Botanical Studies (2010) 51: 553-561.
A synopsis of Codonopsis subg. Pseudocodonopsis (Campanulaceae: Campanuloideae), with description of a new species of uncertain provenance
Thomas G. LAMMERS* and Laura L. KLEIN1
Department of Biology and Microbiology, University of Wisconsin Oshkosh, Oshkosh, WI 54901, USA
(Received February 5, 2010; Accepted May 28, 2010)
ABSTRACT. The species of Codonopsis subg. Pseudocodonopsis are inodorous herbaceous geophytes with an unbranched tuberous root no more than four times longer than thick; twining stems leafy at least toward base; horizontal to erect usually solitary terminal flowers, in which the corolla is rotate and cleft for more than three-fourths its length; pollen grains with 5-10 colpi covering less than one-fourth the distance from pole to pole and spinules 2-3 [m long; and a 3-locular ovary that forms an obconic capsule with a relatively short dehiscent apex, containing numerous smooth or finely striate seeds. The most recent synopsis recognized C. convolvulacea (here designated as the type of the subgenus), C. efilamentosa, C. forrestii, C. grey-wilsonii (as C. nepalensis, nom. illeg.), C. limprichtii (as "C. pinifolia", nom. invalid.), and C. vinciflora. Codonopsis rosulata is also a member of C. subg. Pseudocodonopsis despite its scapose non-twining stem and basally rosulate leaves. Codonopsis hirsuta was previously included in C. convolvulacea, but here is judged to be a distinct species allied to C. limprichtii. Codonopsis mairei was previously included in C. forrestii, but is here judged to be a distinct species allied to C. convolvulacea. Plants known only from cultivation, which have been identified as C. forrestii but differ from all known species in several characters, are described here as C. macrophylla, sp. nov. Yunnan is the center of diversity, harboring eight out of ten species; the exceptions are C. grey-wilsonii, endemic to the eastern Himalaya; and C. macrophylla, of uncertain provenance. From Yunnan, the ranges of various species extend into Sichuan, Guizhou, Tibet (Xizang), and/or Burma (Myanmar).
Keywords: Asia; Campanulaceae; Campanuloideae; Classification; Codonopsis; New species; Pseudocodonopsis; Taxonomy.
INTRODUCTION
1. Plants not fetid; root-tuber no more than 4 times longer than thick, unbranched; colpi covering less than / the distance from pole to pole; fruit an obconic (rarely oblate) capsule, the dehiscent apex relatively short; seeds smooth or finely striate.
2. Flower nodding to pendent; corolla campanulate, lobed for less than / its length; colpi covering % -/ the distance from pole to pole; spinules on pollen grains ca. 1 [m long ....................................................
...........................C. subg. Obconicapsula D. Y. Hong
2. Flower horizontal to erect; corolla rotate, cleft for more than 3A its length; colpi covering less than % the distance from pole to pole; spinules on pollen grains
2-3 [im long..........C. subg. Pseudocodonopsis Kom.
The autonymic subgenus is the largest of the three, with 52 species divided between two sections: C. sect. Codon­opsis [including Leptocodon (Hook. f.) Lem. and Cam-panumoea Blume, s. str.] and C. sect. Erectae (Kom.) D. Y. Hong [including Glosocomia D. Don]. Its distribution extends from Afghanistan to Kamchatka and as far south as Java (Lammers, 2007a, 2007b). There is no modern taxonomic treatment of C. subg. Codonopsis in its entirety, though the floristic account for China (Shen and Hong, 1983) covers the majority of its members.
Codonopsis Wall. (Campanulaceae: Campanuloideae) is endemic to southern and eastern Asia. It is character­ized by a large tuberous root, herbaceous often scandent or climbing stems, flowers usually solitary and terminal, 5-10-colpate pollen, 3-(5)-locular ovary, apically dehis­cent many-seeded capsule (rarely a berry partly subtended by the calyx lobes), and a chromosome number of 2n = 16 (Lammers, 2007a). Three subgenera are recognized (Shen and Hong, 1983), which may be distinguished using the following key:
Key to the Subgenera of Codonopsis
1. Plants commonly fetid; root-tuber much longer than thick, sometimes branched; colpi covering more than 2/3 the distance from pole to pole; fruit an ovoid capsule with elongate apex (rarely a berry partly subtended by the calyx lobes); seeds reticulate.....C. subg. Codonopsis
1Present address: Department of Botany, Miami University,
Oxford, OH 45056, USA.
*Corresponding author: E-mail: lammers@uwosh.edu.
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Codonopsis subg. Obconicapsula comprises but a single species, C. dicentrifolia (C. B. Clarke) W. W. Smith of the Himalaya. As noted by Fletcher (1937) and Grey-Wilson (1990), this species is intermediate in its character­istics and might logically be included in either subgenus. It resembles C. subg. Codonopsis in its floral posture, corolla morphology, and short pollen spinules, but also possesses the compact root-tubers, obconic short-beaked capsule, and smooth or finely striate seeds of C. subg. Pseudocodonopsis; furthermore, its colpi are of intermedi­ate length. Were it not for this species, it would be very tempting to recognize the other two subgenera as distinct genera.
