נובמבר 12, 2023

Campanula, partial treatment for Flora of North America.

  1. CAMPANULA Linnaeus, Sp. Pl. 1: 163. 1753; Gen. Pl. ed. 5, 77. 1754 * [Latin campanula, bell, alluding to bell-shaped corolla of many species]
    Nancy R. Morin
    Herbs, perennial or biennial [annual], 2--130 cm, glabrous or hairy; rhizomatous or with woody caudex. Stems erect, ascending, arching, decumbent, or reclining, branched or unbranched. Leaves alternate, in basal rosette or not, and cauline, petiolate or sessile; proximal leaf blade elliptic, ovate, ovate-oblong, oblong, ovate-cordate, ovate-orbicular, suborbicular, lanceolate, oblanceolate, spatulate, linear, or nearly filiform, base attenuate, acute, cordate, cuneate, truncate, obtuse, to rounded, rarely amplexicaulous, margins entire, dentate, crenate, or serrate, teeth generally glandular. Inflorescences capitate, spikelike, racemelike, paniclelike, cymose, or flowers solitary. Pedicels 0--8 cm. Flowers chasmogamous, erect, ascending, horizontal, nodding, or pendent; hypanthium obconic-cylindric, obconic, turbinate, cup-shaped, hemispheric, to suborbicular, ribbed or not, ribs equal, raised or not, base tapered, acute, rounded, or flat; appendages between sepals present or absent; sepals deltate, deltate-lanceolate, lanceolate-elliptic, lanceolate-attenuate, oblong, oblong-attenuate, linear-deltate, linear-lanceolate, linear, acuminate, setaceous, or subulate, equal in size; corolla pale blue to blue, blue-violet, violet, mauve, rarely pink or white, 6--70 mm, tube funnelform, saucer-shaped, cup-shaped, campanulate, subglobose, tubular, or cylindric, lobes nearly patent to spreading, recurved or tightly rolled back, to erect, obovate to elliptic, oblong-ovate, oblong-deltate, ovate-deltate, ovate-acute, lanceolate, or deltate, 10--100% of corolla length; filaments usually dilated basally, ciliate; ovary 3-locular (5-locular in C. medium); styles straight (curved in C. aurita), papillate in distal 15--95%; stigma lobes 3 (5 in C. medium, sometimes 2 in C. scouleri). Capsules pendent, nodding, erect, or horizontal, subglobose, hemispheric, ovoid, to broadly elliptic, clavate to urceolate, obconic or turbinate, or oblong-obovate, cylindric, dehiscing by basal, medial, or apical pores on sides of hypanthium. Seeds oblong to fusiform, lenticular in cross-section, smooth, reticulate, or striate. x = 17.
    Species 400--500 (22 in the flora): North America, Mexico, Eurasia, n Africa; introduced in South America, Pacific Islands, Australia.
    Based on morphological and phylogenetic evidence, species native in the flora area formerly in Campanula have been placed in other genera: Campanula robinsiae in Protocodon, C. aparinoides in Palustricodon, C. californica in Eastwoodiella, the annual California campanulas in Ravenella, C. prenanthoides and C. wilkinsiana in Smithiastrum, C. reverchonii in Poolea, and C. floridana in Rotanthella (N. R. Morin 2020). See discussion under Campanuloideae for overview of these changes. The ten native species (seven of them endemic) still in Campanula require further study to determine their status. The native species belong to three clades: C. chamissonis, ranging from Alaska to Japan, in the Campanula clade; seven species endemic to montane areas in the flora area, plus one, C. lasiocarpa, which extends into Russia and Japan, in the Cordilleran clade; and the circumpolar Campanula rotundifolia, in a more distant clade (K.-O. Yoo et al. 2018). The arctic-alpine western species in North America underwent repeated periods of isolation as they retreated to refugia, alternating with population expansion, leading to narrow endemics (for example, E. G. DeChaine et al. 2014).
    Some populations of Campanula rotundifolia, C. parryi, and C. divaricata have a distinctive growth habit of long, drooping stems and extremely long, narrow leaves, usually when growing in very wet habitats. If this pattern is correlated with any other morphological features and the pattern is maintained in drier conditions, these populations may warrant taxonomic recognition.
    The high-elevation native species of Campanula are sometimes grown in rock gardens. European and Asian species of Campanula are widely cultivated, some serving as border plants and others in rockeries and alpine houses. Most of the non-native species in North America are garden escapes that may persist in isolated populations. Campanula rapunculoides is the only species that becomes invasive (T. D. Whitson et al. 1996).
    SELECTED REFERENCES DeChaine, E. G., B. M. Wendling, and B. R. Forester. 2014. Integrating environmental, molecular, and morphological data to unravel an ice-age radiation of arctic-alpine Campanula in western North America. Ecol. Evol. 20: 3940--3050. Roquet, C., I. Sanmartin, N. Garcia-Jacas, L. Sáez, A. Susana, N. Wikström, and J. J. Aldasoro. 2009. Reconstructing the history of Campanulaceae with a Bayesian approach to molecular dating and dispersal-vicariance analyses. Molec. Phylogen. Evol. 52: 575--587. Shetler, S. G. 1963. A checklist and key to the species of Campanula native or commonly naturalized in North America. Rhodora 65: 319—337. Wendling, B. M., K. E. Galbreath, and E. G. DeChaine. 2011. Resolving the evolutionary history of Campanula (Campanulaceae) in Western North America. PLoS ONE 6(9): e23559. . Whitson, T. D., L. C. Burrill, S. A. Dewey, D. W. Cudney, B. E. Nelson, R. D. Lee, and R. Parker. 1996. Weeds of the West. Western Society of Weed Science in cooperation with Cooperative Extension Services, University of Wyoming, Laramie, Wyoming. 630 pp. Xu, C. and Hong D. Y. 2021. Phylogenetic analyses confirm polyphyly of the genus Campanula (Campanulaceae s. str.), leading to a proposal for generic reappraisal. J. Syst. Evol. 59: 475-–489. Yoo, K.-O., A. A. Crowl, K.-A. Kim, K-S. Cheon, and N. Cellinese. 2018. Origins of East Asian Campanuloideae (Campanulaceae) diversity. Molec. Phylogen. Evol. 127: 468--474.

