Feature Flowerby David Rogers
As this Spring Equinox edition of the DCQ comes online, most of the local populations of these harbingers of the oncoming wildflower season will be at, if not beyond, the zenith of their annual display. Thus for those who have had the fortune to be there when these showy-flowered herbs are at the peak of their flowering season, the memory of their displays (especially those of large colonies) may linger in the mind for a lifetime. In contrast, those who wait for the weather to become more suitable for hiking will have to settle for perhaps the sight of a late flower or two, for by then most of the plants will be in (if not beyond) their fruiting stage.
The genus Dodecatheon is represented in Santa Lucia Mountains by at least three taxa: two subspecies of the D. clevelandii complex (D. c. sanctarum and D. c. insularis) and D. hendersonii in its typical form. Of these, D. c. sanctarum is by far the most common Dodecatheon taxon in the Santa Lucia Mountains. It is also possible that other lesser taxa of both of these species occur in this region.
The name Dodecatheon, derived from the Greek words dodeca, twelve, and theos, gods, was applied by the Roman naturalist Pliny (the Elder, 23-79 AD) to a primrose or a similar plant with "a yellow root and leaves like the garden lettuce." This plant was said to be under the special protection of the twelve major (Olympian) gods. Although Carolus Linnaeus (1707-1778), the founder of the binomial nomenclature system, did not state why he chose this name for a North American genus, an explanation was provided by the eminent English botanist Sir William Hooker (1785-1865):
The Dodecatheon of modern authors has, however, nothing to do with the individual to which Pliny applied so grand a designation, as it is a native of the New World, and receives its name in allusion to the number of flowers, frequently twelve, which it bears in each umbel (Curtis's Botanical Magazine 64, plate 3622, 1837, as quoted in Ingram, 1963).
The application of Dodecatheon to this genus was also controversial. D. meadia (of eastern North America) had been introduced into English gardens by 1709, and in 1743 Mark Catesby, in his "The Natural History of the Carolinas, Florida and the Bahama Islands" (appendix. 2: 1. pl. 1), named it Meadia meadia, in honor of Richard Mead, a physician who was "a generous encourager of every useful branch of science." In 1751 Linneaus, in his "Nova plantarum genera", renamed the genus Dodecatheon, but in the following year Philip Miller, in his "Gardener's Dictionary," revived the name Meadia. Miller argued that: "the title of this genus was given by to it by Mr. Mark Catesby, F. R. S., in honor of the late Dr. Mead... but being himself no great botanist, Dr. Linnaeus was unwilling [in that] any plant should bear his name" (Miller as quoted in Thompson, 1953). It must be noted that Linnaeus did retain the name meadia for the species, so Mead was not completely ignored.
Linnaeus retained the name Dodecatheon in his "Species plantarum" of 1753, and as, by the accepted rules of botanical nomenclature, the priority of generic and specific epithets date to this text, the name is "official." The controversy was not completely ended, for in 1891 the German botanist Carl Ernst Otto Kuntze (1843-1907), revived the name Meadia in his "Revisio Generum Plantarum." Kuntze was an advocate of strict priority in nomenclature, and in this text he changed the names of more than 30,000 species.
Even though shooting stars are extremely conspicuous and easily recognized even by those who have no interest in plants, for a long time the relationships within genus remained poorly understood. In fact for more than a century Dodecatheon remained a monospecific genus (i.e., a genus comprised of only one species). In 1886, the eminent Harvard University botanist Asa Gray (1810-1888), in the first paragraph of his "Essay Toward a Revision of Dodecatheon," stated that:
Probably every botanist who has turned his attention to this genus has suspected it to be of more than one species. But those who have attempted to deal with the numerous now extant forms have been baffled in their endeavors to distinguish and define them. In the Synoptical Flora of North America I could do no better than to arrange the forms loosely under seven varieties. If I have now done better in the attempted discrimination of five species the credit is largely due to indications and specimens supplied to me by two western correspondents.
The species proposed by Gray were D. meadia, three taxa that had been proposed by other authors, and a new one that Gray named in honor of one of his western correspondents: D. hendersonii. D. hendersonii is one of the taxa that occurs in the Santa Lucia Mountains.