Codonopsis subg. Pseudocodonopsis is endemic to south-central China and adjacent areas. Grey-Wilson (1995) provided a useful summary of the group, in which he recognized six species: C. convolvulacea, C. efilamen­tosa, C. forrestii, C. nepalensis, "C. pimifolia", and C. vinciflora. In examining material for the Flora of China project, however, we recognized four additional species (one unnamed) referable to this subgenus, corrected no-menclatural problems with two of the names, and noted some additional synonymy. For this reason, we here pres­ent a synopsis of the subgenus, to make clear the rationale for changes from Grey-Wilson's treatment that will appear in the Flora of China.
ish or brownish; stems typically branched, slender, up to 3 m long; latex white. Leaves exstipulate, alternate or sometimes opposite, simple, dorsiventral, pinnately veined (dillenid), entire or minutely toothed, flat or revolute, commonly petiolate. Flowers tetracyclic, actinomorphic, perfect, epigynous, relatively large, horizontal to erect, solitary on main stem and branches (in few-flowered axil­lary clusters in C. efilamentosa), chasmogamous, proteran-drous, zoophilous. Calyx synsepalous, adnate to the ovary, forming an appendicular hypanthium; lobes 5, valvate, inserted at rim of hypanthium. Corolla early-sympetalous, rotate, cleft for more than 3A its length, various shades of blue or purple, less often yellow or white, sometimes with a contrasting ring toward center; lobes 5, valvate. Stamens 5, antisepalous, inserted at top of ovary; filaments dilated basally, distinct but proximate, forming a dome over the base of the style; anthers tetrasporangiate, dithecal, intror-sely dehiscent by longitudinal slits, basifixed, coherent; pollen (Morris and Lammers, 1997; Wei, 2001) oblate-spheroidal or suboblate, circular in polar outline, 34-48 [im polar diam. x 38-51 equatorial diam., (4-)6-colpate with colpi that cover less than one-fourth the distance from pole to pole, surface spinules 2-3 [m long, and a nexine less than 0.60 [im thick. Gynoecium syncarpous, 3-locular with axile placentation; ovary inferior, crowned by an an­nular nectary; ovules numerous, small, anatropous, uniteg-mic, tenuinucellate; style solitary, pubescent below apex; stigma large, broadly 3-lobed. Fruit an obconic (oblate in C. grey-wilsonii) capsule, the low-conic apex loculicidal via valves. Seeds usually small, numerous; testa smooth or faintly striate. Chromosome number unknown.
Distribution and ecology: Endemic to south-central China (Yunnan, Sichuan, Guizhou) and adjacent areas of Tibet (Xizang), the eastern Himalaya, and Burma (Myan-mar). All of the species flower at the end of the monsoon season (August-October), and perfect seed in the dry inter­val before winter (Grey-Wilson, 1995).
TAXONOMIC TREATMENT
Codonopsis subg. Pseudocodonopsis Kom., Trudy Imp. S.-Peterburgsk. Bot. Sada 29: 102. 1908.—TYPE [here designated]: Codonopsis convolvulacea Kurz. This and C. vinciflora were the two original species assigned to this subgenus.
Twining geophytes (C. rosulata rosulate and scapose); plants terrestrial, inodorous; roots tuberous, large, no more than 4 times longer than thick, unbranched, whit-
Key to the Species of Codonopsis subg. Pseudocodonopsis
1. Stems procumbent or weakly scandent, 0.1-0.4 cm long; leaves basally rosulate .....................................................1. C. rosulata
1. Stems twining, at least in the upper half, 0.5-3 m long; leaves cauline.
2. Stems decumbent or ascending below the middle, twining above; leaves largely confined to the non-twining portion of stem.
3. Stems hirsute; lamina ovate or narrowly oblong, hirsute abaxially ....................................................................2. C. hirsuta
3. Stems glabrous; leaves linear or linear-lanceolate, glabrous...............................................................................3. C. limprichtii
2. Stems twining; leaves scattered throughout length of stem.
4. Corolla lobes 26-50 mm long.
5. Calyx lobes erect or spreading, 11-24 mm long; corolla glabrous, concolorous, the lobes narrowly elliptic; fila­ments without prominent auricles at summit of the dilated portion; capsules obconic, 24-26 mm long..........................................
............................................................................................................................................................................9. C. forrestii
5. Calyx lobes deflexed, 7-11 mm long; corolla pubescent within, with a darker ring at center, the lobes ovate; fila­ments with a pair of prominent auricles at summit of the dilated portion; capsules oblate, 12-14 mm long ..........
....................................................................................................................................................................10. C. grey-wilsonii
4. Corolla lobes 14-28 mm long.
LAMMERS and KLEIN ― A synopsis of Codonopsis subg. Pseudocodonopsis 555
6. Flowers mostly in axillary clusters; corolla lobes oblong, the apex rounded.................................................8. C. efilamentosa
6. Flowers terminal on main stem and upper branches; corolla lobes elliptic, the apex acute.
7. Petioles 10-13 mm long; calyx lobes 5-8 mm long, in bud free and divergent...............................................7. C. vinciflora
7. Petioles 1-7 mm long; calyx lobes (6-)8-16 mm long, in bud coherent and enveloping the corolla.
8. Lamina ovate, 5.5-7.5 x 3.3-5.1 cm, the apex obtuse; hypanthium 9-13 mm diam.; calyx lobes triangular; corolla lobes 15-18 mm wide ................................................................................................................................... 6. C. macrophylla
8. Lamina linear-lanceolate, lanceolate, or ovate-lanceolate sometimes narrowly elliptic), 1.8-5 x 0.2-3 cm, the apex acuminate or acute; hypanthium 4-10 mm diam.; calyx lobes lanceolate or narrowly triangular; corolla lobes 7-15 mm wide.
10. Lamina linear-lanceolate to lanceolate (sometimes narrowly elliptic), 0.2-1.4 cm wide; hypanthium 3-5 mm long; corolla lobes 14-22 mm long ..................................................................................................... 4. C. convolvulacea
10. Lamina lanceolate to ovate-lanceolate, 1-3 cm wide; hypanthium 5-7 mm long; corolla lobes 22-28 mm long ............................................................................................................................................................. 5. C. mairei
1. Codonopsis rosulata W. W. Sm., Notes Roy. Bot. Gard.
Edinburgh 13: 157. 1921.—TYPE: CHINA. Sichuan:
Mu-li mountains, amongst heavy grass, 28°12'N, 11000 ft, Aug 1918, G. Forrest 16856 (holotype: E-00265625! isotype: K-000575332!).
Icones. Shen and Hong (1983), pl. 7.4.
