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CAMPANULACEAE BELLFLOWER FAMILY

Nancy R. Morin, except as noted

Annual to perennial herb [tree]. Leaf: generally cauline, generally simple, generally alternate, petioled or not; stipules 0. Inflorescence: cyme, panicle, raceme, spike, or flowers 1; terminal or in axils of leaf-like or reduced bracts. Flower: bisexual, cleistogamous or open, radial or bilateral, inverted (pedicel twisted 180°) or not; hypanthium generally present, ± fused to ovary; sepals generally 5; corolla radial to 2-lipped, petals generally fused, tube deeply divided on 1 side or not, lobes generally 5; stamens 5, free or ± fused (anthers, filaments fused into tube or filaments fused above middle); ovary inferior or 1/2 inferior (superior in fruit), chambers 1–3, placentas axile or parietal, ovules many, style generally 1, 2–5-branched. Fruit: generally capsule, open on sides or top by pores or short valves. Seed: many.
± 90 genera, ± 2500 species: worldwide. [Haberle et al. 2008 J Molec Evol 66:350–361] Some cultivated for ornamental (Campanula, Jasione, Lobelia). Subfamilies sometimes treated as families. Positions of flower parts given after flowering inversion, if any. Parishella moved to Nemacladus. —Scientific Editor: Thomas J. Rosatti.
Unabridged references: [Lammers 2007 World Checklist and Bibliography of Campanulaceae. Royal Botanic Gardens, Kew.]

Key to Campanulaceae

DOWNINGIA

Lisa M. Schultheis

Annual, glabrous. Stem: decumbent to erect, (10)20–40 cm. Leaf: cauline, often deciduous before flower, 0.5–2(4) mm wide, lanceolate to awl-like (uppermost wider or not), sessile, generally entire. Inflorescence: spike; terminal flowers often aborted, overtopped by fertile; pedicels 0. Flower: bilateral, generally inverted at full bloom by twisted ovary; corolla generally >> calyx, blue to pink or white, generally with a symmetric white or yellow spot on lower lip, tube entire, limb strongly 2-lipped, upper lip lobes 2, lower lip generally with 2 low ridges, these often with 2 knob-like projections near throat, lobes 3, > upper, generally obovate, obtuse-mucronate; stamens fused (filaments, anthers in tubes), generally 2 smallest anthers each with terminal tuft of bristles, 1 bristle triangular or horn-like, generally 0.2–0.5 mm, others linear, shorter; ovary inferior, long, narrow, ± pedicel-like, chambers 1–2, placentas parietal or axile. Fruit: dehiscent on sides by 3–5 sometimes translucent slits, tardily so or not.
15 species: western North America, Chile. (A.J. Downing, American horticulturist, 1815–1852) [Schultheis 2001 Syst Bot 26:603–621] Flower part positions ("upper" is adaxial; "lower" is abaxial) given at full bloom. Corolla measurements are from base of tube to tip of longest lobe, color including albino for ± all species. Based on geography, interfertility, cytology, and sequences of nuclear as well as chloroplast DNA, Downingia yina Applegate (as recognized in TJM (1993)) split here into 3 morphologically indistinguishable species whose limits need further study, only 2 of which are documented for California: Downingia pulcherrima, Downingia willamettensis.
Unabridged references: [McVaugh 1941 Mem Torrey Bot Club 19:1–57; Weiler 1962 Ph.D. Dissertation, Univ of California, Berkeley; Schultheis 2001 Syst Bot 26:603–621]
Unabridged note: Flower part positions ("upper" is next to stem; "lower" is away from stem) given at full bloom. Corolla measurements are from base of tube to tip of longest lobe, color including albino for ± all species. Based on geography, interfertility, cytology, and sequences of nuclear as well as chloroplast DNA, Downingia yina (as recognized in TJM (1993)) split here into 3 morphologically indistinguishable species whose limits need further study: Downingia yina (as now circumscribed, not in California, although possibly in – yet not documented for – northern Cascade Range), Downingia pulcherrima, and Downingia willamettensis. The present treatment comes after Downingia pulcherrima and Downingia willamettensis had been united under Downingia willamettensis, and that taxon in turn had been transferred to Downingia yina, as Downingia yina var. major (leaving the remainder of that sp. under Downingia yina var. yina).

Key to Downingia

D. bicornuta A. Gray
NATIVE
Flower: corolla 7–19 mm, densely white-hairy in tube, lateral sinuses > upper, appearing hinged, lower lip blue-purple, with central white area including yellow-green spots, base purple with 2 prominent knob-like projections near throat; anthers < 45° to filaments, 2 larger bristles at tips of 2 lower, smaller anthers 0.6–2.7 mm, generally tightly twisted together (or 0 or not twisted); ovary glabrous to minutely spiny-scabrous, 2-chambered, placentas axile. Fruit: 35–65(90) mm, tardily dehiscent, lines not translucent. Seed: with longitudinal lines.
n=11. Occasionally intermediates between Downingia bicornuta var. bicornuta and Downingia bicornuta var. picta in San Joaquin, Stanislaus cos. [Online Interchange]

D. bicornuta var. bicornuta
NATIVE
Flower: corolla 9–19 mm, tube (2.5)3–4(4.5) mm, blue, without prominent veins adaxially, 2 upper lobes erect or recurved, not crossed at tips, lower lip generally ± flat; 2 largest bristles at tips of 2 lower, smallest anthers 0.6–1.5 mm, < anthers.
Vernal pools, roadside ditches, lake margins; < 1700 m. Inner North Coast Ranges, s Cascade Range, n Sierra Nevada, c Sierra Nevada Foothills, Sacramento Valley, n San Joaquin Valley, Modoc Plateau; to southern Oregon, southwestern Idaho, western Nevada. Apr–Jul [Online Interchange]

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Citation for the whole project: Jepson Flora Project (eds.) 2013. Jepson eFlora, http://ucjeps.berkeley.edu/IJM.html, accessed on Oct 23 2014
Citation for this treatment: [Author of taxon treatment] 2013. Downingia, in Jepson Flora Project (eds.) Jepson eFlora, http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=57808, accessed on Oct 23 2014

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Geographic subdivisions indicated for the distribution of Downingia bicornuta var. bicornuta Markers link to CCH specimen records. If the markers are obscured, reload the page [or change window size and reload]. Yellow markers indicate records that may provide evidence for eFlora range revision or may have georeferencing or identification issues.
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map of distribution 1
(Note: any qualifiers in the taxon distribution description, such as 'northern', 'southern', 'adjacent' etc., are not reflected in the map above, and in some cases indication of a taxon in a subdivision is based on a single collection or author-verified occurence).

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Data provided by the participants of the Consortium of California Herbaria.
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CCH collections by month

Duplicates counted once; synonyms included.
Species do not include records of infraspecific taxa.
Blue line denotes eFlora flowering time.