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10. Campanulaceae
桔梗科 jie geng ke
Authors: Deyuan Hong, Prof. Ge Song, Thomas G. Lammers & Laura L. Klein
Herbs, lactiferous, perennial (often with rhizomes or caudices), less often annual, [or shrubs or trees outside of China]. Leaves estipulate, alternate (rarely opposite or whorled), simple, entire or variously toothed or dissected, petiolate or sessile. Flowers tetracyclic, perfect, with a specialized method of proterandrous secondary pollen presentation, resupinate in Lobelioideae, solitary in an axillary or less often terminal position, or aggregated into terminal or axillary inflorescences that appear racemose or paniculate (rarely capitate), pedicellate or rarely sessile; bracts leaflike or reduced, rarely absent; pedicels often bracteolate. Calyx synsepalous, adnate to ovary and forming a hypanthium (free in Cyananthus); lobes (3-)5(-10), valvate, sometimes with a reflexed appendage in each sinus, in some Codonopsis, inserted well below rim of hypanthium. Corolla sympetalous, actinomorphic or zygomorphic, most often some shade of blue or violet, or white; lobes (4 or)5(-10), valvate. Stamens equaling number of corolla lobes, antisepalous, distinct, coherent, or connate; anthers tetrasporangiate, dithecal, introrsely dehiscent by longitudinal slits, basifixed. Gynoecium syncarpous, 2-5(-10)-locular with axile placentation, [rarely 1-locular with parietal, basal, or apical placentation outside of China]; ovary at least partly inferior (superior in Cyananthus), sometimes crowned by an annular or tubular nectary; style solitary, pubescent with pollen-collecting hairs below apex; stigma typically with as many lobes as ovary locules. Fruit a capsule, commonly apically loculicidal or laterally poricidal, or a berry. Seeds usually small, numerous; embryo small, straight; endosperm copious, oily or rarely starchy.
Eighty-six genera and over 2,300 species: cosmopolitan; 16 genera (one endemic, two introduced) and 159 species (77 endemic, two introduced) in China.
Two of us (Lammers and Klein) do not agree with the separation of Leptocodon and Campanumoea from Codonopsis in this treatment. We consider that the single character used in each case, five lamellar glands alternate with the stamens in Leptocodon and baccate fruit in Campanumoea, is not sufficient to justify recognition of a distinct genus. We regard these two genera as embedded within Codonopsis, as was done by Lammers (in C. I. Peng, Phytogeogr. Bot. Invent. Taiwan, 43-61. 1992), Morris and Lammers (Bot. Bull. Acad. Sin. 38: 277-284. 1997), Lammers (in T. C. Huang, Fl. Taiwan, ed. 2, 4: 775-802. 1998), Lammers (in Kubitzki [Kadereit & Jeffrey], Fam. Gen. Vasc. Pl. 8: 26-56. 2007), and Lammers (World Checkl. Bibliogr. Campanulaceae. 262-273. 2007). On the other hand, one of us (Hong) notes that two characters distinguish Leptocodon from Codonopsis. One is the five lamellar glands and the other is the shape of the stigma segments, which are clavate in Leptocodon but orbicular-deltoid in Codonopsis. Campanumoea differs from Codonopsis not only in having baccate fruit but also in its pollen, which is shortly 5-colpate (vs. 5-8-long-colpate in the core group of Codonopsis). Campanumoea is not a very distinct genus, and it seems that it would be equally reasonable to sink it into Codonopsis or to separate it. However, if it were sunk, then it would also be reasonable to sink Adenophora into Campanula because the former genus differs from the latter only in having a nectar disk around the style. Pentaphragma Wallich ex G. Don and Sphenoclea Gaertner were included in Campanulaceae in FRPS but are treated in this volume as Pentaphragmataceae and Sphenocleaceae, respectively. See the notes under those two families. Hong De-yuan, Lian Yong-shan & Shen Lian-dai. 1983. Campanulaceae (Campanuloideae and Lobelioideae). In: Hong De-yuan, ed., Fl. Reipubl. Popularis Sin. 73(2): 1-173. T. G. Lammers. 1998. Campanulaceae. In: Tseng-Chieng Huang, ed., Fl. Taiwan, ed. 2, 4: 775-802.