Distribution. Endemic to south-central China, in north­western Yunnan and southwestern Sichuan. Grassy slopes and margins of pine forests, 2,600-3,600 m.
Discussion. Codonopsis rosulata was not mentioned by Grey-Wilson (1995). Perhaps he was influenced in this by Anthony (1926), who assigned the species to C. subg. Codonopsis. While C. rosulata does differ from the remainder of C. subg. Pseudocodonopsis in its basally rosulate scapose habit, the distinctive corolla and capsule clearly support its inclusion here (Shen and Hong, 1983; Hong and Ma, 1992).
in contrast has longer stems that are leafy and twining throughout. Lammers (2007b) included C. hirsuta within the circumscription of C. limprichtii because of the dis­tinctive habit they shared. However, the differences in pu­bescence and leaf morphology given in the key above are quite pronounced, with no intermediates. For that reason, we follow Hong and Ma (1992) in recognizing them as distinct species.
3. Codonopsis limprichtii Lingelsh. & Borza, Repert. Spec. Nov. Regni Veg. 13: 391. 1914. Codonopsis convolvulacea var. limprichtii (Lingel & Borza) Antho­ny, Notes Roy. Bot. Gard. Edinburgh 15: 178. 1926.— TYPE: CHINA. Yunnan: "Talifu, an schattigen Platzen unter Kiefern vor Ai-ning-po, bei Pu-peng, 2,500 m," 21 Aug 1913, H. W. Limpricht 851 (holotype: WRSL).
Codonopsis graminifolia H. Lev., Cat. Pl. Yun-nan: 24. 1916.—TYPE: CHINA. Yunnan: Paturages des mon-tagnes, partout, 2,400 m, Jul 1911, E. E. Maire s.n. (ho-lotype: E; isotype: E). The holotype includes extraneous material of C. convolvulacea (Chamberlain, 1977).
Codonopsis limprichtii var. pinifolia Hand.-Mazz., Akad.
Wiss. Wien, Math.-Naturwiss. Kl., Anz. 61: 170.
1924. Codonopsis convolvulacea var. pinifolia (Hand.-Mazz.) J. Anthony, Notes Roy. Bot. Gard. Edinburgh
15: 178. 1926. "Codonopsis pinifolia" (Hand.-Mazz.) Grey-Wilson, Plantsman 12: 89 (1990). Nom. invalid. sub Art. 33.4.—TYPE: CHINA. Yunnan: "In pinetis
calide temperatis circa jugum Yenaping ad occid. op-pidi Djientschwan (»Kientschwan«) ubique, s. calceo,
arenaceo, etc., 2,800-3,250 m," 22 Sep 1916, H. Han-
del-Mazzeti 10053 (Syntype: W). Specimens 3713 pro parte and 4370, both from near "Lidjiang" were also cited.
Icones. Grey-Wilson (1990), pg. 76 A [as "C. pinifo-lia"]; Grey-Wilson (1995), pg. 214 [as "C. pinifolia
Distribution. Endemic to south-central China, in north­ern Yunnan, southwestern Sichuan, and western Guizhou. Open grassy slopes, thickets, and pine forests, 1,500-3,300 m.
2. Codonopsis hirsuta (Hand.-Mazz.) D. Y. Hong & L. M. Ma in H. W. Li & Z. Y. Zhu, Fl. Sichuamca 10: 546. 1992. Codonopsis limprichtii var. hirsuta Hand.-Mazz., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Anz. 61: 169. 1924. Codonopsis convolvulacea var. hirsuta (Hand.-Mazz.) J. Anthony, Notes Roy. Bot. Gard. Ed­inburgh 15: 178. 1926.—TYPE: CHINA. Sichuan: "ad terram abruptam calceam reg. temperatae supra vicum Lodjiahosan pr. oppidum Yenyuen, 3,000 m," 30 Sep 1914, H. Handel-Mazzeti 5423 (holotype: W).
Distribution. Endemic to south-central China, in north­western Yunnan and southwestern Sichuan. Open thickets and sunny grassy slopes, 2,400-3,100 m.
Discussion. Codonopsis hirsuta has been treated as a variety (Anthony, 1926; Shen et al., 1975; Shen and Hong, 1983; Huang, 1991) or heterotypic synonym (Nan-nfeldt, 1936) of C. convolvulacea, which is very similar in characters of leaf and flower. However, C. hirsuta has the distinctive habit of C. limprichtii, with the rather short stems divided into a leafy non-twining basal portion and an almost leafless twining apical portion; C. convolvulacea
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Discussion. Members of C. subg. Pseudocodonopsis with the half-twining habit of C. hirsuta (see above) but glabrous much narrower leaves have been called C. con-volvulacea var. pinifolia (Anthony, 1926; Tsoong, 1935;
Shen et al., 1975; Chamberlain, 1977; Shen and Hong, 1983; Huang, 1991). Grey-Wilson (1990) was of the opin­ion that this distinctive taxon merits specific rank. Unfor­tunately, in coining a name for it, he neglected to provide a full and direct reference to the basionym (C. limprichtii var. pinifolia), citing merely "syn. C. convolvulacea var. pinifolia Hand.-Mazz." As a result, "C. pinifolia" was not validly published. The binomial C. graminifolia is a heterotypic synonym of C. convolvulacea var. pini-folia (Anthony, 1926; Tsoong, 1935; Shen et al., 1975; Chamberlain, 1977; Shen and Hong, 1983; Huang, 1991; Lammers, 2007b) and would be the correct name for this variety at specific rank (Hong and Ma, 1992).
