. [9980]
22. 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. [3455]
23. Lloyd, F. E. 1942. The carnivorous plants. Waltham, MA: Chronica Botanica Company. 352 p. [12247]
24. 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. 10 p. [20090]
25. Mellichamp, T. L. 1979. The occurrence of ladyslipper orchids and insectivorous plants Part 1. Darlingtonia californica as it occurs with Cypripedium californicum. Orchid Digest. 43(3): 108-113. [12299]
26. Mellichamp, T. L. 1983. Cobras of the Pacific Northwest. Natural History. 92(4): 46-51. [12298]
27. Munz, Philip A. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. [6155]
28. Naeem, Shahid. 1988. Resource heterogeneity fosters coexistence of a mite and a midge in pitcher plants. Ecological Monographs. 58(3): 215-227. [12309]
29. Nielsen, David W. 1990. Arthropod communities associated with Darlingtonia californica. Annals of the Entomological Society of America. 83(2): 189-200. [12308]
30. Peck, Morton E. 1941. A manual of the higher plants of Oregon. Portland, OR: Binfords & Mort. 800 p. [12444]
31. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
32. Schnell, Donald E. 1976. Carnivorous plants of the United States and Canada. Winston-Salem, NC: John F. Blair. 125 p. [12292]
33. Slack, Adrian. 1979. Carnivorous plants. Cambridge, MA: The MIT Press. 240 p. [12293]
34. Thorne, Robert F. 1976. The vascular plant communities of California. In: Latting, June, ed. Symposium proceedings: plant communities of southern California; 1974 May 4; Fullerton, CA. Special Publication No. 2. Berkeley, CA: California Native Plant Society: 1-31. [3289]
35. Trappe, J. M; Gerdemann, J. W. 1974. A northern extension of the range of Darlingtonia. Madrono. 22(5): 279. [12212]
36. 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]
37. Vankat, John L. 1979. The natural vegetation of North America: an introduction. New York: John Wiley and Sons. 261 p. [25214]
38. Whittaker, R. H. 1954. The ecology of serpentine soils: IV. The vegetational response to serpentine soils. Ecology. 35(2): 275-288. [10397]
39. Whittaker, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs. 30(3): 279-338. [6836]
40. Zembal, Richard. 1990. Riparian habitat and breeding birds along the Santa Margarita and Santa Ana Rivers of southern California. In: Schoenherr, Allan A., ed. Endangered plant communities of southern California: Proceedings, 15th annual symposium; 1989 October 28; Fullerton, CA. Special Publication No. 3. Claremont, CA: Southern California Botanists: 98-114. [21322]
41. Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. Gen. Tech. Rep. PNW-8. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 417 p. [961]
42. Holland, Robert F. 1986. Preliminary descriptions of the terrestrial natural communities of California. Sacramento, CA: California Department of Fish and Game. 156 p. [12756]
43. DeBuhr, Larry Eugene. 1973. Distribution and reproductive biology of Darlingtonia californica. Santa Ana Bot. Gardensr, CA: Claremont Graduate School. 43 p. Thesis. [12433]

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