  2. Flowers with appendages between sepals.
  3. Plants 5--10 cm; appendages 1--2 mm; corollas 30--40 mm; Alaska

  4. Campanula chamissonis



  5. Plants 20--100 cm; appendages 3--12 mm; corollas 40--70 mm; s Canada southwards.

  6. Corollas blue, without dots; stigma lobes 5 11. Campanula medium

  7. Corollas cream-colored or pink, purple-dotted; stigma lobes 3 14. Campanula punctata

  8. Flowers lacking appendages between sepals.

  9. Inflorescences densely capitate, sometimes flowers also in clusters in axils of leaflike bracts.

  10. Plants biennial or short-lived perennial; basal leaf blade bases attenuate; hypanthium bases rounded, sepals ovate-deltate; corollas 10--15 mm 17. Campanula cervicaria

  11. Plants perennial; basal leaf blade bases cordate or rounded; hypanthium bases tapered, sepals linear to lanceolate; corollas 15--25(--40) mm
    1. Campanula glomerata


  12. Inflorescences paniclelike or racemelike, sometimes spikelike or cymose, or flowers solitary.

  13. Basal leaf blade bases cordate.

  14. Corollas 15--20 mm; capsules erect to horizontal.

  15. Stems decumbent to ascending; cauline leaf blades round to broadly lanceolate; hypanthia hemispheric; corolla lobes 60--65% of corolla length
    1. Campanula poscharskyana


  16. Stems erect; cauline leaf blades ovate-lanceolate; hypanthia obconic-cylindric; corolla lobes 20--40% of corolla length 21. Campanula carpatica

  17. Corollas 20--60 mm; capsules declined.

  18. Flowers erect or ascending; sepal margins serrate, sometimes infolded; basal leaf blade bases rounded to cuneate
    1. Campanula latifolia


  19. Flowers pendent or nodding; sepal margins entire; basal leaf blade bases cordate.

  20. Cauline leaf blade bases amplexicaulous; corollas 20--25 mm
    1. Campanula bononiensis


  21. Cauline leaf blade bases cordate, rounded, or truncate; corollas 30--50 mm.

  22. Inflorescences paniclelike, bracts absent or narrowly elliptic
    1. Campanula trachelium


  23. Inflorescences racemelike, spikelike, bracts ovate to oblong-ovate, or lanceolate
    1. Campanula rapunculoides


  24. Basal leaf blade bases attenuate, cuneate, rounded, acute, or obtuse (basal leaves absent in C. divaricata).

  25. Stigma lobes 40--60% of style length.

  26. Plants biennial or short-lived perennial; corollas 15--25 mm; anthers 3--6 mm 20. Campanula patula

  27. Plants perennial; corollas 30--50 mm; anthers 7--11 mm
    1. Campanula persicifolia


  28. Stigma lobes 4--30% of style length.

  29. Styles longer than corollas.

  30. Corolla lobes 25--45% of corolla length, tubes cylindric to subglobose to broadly campanulate; stigma lobes 4--6% of style length
    1. Campanula divaricata


  31. Corolla lobes 50--60% of corolla length, tubes broadly funnelform; stigma lobes 10--13% of style length 6. Campanula scouleri

  32. Styles equal to or shorter than corollas (appearing exserted in C. aurita due to rotate corolla).
    [16. Shifted to left margin.---Ed.]

  33. Plants 4--90 cm; hypanthia glabrous (rarely puberulent or hirtellous).

  34. Corolla tubes campanulate, funnelform, or tubular; capsules pendent, pores basal
    1. Campanula rotundifolia


  35. Corolla tubes broadly funnelform or shallowly cup-shaped to saucer-shaped; capsules erect, pores apical.

  36. Corolla tubes broadly funnelform, lobes 33--66% of corolla length; styles straight; stigma lobes 20--30% of style length 1. Campanula parryi

  37. Corolla tubes shallowly cup-shaped to saucer-shaped, lobes 88--100% of corolla length; styles abruptly curved upward distally; stigma lobes 10% of style length 8. Campanula aurita

  38. Plants 3--12 cm; hypanthia woolly-villous, scabrous-puberulent, scabrellous, or scabrous-hispidulous.

  39. Leaf blade margins entire; hypanthia scabrous-puberulent; sepal margins entire
    1. Campanula scabrella


  40. Leaf blade margins sharply and coarsely serrate, dentate or serrate-dentate, or with widely spaced, abrupt callous teeth; hypanthia densely woolly-villous, hirtellous, scabrellous, hispidulous, scabrid-hispidulous, or glabrous; sepal margins entire or with 1--several prominent teeth or dentate.

  41. Plants glabrous except long hairs scattered on stem angles and leaf margins; hypanthia moderately to densely woolly-villous; corollas 15--30 mm; capsule pores apical
    1. Campanula lasiocarpa


  42. Plants glabrous, sparsely hirtellous or scabrid-hispidulous; hypanthia densely scabrellous or scabrous-hispidulous; corollas 7--16 mm; capsule pores +/- at middle.

