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Darlingtonia californica:

Correcting Errors Perpetuated in the Literature

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Drawing of Darlingtonia californica, also known as the California Pitcher Plant or Cobra Lily. From: John Torrey. On the Darlingtonia Californica, A New Pitcher Plant, from Northern California. In: Smithsonian Contributions to Knowledge, April, 1853. - plant leaf is hollow with mustache appendage and flower is on long thin stem with 10 petals Despite efforts to achieve accuracy, several facts regarding the discovery, distribution, and characteristics of the substrate in which Darlingtoniagrows have been misstated then repeated in the literature. This paper will discuss some of the mistakes encountered in the literature regarding Darlingtonia californica. The elevation, slope, calcium and magnesium levels of the water samples, and the pH of the water in which Darlingtonia was found growing on Mt. Eddy, California will be provided.

Darlingtonia californica is a well-known insectivorous pitcher plant belonging to the American pitcher plant family, Sarraceniaceae. It was first collected by William Brackenridge during the overland portion of the U.S. Exploring Expedition in early October, 1841. The three genera in this family have a wide geographic distribution: Sarracenia occur in the eastern United States; Heliamphora are found in northern South America; and Darlingtonia, a monotypic genus, grows in wet areas in western Oregon and northern California.

The most captivating feature of Darlingtonia is its pitcher, hence the common name, California Pitcher Plant. This modified leaf is composed of a twisted tube which enlarges into an inflated head that bends down like a hood. Near the opening on the underside of the hood is a forked appendage often referred to as a "fishtail appendage." The cobra-like shape of the pitcher and its fang-like appendage suggest its other common name, Cobra Lily.

Study Area / Methods

During the summers of 1992 and 1993 I conducted a study of the distribution of Darlingtonia on Mt. Eddy, which straddles Siskiyou and Trinity counties in northern California. Mt. Eddy is the highest peak in the Klamath Mountains. The study area ranges in elevation from 1036 to 2751 meters and is in the central portion of Darlingtonia's known geographical range (Figure 1). When I found Darlingtonia on Mt. Eddy, its location was mapped, the pH of the water was recorded, and water samples were collected to analyze the magnesium and calcium content of the water in which Darlingtonia was found growing. The methods and materials may be found in the thesis Distribution of Darlingtonia californica on Mt. Eddy, California (Freeman 1994), available at the CSU Chico library and the College of the Siskiyous library.

Map of California and Oregon showing known distribution of the California Pitcher Plant.
Figure 1. The study area in relation to the distribution of Darlingtonia californica.
Source: L. E. De Buhr, 1973 (adapted).

Review of the Literature

The literature concerning Darlingtonia is relatively scant, with about 25 known publications dealing specifically with Darlingtonia (Elder 1994). Most of the related literature deals with carnivorous plants in general or discusses a region in which Darlingtonia is found, without emphasizing Darlingtonia. About one-third of the publications which focus on Darlingtonia are encountered in the popular press, while another third were written in the 1800s. It should be noted that although some information found in the literature is herein criticized, this does not necessarily mean the source is not of high quality. For instance, The Carnivorous Plants (Juniper, et. al. 1989) is well-written, informative, and enjoyable.

Both the collector's name and date concerning the discovery of Darlingtonia have been incorrectly stated in the literature (Torrey 1853; Lloyd 1942; Schnell 1976; Slack 1979; Juniper et. al. 1989). This is due, in part, to John Torrey's 1853 article in Smithsonian Contributions to Knowledge. Torrey writes, "This new Pitcher-plant was first detected by Mr. J. D. [sic] Brackenridge...in the year 1842 [sic]." The errors relating to William D. Brackenridge's name and the date of discovery are most likely due to transcription or typographical errors. Now that these mistakes have entered the scientific literature, they are often accepted as facts.

A chased-by-Indians story is often associated with the discovery of Darlingtonia, apparently to explain the reason for the less-than-ideal quality of the specimen collected (Schnell 1976; Mellichamp 1983; Juniper, et. al. 1989, Rondeau 1991). This story has it that Brackenridge was unable to make a decent collection of Darlingtonia due to the "fact" that he was being pursued by Indians at the time. Eastwood (1945) and Maloney (1945) wrote articles that accompanied the first publication of Brackenridge's journal. Eastwood questioned the validity of this story and challenged anyone to provide evidence of the Indian-pursuit story related to the discovery of Darlingtonia. Eastwood finally found the story in print in The Life of James Dwight Dana by Daniel Coit Gilman (1899). Maloney figured Dana, who was on the expedition with Brackenridge, told the story to Gilman and considered the problem solved. However, Miesse (1993) points out that Gilman did not credit Dana with the story. Gilman may have had access to information which supported this story but he did not supply evidence of this in his account. Further, Brackenridge himself did not include any reference to this incident in his journal. The perpetuation of this story in the literature appears to be a literary device rather than an historical fact.