Shen et al. (1975) expanded the circumscription of C. convolvulacea var. pinifolia by adding to it C. c. var. lim-prichtii (Anthony, 1926; Shen and Hong, 1983; Huang, 1991). At varietal rank, the former name has priority. Grey-Wilson (1995) similarly could discern no way by which to distinguish the two. He, however, wished to rec­ognized the resulting taxon at specific rank. In doing so, he maintained the invalid name "C. pinifolia" and cited C. limprichtii as a heterotypic synonym. The correct name under Art. 11.4 for the species thus circumscribed is C. limprichtii. Lammers (2007b) included C. limprichtii var. hirsuta as a heterotypic synonym of this species, but as noted above, it is here considered a distinct species, fol­lowing Hong and Ma (1992).
feldt (1936) felt it was "pointless" to distinguish these va­rieties (as well as C. forrestii), they were accepted by Shen and Hong (1983) and Huang (1991), who added to them C. c. var. efilamentosa, C. c. var. forrestii and C. c. var. vinci­flora. In this way, the entire subgenus save non-twining C. rosulata was reduced to a single species. As Grey-Wilson (1990, 1995) pointed out, however, there are consistent morphological and ecological differences among these groups of populations, which support their recognition at specific rank. Consequently, we adopt here a very narrow circumscription of C. convolvulacea, encompassing only the plants called C. c. var. convolvulacea by earlier authors (Anthony, 1926; Shen and Hong, 1983; Huang, 1991).
Codonopsis forrestii var. heterophylla has been treated as a heterotypic synonym of C. vinciflora by most au­thors (Shen and Hong, 1983; Huang, 1991; Hong and Ma, 1992). However, careful study of the photograph of the holotype included in the protologue indicates to us that this is not an appropriate disposition for the name. That photo shows a plant with lanceolate acuminate leaves on very short petioles and relatively long calyx lobes envelop­ing the flower in bud; in contrast, C. vinciflora has ovate or deltate acute or obtuse leaves on rather long petioles and relatively short divergent calyx lobes. Lammers (2007b) treated C. forrestii var. heterophylla as a heterotypic syn­onym of C. forrestii, but that assignment is likewise unten­able, as the corolla seen in the photo is far smaller than in C. forrestii. There is nothing in either the photo or the diagnosis that precludes identifying the specimen as C. convolvulacea.
4. Codonopsis convolvulacea Kurz, J. Bot. 11: 195. 1873.
—TYPE: "Yunan, Hotha".
Codonopsis forrestii var. heterophylla C. Y. Wu, Rep.
Yunnan Trop. Subtrop. Fl. Res. Inst. 1: 80. 1965.—
TYPE: T. T. Yu 10173.
Icones. Oliver (1895), pl. 2385; Tsoong (1935), fig. 6; Wu and Li (1965), pl. XXX.1 [as C. forrestii var. hetero-
phylla]; Grey-Wilson (1990), pg. 76 right; Huxley (1992), pg. 667(i); Grey-Wilson (1995), pg. 221 R; Morris and
Lammers (1997), figs. 7-9 [pollen]; Wei (2001), pl. II.2-3 [pollen].
Distribution. Endemic to southern Yunnan and adjacent Burma (Myanmar); reports from other regions (e.g., by
Prain, 1908; Limpricht, 1922; Tsoong, 1935; Cox, 1936;
Fletcher, 1937; Finlay, 1972; Hara, 1982) refer to various other members of the subgenus. Forests, open woods, grassy slopes, pastures, and thickets, 1,000-4,600 m.
Discussion. The circumscription of this, the first mem­ber of C. subg. Pseudocodonopsis to be described, has varied dramatically over the years, depending upon which of the other taxa have been subordinated to it as varieties or heterotypic synonyms. Anthony (1926) recognized four varieties: C. convolvulacea var. convolvulacea (as "var. typica," nom. invalid. sub Art. 24.3), C. c. var. limprichtii, C. c. var. hirsuta, and C. c. var. pinifolia. Although Nann-
5. Codonopsis mairei H. Lev., Cat. Pl. Yun-nan: 24. 1916. —TYPE: CHINA. Yunnan: Haies, brousse, paturages des mont. a Ta-Hai, 3,200 m, Jul 1911, E. E. Mai-re s.n.
(holotype: E-00265629!).
Codonopsis retroserrata Z. T. Wang & G. J. Xu, Acta
Phytotax. Sin. 31: 186. 1993.—TYPE: CHINA.
Sichuan: Yan yuan, 2,400 m, 5 Sep 1986, Z. T. Wang
869382 (holotype: CPU; isotype: PE).
Icon. Wang and Xu (1993), fig. 2 [as C. retroserrata].
Distribution. Endemic to south-central China, in north­western Yunnan and southwestern Sichuan. Meadows and
thickets, 2,400-3,200 m.
Discussion. Codonopsis mairei was not mentioned by Grey-Wilson (1995). Most authors have treated it as a heterotypic synonym of C. forrestii (Anthony, 1926; Bal-
lard, 1939; Shen et al., 1975; Chamberlain, 1977; Shen and Hong, 1983; Huang, 1991; Hong and Ma, 1992; Lam-
mers, 2007b). It does resemble that species vegetatively, but differs in its smaller flowers: the hypanthium is just 5-7 x 4-7 mm (vs. 9-17 x 8-13 mm in C. forrestii), the calyx lobes are just 8-12 mm long (vs. 12-24 mm), and the corolla lobes are 22-28 x 8-15 mm (vs. 30-50 x 10-20 mm). In addition, the leaves of C. mairei tend to be more ovate, more acuminate at the apex and more rounded at the base, with somewhat shorter petioles. In all these features, C. mairei more nearly resembles C. convolvulacea, from
LAMMERS and KLEIN ― A synopsis of Codonopsis subg. Pseudocodonopsis                                                                             557
which it can be distinguished by the characters given in the key.
Although we have not able to examine type material of C. retroserrata, its original description conforms completely with the type of C. mairei and the illustration that accompanies the protologue almost seems as though it could have been modeled on the same specimen. It may have been the fact that C. mairei was hiding in the synonymy of C. forrestii at that time that led Wang and Xu to conclude that they had an undescribed species in hand.
Distribution. Not known from nature, described from cultivated material.