  43. Capsules subglobose to broadly elliptic to cylindric, pores just proximal to middle; anthers 3.5--5 mm 4. Campanula piperi

  44. Capsules broadly to narrowly cylindric or urceolate, pores near or just above middle, ultimately entire panel opens; anthers 2.5--3 mm 5. Campanula shetleri


  45. Campanula parryi A. Gray in A. Gray et al., Syn. Fl. N. Amer. ed. 2, 2(1): 395. 1886 * Parry’s bellflower E
    Herbs perennial, 4--40 cm, glabrous throughout or sometimes sparsely pubescent proximally; cespitose from slender, branching rhizome. Stems erect, unbranched or few-branched. Leaves: basal not in rosette, petiolate, blade oblong-elliptic, oblanceolate, to spatulate or linear, 10--80 mm, base acute or attenuate, margins eciliate or coarsely ciliate proximally, entire, callous-toothed or minutely denticulate, apex rounded or abruptly acute to attenuate; cauline sessile, blade linear-attenuate or filiform to lanceolate, elliptic, obovate, or oblanceolate, 10--80 mm, base attenuate, margins entire or remotely toothed, apex acute or acuminate, surfaces glabrous, becoming linear and bractlike distally. Inflorescences: flowers usually solitary, sometimes 1--3-flowered on axillary branches; bracts absent or linear. Pedicels 0.2--8 cm. Flowers somewhat nodding in bud or not, becoming erect; hypanthium obconic to cup-shaped, base tapered, surface glabrous or with a few hairs, or papillate; appendages absent; sepals spreading, broadly to narrowly lanceolate or filiform-linear, 2--18 mm, margins with a few callous teeth or entire; corolla pale to deep blue or mauve, 9--23 mm, tube broadly funnelform, lobes spreading, broadly elliptic to obovate, 33--66% of corolla length; anthers 4--6 mm, pollen white; styles , white or pale to deep blue,7--22 mm, papillate in distal 50--60%; stigma lobes white, 20--30% of style length. Capsules erect, clavate to obconic or oblong-obovate, base narrowed, pores apical. 2n = 34.
    Varieties 3 (3 in the flora): w United States, Canada{{Reviewers please confirm}}.
    Campanula parryi and C. rotundifolia are often mistaken for each other. Campanula rotundifolia has flowers arranged in racemes, seldom solitary, and nodding, turbinate capsules opening by basal pores, whereas C. parryi has flowers generally solitary, and erect, obconic to clavate capsules opening by apical pores.
    The relationships among Campanula rotundifolia, C. scabrella, and C. parryi are complex. McVaugh’s description of C. parryi var. idahoensis expanded the concept of C. parryi to include plants with hypanthia that are cup-shaped or campanulate, rounded at base, and sparsely or densely hairy, the hairs short, all of which are also found in C. scabrella. S. G. Shetler (1963) suggested at least some of the specimens identified as var. idahoensis represented a hybrid series between C. parryi and C. scabrella or that this variety was being used for several discordant elements, potentially including undescribed species. At least some of the plants identified as var. idahoensis from Washington have campanulate corollas with shallow lobes typical of C. rotundifolia. B. M. Wendling et al. (2011) concluded that var. idahoensis and var. parryi were not sister taxa, based on cpDNA analysis of two populations of var. idahoensis, one in Montana and one in Washington. E. G. DeChaine et al. (2014) provided morphological measurements for one of the Washington populations indicating the corolla tube was tubular-campanulate and the lobes were about 33% of total corolla length, character states typical of C. rotundifolia. DeChaine et al. reported that plants identified as var. idahoensis from a Montana population had lobes generally 40% of corolla length, typical of C. parryi. DeChaine et al. concluded that var. idahoensis was most closely related to C. scabrella and that the two taxa split about 240,000--500,000 years ago. Variety idahoensis is recognized here with the caveat that this group needs further study.
    The Navajo people have used the dried plant of Campanula parryi to dust sores (P. A. Vestal 1952), and the Zuni made chewed root poultices for bruises (M. C. Stevenson 1908).
    SELECTED REFERENCES Stevenson, M. C. 1908. Ethnobotany of the Zuni Indians. Rep. (Annual) Bur. Amer. Ethnol. 30: 44. Vestal, P. A. 1952. The Ethnobotany of the Ramah Navajo. Papers of the Peabody Museum of American Archaeology and Ethnology 40: 1--94.


  46. Cauline leaves linear-attenuate to filiform, apices abruptly acute to attenuate; sepals filiform-linear 1c. Campanula parryi var. arizonica


  47. Cauline leaves elliptic, oblanceolate, or obovate, apices rounded; sepals broadly to narrowly lanceolate.


  48. Sepal and some leaf margins with evident teeth; main pedicels 3--5 cm; corollas (9--)15--23 mm 1a. Campanula parryi var. parryi


  49. Sepal and leaf margins entire or minutely callous-denticulate; main pedicels0.2--1.5(--5) cm; corollas 9--15 mm
    1b. Campanula parryi var. idahoensis


  50. 1a. Campanula parryi A. Gray var. parryi E
    Cauline leaf blades elliptic, oblanceolate, or obovate, 10--50 mmm, margins callous-toothed, teeth evident, small, abrupt, coarsely ciliate proximally, hairs stiff, white, apex rounded. Inflorescences: main pedicel 3--5 cm. Flowers: mature sepals broadly to narrowly lanceolate, (6--)10--15 mm in fruit, margins with evident teeth, with stiff, white cilia; corolla (9--)15--23 mm, lobes 50--66% of corolla length; styles 10--22 mm. Capsules 7--11 mm, ribs thick.
    Flowering Jun--Aug. Mountain and subalpine meadows, stream banks, in wet soils; 1800--4100 m; Ariz., Colo., Idaho, Mont., N.Mex., Utah, Wash., Wyo.

