Dodecatheon (Primulaceae) of North America north of Mexico

Revision of Dodecatheon (Primulaceae)

James L. Reveal

Adjunct Professor, Cornell University,
Ithaca, New York 14853-4301, U.S.A.
Professor Emeritus, University of Maryland
College Park, Maryland 20742-5815, U.S.A.
Honorary Curator, The New York Botanical Garden,
Bronx, New York, 10458-5026, U.S.A.

Lectotype of Dodecatheon jeffreyi Van Houtte
Fl. Serres Jard. Eur. 16: t. 1662. 1867
(Image courtesy of GH with thanks to Susan L. Kelley)

5. DODECATHEON Linnaeus, Sp. Pl. 1: 144. 1753; Gen. Pl. ed. 5, 71. 1754 · Shootingstar [Greek dodeca, twelve, and theos, god, a fanciful name given by Pliny to a primose protected by the gods.]

       Meadia Miller, nom. illeg.; Primula L. subg. Auriculastrum Schott sect. Dodecatheon (L.) A. R. Mast & Reveal

Herbs perennial; scapes erect or nearly so, glabrous or glandular-pubescent or -puberulent.   Caudex generally short-lived and not obvious at anthesis being short, fleshy, slender and erect to slightly spreading, or obvious at anthesis and elongate, thick and nearly horizontal, sometimes long-lived, woody and horizontal; roots arising anew annually from terminal buds on an obscure caudex, from small adventitious buds on the roots persisting on caudex, or persistent but replaced annually; bulblets sometimes present. Leaves in basal rosettes, often decurrent; petiole slender or winged for all or much of its length; blade linear to oval, simple, entire or variously dentate or crenulate, sometimes undulate, glabrous or glandular-pubescent or -puberulent, abruptly or gradually tapering to petiole, rarely cordate. Inflorescences generally umbellate, terminal, occasionally reduced to a single flower; bracts mostly lanceolate; pedicels slender, recurved in flower, generally straight and longer in fruit, glabrous or variously glandular. Flowers 1–25(–125) per umbel, 4–5-merous, monomorphic, nodding at anthesis; calyx tubular, lobes spreading in early anthesis becoming forcibly reflexed at anthesis and often remaining so in fruit, glabrous or glandular-pubescent or -puberulent; corolla glabrous or occasionally minutely glandular abaxially, tube short, lobes long and strongly reflexed, white to pink or violet, or magenta to purple with a yellowish and/or whitish base, often with a purple-, maroon- or reddish-ring at point of reflection, ring sometimes absent; stamens 5, exserted, connivent, filaments short, broad, free or partially to fully fused and forming a tube, connective smooth to rugose, generally purplish, maroon or blackish, occasionally yellow, anthers basifixed, anther sacs purplish to maroon or yellow, infrequently reddish or purplish, sometime with reddish or maroon speckles, dehiscent longitudinally on adaxially surface; ovary superior, style filiform, usually exserted beyond stamens, stigma capitate, sometimes up to twice or more diameter of style. Capsules 1-celled, valvate or operculate with a small lid, glabrous or glandular-puberulent (at least apically), usually 5-parted but often further subdivided into 10 or more erect and inwardly curved to outwardly spreading, greenish-yellow to yellow, reddish-brown or purple teeth-like distal segments; seeds 50–200, small, dark brown to black, globose to ovoid or quadrate, irregularly aveolate formed by collapse of minute bulbous cells, with or without a thin membranous margin. x = 22.

     Species 17 (17 in the flora): North America and ne Asia (Russian Far East).

     Dodecatheon is notoriously difficult taxonomically. Members of the genus are widespread throughout much of North America, extending from northwestern Mexico to the Arctic in Alaska and northwestern Canada. The taxonomic boundaries between species are sometimes blurred, and the variation within the more widespread species (such as the eastern D. meadia and the western D. pulchellum) can be bewildering. Nearly all recognized species are replete with an array of synonyms, and many names, used in the past, have proven to be not legitimate or misapplied adding to the nomenclatural morass.

     The genus can be subdivided into two groups, but not the three recognized by H. J. Thompson (1953), based primarily on the rugose (Sect. Purpureo-tubulosa R. Knuth) versus smooth (Sect. Dodecatheon) nature of the anther connective (A. R. Mast et al. 2004). Those species with an enlarged stigma (notably Dodecatheon jeffreyi, the type of Sect. Capitata H. J. Thompson) falls into the latter taxon. Even so, as is noted in the key, both D. hendersonii and D. subalpinum occasionally have smooth connectives, and D. poeticum, a member of Sect. Dodecatheon, has rugose connectives.

     Recognition of Dodecatheon creates a paraphyletic Primula (M. Källersjö et al. 2000; A. R. Mast et al. 2001, 2004; L. Martins et al. 2003). Dodecatheon falls within Primula subg. Auriculastrum Schott, and is seemingly allied with the Sierra Nevada endemic P. suffrutescens A. Gray. The two share an involute leaf vernation. While Primula has a base number of x = 11, Dodecatheon is x = 22 as H. J. Thompson (1953) has shown that 2n = 66 plants are