Discussion. In his synopsis, Grey-Wilson (1995) point­ed out that at least some of the material historically culti­vated as C. forrestii (or C. convolvulacea var. forrestii) did not in fact conform to the characterization of that taxon. Specifically, he pointed to the specimen upon which Bal­lard's (1939) plate was based. This plant differed from typical C. forrestii in its ovate (vs. linear-lanceolate to ovate-lanceolate) leaves, 33-51 mm wide (vs. 7-35 mm wide), and subcordate (vs. cuneate or rounded) at base; hypanthium 6-9 (vs. 9-17) mm long; calyx lobes triangular (vs. lanceolate or elliptic); and corolla lobes 26-28 (vs. 30­50) mm long, light lavender blue with darker veins and a dark reddish purple transverse band at base (vs. blue to lavender blue and unmarked). Grey-Wilson (1995: 219) stated that this plant "does not match anything collected in the field and may represent another new species." He de­clined to christen it, however, citing the need for additional study.
Although no additional material has come to hand in the past 15 years, it is our opinion that the best way to make collectors aware of this species is to name it formally. If the plant has a name, it can be included in Flora of China and other regional reference works, thus increasing the chance that it will be rediscovered in its homeland.
As to where that homeland might be, little can be said for
certain. On 19 September 1922, A. K. Bulley (the founder and owner of Bee's Seeds in Neston, Cheshire) exhibited before the Floral Committee of the Royal Hor­ticultural Society a plant he identified as "C. tibetica", which he had grown from material collected in nature by George Forrest (Anonymous, 1922, 1923). On 15 August 1925, Director John W. Besant reported that the National Botanic Gardens at Glasnevin, Ireland, had C. forrestii in bloom, courtesy of A. K. Bulley (Besant, 1925); how­ever, he made no mention of "C. tibetica" or the Award of Merit. The specimen upon which Ballard's (1939) plate was based was sent from Glasnevin to Kew in 1937 by Di­rector Besant, who was said to have obtained it from Bul-ley. Ballard's (1939) comments regarding the culture at Glasnevin of the plant shown in his plate very closely par­allel Besant's (1925) comments on the same topic. This permits us to infer continuity between Besant's plant and Ballard's. Furthermore, to judge from the published pho­tographs and descriptions, Bulley's "C. tibetica" of 1922 is identical to Besant's C. forrestii of 1925 and Ballard's C. convolvulacea var. forrestii of 1937 (cf. Fletcher, 1937, 1949). If Bulley had accurate information on the source of the material he grew, the fact that he coined the epithet "ti-betica" for it could be an indication that the homeland of C. macrophylla lies somewhere in that region.
Grey-Wilson (1995) was unaware of the survival of any living plants of this species, nor have we located any. The most recent indication of its survival that we have seen is the photograph published by Finlay (1972), showing plants in cultivation at Keillour in Scotland. They are a match
6. Codonopsis macrophylla Lammers & L. L. Klein, sp.
nov.—TYPE: IRELAND. Dublin, "Cult. Glasnevin Bot. Gard., Figured Bot. Mag. t. 9581," 11 Aug 1937,
"presented on behalf of the Royal Horticultural Society by the Editor of the Botanical Magazine" (holotype: K!). Figure 1
"Codonopsis tibetica" Anonymous, Gard. Chron. (ser.
3) 72: 184. 1922. Nom. invalid. sub Art. 32.3. Name
applied to a living plant "collected by Forrest. Shown by Mr. A. K. Bulley, Ness, Neston, Cheshire," which received an Award of Merit from the Floral Committee of the Royal Horticultural Society.
Planta culta patriae ignotae a ceteras speciebus Codonopsidis subg. Pseudocodonopsidis lamina ovata 5.5-7.5 cm longa et 3.3-5.1 cm lata cum apice obtuso, hy-panthio 9-13 mm diametro, et corolla lazulina cum annulo atrovinoso prope basin, corollae lobis ellipticis obtusis 15-18 mm latis distinguenda.
Stems twining counter-clockwise throughout their length, sparingly branched, 0.3-1.5 m long, suffused pur­ple, glaucous. Leaves evenly distributed along stems, gla­brous; lamina coriaceous, ovate, 5.5-7.5 cm long, 3.3-5.1 cm wide, adaxially dark green, abaxially whitish green; apex obtuse; base subcordate; margin denticulate and sparsely ciliate, dark purple, flat; petiole 4.5-5.5 mm long. Flowers solitary and terminal on main stems and branches, the portion above the last leaf 9-14 cm long. Hypanthium obconic, 6-9 mm long, 9-13 mm diam., glabrous. Calyx lobes triangular, 9-16 mm long, 4.5-6 mm wide, glabrous, in bud coherent and enveloping the corolla; apex acumi­nate or acute; margin entire; sinus acute. Corolla rotate, light lavender-blue with darker veins and a dark reddish purple ring at base; lobes elliptic, 2.6-2.8 cm long, 1.5-1.8 cm wide, connate for only 2-3 mm at base, the apex obtuse and subapiculate. Filaments 4-5 mm long, dilated into a broad triangular base, pilose on inner surface and margin and appressed to the top of the ovary, abruptly narrowed and reflexed at apex; anthers oblong, 4-6 mm long, coher­ent. Style 4 mm long. Seeds oblong, somewhat com­pressed, 1.5 x 0.7 mm, smooth, shining chestnut-brown.
Icones. Figure 1; Anonymous (1922), fig. 78 [as "C. tibetica"]; Besant (1925), fig. 46 [as C. forrestii]; Bal-lard (1939), tab. 9581 [as C. convolvulacea var. forrestii]; Finlay (1972), fig. 37 [as C. convolvulacea var. forrestii];
Grey-Wilson (1990), pg. 80 [as C. forrestii].
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for Bulley's "C. tibetica" and Ballard's plate, and could conceivably stem from the same original introduction.