    1b. Campanula parryi A. Gray var. idahoensis McVaugh, Bull. Torrey Bot. Club 69: 241. 1942 E
    Proximal cauline leaf blades elliptic, oblanceolate, or obovate, 15--45 mm, margins entire or minutely callous-denticulate, teeth not visible without magnification, coarsely ciliate proximally, hairs stiff, white, apex rounded. Inflorescences: primary pedicel 0.2--1.5 (--5) cm. Flowers: mature sepals narrowly lanceolate, 2--6 (--9) mm, margins entire, ciliate or eciliate; corolla 9--15 mm, lobes 33--50% of corolla length; styles 7--13 mm. Capsules 5--11 mm, ribs thin.
    Flowering Jul--Aug. Open areas, sandy or gravelly soil, scree, streamsides, meadows, in Douglas fir, Ponderosa pine, or juniper woodlands; 1100--3000 m; Alta.{{Reviewers please confirm}}; Idaho, Mont., Wash.

    1c. Campanula parryi A. Gray var. arizonica Morin ined. E
    Proximal cauline leaf blades linear-attenuate to filiform, 60--80 mm, margins entire, eciliate, apex abruptly acute to attenuate. Inflorescences: primary pedicel 4--8 cm. Flowers: mature sepals filiform-linear, (8--)13--18 mm, margins entire, eciliate{{?}}; corolla 10--18 mm, lobes 66% of corolla length; styles 10 mm. Capsules 8--11 mm, ribs very thin.
    Flowering Jul--Aug. Hanging gardens, wet rock ledges in steep cliffs; 1400--2100 m; Ariz.
    Variety arizonica occurs in Apache and Coconino counties. In addition to the very narrow leaves and sepals, var. arizonica has elongated capsules. {{Regional reviewers: var. arizonica may get into similar habitat in Glen Canyon, Utah?}} Also occurring in Arizona and having elongated capsules are compact plants with obovate to lanceolate-elliptic proximal leaves that have margins with small abrupt callous teeth and shorter corolla lobes that are 25% of the corolla length. More study is needed to determine whether these plants belong in var. arizonica.

    1. Campanula lasiocarpa Chamisso, Linnaea 4: 39. 1829 * Mountain bellflower or harebell, campanule tomenteuse
      Herbs perennial, 3--20 cm, glabrous except long hairs scattered on stem angles and leaf margins ; cespitose from branching rhizomes. Stems erect or ascending, unbranched. Leaves: basal in prominent primary and axillary rosettes, petiolate or sessile, blade elliptic to broadly oblanceolate or subrhombic, 10--30 mm, base attenuate, margins coarsely dentate, teeth sharp, glandular, straight, apex acute; cauline sessile, blade elliptic to oblanceolate, 10--20 mm, , base attenuate, margins with widely spaced abrupt callous teeth, sometimes laciniate, apex acute, surfaces glabrous. Inflorescences: flowers solitary, rarely cymose with 2 flowers; bracts linear. Pedicels 1--4 cm. Flowers nodding, becoming erect; hypanthium obconic, base rounded, surface moderately to densely woolly-villous; appendages absent; sepals erect to spreading, , lanceolate-attenuate, 5--18 mm, margins dentate, teeth distant, slender, large, ciliate; corolla deep blue, sometimes nearly black immediately adjacent to filament bases, 15--30 mm, tube broadly funnelform to oblong-campanulate, lobes erect to spreading, ovate-acute, 33% of corolla length; anthers 3--5 mm, pollen blue; styles blue, 10--14 mm, papillate in distal 75% ; stigma lobes white, 10--15% of style length. Capsules erect, obconic to cylindric, base rounded, pores apical. 2n = 34.
      Flowering Jun--Sep. Alpine or subalpine slopes and meadows, rocky or gravelly sites or loamy soil, noncalcareous soils; 0--2500 m, Alta., B.C., N.W.T., Yukon; Alaska, Wash.; Asia (Japan, Russia).
      According to L. H. Bailey (1953), the flowers of Campanula lasiocarpa are fragrant, which is unusual for Campanula. Campanula lasiocarpa occurs in the Cascade and Selkirk ranges, north to Alaska, through the Aleutian Islands to Kamchatka and Hokkaido, Japan, and northeastern Russia. Its southernmost distribution is in Snohomish and King counties, Washington. K.-O. Yoo et al. (2018) suggested that C. lasiocarpa migrated from western North America into eastern Asia, the only species of Campanula to have done so.
      SELECTED REFERENCE Yoo, K.-O., A. A. Crowl, K.-A. Kim, K.-S. Cheon, and N. Celinese. 2018. Origins of East Asian Campanuloideae (Campanulaceae) diversity. Molec. Phylogen. Evol. 127: 468--474.