triploids, not hexaploids. The morphological differentiation of the monophyletic Dodecatheon clade is specialization associated with the evolution of buzz-pollinated flowers (e.g., similar to that found in Solanum) coupled with a monomorphic rather than the heterostylous floral condition typical of Primula (A. R. Mast et al. 2004). In addition was fixation of recessive alleles at the heterostyly linkage group (pin phenotype) and at least six other traits that likely arose with the origin of Dodecatheon. One major change preceded its origin (flower coloration, a transfer exaptation in Dodecatheon), and another followed it (rugose anther connectives, an adaptation to buzz pollination). The first accounts for the shared floral colors among P. suffrutescens and Dodecatheon. The second, significantly, provides “footing” for the pollinator while buzzing the flower. In general, anthers with a rugose connective are larger than those with a smooth connective, and the anthers of Dodecatheon are considerably larger than those of Primula. Also related to these changes are the generally fused filaments forming a tube, thick connectives, and elongated anthers (L. D. Harder & R. M. R. Barclay 1994). These data have resulted in the transfer of all species of Dodecatheon to Primula (A. R. Mast & J. L. Reveal 2006), and the establishment of Primula sect. Dodecatheon (L.) A. R. Mast & Reveal within subg. Auriculastrum. For those wishing to adopt this concept, the appropriate names are provided in synonymy.

     Pollination studies in Dodecatheon are limited as only two species (D. amethystinum and D. meadia) have been examined in detail (L. W. Macior 1964, 1970). According to H. J. Thompson (1953: p. 77) flowers not visited by a pollinator can self-pollinate.

     Use of the taxonomic rank of variety, rather than subspecies, was discussed fully by N. H. Holmgren (1994).

     Well-preserved flowers of Dodecatheon are critical for identification. The nature of the anther (especially whether the connective is smooth or rugose) and the color patterns of the corolla are important observations that should be made in the field as the flowers can lose color when dried. In particular one should check for small bulblets (about the size of grains of rice) that are produced among the roots of a few species at anthesis. Vegetative plasticity in response to both moisture and time of season contributes to extremes in variation, especially height and robustness of plants, and length and breath of leaves. As some species of Dodecatheon tend to flower in moist soils of grassy meadows (or even in running water), but only in places that tend to dry out, the length of time a particular site is wet can be a significant factor in determining the overall size of the plant and leaves. To what extent some populations within D. pulchellum var. zionense are products of their local environment, rather than being genetically distinct from other expressions of the species, remains to be shown. In many species one can find an ecological gradient with some plants near a stream bank having longer, broader leaves than those on the drier slopes away from the stream. As there is often a continuum, it is easy to notice in the field if alert to variation within the population. In others, such as D. jeffreyi, which tends to be in moist places throughout most of its growing cycle, elevation is a factor that seemingly plays a role in the vegetative plasticity. Small, slender, high-elevation plants of D. jeffreyi found mainly above 3500 m in the southern Sierra Nevada of California are sometimes differentiated as subsp. pygmaeum (H. M. Hall) H. J. Thompson. Their assignment to D. jeffreyi is by virtue of the plants being glandular even though they closely approach D. alpinum in stature and the size and shape of its leaves. The same is true for the watsonii phase of D. pulchellum. Both are assigned to synonymy here.

     Adding to the complexity is the need to observe the valvate or operculate dehiscence of the capsule, and the degree of firmness of the capsule wall. Dehiscence is clearly valvate in some species (e.g. Dodecatheon pulchellum) with no hint of a line of separation. In other species (e.g. D. jeffreyi) the capsule opens on a transverse line near the top of the fruit, shedding a small cap (operculum) often with an intact style. Then there are specimens that have both valvate and operculate dehiscence even on the same plant (e.g., D. clevelandii). The distinction is further complicated by the “line of separation” which is often distinguished by the upper portion of the capsule being of a darker color that can also (in some) be glandular. The small operculum may consist of little more than the base of the style and be apical of the “line” that is indicative of the depth to which the capsule will split into five, ten or more teeth-like segments. With age, the (usually) inwardly curved teeth shed the operculum and then fall away resulting in what appears to be a toothless, circumscissile capsule. In valvate capsules, the teeth form at the apex of the fruit resulting (often) in a splitting of the style into parts. With age, the (usually) outwardly curved teeth shed the fragments of the style. The teeth usually remain attached to the body of valvate capsules, but not always resulting again in what appears to be a toothless, circumscissile capsule.

     Finally, be aware that occasionally two or more species may occur in close proximity. As the distinguishing features used here to recognize species can be difficult to observe without a critical examination of the flower, and in some cases the root system and capsules, a seemingly variable population may, in fact, be a mixture of plants of different species with an occasional sterile hybrid added to the mix. This can result in herbarium collections composed of two entities adding even more difficulty to the identification of these plants.

     Shootingstars are widely cultivated, and numerous cultivars have been developed. Essentially all species are found in nurseries, although several are misidentified. Most can be grown is sunny to partially shaded places in dampish soil (when flowering) that slowly dries (when dormant). Most flower during the spring months and are especially attractive when planted in masses. None of the species has much of a history of medicinal use by Native Americans. However, flowers of various species were used decoratively, to attract men, and to aid youngsters to sleep. Leaves were occasionally used as an eye wash or as an oral gargle.

