CAMPANULACEAE 桔梗科 jie geng ke Hong Deyuan (洪德元)1, Ge Song (葛颂)1; Thomas G. Lammers2, Laura L. Klein2 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 tet- racyclic, 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, dis- tinct, coherent, or connate; anthers tetrasporangiate, dithecal, introrsely dehiscent by longitudinal slits, basifixed. Gynoecium syn- carpous, 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. Rei- publ. Popularis Sin. 73(2): 1–173. T. G. Lammers. 1998. Campanulaceae. In: Tseng-Chieng Huang, ed., Fl. Taiwan, ed. 2, 4: 775–802. 1a. 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). 2a. 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 ......................... 15. Lobelia 2b. 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 ......................................................... 16. Hippobroma 1b. 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). 3a. Fruit a berry; ovary and fruit apically flat. 4a. Herbs twining; calyx lobes ovate-deltoid or ovate-lanceolate, margin entire ............................................ 4. Campanumoea 4b. Herbs erect; calyx lobes linear or linear-lanceolate, margin dentate, rarely entire ......................................... 5. Cyclocodon 3b. Fruit a capsule; ovary and fruit apically conical. 5a. Fruit dehiscing apically (i.e., above calyx lobes). 6a. Calyx completely free from ovary, inserted below it; corolla and stamens inserted below ovary .......... 1. Cyananthus 6b. Calyx tube adnate to ovary, forming a hypanthium (though lobes sometimes apparently inserted 1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 2 Biology and Microbiology Department, University of Wisconsin Oshkosh, Oshkosh, Wisconsin 54901–8640, U.S.A. 505 506 CAMPANULACEAE at base or middle rather than its rim); corolla and stamens inserted on rim of hypanthium (i.e., atop ovary). 7a. Glands 5, alternate with stamens, orbicular, compressed; herbs twining .......................................... 2. Leptocodon 7b. Glands absent; herbs erect, climbing, or twining. 8a. Taproot greatly thickened, carrotlike or tuberous; flowers large, corolla 15–50 mm; stigmas ovoid or orbicular, ± as long as broad. 9a. Plants not malodorous; stems erect or ascending; locules 5, alternating with calyx lobes ... 7. Platycodon 9b. Plants often malodorous; stems often scandent, climbing, or twining; locules 3 ................ 3. Codonopsis 8b. Taproot fibrous or but scarcely thickened; flowers small, corolla 2–12 mm; stigmas filiform or linear, much longer than broad. 10a. Leaves pinnately lobed or parted; calyx lobes echinate; corolla lobes and stamens typically 4; pollen 4- or 5-colpate; seeds triquetrous ....................................................... 6. Echinocodon 10b. Leaves entire or toothed; calyx lobes entire or toothed; corolla lobes and stamens typically 5; pollen 3-porate; seeds terete or compressed ................................................ 8. Wahlenbergia 5b. Fruit dehiscing laterally (i.e., below calyx lobes). 11a. Corolla rotate, lobed for 1/2 or more its length. 12a. Plants annual, roots fibrous; flowers both chasmogamous and cleistogamous, latter often predominating; corolla lobes cut for 1/2–3/4 their length, lanceolate ............................................ 13. Triodanis 12b. Plants perennial, root carrotlike; flowers all chasmogamous; corolla lobes divided to base, linear ............................................................................................................................................... 14. Asyneuma 11b. Corolla campanulate, lobed less than 2/5 its length. 13a. Flowers large, corolla typically 15–75 mm; fruit dehiscing via 2 or 3 definite pores in firm sclerenchymatous pericarp. 14a. Flowers without an obvious nectar disk .............................................................................. 9. Campanula 14b. Flowers with a conspicuous tubular (rarely annular) nectar disk ..................................... 10. Adenophora 13b. Flowers small, corolla 5–12 mm; fruit dehiscing irregularly via ruptures and tears of membranous or papery pericarp. 15a. Plants perennial via tuberiferous stolons; stems terete; flowers long pedicellate; seeds > 1.5 mm, testa finely striate ...................................................................................... 11. Peracarpa 15b. Plants annual; flowers sessile; stems 3-winged; seeds < 1 mm, testa reticulate .............. 12. Homocodon 1. CYANANTHUS Wallich ex Bentham in Royle, Ill. Bot. Himal. Mts. 1: 309. 1836, nom. cons., not Rafinesque (1815). 蓝钟花属 lan zhong hua shu Hong Deyuan (洪德元); Thomas G. Lammers, Laura L. Klein Herbs, annual or perennial, caudices of latter thick, lignified, covered with numerous scales, usually branched, stems thus caespitose. Leaves alternate, but sometimes 4 or 5 leaves aggregated below pedicel, appearing whorl-like. Flowers solitary, terminal on main stems and branches, rarely in a dichasium and dichasia aggregated on top of stem and branches, pedicellate. Calyx tubular or tubular-campanulate, 5- or rarely 4-lobed. Corolla tubular-campanulate, (3–)5-lobed. Stamens 5, often aggregated and surrounding ovary at upper part. Ovary superior, conical, mostly 5-locular, less frequently 4- or 3-locular. Fruit a capsule, loculicidal. Seeds numerous, brown-red or brown-black. Eighteen species: Himalaya