    2. Campanula divaricata Michaux, Fl. Bor.-Amer. 1:109. 1803 * Southern harebell, Appalachian bellflower E F
      Campanula flexuosa Michaux
      Herbs perennial, 20--90 cm, glabrous; with +/- woody caudex and slender rhizomes. Stems erect to arching, many-branched. Leaves: basal absent; cauline sessile or short-petiolate, blade broadly linear to linear-lanceolate to broadly lanceolate, 20--80 mm, base cuneate, margins sharply and coarsely dentate, almost laciniate, to smooth with minute callous teeth, apex attenuate to acute, surfaces glabrous. Inflorescences: paniclelike, often with many branches, ; bracts linear to subulate. Pedicels 0.5--2.5 cm. Flowers nodding ; hypanthium turbinate, base tapered, surface glabrous; appendages absent; sepals erect and appressed to corolla, widely spreading, or recurved, subulate to narrowly deltate, , 1--4 mm, margins entire or with 2--4 small calloused teeth; corolla deep to pale blue, violet, or rarely white, 6--9 mm, tube cylindric to subglobose to broadly campanulate, lobes usually tightly rolled back, sometimes widely spreading, broadly to narrowly deltate, 25--45% of corolla length; anthers 2--2.5 mm, pollen dark or pale mauve, pink, or white; styles deep or sometimes pale blue proximally, pale blue to cream-colored distally, 10--12 mm, papillate in distal 15%, ; stigma lobes cream-colored or pale green, 4--6% of style length. Capsules nodding, turbinate, base truncate, pores basal. 2n = 34.
      Flowering (Mar--)Jun--Oct(--Dec). Rocky woods and cliffs, talus slopes, woodlands, balds; 200--1900 m; Ala., Ga., Ky., N.C., S.C., Tenn., Va., W.Va.
      The relationship of Campanula divaricata to other species in the Rapunculus clade of Campanula is unclear. B. M. Wendling et al. (2011) and C. Xu and Hong D. Y. (2020) placed it with what they called the Cordilleran clade based on cpDNA analysis; Wendling et al. (2011) noted that it fell in the Rapunculus 2B clade, sister to Triodanis leptocarpa, based on ITS analysis.
      Campanula divaricata grows primarily in the central and southern Appalachian Mountains. Its long-exserted style and small, campanulate corolla make it easily recognizable. Populations with extremely long, narrow leaves, some of which have barely discernible callous teeth, seem to occur in the western portion of the range; populations with shorter, broadly lanceolate, sharply and deeply toothed leaves are more common. Corollas range from straight-sided, narrow trumpets with flared lobes to balloonlike bells with tightly curled lobes; corolla color and style color are similarly diverse. A comprehensive study is needed to determine whether these different character states are correlated in any way or have any geographic or habitat patterns.
      In most campanulas, the anthers fall away, or curl back, at anthesis; in Campanula divaricata, the narrow portion of the filament remains erect, holding the anthers up so they are generally visible at the base of the corolla lobe sinuses. The chromosome number of 2n = 34 was reported by Kovanda (1978) and Gadella (1964). M. Kovanda (1978) suggested that a count of 2n = 40 (C. D. Darlington and E. K. Janaki-Ammal 1945; A. E. Radford et al. 1968) was incorrect.
      SELECTED REFERENCES Darlington, C. D. and E. K. Janaki-Ammal. 1945. Chromosome Atlas of Cultivated Plants. London. Gadella, T. W. J. 1964. Cytotaxonomic studies in the genus Campanula. Wentia 11: 1--104. Kovanda, M. 1978. Chromosome numbers of miscellaneous United States Dicotyledons. Rhodora 80: 431--440. Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. Chapel Hill. Wendling, B. M., K. E. Galbreath, and E. G. DeChaine. 2011. Resolving evolutionary history of Campanula (Campanulaceae) in western North America. PLoS ONE 6(9): e23559. Doi: 10.1371/journal.pone.0023559.

    3. Campanula piperi Howell, Fl. N. W. Amer. (Howell) 409. 1901 * Olympic bellflower E
      Astrocodon piperi (Howell) A. P. Khokhrjakov
      Herbs perennial, 5--11 cm, glabrous or sparsely hirtellous; cespitose, with multi-branched caudex and slender rhizomes. Stems erect, unbranched. Leaves: basal in rosette , petiolate, blade elliptic, 10--30 mm, base attenuate, margins coarsely dentate, teeth sharp, glandular, sometimes hooked, apex obtuse; cauline sessile, blade cuneate to spatulate, 6--9 mm, base attenuate, margins coarsely serrate-dentate, , apex broadly acute, surfaces glabrous, . Inflorescences: flowers solitary, sometimes cymose with 2--5 flowers; bracts lanceolate, . Pedicels 0.2--1 cm. Flowers erect; hypanthium , broadly cup-shaped or obconic, base rounded or flat, surface sparsely to densely scabrellous or scabrous-hispidulous; appendages absent; sepals spreading, oblong-attenuate to linear-lanceolate, 5--10 mm, margins with (1--)2--3 pairs of slender teeth; corolla blue to lavender blue, rarely white, 7--16 mm, tube shallowly saucer-shaped, lobes patent to nearly patent, spreading, elliptic, 66% of corolla length; anthers 3.5--5 mm, pollen dark purple or dark red; styles blue, 7--10 mm, papillate in distal 50--60%; stigma lobes white, 10--15% of style length. Capsules erect, subglobose to broadly elliptic to cylindric, base +/- flat, pores just proximal to middle. 2n = 34.
      Flowering Jul--Aug. Rock crevices in openings of subalpine forest; 1500--2000 m; Wash.
      Campanula piperi is endemic to the Olympic Mountains. The plants can be almost completely covered with flowers, and the combination of a lavender blue corolla, white filament bases, deep blue styles, and purple or dark red pollen is striking. Although it is not considered threatened or endangered, there are few populations. S. G. Wershow and E. G. DeChaine (2018) found that C. piperi was more widely distributed than other endemics of the Olympic alpine zone and suggested it would experience less habitat reduction as the climate warms than the other endemics.
      SELECTED REFERENCE Wershow, S. G. and E. G. DeChaine. 2018. Retreat to refugia: Severe habitat contraction projected for endemic alpine plants of the Olympic Peninsula. Amer. J. Bot. 105: 760--778.