SELECTED REFERENCES   Holmgren, N. H. 1994. Redefinition of Dodecatheon dentatum (Primulaceae) and rationale for use of varietial rank. Brittonia 46: 87–94.   Källersjö, M., G. Bergqvist and A. A. Anderberg. 2000. Generic realignment in primuloid families of the Ericales s.l.: A phylogenetic analysis base on DNA sequences from three chloroplast genes and morphology. Amer. J. Bot. 87: 1325–1341.   Martins, L., C. Oberprieler and F. H. Hellwig. 2003. A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data. Pl. Syst. Evol. 237: 75–85.   Mast, A. R., S. Kelso, A. J. Richards, D. J. Lang, D. M. S. Feller and E. Conti. 2001. Phylogenetic relationships in Primula L. and related genera (Primulaceae) based on noncoding chloroplast DNA. Int. J. Pl. Sci. 162: 1381–1400.   Mast, A. R., D. M. S. Feller, S. Kelso and E. Conti. 2004. Buzz-pollinated Dodecatheon originated from within the heterostylous Primula subgenus Auriculastrum (Primulaceae): A 7-region cpDNA phylogeny and its implications for floral evolution. Amer. J. Bot. 91: 926–942. Mast, A. R. & J. L. Reveal. 2006. Transfer of Dodecatheon to Primula (Primulaceae). Brittonia 00: 000–000.   Thompson, H. J. 1953. The biosystematics of Dodecatheon. Contr. Dudley Herb. 4: 73–154.   Trift, I., M. Källersjö and A. A. Anderberg. 2002. The monophyly of Primula (Primulaceae) evaluated by analysis of sequences from the chloroplast gene rbcL. Syst. Bot. 27: 396–407.

1. Connective transversely rugose.

     2. Stigma enlarged, at least twice the diameter of style; seeds with a thin membrane.

          3. Flowers 4-merous; capsules valvate; leaves linear to linear-oblanceolate, glabrous; inflorescences glabrous, occasionally with pedicels and bracts sparsely glandular-puberulent . . . . . . . . . . . . 7. Dodecatheon alpinum

          3. Flowers 4–5-merous; capsules operculate or valvate; leaves narrowly oblanceolate to oblanceolate or spatulate, glabrous or glandular-pubescent; inflorescences generally glandular-pubescent.

               4. Flowers 5-merous; capsules valvate; leaves and inflorescences glandular-puberulent; anther tips acute; corolla tube covering base of anthers, yellow . . . . . . . . . . . . . . . . . . . . . . . . . 5. Dodecatheon redolens

               4. Flowers 4–5-merous; capsules operculate or infrequently valvate (often on same plant); leaves and inflorescences glabrous or glandular-puberulent; anther tips truncate to obtuse; corolla tube not covering base of anthers, white or rarely yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Dodecatheon jeffreyi

     2. Stigma not enlarged, not much broader than the style; seeds without a thin membrane.

          5. Plants generally producing bulblets at anthesis.

               6. Leaves strongly decurrent, petioles winged or nearly so; flowers 4–5-merous (often on same plant); plants (0.7–)1–5(–5.5) dm; corolla lobes 0.6–2.5(–2.8) cm; below 2100 m . . . . . 2. Dodecatheon hendersonii

               6. Leaves slightly decurrent, petiole generally not winged; flowers 5-merous; plants 0.7–1.5(–2.5) dm; corolla lobes 0.5–0.9(–1.2) cm; above 2100 m . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Dodecatheon subalpinum

          5. Plants not producing bulblets at anthesis.

               7. Stamens free, filaments rarely partially fused; plants glabrous, or if minutely glandular-puberulent, then of southeastern British Columbia, southwestern Alberta, Saskatchewan, eastern Washington, northern Idaho and western Montana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Dodecatheon conjugens

               7. Stamens not free, filaments fused into a tube; plants glabrous, or if glandular-puberulent, then of south-central Washington and north-central Oregon.

                    8. Leaf-blades gradually tapering to petiole; plants glandular-puberulent throughout; south-central Washington and adjacent north-central Oregon . . . . . . . . . . . . . . . . . . 14. Dodecatheon poeticum

                   8. Leaf-blades narrowing abruptly to petiole; plants not glandular-puberulent throughout; California.

                        9. Pedicels and calyx glabrous . . . . . . . . . . . . . . . . . . . . . . . 2. Dodecatheon hendersonii (in part)

                        9. Pedicels and calyx glandular-puberulent . . . . . . . . . . . . . . . . . . . . . . . 4. Dodecatheon clevelandii

1. Connective smooth, occasionally longitudinally wrinkled.

    10. Caudex horizontal, often woody; roots reddish; inland arctic or subarctic regions . . 8. Dodecatheon frigidum

    10. Caudex vertical to slightly horizontal, not woody, sometimes lacking; roots whitish; not of inland arctic or subarctic regions.

           11. Plants generally producing bulblets at anthesis.

                  12. Leaves strongly decurrent; petiole winged or nearly so; flowers 4–5-merous (often on same plant); plants (0.7–)1–5(–5.5) dm; corolla lobes 0.6–2.5(–2.8) cm; below 2100 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Dodecatheon hendersonii (in part)

                  12. Leaves slightly decurrent; petiole generally not winged; flowers 5-merous; plants 0.7–1.5(–2.5) dm; corolla lobes 0.5–0.9(–1.2) cm; above 2100 m . . . . . . . . . . . . . . 3. Dodecatheon subalpinum (in part)

           11. Plants not producing bulblets at anthesis.

                  13. Wall of capsule thin, flexible.

                         14. Leaf-blades gradually tapering to petiole; corolla lobes magenta to lavender or, if white, then plants of eastern North America.