The further dividing of Dodecatheon was continued by other botanists, and one of most prominent in this regard was Edward E. Greene (1843-1915), the first staff botanist at the University of California. Over a period of a few decades, Greene described about 18 species and varieties among the Dodecatheon of western North America. These taxa included D. clevelandii, D. sanctarum and D. patulum, which would later be reclassified as subspecies of the four-fold D. clevelandii complex. D. clevelandii sanctarum is by far the most common shooting star in the Santa Lucia Mountains.
The first text to address Dodecatheon in a comprehensive manner was Henry J. Thompson's "The Biosystematics of Dodecatheon," which was published in 1953. In the introductory paragraph Thompson stated the following:
Although Dodecatheon is a genus of common plants that have long been the object of botanical collection and description of, little is known of their systematics. This is perhaps due to two factors. First, being relatively simple plants with a short underground caudex, a rosette of leaves, and occasionally a scapose inflorescence, they do not possess a complex morphology. The leaves are readily modified environmentally so as to render them virtually barren of taxonomic characteristics. Thus only the simple flower and fruit have afforded the taxonomist morphological characters with which to mark the evolutionary units. Secondly, the variation, both within the local populations and between populations, has made the species puzzling and difficult to classify... The modern taxonomist, supported by data from extensive collections as well as chromosome numbers and genetic analysis, still finds a scarcity of reliable taxonomic characters, and there is great difficulty in locating taxonomic characters to delimit demonstrable evolutionary units morphologically.
Although this statement provides more information than that of the equivalent statement by Gray, the conclusions concerning the confusing characteristics within the genus are nearly the same. From the more than 170 taxa proposed by 1953, Thompson was able to reduce this number to 27 (14 species and 13 subspecies). This number is similar to that listed on the USDA Natural Resources Conservation Service website (15 species and 15 subspecies), although there are differences in the taxa that are accepted. The USDA list can be found at http://plants.usda.gov.
In its natural distribution Dodecatheon is nearly endemic to the temperate and arctic regions of North America, for only one species, D. frigidum of northwestern Canada and Alaska, also extends to the eastern tip of Siberia. The genus is also primarily western North American, for most of the taxa occur west of the Great Plains.
The local representatives of Dodecatheon belong to the Purpureo-Tubulosa section of the genus, which Thompson divided into two subsections: the D. frigidum and D. hendersonii complexes. The former represents species that are restricted to the often cold environments of high interior mountains and arctic regions, while the latter is comprised of plants that occur in the more mild conditions of the Pacific Slope of temperate North America. One of the most noteworthy aspects of the D. hendersonii complex is its diversification in the southern portion of its range; it is here that it extends into the California Floristic Province, one of the world's greatest centers of plant evolution.
All shooting stars are perennial herbs with a short caudex and fleshy-fibrous roots. The leaves are strictly basal and produced in rosettes, and the showy flowers are produced on erect stems that terminate in umbels (i. e., an inflorescence in which the pedicles radiate from a single point, like the spokes of an umbrella). The umbels are few to many flowered and are subtended by bracts. The pedicels are erect when budding, but outwardly they turn downward before the maturation of the flower. The flower parts are in fives (or sometimes fours). The calyx is deeply cleft, and the segments are reflexed while the flowers are blooming, but become erect later on. The reflexed corollas are five-lobed, and have short tubes and dilated throats. The exserted stamens are produced on short, broad and often united filaments. The slender styles are singular and extend a short distance beyond the stamens. After anthesis, the pedicels once again become straight, so that the 5-valved or circumscissle seed capsules are held in an erect position. The small ovoid or angled seeds are many. The flowering season of the local plants begins as early as January, and may last until May. Although the plants are perennial, all the above ground features whither away after the maturation of the fruits, and new roots, leaves and flowering stems are produced with the next growth cycle.
One of the most interesting characteristics of Dodecatheon species is that they reproduce not only from seeds, but also vegetatively via the roots. New plants are continuously cloned from larger roots that become detached from the caudex (as in D. clevelandii), or by rice grain-like bulblets that detach from the mother plant during the dormant season (as in D. hendersonii). It is this mode of reproduction that is responsible for the dense clumps of plants that are often encountered in the field.
The local representatives of Dodecatheon are as follows:
This is by far the most common Dodecatheon in the Santa Lucia Mountains, and, as evidenced by many herbarium specimens, it is rather evenly distributed in suitable habitats from the Monterey Peninsula southward. Northward D. c. sanctarum extends to western Santa Clara County and southwestern San Mateo County, and southward to the far western Transverse Ranges of Santa Barbara County, and also in the mountains of southeastern Ventura County.