7. Codonopsis vinciflora Kom., Trudy Imp. S.-Peter-burgsk. Bot. Sada 29: 103. 1908. Codonopsis convol-vulacea var. vinciflora (Kom.) L. T. Shen, Fl. Reipubl. Popularis Sin. 73(2): 68. 1983. Codonopsis convolvu-lacea subsp. vinciflora (Kom.) D. Y. Hong in C. Y. Wu, Fl. Xizang. 4: 582. 1985.—TYPE: CHINA. Sichuan: "inter Va-sy-kou et Za-li in valle fl. Siao-tzin-che," 15 Jul 1893, G. N. Potanin s.n. (Syntype: LE). A specimen grown in the botanical garden at St. Petersburg from seed collected by Potanin was also cited (cf. Fletcher 1949).
Icones: Komarov (1908), tab. II.IV; Wilkie (1937), fig.
XXI; Fletcher (1949), tab 59; Finlay (1972), fig. 35; Grey-
Wilson (1990), pg. 97; Grey-Wilson (1995), pg. 221 F-K, 223.
Distribution: Endemic to south-central China, in northwestern Yunnan, southwestern Sichuan, and southeastern Tibet (Xizang). Thickets and forests, 1,800­4,600 m.
Discussion. Codonopsis vinciflora has been referred to C. convolvulacea as a heterotypic synonym (Chipp, 1908), variety (Shen and Hong, 1983; Huang, 1991), or subspecies (Hong, 1985; Hong and Ma, 1992). However, as noted by Grey-Wilson (1995), C. vinciflora consistently differs from that species in its wider leaves (ovate or del-tate vs. linear lanceolate or lanceolate) of thinner texture with more obtuse apex and more prominently toothed
Figure 1. Codonopsis macrophylla Lammers & L. L. Klein (from the holotype).
LAMMERS and KLEIN ― A synopsis of Codonopsis subg. Pseudocodonopsis
559
margin, in its much longer petioles (10-13 vs. 1-3 mm), and its shorter calyx lobes (5-8 vs. 6-13 mm) that do not envelop the flower in bud. We therefore follow Anthony (1926) and several subsequent authors (Nannfeldt, 1936; Wilkie, 1937; Fletcher, 1949; Finlay, 1972; Gray-Wilson, 1990, 1995; Lammers, 2007b) who recognized this taxon at specific rank.
Codonopsis forrestii var. heterophylla has been treated as a heterotypic synonym of C. vinciflora (Shen and Hong, 1983; Huang, 1991; Hong and Ma, 1992). As discussed above, its type specimen is better referred to C. convolvu-lacea.
nfeldt, 1936), variety (Finlay, 1972; Shen and Hong, 1983; Huang, 1991), or subspecies (Hong and Ma, 1992). However, C. forrestii differs from that species in its larger leaves (18-46 x 2-14 vs. 25-105 x 7-35 mm) on longer (3-13 vs. 1-3 mm) petioles and in its larger flowers: hy-panthium 9-17 (vs. 3-5) mm long, calyx lobes 12-24 (vs. 6-13) mm long, and corolla lobes 30-50 (vs. 14-22 mm long). We therefore follow Anthony (1926) and several subsequent authors (Tsoong, 1935; Gray-Wilson, 1990, 1995; Lammers 2007b) who recognized this taxon at spe-cific rank.
Codonopsis forrestii var. hirsuta differs only in some minor pubescence on the abaxial leaf surface. All sub­sequent authors (Shen and Hong, 1983; Huang, 1991; Hong and Ma, 1992; Lammers, 2007b) have included it under this species as a heterotypic synonym. Codonopsis forrestii var. heterophylla has also been included here as a heterotypic synonym (Lammers, 2007b), but as noted above, its type is indistinguishable from C. convolvulacea.
Codonopsis mairei was treated as a heterotypic syn­onym of C. forrestii by numerous authors (Anthony, 1926;
Ballard, 1939; Shen et al., 1975; Chamberlain, 1977;
Shen and Hong, 1983; Huang, 1991; Hong and Ma, 1992; Lammers, 2007b); it is here recognized as distinct for the reasons given above. In a similar vein, the plants here described as C. macrophylla were long identified as this
taxon (Besant, 1925; Fletcher, 1937, 1949; Ballard, 1939; Finlay, 1972; Grey-Wilson, 1990).
8. Codonopsis efilamentosa W. W. Sm., Notes Roy. Bot. Gard. Edinburgh 8: 107. 1913. Codonopsis convolvulacea var. efilamentosa (W. W. Sm.) L. T.
Shen, Fl. Reipubl. Popularis Sin. 73(2): 69. 1983.—
TYPE: CHINA. Yunnan: Eastern flank of the Lichiang Range, 27°30' N, on scrub in dry situations, 10,000 ft,
Jul 1910, G. Forrest 6258 (holotype: E-00265632!).
Distribution. Endemic to northwestern Yunnan and ad­jacent Burma (Myanmar).
Discussion. Codonopsis efilamentosa has been treated as a variety of C. convolvulacea by Shen and Hong (1983) and Huang (1991). However, C. efilamentosa consistently differs from that species in its clustered axillary (vs. soli­tary terminal) flowers, wider leaves (15-33 vs. 2-14 mm) on longer petioles (5-22 vs. 1-3 mm), longer hypanthium (6-10 vs. 3-5 mm), and oblong corolla lobes rounded at apex (vs. elliptic and acute or obtuse). We therefore follow Anthony (1926) and several subsequent authors (Tsoong, 1935; Nannfeldt, 1936; Fletcher, 1937; Gray-Wilson, 1990, 1995; Lammers, 2007b) who recognized this taxon at specific rank.