                                15. Filaments free; coastal mountains, sw Washington and nw Oregon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Dodecatheon austrofrigidum

                                15. Filaments fused into a tube; widespread.

                                       16. Pollen sacs reddish; midwestern and mid-Atlantic North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Dodecatheon amethystinum

                                       16. Pollen sacs maroon to black or yellow; western North America.

                                             17. Filament-tubes yellow, or if purplish then connective smooth or merely longitudinally wrinkled at full anthesis . . . . . . . . . . . . . . . . . . . . . . . . 13. Dodecatheon pulchellum

                                             17. Filament-tubes maroon, generally rugose . . . 14. Dodecatheon poeticum (in part)

                         14. Leaf-blades abruptly taping to petiole; corolla lobes white or rarely pale lavender to pink.

                                18. Filaments and basal portion of connective yellow; corolla white, rarely lavender; Arizona and New Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Dodecatheon ellisiae

                                18. Filaments and connective dark maroon to black; corolla white or pale lavender to pink.

                                       19. Corolla lobes white; Pacific Northwest . . . . . . . . . . . . . . . . 9. Dodecatheon dentatum

                                       19. Corolla lobes pale lavender to pink; Utah . . . . . . . . . . . . . . 11. Dodecatheon utahense

                  13. Wall of capsule thick, firm.

                         20. Filament-tube maroon; Pacific Northwest . . . . . . . . . . . . . . 14. Dodecatheon poeticum (in part)

                         20. Filament-tube yellow; eastern United States.

                                21. Leaf-blade abruptly tapering to a petiole . . . . . . . . . . . . . . . . . . 16. Dodecatheon frenchii

                                21. Leaf-blade gradually tapering to a petiole . . . . . . . . . . . . . . . . . . 17. Dodecatheon meadia

1. Dodecatheon conjugens Greene, Erythea 3: 40. 1895 · Bonneville shootingstar E

Primula conjugens (Greene) A. R. Mast & Reveal

Plants 0.5–3(–4) dm, glabrous or glandular-puberulent. Caudex not obvious at anthesis; roots whitish; bulblets absent. Leaves 3–13(–18) ´ 0.7–2.5(–4) cm, generally not decurrent to base; blade usually abruptly tapering to a slender (at least basally) petiole, narrowly oblanceolate to spatulate or obovate, glabrous or glandular-puberulent, margin entire. Inflorescences 1–7(–10)-flowered; bracts lanceolate to broadly lanceolate, 3–10 mm; pedicels 1–5 cm at anthesis, glabrous or glandular-puberulent. Flowers 5-merous; calyx 5–12 mm, light green to yellowish, occasionally finely purple-speckled or -dotted, glabrous or glandular-puberulent, tube 2–6 mm, lobes 3–7 mm; corolla lobes 0.7–2.5(–3.5) cm, magenta, occasionally white, tube yellowish with a thin, wavy, purplish-red ring; filaments free and 0.5–1.5 mm, infrequently fused and tube 0.5–1.5 ´ 1.5–5 mm, yellowish or dark maroon; anthers 5–9 mm, pollen sacs maroon or yellow, sometimes yellowish and speckled maroon, rarely with reddish-purple to purple speckles, connective rugose, maroon to purple, infrequently yellowish or even light blue to whitish; stigma not enlarged. Capsules cylindric-ovoid, 8–17(–22) ´ 4–6(–8) mm, tan, often striped with purple, glabrous, operculate or rarely valvate; walls thin. 2n = 44.

     Varieties 2 (2 in the flora): w North America.

     Both Dodecatheon conjugens and D. poeticum occur in close proximity in the Columbia River Gorge of Oregon and Washington. Some specimens here assigned to var. conjugens may have a few, scattered, minute glands on the pedicels that might indicate past intergradation with D. poeticum (e.g., G. N. Jones 6286, ORE; R. R. Halse 3790, OSC, WTU). Dodecatheon poeticum is densely glandular not only on the pedicels, but on the calyx and scape. The type of minute glandular-puberulent seen on var. conjugens found along the Columbia River west of The Dalles is somewhat similar to that seen on var. viscidum in western Montana and Canada. Some plants referred here to D. conjugens have slightly fused filaments which may indicate some intergradation with D. pulchellum var. pulchellum. This suggestion is supported by the tendency in the same plants to have narrower leaves.

     Some newly immerged flowers tend to have connectives that are less rugose than normal. This is particular true of some populations in southern Alberta and, to a lesser degree, in Saskatchewan.

1. Leaves, scapes and pedicels glabrous . . . . . . . . . . . . . . . . . . . . . . 1a. Dodecatheon conjugens var. conjugens

1. Leaves, lower scapes and pedicels glandular-puberulent . . . . . . . . . . . 1b. Dodecatheon conjugens var. viscidum

1a. Dodecatheon conjugens Greene var. conjugens · Bonneville shootingstar E

Leaves: blades glabrous. Scapes glabrous. Inflorescences: pedicels glabrous. Flower: calyx glabrous; connective maroon. 2n = 44.

     Flowering spring-early summer. Moist slopes and meadows, often in sagebrush communities and conifer woodlands or in alpine meadows; 50–2900(–3200) m.; Calif., Idaho, Mont., Nev., Oreg., Wash., Wyo.

     Bonneville shootingstar is widely scattered east of the Cascade Ranges from northeastern California (Modoc County) and northwestern Nevada (northern Washoe County) northward through Oregon to Washington, then eastward into the Wallowa Mountains of northeastern Oregon. In the Rocky Mountains the variety is found in central and northern Idaho eastward into western Montana and the northern two-thirds of Wyoming as far east as the western edge of the Great Plains. High elevation plants of var. conjugens in western Wyoming approach var. viscidum in sometimes having a few minute glands on the pedicels, making a distinction between the two rather arbitrary. Generally the lower scape of var. viscidum is also glandular-puberulent.