    4. Campanula shetleri Heckard, Madroño 20: 231, fig. 1. 1970 * Castle Craggs harebell C E F
      Herbs perennial, 3--5 cm, scabrous-hispidulous throughout, hairs slightly retrorse; mat-forming, cespitose, with slender rhizomes. Stems erect, unbranched. Leaves: basal in rosette, , subsessile, blade short-spatulate (roughly hexagonal), 6--7 mm, base cuneate, margins coarsely dentate, , apex cuneate, acute to obtuse, ; cauline sessile, blade oblanceolate or oblong, 5--7 mm, base attenuate, margins with 2 pairs of sharp teeth, apex acute, surfaces hispidulous. Inflorescences: flowers solitary, or sometimes cymose with 2(--4) flowers, axillary appearing later; bracts oblong to linear. Pedicels 0.1--0.4 cm. Flowers erect; hypanthium cup-shaped, base rounded, , surface densely scabrous-hispidulous; appendages absent; sepals ascending to erect (erect in fruit), subulate, 4--5 mm (in fruit 5--6 mm), margins entire, ; corolla pale blue to white, 7--10.5 mm, tube broadly funnelform, lobes recurved to spreading, ovate-deltate, 50% of corolla length, ; anthers 2.5--3 mm, pollen blue; styles blue, 7–-8 mm, papillate in distal 33%; stigma lobes white, 15% of style length. Capsules erect, broadly to narrowly cylindric or urceolate, base 3-lobed, pores near or just above middle, ultimately entire panels open. 2n = 34.
      Flowering Jun--Aug. Granitic detritus and soil in crevices of steep to vertical north- to northeast-facing granite cliffs in yellow pine forest; of conservation concern; 1200--1900 m; Calif.
      Campanula shetleri is known only from two clusters of populations on Castle Crags, one on the northwest slope in Siskiyou County, the other on the southwest slope in Shasta County. Campanula shetleri is similar to C. piperi in habit, but all parts are smaller, the sepals are subulate rather than deltate, and the leaf blades bear only 2 pairs of teeth rather than 3--6 pairs.

    5. Campanula scouleri Hooker ex A. de Candolle, Monogr. Campan. 312. 1830 * Scouler’s harebell, pale bellflower E
      Herbs perennial, 10--40 cm, glabrous or hairs short; with slender rhizomes. Stems reclining to erect, unbranched. Leaves: basal not in rosette, petiolate, blade elliptic to broadly lanceolate to nearly round, 5--20(--40) mm, base rounded to cuneate or obtuse, margins shallowly toothed or coarsely dentate, , apex acute; cauline petiolate or sessile, blade nearly round to lanceolate-elliptic to elliptic, 10--60(--80) mm, base attenuate, margins shallowly apiculate-glandular-toothed, apex acute, surfaces glabrous or with short hairs. Inflorescences racemelike or flowers solitary; bracts linear. Pedicels 0.5--2 cm. Flowers nodding to horizontal, rarely erect; hypanthium hemispheric to obconic, base flat or rounded, surface glabrous; appendages absent; sepals nearly patent to ascending, linear to narrowly deltate, 4--10 mm, margins entire; corolla white to pale blue or pale pink, 8--18 mm, tube broadly funnelform, lobes reflexed, oblong-ovate, deltate distally, 50--60% of corolla length; anthers 4--6 mm, pollen lavender blue to pink; styles , cream-colored or pale blue or pale pink, 12–-25 mm, papillate in distal 20–-40%; stigma lobes <2 or 3>, cream-colored, 10--13% of style length. Capsules erect, obconic to hemispheric, base +/- flat, pores near middle.
      Flowering Jun--Aug. Openings in mixed conifer forest, rock outcrops; 50--2000 m; B.C.; Alaska, Calif., Oreg., Wash.
      Campanula scouleri occurs mostly west of the Cascade Range; it is also in the northern Sierra Nevada. Sometimes mistaken for Smithiastrum prenanthoides, C. scouleri can be distinguished from that species by its petiolate proximal leaves with broad blades and its oblong-ovate corolla lobes. Campanula scouleri is unusual in having a thick, clublike style that sometimes curves up.

    6. Campanula scabrella Engelmann, Bot. Gaz. 6: 237. 1881 * Rough harebell E
      Herbs perennial, 2--12 cm, densely minutely scabrous-puberulent, appearing velvety throughout; cespitose, with taproot and branched caudex. Stems ascending to erect, unbranched. Leaves: basal in rosette, , petiolate, blade oblanceolate, 5--40 mm, <+/- fleshy>, base attenuate, margins entire, apex apiculate; cauline sessile; blade lanceolate to oblanceolate, 20--30 mm, , base attenuate, margins entire, apex acute to obtuse, surfaces densely minutely scabrous-puberulent. Inflorescences: flowers solitary; bracts narrowly lanceolate or linear. Pedicels 0.3--1.8 cm. Flowers erect; hypanthium obconic to broadly cup-shaped, base rounded, , surface scabrous-puberulent; appendages absent; sepals ascending, narrowly to broadly deltate, 2--6 mm, margins entire; corolla powder blue to lavender blue, 6--12 mm, , tube funnelform, lobes spreading to nearly patent, elliptic, <+/- boat-shaped>, 50--80% of corolla length; anthers 3--5 mm, pollen dark blue; styles blue, 5.5--12 mm, papillate in distal 50%; stigma lobes white, 15--20% of style length. Capsules erect, cylindric-obconic, , base +/- rounded, pores apical.
      Flowering Jun--Aug. Alpine fellfields, stony fields, talus on noncalcareous soil; 1500--3100 m; Calif., Idaho, Mont., Wash.
      Campanula scabrella occurs in the Wenatchee Mountains and on Mount Adam in the Washington Cascade Range, on Mount Shasta and Mount Lassen in northern California, and in the Rocky Mountains of northern Idaho and western Montana. Plants of Campanula parryi var. idahoensis are similar to and sometimes mistaken for C. scabrella. Campanula parryi var. idahoensis plants are taller, with obconic to cup-shaped, papillate hypanthia, narrowly lanceolate sepals, and corolla lobes less than half the length of the corolla. See also the discussion under 1. C. parryi.