Literature pertaining to the distribution of Darlingtonia contains small errors which are mostly due to the age of the source consulted and the language style used. Over time, there has been an increase in knowledge of plant sites as herbarium collections have been made. Darlingtonia's distribution has become better known as specimens from several herbaria are studied as a group (e.g. De Buhr 1973). Articles that describe the distribution of Darlingtonia based on early sources may be incorrect in light of new evidence. Minor errors can also result when authors use directional or geographical designations that, if taken literally, do not describe the intended distribution accurately. For example, Cheatham (1976) states that Darlingtonia has a "...limited distribution to scattered locations in northern California and the southern half of Oregon." It would be more accurate to state that Darlingtonia grows in western Oregon, rather than southern Oregon, as it is not found in southeastern Oregon and it is found in northwestern Oregon in Linn, Tillamook, and Lincoln counties (DeBuhr 1973).

The upper and lower elevational limits of Darlingtonia differ slightly among various sources. The range in elevation is approximately sea level to about 2600 meters. Schnell (1976) and others list the range as sea level to 2800 meters. The highest peak in Darlingtonia's known geographic range is Mt. Eddy (2751 meters) and the next highest peak is Mt. Thompson in the Trinity Alps (2741 meters). Not only are both peaks less than 2800 meters, but Darlingtonia needs water from runoff and therefore will not be found growing on the peaks themselves. The highest elevation at which Darlingtonia is found on Mt. Eddy is 2390 meters, but it was most commonly found between 1700 and 2100 meters (Freeman 1994). The highest it has been recorded growing in the southern Klamath Ranges is 2587 meters (Rondeau 1991).

An early study of Darlingtonia by Mary Austin (Juniper et. al. 1989) suggested that Darlingtonia was confined to south- and southwest-facing slopes. While Darlingtonia may, in fact, have been limited to these slopes in the earlier study, that is no reason to believe that Darlingtonia is confined to south- and southwest-facing slopes throughout its range. Darlingtonia was found growing on virtually all slope aspects on Mt. Eddy, indicating that there are no apparent slope-orientation requirements for the California Pitcher Plant.

Differences of opinion exist regarding the pH of the substrate in which Darlingtonia grows. Part of the confusion stems from the fact that Darlingtonia grows in wet areas, which are often referred to as bogs. Since bogs are commonly thought to be acidic, the inference has been made that Darlingtonia grows in an acidic medium (e.g. Naeem and Dusheck 1985; Schoenherr 1992). However, the marshy areas in which Darlingtonia is found growing are more accurately called seeps (Sawyer 1986); using this term could prevent one from concluding that the water is acidic. Many sources (DeBuhr 1973; Cheatham 1976; Schnell 1976; Naeem and Dusheck 1985; Sawyer 1986; Juniper, et. al. 1989; Schoenherr 1992) point out the association of Darlingtonia with serpentine or ultramafic soils, which tend to produce alkaline runoff. Only two of the cited studies (Rondeau 1991; Juniper, et. al. 1989) actually give the pH values (5.1- 7.0 and 7.5-9.5 respectively). It appears that the pH of the water or soil in which Darlingtonia grows has not been tested throughout its range. In the study area Darlingtonia is more commonly found in association with water having a pH from 7 to 8 but it grows in both acidic and basic waters, with the pH ranging from 5.2 to 8.5.

The substrate in which Darlingtonia grows needs further study. Several articles (Cheatham 1976; Schnell 1976; Mellichamp 1983; Naeem and Dusheck 1985) state that the California Pitcher Plant grows in a nutrient-deficient medium, but they do not provide evidence which supports this claim. Many sources (Cheatham 1976; Mellichamp 1983; Naeem and Dusheck 1985) imply that the soil or water where Darlingtonia grows is high in magnesium, nickel, and chromium, but values for these metals are often not given nor is the term "high" defined. The magnesium levels of the water samples from the study area ranged from 44 mg/l to 223 mg/l while the calcium levels ranged from 2 mg/l to 18 mg/l. The average concentration of calcium in river water is 13.4 to15 mg/l and the average concentration of magnesium is 3.35 to 4.1 (Hem, 1989). Since magnesium hinders calcium absorption by plants, these data add support to statements made in the literature regarding the "poor" quality of the substrate in which Darlingtonia grows, but they do not verify these statements.