1b. Dodecatheon conjugens Greene var. viscidum (Piper) H. Mason ex H. St. John, Fl. SE. Washington, 311. 1937. ·    Sticky shootingstar E

Dodecatheon viscidum Piper, Bull. Torrey Bot. Club 28: 43. 1901; D. conjugens var. beamishiae B. Boivin; D. conjugens subsp. viscidum (Piper) H. J. Thompson; Primula conjugens var. viscidum (Piper) A. R. Mast & Reveal

Leaves: blades generally glandular-puberulent at least marginally. Scapes generally glandular-puberulent at least basally. Inflorescences: pedicels glandular-puberulent. Flower: calyx glabrous, occasionally glandular; connective yellow at least apically, infrequently maroon. 2n = 44.

     Flowering spring-early summer. Moist slopes and meadows in sagebrush communities and in conifer woodlands; (500–)1100–2800 m; Alta., Sask., B.C., Idaho, Mont., Wash.

     Sticky shootingstar is found throughout the northern range of the species from southeastern British Columbia, southern Alberta and southwestern Saskatchewan south into eastern Washington (Lincoln and Spokane counties), northern Idaho (as far south as Fremont County) and western Montana to the edge of the Great Plains. The variety appears to be disjunct near Tompkins, Saskatchewan, Canada (Looman 10304, OSC, UBC, UC, UTC) but it approaches Dodecatheon pulchellum var. cusickii (now otherwise known from here) as plants with both smooth and rugose connectives have been seen. The distribution of the glandular condition varies on the plant itself. Most have at least the lower portion of the scape glandular-puberulent, but the leaves and pedicels can be either glandular or infrequently glabrous. Only rarely is the entire scape glandular and then the plants are confined mainly to the Waterton Lakes National Park area but can be found occasionally elsewhere in Alberta. The calyx is rarely glandular. Even within the established range of var. viscidum, some (but by no means all) populations may consist of both glandular and a few non-glandular individuals. In some instances, only the pedicels have small, sparse glands; in Wyoming such plants are assigned arbitrarily to var. conjugens. Only those plants with distinctly rugose connectives are termed here var. viscidum. H. J. Thompson (1953) assigned a few Montana specimens with smooth or horizontally wrinkled connectives to var. viscidum but here they are assigned to D. pulchellum var. cusickii.

2. Dodecatheon hendersonii A. Gray, Bot. Gaz. 11: 233. 1886 · Mosquito-bill E

Dodecatheon hansenii (Greene) H. J. Thompson; D. hendersonii subsp. cruciatum (Greene) H. J. Thompson; D. hendersonii var. hansenii Greene; D. hendersonii subsp. parvifolium (R. Knuth) H. J. Thompson; Primula hendersonii (A. Gray) A. R. Mast & Reveal

Plants (0.7–)1–5(–5.5) dm, glandular at least distally or glabrous. Caudex not obvious at anthesis; roots usually whitish; bulblets present or absent. Leaves 0.5–14(–16) ´ (1–)1.5–6(–7) cm, somewhat decurrent nearly to base; blade abruptly tapering to a generally slender (at least basally) petiole, oblanceolate to elliptic or spatulate, sometimes ovate to nearly rounded, glabrous, margin entire. Inflorescences 2–17-flowered; bracts narrowly to broadly lanceolate, 3–10(–15) mm; pedicels 2–7 cm at anthesis, glandular or glabrous. Flowers 4–5-merous (often on same plant); calyx 5–10 mm, green or greenish with reddish or purple speckles, glandular-puberulent or glabrous, tube 1.5–3 mm, lobes 3–8 mm; corolla lobes 0.6–2.5(–2.8) cm, magenta to lavender, occasionally white, tube yellow or whitish with a thick, wavy, reddish to reddish-purple ring; filaments fused, tube 1–3.5 ´ 1–4 mm, dark maroon; anthers 2.5–6 mm, pollen sacs deep red to purple or maroon, sometimes yellow and often speckled with red or maroon, connective generally rugose, dark maroon to black; stigma not enlarged. Capsules cylindric-ovoid, 7–17(–19) ´ 4–7(–9) mm, green or greenish, sometimes speckled purple or reddish, glabrous or glandular-puberulent, operculate or valvate; walls thin. 2n = 44, 66, 88, 132.

      Flowering winter–early summer. Grassland communities and oak and conifer woodlands, in sunny or more often shady places; 0–1900(–2100) m; B.C.; Calif., Oreg., Wash.

     Mosquito-bill occurs from southern Vancouver Island southward in the coastal ranges to west-central California (as far south as San Benito County), and then disjunct onto the San Bernardino Mountains in southern California. To the east, the species is found on the Siskiyou Mountains and in the Sierra Nevada of California as far south as Tulare County.