    7. Campanula aurita Greene, Pittonia 1: 221. 1888 * Yukon bellflower E
      Astrocodon auritus (Greene) A. P. Khokhrjakov
      Herbs perennial, 4--9(--30) cm, glabrous; with a much-branched rhizome. Stems erect, unbranched. Leaves: basal crowded but not in rosette, or absent; basal and cauline sessile or subsessile, blade oblanceolate to linear-lanceolate, 10--40 mm, base attenuate, margins entire or with distant, slender, glandular teeth, , apex acute, surfaces glabrous. Inflorescences racemelike or flowers solitary; bracts linear to linear-lanceolate. Pedicels 0.5--2.5(--6) cm. Flowers erect in bud, becoming horizontal or slightly nodding; hypanthium broadly obconic, base rounded, surface glabrous; appendages absent; sepals spreading, lanceolate-attenuate, 4--8 mm, margins entire or with 1--several prominent teeth near base; corolla deep blue, 12--18 mm, tube shallowly cup-shaped or saucer-shaped, lobes recurved, lanceolate, 88--100% of corolla length; anthers 6--9 mm, pollen dark purple; styles , very dark purple, 10--12 mm, papillate in distal 75%; stigma lobes cream-colored, bluish or greenish, 10% of style length. Capsules erect, subcylindric to obconic, base rounded or tapered, pores apical. 2n = 34.
      Flowering Jun--Aug. Alpine regions, steppe community, on calcareous soil, serpentine soil, grassy openings, mixed forest, in creeks or run-off runnels, talus slopes, cliffs; 150--2100 m; B.C., N.W.T., Yukon; Alaska.
      Campanula aurita occurs in the Brooks Range, Porcupine, Black, Yukon, and Fortymile river drainages of Alaska, Mackenzie and Richardson mountains, to west-central and southern Yukon Territory, with disjunct populations in northern British Columbia.

    8. Campanula rotundifolia Linnaeus, Sp. Pl. 1: 163. 1753 * Harebell, campanule à reuilles rondes
      Campanula alaskana (A. Gray) Wight ex J. P. Anderson & Hultén; C. gieseckeana Vest; C. gieseckeana var. arctica (Lange) Böcher; C. gieseckeana subsp. groenlandica (Berlin) Böcher; C. intercedens Witasek; C. lasiocarpa Chamisso subsp. latisepala (Hultén) Hultén; C. lasiocarpa var. latisepala Hultén; C. latisepala Hultén; C. petiolata A. de Candolle; C. rotundifolia subsp. groenlandica (Berlin) Á. Löve & D. Löve; C. rotundifolia var. alpina Tuckerman; C. rotundifolia var. arctica Lange; C. rotundifolia var. intercedens (Witasek) Farwell; C. rotundifolia var. petiolata (A. de Candolle) J. K. Henry; C. sacajaweana M. Peck
      Herbs perennial, 5--90 cm, glabrous or with short hairs on petioles and stems; with taproot and rhizomes. Stems erect to decumbent, unbranched or branched. Leaves: basal in rosette, , petiolate; blade suborbicular to oblanceolate or ovate-cordate to ovate-elliptic, 10--30 mm, base attenuate, margins crenate-serrate to coarsely toothed, apex acute; cauline sessile, blade linear-lanceolate to nearly filiform, rarely petiolate and broadly lanceolate or ovate, 15--100 mm, base attenuate, margins entire to remotely serrulate, apex acute, surfaces glabrous or with short hairs on petioles. Inflorescences racemelike or paniclelike, or flowers sometimes solitary; bracts linear-subulate. Pedicels 1--2 cm. Flowers erect in bud, nodding when open; hypanthium turbinate or hemispheric, base acute, surface glabrous or rarely puberulent or hirtellous, ; appendages absent; sepals ascending to spreading or reflexed, linear-deltate, subulate, or setaceous, 2--21 mm, , margins entire; corolla pale to intense blue or blue-violet, rarely white, 9--35 mm, tube , campanulate, funnelform, or tubular, lobes erect or spreading, sometimes widely flaring, ovate or ovate-deltate, 25--55% of corolla length; anthers 3.5--6.5 mm, pollen colorless, gray, pink, or pale purple; styles pale to intense blue or blue-violet, rarely white, 9--17 mm, papillate in distal 50--60%; stigma lobes white, 15% of style length. Capsules nodding to pendent, hemispheric, base rounded, weakly ribbed, pores basal. 2n = 34, 68, 102.
      Flowering Jun--Oct. Rocky or sandy shores, cliffs, open woods, prairies, alpine and arctic meadows, fell-fields, screes, talus, alluvial fans, moraines, roadsides, pastures; 0--11400 m; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont. P.E.I., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Conn., Del., D.C., Idaho, Ill., Ind., Iowa, Kans., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nebr., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Oreg., Pa., R.I., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico (Coahuila, Nuevo Léon, Tamaulipas); Eurasia, Atlantic Islands (Iceland); introduced in Africa (South Africa), Pacific Islands (New Zealand).
      Campanula rotundifolia is extremely variable throughout its range. S. G. Shetler (1982) studied the species comprehensively as it occurs in North America. The following is a simplified summary of a very complicated situation, which Shetler discussed at length. He could discern four patterns that he called “races.” The “Alaskan Race” (somewhat equivalent to C. alaskana) has reclining or pendent stems to 77 cm, cauline leaves lanceolate or ovate, 3--4 flowers per stem on average, large corollas to 35 mm and 36 mm in diameter, occurring on rocky shores. The “Arctic Race” (somewhat equivalent to C. gieseckeana) has dwarf plants with erect stems to 50 cm, linear or linear-lanceolate cauline leaves, 1--2 flowers per stem on average, corollas to 26 mm and 27 mm in diameter, occurring in North Atlantic tundra and fjords. The “Eastern Race” (C. intercedens) has decumbent or pendent stems to 91 cm, cauline leaves linear, falcate, ribbonlike, with 7--8 flowers per stem on average (but as many as 33), corollas to 21 mm and 23 mm in diameter, occurring in lowlands and ravines in the Appalachian-Great Lakes region. The “Cordilleran Race” (C. petiolata) has rigid, erect stems to 71 cm, cauline leaves narrowly lanceolate, 5--6 flowers on average (but as many as 21), corollas 9--26 mm, 11--29 mm in diameter, occurring in central and western mountains. In addition to this diversity in vegetative morphology and inflorescences, corollas have a wide range of shapes from narrowly infundibular to deeply or shallowly campanulate. Shetler noted that in any particular location plants may be found with any combination of morphologies mentioned above, or populations with the features of one or more “races” may occur together or in the same general area; however, Campanuloideae taxa often occur sympatrically (N. R. Morin 2020), so this circumstance does not preclude the possibility that these races should be recognized at some rank.
      B. L. Sutherland and L. F. Galloway (2018), based on an analysis of genome duplication, determined that Campanula rotundifolia originated in south-central Europe. Hexaploids from western Europe radiated into North America about 114,000 years ago. Post-glacial colonization from a midwestern refugium occurred in an Eastern/Southern lineage and a northwestern North American lineage, the latter then split into a Cordilleran group and a Washington to Alaska group. These conclusions are consistent with Shetler’s concept of races. Shetler noted that diploids, tetraploids, and hexaploids occur in a matrix, in which the tetraploids form what he called the “groundmass,” and the others might turn up anywhere; however, he was able to discern two classes within the diploids: arctic-montane (many considered by European botanists as species separate from C. rotundifolia) and temperate lowland. Tetraploids in western North America and eastern North America can similarly be separated into several classes each. R. A. Bingham and T. A. Ranker (2000) found that arctic-alpine populations of tetraploid C. rotundifolia were as genetically diverse as lowland populations and attributed this to the effectiveness of bumblebees, which are their main pollinator. Morphometric and genetic analyses might shed light on species circumscriptions and affinities within the Campanula rotundifolia complex.
      SELECTED REFERENCES Bingham, R. A. and T. A. Ranker. 2000. Genetic diversity in alpine and foothill populations of Campanula rotundifolia (Campanulaceae). Int. J. Pl. Sci. 161: 403--411. Morin, N. R. 2020. Taxonomic changes in North American Campanuloideae (Campanulaceae). Phytoneuron 2020-49: 1--46. Shetler, S. G. 1982. Variation and Evolution of the Nearctic Harebells (Campanula subsect. Heterophylla). 516 pp. J. Cramer. Sutherland, B. L. and L. F. Galloway. 2018. Effects of glaciation and whole genome duplication on the distribution of the Campanula rotundifolia polyploid complex. Amer. J. Bot. 105: 1760--1770.