10. Codonopsis grey-wilsonii J. M. H. Shaw, New Plants-man 3: 93. 1996. Codonopsis nepalensis Grey-Wilson, Plantsman 12: 99. 1990. Nom. illeg. sub Art. 53.1. Non Codonopsis nepalensis H. Hara, J. Jap. Bot. 53:
139. 1978.—TYPE: NEPAL. Kali Gandaki Valley, W.
of Marpha, woodland fringes, ca. 3,500 m, Sep 1973, C. Grey-Wilson & Phillips 777 (holotype: K; isotype:
K).
Icones. Prain (1908) tab. 8178 [as C. convolvulacea];
Grey-Wilson (1990), pg. 87; Grey-Wilson (1995), pg. 215
[as C. nepalensis 'Himal Snow'], pg. 221 A-E [as C. nep­alensis], pg. 222 [as C. nepalensis 'Himal Snow'].
Distribution. Endemic to the eastern Himalaya. Shrubberies and forest margins, 2,400-3,600 m.
Discussion. The Himalayan representative of C. subg. Pseudocodonopsis has been known for over a century, but until Grey-Wilson (1990) pointed out its distinguishing features, it had been identified as C. convolvulacea (Prain, 1908; Finlay, 1972; Hara, 1982) or C. vinciflora (Fletcher, 1949). Codonopsis grey-wilsonii differs from both those species by its longer (3-13 vs. 1-3 mm) petioles and larger flowers, with corolla lobes 30-45 (vs. 14-23) mm long and pubescent (vs. glabrous) on the lower third internally. The only other member of the subgenus with flowers so large is C. forrestii (with which it has sometimes been confused; cf. Grey-Wilson, 1995); these two species may be distin­guished by the characters given in the key above.
9. Codonopsis forrestii Diels, Notes Roy. Bot. Gard. Edinburgh 5: 171. 1912. Codonopsis convolvulacea
var. forrestii (Diels) Ballard, Bot. Mag. 162: tab. 9581.
1939. Codonopsis convolvulacea subsp. forrestii (Diels) D. Y. Hong & L. M. Ma in H. W. Li & Z. Y.
Zhu, Fl. Sichuanica 10: 546. 1992.—TYPE: CHINA.
Yunnan: common on moist places on the banks of
the Yangtse, 5,000-7,000 ft, Sep 1907, G. Forrest 48 (holotype: E-00265630!).
Codonopsis forrestii var. hirsuta P. C. Tsoong & L. T.
Shen, Acta Phytotax. Sin. 13(3): 55. 1975.—TYPE: CHINA. Sichuan: "Wobosego."
Icones. Grey-Wilson (1995), pg. 218, pg. 221 L-Q. Other illustrations purporting to show this taxon (Besant,
1925; Ballard, 1939; Finlay, 1972; Grey-Wilson, 1990) ac­tually depict C. macrophylla.
Distribution. Endemic to south-central China, in north­ern Yunnan, southwestern Sichuan, and western Guizhou. Open woods and thickets, 1,800-3,900 m.
Discussion. Codonopsis forrestii has been referred to C. convolvulacea as a heterotypic synonym (Nan-
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Acknowledgments. Support for our research on the Flora of China during 2008-2009 was provided by the National Science Foundation via ROA Supplement DEB-0810237 (Ihsan Al-Shehbaz, Principal Investigator). We thank Alex Sanford, a recent graduate of the University of Wisconsin Oshkosh, for expertly producing the illustration of the new species; and the authorities at Kew and Edinburgh for the loan of relevant herbarium specimens.
Press, Beijing, pp. 515-586.
Huang, S. H. 1991. Campanulaceae nom. conserv. In C.Y. Wu & J. Chen (eds.), Flora Yunnanica, vol. 5. Science Press,
Beijing, pp. 452-509.
Huxley, A.J. (ed.) 1992. Codonopsis Wallich. In The New Royal Horticultural Society Dictionary of Gardening, vol. 1. MacMillan, London, pp. 666-668.
Komarov, V. L. 1908. Revisio critica specierum generis Codonopsidis Wall. Trudy Imp. S.-Peterburgsk. Bot. Sada 29: 98-124 + tab. II.
Lammers, T.G. 2007a. Campanulaceae. In K. Kubitzki (ed.), The Families and Genera of Vascular Plants, Vol. 8, Asteridae. Springer-Verlag, Berlin, pp. 26-56.
Lammers, T.G. 2007b. World Checklist and Bibliography of Campanulaceae. Royal Botanic Gardens, Kew.
Limpricht, W. 1922. Botanische Reisen in den Hochgebirgen Chinas und Ost-Tibets. Repert. Spec. Nov. Regni Veg.,
Beih. 12: i-viii, 1-515, pl. I-XVI, maps 1-9.
Morris, K.E. and T.G. Lammers. 1997. Circumscription of Codonopsis and the allied genera Campanumoea and Leptocodon (Campanulaceae: Campanuloideae). I.
Palynological data. Bot. Bull. Acad. Sin. 38: 277-284.
Nannfeldt, J.A. 1936. Campanulaceae. In H. Handel-Mazzeti, Symbolae Sinicae, vol. 7. Julius Springer, Vienna, pp.
1067-1084.
Oliver, D. 1895. Codonopsis convolvnlacga, Kurz. Hooker's Icon. Pl. 24: pl. 2385 + 1 pg.
Prain, D. 1908. Codonopsis convolvulacea. Bot. Mag. 134: tab. 8178 + 3 pp.
Shen, L.D., S.Y. Tang, S.G. Yuen, and C.K. Hsieh. 1975. The
genus Codonopsis Wall. in Szechuan. Acta Phytotax. Sin. 13(3): 51-68.
Shen, L.D. and D.Y. Hong. 1983. Codonopsis Wall. In D. Y.
Hong (ed.), Flora Reipublicae Popularis Sinicae, Vol. 73, No. 2. Science Press, Beijing, pp. 32-69.
Tsoong, P.C. 1935. Preliminary study on Chinese Campanulaceae. Contrib. Inst. Bot. Natl. Acad. Peiping 3: 61-118 + pl. IX-XIV.