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| 1 |
Flowers resupinate at anthesis; odd (unpaired) calyx lobe in a ventral (anterior) position prior to resupination; corolla zygomorphic (sometimes only slightly so); stamens connate, forming an asymmetric tube; locules and stigmas 2 (Lobelioideae) |
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(2) |
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Flowers oriented normally; odd (unpaired) sepal in a dorsal (posterior) position; corolla actinomorphic; stamens distinct or coherent into a symmetric tube at anthesis but eventually separating; locules and stigmas (2 or)3-6 (Campanuloideae) |
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(3) |
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| 2 (1) |
Corolla strongly zygomorphic, unilabiate or bilabiate, lobes ± as long as tube or slightly shorter, tube cleft to base on dorsal surface; staminal column free from corolla, exserted from dorsal fissure of corolla. |
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15 Lobelia |
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Corolla scarcely zygomorphic, salverform, lobes less than 1/3 as long as tube, tube entire; staminal column adnate to corolla below middle, scarcely exserted from mouth of corolla. |
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16 Hippobroma |
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| 3 (1) |
Fruit a berry; ovary and fruit apically flat |
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(4) |
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Fruit a capsule; ovary and fruit apically conical |
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(5) |
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| 4 (3) |
Herbs twining; calyx lobes ovate-deltoid or ovate-lanceolate, margin entire. |
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4 Campanumoea |
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Herbs erect; calyx lobes linear or linear-lanceolate, margin dentate, rarely entire. |
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5 Cyclocodon |
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| 5 (3) |
Fruit dehiscing apically (i.e., above calyx lobes) |
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(6) |
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Fruit dehiscing laterally (i.e., below calyx lobes) |
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(11) |
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| 6 (5) |
Calyx completely free from ovary, inserted below it; corolla and stamens inserted below ovary. |
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1 Cyananthus |
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Calyx tube adnate to ovary, forming a hypanthium (though lobes sometimes apparently inserted at base or middle rather than its rim); corolla and stamens inserted on rim of hypanthium (i.e., atop ovary) |
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(7) |
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| 7 (6) |
Glands 5, alternate with stamens, orbicular, compressed; herbs twining. |
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2 Leptocodon |
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Glands absent; herbs erect, climbing, or twining |
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(8) |
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| 8 (7) |
Taproot greatly thickened, carrotlike or tuberous; flowers large, corolla 15-50 mm; stigmas ovoid or orbicular, ± as long as broad |
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(9) |
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Taproot fibrous or but scarcely thickened; flowers small, corolla 2-12 mm; stigmas filiform or linear, much longer than broad |
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(10) |
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| 9 (8) |
Plants not malodorous; stems erect or ascending; locules 5, alternating with calyx lobes. |
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7 Platycodon |
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Plants often malodorous; stems often scandent, climbing, or twining; locules 3. |
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3 Codonopsis |
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| 10 (8) |
Leaves pinnately lobed or parted; calyx lobes echinate; corolla lobes and stamens typically 4; pollen 4- or 5-colpate; seeds triquetrous. |
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6 Echinocodon |
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Leaves entire or toothed; calyx lobes entire or toothed; corolla lobes and stamens typically 5; pollen 3-porate; seeds terete or compressed. |
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8 Wahlenbergia |
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| 11 (5) |
Corolla rotate, lobed for 1/2 or more its length |
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(12) |
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Corolla campanulate, lobed less than 2/5 its length |
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(13) |
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| 12 (11) |
Plants annual, roots fibrous; flowers both chasmogamous and cleistogamous, latter often predominating; corolla lobes cut for 1/2-3/4 their length, lanceolate. |
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13 Triodanis |
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Plants perennial, root carrotlike; flowers all chasmogamous; corolla lobes divided to base, linear. |
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14 Asyneuma |
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| 13 (11) |
Flowers large, corolla typically 15-75 mm; fruit dehiscing via 2 or 3 definite pores in firm sclerenchymatous pericarp |
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(14) |
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Flowers small, corolla 5-12 mm; fruit dehiscing irregularly via ruptures and tears of membranous or papery pericarp |
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(15) |
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| 14 (13) |
Flowers without an obvious nectar disk. |
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9 Campanula |
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Flowers with a conspicuous tubular (rarely annular) nectar disk. |
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10 Adenophora |
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| 15 (13) |
Plants perennial via tuberiferous stolons; stems terete; flowers long pedicellate; seeds > 1.5 mm, testa finely striate. |
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11 Peracarpa |
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Plants annual; flowers sessile; stems 3-winged; seeds < 1 mm, testa reticulate. |
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12 Homocodon |
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List of lower taxa
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