    9. Campanula chamissonis Federov, Fl. URSS 24: 279. 1957 * Hairyflower bellflower
      Campanula dasyantha M. Bieberstein subsp. chamissonis (Federov) Victorov
      Herbs perennial, 5--10 cm, glabrous or with a few stiff hairs on leaf margins and stem angles; cespitose, with taproot. Stems erect, unbranched. Leaves: basal in rosette, long-petiolate, blade broadly spatulate to obovate, 20--30 mm, base attenuate, margins crenate, apex obtuse to broadly acute; cauline petiolate, blade broadly elliptic, 10--20 mm, base attenuate, margins crenate to serrate, apex obtuse, surfaces pubescent. Inflorescences: flowers solitary; bracts elliptic to narrowly lanceolate. Pedicels 0.5--1 cm. Flowers erect to nearly horizontal; hypanthium broadly obconic, base broadly acute, surface densely hairy; appendages reflexed, auricular, 1--2 mm, ; sepals ascending, ovate-lanceolate, broad, 10--15 mm, margins entire or crenate-serrate; corolla blue, 30--40 mm, tube , broadly cylindric to funnelform, lobes erect to spreading, ovate, 33%--50% of corolla length, ; anthers 5 mm, pollen white; styles white, 15--20 mm, papillate in distal 90%; stigma lobes white, 10--15% of style length. Capsules pendent, broadly obconic, base rounded, pores basal. 2n = 34.
      Flowering Jul--Aug. Gravelly and sandy alpine slopes, dry scree slopes, muddy, gravelly areas, Empetrum nigrum-Racomitrium lanuginosum heath; 0--1400 m; Alaska (Aleutian Islands); Russia (Sakhalin), Japan.
      In Alaska, Campanula chamissonis is known from Agattu, Adak, Amchitka, Atka, Fox, Pribilof, and Unalaska islands. Campanula chamissonis is the only native campanula that phylogenetic studies place in the Campanula s.s. clade (Wendling et al. 2011), close to C. punctata and C. collina M. Bieberstein (C. Roquet, et al., 2009). Some authors consider C. chamissonis a subspecies of C. dasyantha (for example, Wendling et al. 2011), which is native in northern Asia. Campanula dasyantha differs in having basal leaves oblong-elliptic, lanceolate, or linear-lanceolate, 2--12 cm, all leaves +/- long-haired, and flowers declined or held horizontally.

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