     The distinction between var. hendersonii and var. hansenii is not attempted here although it is probably justified. The former may be broadly characterized as plants bearing bulblets at anthesis with sparsely glandular scapes, pedicels and sometimes calices. The calyx of var. hendersonii is usually greenish with purple or reddish speckles. The most distinctive character of this phase is a filament-tube that is 1–2.5 mm wide with apically acute anthers. This expression is found mainly along the coast from British Columbia to southern Oregon, and then in scattered locations in coastal California, with disjunct populations in the foothills of the central Sierra Nevada, and in the mountains of southern California. The var. hansenii is glabrous, lacks bulblets, and the calyx typically is green; the taxon generally is found inland in the Siskiyou Mountains and the Sierra Nevada, but scattered populations occurs in the coastal ranges of northern California. The filaments on var. hansenii are broader, being 1.5–4 mm wide, with apically obtuse anthers. Capsules of var. hendersonii are usually operculate while those of var. hansenii appear to be consistently valvate. Known chromosome numbers for var. hendersonii are 2n = 44, 66, 132 while that of var. hansenii is only 2n = 88. The 2n = 66 phase appears to be primarily individuals that produce little or no pollen. Inasmuch as bulblets and mature capsules are rarely collected it is difficult to clearly distinguish between the two. H. J. Thompson (1953) recognized Dodecatheon hansenii whereas A. F. Cholewa and D. M. Henderson (1993) accepted only D. hendersonii. In their molecular study, A. R. Mast et al. (2004) considered the two entities distinct, albeit closely related species. Clearly more study is needed especially in areas where the two seemingly have overlapping ranges in the Bay Area and Sierra Nevada foothills of California.

     A Macoun (s. n.., DAO) specimen supposedly gathered at Yale, British Columbia, is considered here to be a doubtful location (Beamish 1951) as all other known localities are from Vancouver Island.

Dodecatheon hendersonii

Detail of habit

Inflorescence

Anthers and connectives

Leaves

3. Dodecatheon subalpinum Eastwood, Leafl. W. Bot. 2: 37. 1937 ·   Sierran shootingstar E

Primula subalpina (Eastwood) A. R. Mast & Reveal

Plants 0.7–1.5(–2.5) dm, glabrous. Caudex not obvious at anthesis; roots reddish, bulblets generally present. Leaves (2.5–)3–8(–10) ´ 0.5–1.5(–1.8) cm, generally slightly decurrent to base; blade gradually tapering to generally a non-winged petiole, oblanceolate to narrowly spatulate, glabrous, margins entire, sometimes slightly undulate. Inflorescences 1–5(–8)-flowered; bracts linear to narrowly lanceolate, 2–6 mm; pedicels 1–2(–3.5) cm at anthesis, glabrous. Flowers 5-merous; calyx 3.5–6 mm, green, glabrous, tube 2–3 mm, lobes 2.5–4.5 mm; corolla lobes 0.5–0.9(–1.2) cm, magenta, occasionally white, tube yellow or infrequently white with a thick, wavy, dark maroon ring; filaments fused, tube 2–3.5 ´ 1–1.5 mm, dark maroon; anthers 3–4 mm, pollen sacs yellow, streaked with purple, connective rugose, dark maroon; stigmas not enlarged. Capsules cylindric-ovoid, 6–10(–13) ´ 3–4.5 mm, glabrous, valvate; walls thin. 2n = 66.

     Flowering summer. Moist slopes mainly of shady places in conifer woodlands or in meadows and along stream banks; 2100–4000 m; Calif.

     Sierran shootingstar is confined to the high western slope of central and southern Sierra Nevada from Tuolumne County to Tulare County. This high elevation ecotype might better be considered a variety of Dodecatheon hendersonii, for which the name var. yosemitanum H. L. Mason is available.

4. Dodecatheon clevelandii Greene, Pittonia 1: 213. 1888 (as clevelandi)

Primula clevelandii (Greene) A. R. Mast & Reveal

Plants (0.7–)1–4 dm. glabrous, generally glandular-puberulent apically. Caudex not obvious at anthesis; roots tannish; bulblets absent. Leaves (1–)3–18(–20) ´ 0.5–4(–5.5) cm, generally not decurrent or only so to near base; blade narrowing abruptly to a generally slender petiole, oblanceolate to spatulate, glabrous, occasionally with a few sessile glands, margins entire or rarely finely denticulate. Inflorescences (1–)5–16-flowered; bracts narrowly lanceolate to lanceolate or rarely oblanceolate, 3–22 mm; pedicels 2–5 cm at anthesis, sparsely to moderately glandular-puberulent. Flowers 5-merous; calyx 5.5–8.5, light greenish, glandular-pubescent abaxially, tube 1.5–2.5 mm, lobes 3–6 mm; corolla lobes 0.6–2.5(–3) cm, magenta or white, tube yellow with a thick, often wavy, dark maroon ring; filaments fused, tube 2.5–4 ´ 3–4 mm, yellow or dark maroon to black; anthers 3–5 mm, pollen sacs yellow, connective rugose, yellow or maroon to black; stigmas not enlarged. Capsules cylindric-ovoid, 8–16 ´ 4–7 mm, yellowish or reddish tan, often suffused with purple, glabrous or glandular-puberulent, valvate or operculate; walls thin. 2n = 44, 66, 88.

     Varieties 4 (4 in the flora): w North America.

1. Connective yellow; filament-tube without a yellow or white spot below each anther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. Dodecatheon clevelandii var. clevelandii

1. Connective maroon to black; filament-tube with or without a yellow or white spot below each anther.

     2. Filament-tube without a yellow spot below each anther . . . . . . . . 4b. Dodecatheon clevelandii var. insulare

     2. Filament-tube with a yellow spot below each anther.

          3. Pollen sac usually yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4c. Dodecatheon clevelandii var. gracile

          3. Pollen sac usually dark purple . . . . . . . . . . . . . . . . . . . . . . . . 4d. Dodecatheon clevelandii var. patulum

4a. Dodecatheon clevelandii Greene var. clevelandii · Padre’s shootingstar

Inflorescences (1–)5–16-flowered. Flowers: filament-tube without a yellow or white spot below each anther; pollen sacs yellow; connective yellow. 2n = 44.