Conclusion

This paper points out some mistakes commonly encountered in the literature. The botanist who first collected this plant was William Dunlop Brackenridge who found the California Pitcher Plant in early October 1841 during the U. S. Exploring Expedition. Despite statements to the contrary, there is no recorded evidence which supports the viewpoint that Brackenridge collected a poor specimen because he was being chased by Indians. Darlingtonia is found in wet areas in northern California and western Oregon between sea level and 2600 meters. Darlingtonia is frequently, but not exclusively, found on serpentine soils, which tend to have high concentrations of magnesium and low amounts of calcium. According to the literature, Darlingtonia grows in a substrate with a pH ranging between 5.1 and 9.5 but this needs further testing throughout its range.

Regardless of the minor errors, the literature pertaining to Darlingtonia is interesting and informative; however, this unusual plant needs further research. As the distribution and habitat of Darlingtonia become more thoroughly documented throughout its range, biogeographers will be better able to explain the reasons for its distribution patterns.

Acknowledgements

Many thanks to Ken Goehring and Nancy Shepard for reviewing this article and to my thesis advisors Dr. Christine Rodrigue, Dr. Susan Place, and Dr. Kenneth Beatty.

References

Cheatham, Norden H. "Butterfly Valley botanical area." Fremontia Volume 4 No. 2 (July 1976): 3-8.

De Buhr, L.E. 1973. Distribution and reproductive biology of Darlingtonia californica. M.A. thesis, Claremont Graduate School.

Eastwood, Alice. 1945. "An account and List of the Plants in the Brackenridge Journal." California Historical Society Quarterly Volume 24 No. 4:337- 342.

Elder, Christine. 1994. "Darlingtonia californica: A complete list of literature as of 1994." In Reproductive Biology of Darlingtonia californica: A progress report and comprehensive bibliography. http://www.hpl.hp.com/botany/public_html/cp/pictures/darlingt/0074.htm (last updated 16 July 1996).

Freeman, Linda E. 1994. Distribution of Darlingtonia californica on Mt. Eddy, California. M.A. thesis, CSU Chico.

Gilman, Daniel Coit . 1899. The Life of James Dwight Dana. New York: Harper and Brothers.

Hem, John D. 1989. Study and Interpretation of the Chemical Characteristics of Natural Water. USGS.

Juniper, B.E., R.J. Robins, and D.M. Joel. 1989. The Carnivorous Plants. San Diego, California: Academic Press.

Lloyd, Francis Ernest. 1942. The Carnivorous Plants. Waltham, Massachusetts: Chronica Botanica Co.

Maloney, Alice Bay. 1945. "A Botanist on the Road to Yerba Buena." California Historical Society Quarterly Volume 24 No. 4: 321-325.

Mellichamp, T. Lawrence. 1983. "Cobras of the Pacific Northwest." Natural History (April 1983): 47-50.

Miesse, William C. 1993. Mount Shasta: An Annotated Bibliography. Weed, California: College of the Siskiyous.

Naeem, S. and J. Dusheck. 1985. "Plumbing the deathly depths of the California Pitcher Plant." Pacific Discovery (April/June 1985): 26-31.

Rondeau, J. Hawkeye. 1991. Carnivorous Plants of California. San Jose, California: By the author.

Sawyer, John. "Darlingtonia seeps." Fremontia Vol. 14 No. 2 (July 1986): 18.

Schnell, D.E. 1976. Carnivorous Plants of the United States and Canada. Winston-Salem, N. Carolina: John F. Blair.

Schoenherr, Allan A. 1992. A Natural History of California. Berkeley: University of California Press.

Slack, A.A.P. 1979. Carnivorous Plants. London: Ebury Press.

Torrey, John. 1853. "On the Darlingtonia californica, a new pitcher plant from Northern California." Smithsonian Contributions to Knowledge 6: 1-8.

Note: This unpublished paper was written in 1996 by Linda Freeman.


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