The type specimen was collected by Reason Alpha Plaskett in the Pacific Valley area of the Big Sur Coast in February of 1898, and was one of a large group of specimens that Plaskett sent to Alice Eastwood, who was then the curator of botany at California Academy of Sciences. Mr. Plaskett was a member of a large family of early settlers in the Pacific Valley region, and his family hosted Eastwood on two of her botanical expeditions to the Santa Lucia Mountains.
This taxon was first described, as D. sanctarum, by Edward Greene in 1903 (Pittonia 5: 113). The only additional information supplied by Greene was: "Santa Lucia Mountains, California. R. A. Plaskett, Feb., 1898; distributed for D. clevelandii, but not related to that; much more like D. hendersonii." Although Greene failed to state why he chose the name sanctarum, derived from the Latin name for holy, it seems that it had no connection with the common name for D. c. clevelandii: "Padres Shooting Star." "Padres Shooting Star" appears to have first been used by Jepson in 1939. Jepson also thought that Greene's D. sanctarum was perhaps more closely related to D. hendersonii than to D. clevelandii.
Greene also described other taxa that would later become synonyms for D. c. sanctarum. In 1895 he described D. patulum var. gracile (Erythea 3: 72), which was based on a plant at the U. C. Botanical Gardens that had been grown from roots collected by Joseph Burtt Davy on the slopes of Loma Prieta, Santa Clara County. Greene noted that the "segments of the corolla [were] white, linear, narrow, [and] elegantly twisted," and that it was "an exceedingly beautiful variety, if a mere variety, of the species so common on the low plains of the interior of the country." Jepson (1925) assigned this variety to D. hendersonii. In 1903 Greene also described D. laetiflorum (Pittonia 5: 112-113), the type of which was collected in "groves of oak at Gilroy, Santa Clara County" by Carl F. Baker (#1945) in April of that year. Jepson (1925) considered this to represent a variety of D. patulum. Another synonym, in part, is D. meadia var. macrocarpum A. Gray (Botany, vol. 1, Geological Survey of California, 1876). Gray had grouped a specimen, collected by William H. Brewer in the Santa Suzana Mountains of Ventura County (Brewer #217), with those of plants that extended "from Alaska southward."
D. c. sanctarum is comprised of plants that range from about 1.2 to 4 dm. (5-16") tall. The leaves are about 2 to 6 cm. long (inclusive of the petioles), and the blades are broadly oblanceolate to spatulate or nearly elliptic. The margins are mostly crisped and/or irregularly crenate. The umbels are comprised of about 3 to 7 flowers on that are on pedicels about 2 to 5 cm. long. The corolla lobes, which are generally oblong-acute and about 1 to 2 cm. long, are rose-lavender or white, while the tubes are marked with three bands of color: the upper is white, the central is yellow, and the lower is blackish-purple. The filament tubes and connectives are dark and usually have a yellow or whitish spot at the base of each anther, and the pollen is usually yellow. The seed capsules are about 8 to 13 mm. long. The plants are usually found below about 4,000 ft.
Although specimens representative of this taxon had been collected at an early date, it was not until 1953 that they were recognized as representing a distinct evolutionary unit. An example is Asa Gray's assignment of a specimen from Guadalupe Island (Palmer #55, GH) to D. meadia in 1876, which, in 1886, he reassigned it to D. jefferyi. D. clevelandii insularis is the only shooting star that occurs on Guadalupe Island.
Island Shooting Stars are the most robust members of the D. clevelandii complex. The leaves, inclusive of the petiole, range from about 6 to 18 cm. long. The oblanceolate to spatulate blades range from 1 to 6 cm. wide, and gradually (or sometimes abruptly) taper to the petiole. The leaf margins are almost always crisped and/or toothed. The flowering scapes range from 12 to 45 cm. tall, and the umbels are 5 to 19 flowered. The pedicels are about 2 to 10 cm. long while in flower, but become longer with the maturation of the seed capsules. The corolla lobes are rose-lavender or white, and are subtended by bands of white, yellow and dark maroon. The filament tubes, connectives and anthers are uniformly dark maroon to black. D. clevelandii subsp. insularis occurs mostly in grasslands and open woodlands (but sometimes chaparral), and the peak of it flowering season extends from early March to mid-April.