     Flowering winter-spring. Dry or sometimes moist slopes and flats in coastal scrub communities, and in oak and/or conifer woodlands; 0–700 m.; Calif.; Mexico (n. Baja Calif.).

     Padre’s shootingstar is confined in our area mainly to the low mountains of the Transverse and Peninsular ranges of western Los Angeles, Orange, western Riverside and San Diego counties in southwestern California. In Baja California the variety is in the mountains as far south as the Sierra San Pedro Mártir, and in widely scattered locations along the coast to near El Rosario.

4b. Dodecatheon clevelandii Greene var. insulare (H. J. Thompson) Reveal, Sida 22: 863. 2006. · Island shootingstar

Dodecatheon clevelandii subsp. insulare H. J. Thompson, Contr. Dudley Herb. 4: 134. 1953 (as insularis); Primula clevelandii var. insularis (H. J. Thompson) A. R. Mast & Reveal

Inflorescences 5–9-flowered. Flowers: filament-tube without a yellow or white spot below each anther; pollen sacs yellow with maroon speckles; connective maroon to black. 2n = 44.

     Flowering late winter–spring. Dry slopes and flats in coastal scrub communities and mixed oak and conifer woodlands; 0–800 m.; Calif.; Mexico (Baja Calif.).

     Island shootingstar is confined mainly to the low mountains of Monterrey County south to Santa Barbara County, California, and on the Channel Islands. In Mexico it is disjunct to the immediate coast and off-shore islands near Ensenada, and on Guadalupe Island off the coast of west-central Baja California. The leaves on this phase may be up to 5.5 cm wide.

4c. Dodecatheon clevelandii Greene var. gracile (Greene) Reveal, Sida 22: 863. 2006. · Mission shootingstar E

Dodecatheon patulum Greene var. gracile Greene, Erythea 3: 72. 1895; D. clevelandii subsp. sanctarum (Greene) Abrams; Primula clevelandii var. gracilis (Greene) A. R. Mast & Reveal

Inflorescences 3–7(–12)-flowered. Flowers: filament-tube with a yellow or white spot below each anther; pollen sacs usually yellow with reddish or purplish to maroon speckles; connective maroon to black. 2n = 44, 66, 88.

     Flowering spring. Dry slopes and flats mainly in oak woodlands; 0–700 m.; Calif.

     Mission shootingstar is found mainly in the coastal and western Transverse Ranges from San Francisco County south to Ventura County.

4d. Dodecatheon clevelandii Greene var. patulum (Kuntze) Reveal, Sida 22: 863. 2006. · Valley shootingstar E

Meadia patula Kuntze, Revis. Gen. Pl.: 398. 1891; Dodecatheon patulum (Kuntze) Greene; D. clevelandii subsp. patulum (Kuntze) H. J. Thompson; Primula clevelandii var. patula (Kuntze) A. R. Mast & Reveal

Inflorescences 1–10(–18)-flowered. Flowers: filament-tube with a yellow or white spot below each anther; pollen sacs usually dark purple; connective maroon to black. 2n = 44, 88.

     Flowering spring. Moist flats and slopes usually on serpentine or alkaline soils in grassland communities, and in oak and conifer woodlands; 0–700 m.; Calif.

     Valley shootingstar is found mainly in central and northern California in the Central Valley, on the inner coastal ranges, and on the western foothills of the Sierra Nevada. The variety is disjunct to the Santa Monica Mountains, Los Angeles County. This and the hansenii phase of Dodecatheon hendersonii can be difficult to differentiate in the northern Sierra Nevada. By and large, the anthers of var. patulum are much shorter than those of D. hendersonii. Furthermore, the connective of var. patulum tends to be broadly triangular in shape and heavily rugose whereas that of D. hendersonii is narrower, longer and decidedly less rugose.

5. Dodecatheon redolens (H. M. Hall) H. J. Thompson, Contr. Dudley Herb. 4: 143. 1953 · Scented shootingstar E

Dodecatheon jeffreyi Van Houtte var. redolens H. M. Hall, Bot. Gaz. 31: 392. 1901; Primula fragrans A. R. Mast & Reveal

Plants 2.5–8 dm, generally glandular-pubescent throughout, often sticky. Caudex not obvious at anthesis or more commonly short, thick, generally horizontal; roots generally tan to dark reddish brown; bulblets absent. Leaves 20–50(–60) ´ 2.5–6 cm, decurrent to base; blade gradually tapering to a generally winged petiole, oblanceolate, minutely glandular-pubescent, margins entire. Inflorescences 5–15-flowered; bracts lanceolate to broadly lanceolate, 5–17 mm; pedicels 2–9 cm at anthesis, glandular-pubescent. Flowers 5-merous; calyx 10–18 mm, light green, glandular-pubescent, tube 3–10 mm, lobes 5-12 mm; corolla lobes 1.4–3(–3.5) cm, magenta to lavender, tube yellow with a thick, often wavy, maroon ring, ring rarely absent; filaments free, 0.2-0.8 mm, dark maroon to black, usually concealed by corolla tube; anthers 7–11 mm, acute apically, pollen sacs maroon, connective rugose, dark maroon to dark purple; stigma enlarged, generally more than twice diameter of style. Capsules ovoid, 8–17 ´ 5–9 mm, light brown, often reddish-brown apically, glandular-puberulent or glabrous except for glandular-puberulent teeth, valvate; walls thin.