Wang, Z.T. and G.J. Xu. 1993. Two new species of Codonopsis
from China. Acta Phytotax. Sin. 31: 184-187.
Wei, Z.X. 2001. Pollen morphology of some Codonopsis and relafed [sic] species of Campanulaceae. Acta Bot. Yuanica 23: 335-338 + pl. I-II.
Wilkie, D. 1937. New and interesting plants. Codonopsis vinciflora (Kom [sic]). New Fl. & Silva 10: 58-59 + fig.
XXI.
Wu, C.Y. and H.W. Li. 1965. Campanulaceae. In Yunnan Redai Yaredai Zhiwa Quxi Yanjiu Baogao. Science Press, Beijing, pp. 57-97 + pl. XXVI.3- XXXIII.1.
LITERATURE CITED
Anonymous. 1922. Societies. Royal Horticultural. Floral Committee. Awards of Merit. Gard. Chron. (ser. 3) 72: 184, 193.
Anonymous. 1923. Royal Horticultural Society. Floral Committee. Award of Merit. J. Roy. Hort. Soc. London, Proc. 48: lxxxvii.
Anthony, J. 1926. A key to the genus Codonopsis, Wall., with an account of two undescribed species. Notes Roy. Bot. Gard. Edinburgh 15: 173-190 + pl. ccxvi -ccxvii.
Ballard, F. 1939. Codonopsis convolvulacea var. forrestii. Bot. Mag. 162: tab. 9581 + 4 pp.
Besant, J.W. 1925. Notes from Glasnevin. Codonopsis forrestii.. Gard. Chron. (ser. 3) 78: 125, 128.
Chamberlain, D.F. 1977. Catalogue of the names published by Hector Leveille: X. Notes Roy. Bat. Gard. Edinburgh 35:
247-264.
Chipp, T.F. 1908. A revision of the genus Codonopsis, Wall. J.
Linn. Soc., Bot. 38: 374-391.
Cox, E.H.M. 1936. Codonopsis in cultivation. New Fl. & Silva 9: 25-30 + fig. VI-XLI.
Finlay, M.K. 1972. On Codonopsis which grow at Keillour. J. Roy. Hort. Soc. 97: 82-87 + figs 33-37.
Fletcher, H.R. 1937. The genus Codonopsis in cultivation.
Gard. Chron. (ser. 3) 102: 80-83, 120-121.
Fletcher, H.R. 1949. Codonopsis vinciflora. Bot. Mag. 166: tab. 59 + 4 pp.
Grey-Wilson, C. 1990. A survey of Codonopsis in cultivation.
Plantsman 12(2): 65-99.
Grey-Wilson, C. 1995. Codonopsis convolvulacea and its allies.
New Plantsman 2: 213-225.
Hara, H. 1982. Campanulaceae. In H. Hara, A. O. Chater and L. H. J. Williams, An Enumeration of the Flowering Plants of Nepal, vol. 3. British Museum (Natural History), London,
pp. 49-54.
Hong, D.Y. 1985. Campanulaceae. In C.Y. Wu (ed.) , Flora Xizangica, vol. 4. Science Press, Beijing, pp. 560-601.
Hong, D.Y. and L.M. Ma. 1992. Campanulaceae. In H. W. Li
and Z. Y. Zhu (eds.), Flora Sichuanica, vol. 10. Science
LAMMERS and KLEIN ― A synopsis of Codonopsis subg. Pseudocodonopsis
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黨參屬輻冠黨參亞屬（桔梗科桔梗亞科）之分類大綱，兼報導
產地不詳的一新種：闊葉黨參
Thomas G. LAMMERS and Laura L. KLEIN
Department of Biology and Microbiology, University of Wisconsin Oshkosh, Oshkosh, WI 54901, USA
黨參屬輻冠黨參亞屬的物種爲無氣味的草本地下芽植物，具有不分枝且長度小於直徑四倍長的塊
根；纏繞莖至少靠近基部具有多枚葉子；花通常單朵頂生，其花冠爲輻狀且深裂超過其長度的四分之
三；花粉粒5-10溝，溝的長度短於花粉兩極距離的四分之一，表面具有2-3 長的小突刺；子房三
室；果實爲倒錐形蒴果，先端具有短開裂口 ，內含許多具細紋的種子。輻冠黨參亞屬最近的分類處理包
含了雞蛋參（C. convolvulacea '本文指定爲此亞屬的模式種)、心葉珠子參(C. efilamentosa)、珠子參(C.
forrestii)、尼泊爾黨參(C. grey-wilsonii ；原用C. nepalensis爲不合法名)、直立雞蛋參(C. limprichtii ；原
用"C. pinifolia"爲無效名）與輻冠黨參(C. vinciflora)。蓮座狀黨參(C. rosulata)雖然莖不具纏繞性且具
蓮座狀基生葉，亦爲輻冠黨參亞屬的成員。毛葉雞蛋參（C. hirsuta)之前被倂入雞蛋參（C. convolvulacea)
之中，但本文處理認爲是與直立雞蛋參（C. limprichtii)較近緣的獨立物種。梅氏黨參（C. mairei)之前被
倂入珠子參(C. forrestii)但本硏究認爲是與雞蛋參（C. convolvulacea)近緣的獨立物種。過往有一栽培植
物被鑑定爲珠子參(C. forrestii)，但是有一些特徵與其他所有已知的種都不相同，本文發表爲新種-闊葉
黨參（C. macrophylla, sp. nov.)。雲南爲輻冠黨參亞屬的多樣性中心，孕育了十種之中的八種，東喜馬拉
雅特有的尼泊爾黨參（C. grey-wilsonii)及產地不明的闊葉黨參（C. macrophylla)除外；若干種類的地理分
布由雲南延伸至四川、貴州、西藏及/或緬甸。
關鍵詞：亞洲；桔梗科；桔梗亞科；黨參屬；闊葉黨參；新種；輻冠黨參亞屬；分類學。