The distribution of D. clevelandii insularis is curious, for to the south it is restricted to the islands off the coast of Southern California and Baja California, while to the north it occurs on the mainland and away from the coast. To the south it reaches Guadalupe Island, which is about 200 miles off the coast of central Baja California. This remote island represents the far southern outpost of the California Floristic Province. To the north it extends to Monterey County and probably San Benito County.
As this is the most widespread Dodecatheon of the Pacific Slope of temperate North America, I have taken the liberty to call it "Pacific Shooting Stars." Its range extends from southern Vancouver Island, British Columbia, to the mountains of California. In the Coast Ranges it reaches its most southern limit in San Benito and Monterey Counties, while in the Sierra Nevada, inclusive of its subspecies, it extends to Kern County. There is also a remote population of the typical species far to the south in the Bear Valley region of the San Bernardino Mountains.
The southernmost distribution of D. hendersonii in the Coast Ranges is curious. Most of the many other species that reach their most southern limit in this region become more restricted to coast, but D. hendersonii, after occurring along the coast in the San Francisco region, veers inland. There are a number of herbarium specimens from the inner Coast Ranges of San Benito County, and the southernmost station may be in the San Antonio Valley of Monterey County. I have not had the time to check the holdings of herbaria other than the University of California, Berkeley, so it is possible that I may be mistaken in this regard. In any case, the most southern documented site that I am aware of is near the San Antonio River in Fort Hunter Liggett. The specimen (Rimo Bacigalupi #7570, JEPS, 3/22/1961) was collected along the road to The Indians, 2 miles south of the upper ford across the San Antonio River, just south of China Gulch. Reports in literature about the occurrence of D. hendersonii on the Monterey Peninsula are based on Jepson, 1939, which, in turn, was based on a misidentified specimen. The specimen, collected by N. K. Berg at Monterey in February of 1904, actually represents D. clevelandii sanctarum. Also, in 1888 Edward Greene stated that D. cruciatum (D. hendersonii var. cruciatum) "is the common species at and about San Francisco, extending southward to Monterey, perhaps Santa Barbara, and eastward to Mt. Diablo." The occurrence of this taxon south of the Santa Cruz Mountains was never verified.
Asa Gray named this species for Louis Forniquet Henderson (1853-1942), an early plant collector in the Pacific Northwest, who sent to Gray "excellent specimens of this peculiar species." According to Love (1999), Henderson was at that time a high school teacher in Portland, but he later served as the first professor of botany at the University Idaho (from 1893 to 1911), and as the curator of the University of Oregon Herbarium (from 1924 to 1936).
D. hendersonii is a fairly robust species. The leaves, inclusive of the petioles, are about 5 to 15 cm. long and 2 to 6 cm. wide, and the blades are spatulate to elliptic and obtuse to truncate at the apex. The blades abruptly taper to the petiole, and the margins are entire or sometimes slightly toothed. The flowering scapes range from about 12 to 48 cm. tall, and the umbels produce from 3 to 17 flowers. The corolla lobes range from about 6 to 23 mm. long, and vary from magenta to deep lavender or white. The tubes are maroon at the base and have a narrow band of yellow or white (or sometimes both) above. The filament tube, connectives and anthers are dark maroon to black. The capsules are about 8 to 15 mm. long. The plants usually occur on north facing woodland slopes, and the flowering period is from February to May.
1a. Leaf blades mostly broadly obovate to ovate or elliptic, obtuse to nearly truncate at the apex, and abruptly tapering to the petiole, the margins entire or sometimes slightly toothed. Filament tubes 1.5 to 2.5 mm. wide, the tube, connectives and anthers uniformly dark maroon to black. Flowering plants producing rice grain-like bulblets among the roots. Plants generally of shady, north facing slopes...
1b. Leaf blades mostly oblanceolate to spatulate and gradually tapering to the petiole, the margins usually crisped and irregularly toothed. Filament tubes 3 to 4 mm. wide, the tube, connectives and anthers uniformly dark maroon to black (in D. clevelandii insularis) or in some way yellow or white. Bulblets not produced among the roots. Plants generally of sunny habitats:
2a. Connectives yellow, filament tube lacking a yellow or whitish spot below each anther...