     Flowering late spring-summer. Moist meadows and stream banks mainly in montane conifer woodlands; 2300–3600 m; Calif., Nev., Utah.

     Scented shootingstar occurs mainly in the high mountains of the southern Sierra Nevada, with scattered populations in the San Jacinto and San Bernardino mountains of southern California. Elsewhere the species occurs on scattered desert ranges in the northern Mojave Desert. In the Intermountain West it is found occasionally in Inyo and Mono counties on the eastern foothills of the Sierra Nevada, the White Mountains, and on and near Glass Mountain. The plant is found across central Nevada to the Independence and Ruby mountains of Elko County, and the Deep Creek Range of west-central Utah. A good feature, although hard to see, is that the corolla tube usually covers the filament-tube and lower end of the anthers. In both Dodecatheon jeffreyi and D. alpinum, the corolla tube does not cover the base of the anthers. In addition, the tips of the anthers in the latter two species are truncate to obtuse, whereas in D. redolens the anther tips are acute.

6. Dodecatheon jeffreyi Van Houtte, Fl. Serres Jard. Eur. 16: 99, t. 1662. 1867 · Tall mountain shootingstar E F

Dodecatheon jeffreyi subsp. pygmaeum (H. M. Hall) H. J. Thompson; Primula jeffreyi (Van Houtte) A. R. Mast & Reveal

Plants 1–6(–7.5) dm, glandular-pubescent at least in part, not sticky. Caudex not obvious at anthesis or more commonly short, thick, generally horizontal, occasionally stout, elongate and horizontal; roots generally white; bulblets absent. Leaves (2.5–) 7–40(–53) ´ (0.5–) 1–6(–7.5) cm, decurrent to base; blade usually gradually tapering to a winged petiole, narrowly oblanceolate or more commonly broader to spatulate, glabrous or glandular-pubescent, margins entire or crenate to serrulate. Inflorescences 3–20-flowered; bracts lanceolate to broadly lanceolate, 3–17 mm; pedicels 2–7 cm at anthesis, glandular-pubescent, rarely glabrous. Flowers 4–5-merous; calyx 7–12(–15) mm, glandular-pubescent, rarely glabrous, tube 2–4 mm, lobes 4.5–8(–12) mm; corolla lobes 1–2.5(–2.7) cm, magenta to lavender or light yellow to whitish, tube cream or (rarely) yellow with a thin to thick, often wavy, reddish to purplish ring, ring rarely absent; filaments free or partially fused, usually 1–1.5 mm, dark maroon to black; anthers 6.5–11 mm, truncate to obtuse apically, pollen sacs yellow or maroon, connective rugose, purplish; stigma enlarged by no more than twice diameter of style. Capsules ovoid, 7–11(–15) ´ 4.5–7(–10) mm, yellowish-tan to reddish brown, glabrous or teeth occasionally sparsely glandular-puberulent, operculate or valvate, occasionally both on same plant; walls thin. 2n = 42, 44, 66, 86.

     Flowering summer. Dry to moist stream banks, lakeshores, bogs, and meadows mainly in montane conifer woodlands; 0–3000 m; B.C.; Alaska; Calif., Idaho, Mont., Oreg., Wash., Wyo.

     Tall mountain shootingstar is found in widely scattered montane places in the Sierra Nevada of California and extreme western Nevada and on the northern coastal ranges and Siskiyou Mountains of northern California and southwestern Oregon. The species occurs northward in the Cascade Ranges of Oregon, Washington and British Columbia to the Kenai Peninsula region of south central Alaska, often near the coast especially on many of the off-shore islands. Inland in the United States the plant is widely scattered in the mountains of northeastern Oregon, central and northern Idaho, and western Montana. Isolated stations occur on the Olympic Peninsula of Washington. A single collection (J. Major 2927, GTNP) from Moose Basin, Grand Teton National Park, is the only record from Wyoming.

     Dodecatheon jeffreyi is usually readily recognized, but in portions of California, the delimitation of it from both D. alpinum and D. redolens can be somewhat arbitrary. Whether this is a breakdown of species boundaries due to hybridization or a shift in their respective morphologies due to overlapping ecological settings is uncertain. At least in a few instances, intermediate plants seem to occur in areas where two of the species occur in close proximity. In general, the corolla tube of D. jeffreyi is white except near the ring where it is yellow. In D. redolens, the entire corolla tube is yellow.

     This species is known universally as Dodecatheon jeffreyi, although the plant was named a year earlier as D. jeffreyanum K. Koch. To avoid the introduction of a name that has never been used for this widespread and frequently cultivated plant, D. jeffreyanum has been proposed for rejection (J. F. Veldkamp, J. L. Reveal & K. Gandhi 2007).

Dodecatheon jeffreyi

Revision of Dodecatheon (Primulaceae)

James L. Reveal

Adjunct Professor, Cornell University, Ithaca, New York 14853-4301, U.S.A. Professor Emeritus, University of Maryland College Park, Maryland 20742-5815, U.S.A. Honorary Curator, The New York Botanical Garden, Bronx, New York, 10458-5026, U.S.A.

Adjunct Professor, Cornell University,
Ithaca, New York 14853-4301, U.S.A.
Professor Emeritus, University of Maryland
College Park, Maryland 20742-5815, U.S.A.
Honorary Curator, The New York Botanical Garden,
Bronx, New York, 10458-5026, U.S.A.