2b. Connectives maroon to black, filament tube with or without a yellow or whitish spot at the base of each anther:
3a. Filament tube and anthers uniformly dark. Plants 12 to 45 cm. tall, umbels 5 to 19 flowered, leaves 6 to 18 cm. long...
3b. Filament tube with a yellow or whitish spot at the base of each anther. Plants 4 to 38 cm. tall, umbels 1 to 9 flowered, leaves 2 to 6 cm. long:
4a. Anthers 3 to 4.2 mm. long, usually yellow or whitish, the tips acute to obtuse. Plants 15 to 38 cm. tall, flowers 3 to 9 per umbel...
4b. Anthers 1.5 to 3 mm. long, dark maroon or purple, the tips obtuse to notched. Plants 4 to 20 cm. tall, flowers 1 to 4 per umbel...
The preceding key was extended to include all the members of the D. clevelandii complex because it is possible that more of these occur in the Santa Lucia Mountains. I came across a report stating that D. clevelandii subsp. clevelandii, Padres Shooting Stars, occurs in Monterey County, but I suspect that it was based on specimens that were not identified to a subspecific level. According to the CalFlora web site, a specimen was collected at the "San Antonio River (Salinas River-Pacific Ocean Drive), US route 101 bridge, Camp Roberts" by Larry M. Page in March of 1994. The specimen is at the Illinois Natural History Survey Herbarium. The CalFlora web site also states that this taxon is "reported as probably present" in Monterey County in literature, and gives as its references Lum/Walker/Munz68/Calflora/JM93. Two of these, Munz68 (Philip Munz' "A California Flora") and JM93 (the "Jepson Manual" of 1993), do not report this taxon as probably present in Monterey County. The Lum and Walker texts are both unpublished master's theses, and "CalFlora," I assume, is the site itself.
D. clevelandii was named for Daniel Cleveland (1838-1929), a San Diego lawyer, amateur botanist and a co-founder of the San Diego Society of Natural History. D. clevelandii subsp. clevelandii is a fairly robust species (the scapes are up to 40 cm. tall and the umbels can have as many as 16 flowers) that occurs in coastal Southern California from Los Angeles County southward, and extends to around Rosario on the coast of Northern Baja California.
D. clevelandii subsp. patulum (Greene) H. J. Thompson, Lowland Shooting Stars, occurs in Monterey County, but to the best of my knowledge it has been reported only from the Gabilan Mountains. In any case, the taxon is close enough to have a genetic influence on local plants. For example, plants from the Jolon area exhibit characteristics that are intermediate between this D. c. patulum and D. c. insularis (Thompson 1953). This is the smallest member of the D. clevelandii complex, and it occurs in the Central Valley and adjacent Sierra Nevada foothills from Tehama County to Kern County, and from the San Francisco Bay Area to San Benito and Monterey Counties in the Coast Ranges.
Dodecatheon belongs to the Primrose Family (Primulaceae), and it is closely related to two other showy-flowered genera, Primula (the primroses) and Cyclamen. The many varieties of both of these genera rank among the most common of all garden plants, and are most often seen growing in pots or along the borders of landscaped areas. The resemblance between Dodecatheon and Cyclamen is readily evident, for both have flowers that are upside down and inside out. In Cyclamen, however, the flowers are produced singularly on the scapes, while in Primula the flowers are often produced in umbels, as they are in Dodecatheon.
Although Primula and Cyclamen have long been standards of horticulture, Dodecatheon has also entered the trade. I have yet to see a shooting star in a private garden, but the large number of commercial web sites advertising Dodecatheon species indicates that there is much interest. Shooting stars seem to be particularly popular in Europe, and D. hendersonii (along with D. meadia and D. pulchellum) is included in the Royal Horticultural Society's "Award of Garden Merit Plants" (London, 1994). According to Ingram (1963) D. clevelandii is also "fairly common in cultivation," but as this is an older text, perhaps it is even more so at the present time.
One last note. According to Mitchem (1991), the leaves and roasted roots of D. hendersonii were eaten by some Indian tribes as a survival food. Indians in British Columbia used the flowers of D. jefferyi "as a charm '... to obtain wealth and make people give presents' and also as a love potion used by women '... to obtain the love of men and help them control men.'"