Nordic Journal of Botany 28: 656 660, 2010 doi: 10.1111/j.1756-1051.2010.00872.x, # 2010 The Author. Nordic Journal of Botany # 2010 Nordic Society Oikos Subject Editor: Martin Cheek. Accepted 5 November 2010
Three new species of Dactylicapnos (Fumariaceae) and a synopsis of the D. macrocapnos complex
Magnus Lide´n
Magnus Lide´n ([email protected]), Uppsala Univ. Botanic Gardens, Villava¨gen 8, SE-75236 Uppsala, Sweden.
The Dactylicapnos macrocapnos complex is revised, and D. platycarpa Lide´n, D. odontocarpa Lide´n and D. macrocapnos subsp. echinosperma Lide´n are recognised as new taxa. The complex consists of a chain of 4 vicariant PECIAL ISSUE taxa from northwestern India (Garhwal) to western Bhutan (Thimphu). Dactylicapnos cordata Lide´n (eastern Nepal,
S Darjeeling) is described and contrasted with its close relative, the geographically disjunct D. burmanica (western Yunnan, Burma).
Dactylicapnos Wallich is a very natural and easily circum- flowers are held apex upwards, although they are actually scribed genus, recognised by i.a. the scandent habit with pendent. the terminal leaflet (except of the basalmost leaves)
AXONOMY transformed into a branched tendril,9 corymbose pendent T inflorescences, yellow bisymmetric flowers and unique The Dactylicapnos macrocapnos complex stigma, stamen and seed structures. It has at times been (Fig. 1) included in Dicentra Bernh. (Khanh 1972), solely based on the bisymmetric flowers which is a plesiomorphy at family
LANT The Dactylicapnos scandens group is characterised by having level. Morphological and molecular data show that P two alternate lateral primary leaflets (opposite in other Dactylicapnos is not closely related to Dicentra, but rather perennial species of the genus) and comparatively thin and the sister group to the genus Corydalis DC (Lide´n et al. soft leaflets (Lide´n 2008). The variable species D. scandens 1997). (D. Don) Hutchinson is vegetatively similar to the Dactylicapnos is distributed in the Sino-Himalayan area D. macrocapnos complex. The latter is nevertheless easily from Garhwal to southern China with some annual distinguished from D. scandens when flowering or fruiting species extending north as far as Sichuan. There are 15 or ASCULAR 16 species in two sections: sect. Dactylicapnos (perennials) by the slightly oblique inflorescence with basally slightly reflexed (vs not reflexed) pedicels, sepals that are abruptly V and sect. Minicalcara (annuals) (Khanh 1972, Lide´n 2007). broadened at the base (vs lanceolate), the strongly up- The perennial species (sect. Dactylicapnos) fall into four curved nectary with long thin apex and usually with a groups: a) the D. grandifoliolata group (four spp.) swelling or wart 1/3 to 2/3 from the base (vs short sigmoid characterised by two opposite primary leaflets with few evenly thick, only shortly acute, not up-curved), lanceolate SIATIC firm strongly veined ultimate leaflets, fleshy fruits opening flat membranous green capsules (vs [when fresh] ovoid
A with valves, persistent placentas with tooth-like funiculae, thick fleshy, red yellow or purple), and low broad flat very large easily caducous elaiosomes made up of funiculi (vs thick annular). free tubular cells, and a branched narrow rhizome; Dactylicapnos scandens s.l. is found from central Nepal to b) D. schneideri with two pairs of opposite primary leaflets, southern China (Guizhou) whereas the Dactylicapnos narrow fruits with persistent spongy placentas and seeds macrocapnos complex has an overlapping, but more western with small elaiosomes; c) the D. scandens group (four or distribution from northwestern India (Garhwal) to western five spp, incl. the D. macrocapnos complex, the main Bhutan. subject of this paper); d) the D. burmanica group with two There is a clearcut geograpical variation within the species (one described here). D. macrocapnos complex, with four vicariant taxa along The analysis presented here is based on material at Himalaya. This pattern is not recognised in the excellent ASSAM, BM, CAL, E, K, KATH, PE, TI and UPS. Note monograph by Khanh (1972), as it was revealed by recent that, in the description of floral parts, it is assumed that the collections from eastern Nepal, India and Bhutan.
656 A SIATIC
Figure 1. Fruits, seeds, stigmas and flowers of the Dactylicapnos macrocapnos complex (note that flowers and fruits are actually pendent, V but here illustrated with the pedicel attachment at page bottom). (A) D. macrocapnos supsp. macrocapnos (fruit and seed), (B) D. ASCULAR macrocapnos subsp. echinosperma, (C) D. odontocarpa (fruit, seed and stigma), (D) D. platycarpa.
Key to the Dactylicapnos macrocapnos complex usually with a median wart below. Capsule narrowly lanceolate, flattened, 25 35 6 7 mm; style 5 6 mm. 1. Margin of fruit grossly dentate; seeds faintly colliculate, Seeds colliculate to aculeate, 1.6 1.8 mm; elaiosome small.
1.2mm ...... D. odontocarpa Flowering time: Apr Nov, fruiting time: Jul Nov. P
Margin of fruit even; seeds sharply colliculate to Ecology: climber in forest understories at an altitudinal LANT aculeate, 1.6 1.8mm...... 2 range of 900 2700 m a.s.l. 2. Capsule twice as long as broad ...... D. platycarpa Capsule 3.5 5.0 times as long as broad ...... 3 3. Seeds spinulose/aculeate . . . . D. macrocapnos subsp. Dactylicapnos macrocapnos subsp. macrocapnos T echinosperma Margin of fruit usually without or with indistinct rim; seeds AXONOMY Seeds colliculate, but not spinulose . . . . D. macro- 1.6 1.7 mm, sharply colliculate. capnos subsp. macrocapnos Distributed in Garhwal, Kumaon and extreme west of Nepal.
Dactylicapnos macrocapnos (Prain) Hutchinson (1921, p. 105). Dactylicapnos macrocapnos subsp. echinosperma Lide´n
subsp. nov. S
Basionym: Dicentra macrocapnos Prain (1896, p. 12). Type: A D. macrocapno subsp. macrocapno seminibus aculeatis ISSUE PECIAL India, Garhwal, above Ghat, 2100 2400 m a.s.l., 20 Sep differt. 1885, J. F. Duthie s.n. (lectotype: DD, isotype: LE; fide Khanh 1972). Type: Nepal, Bakhri Kharka north of Pokhara, 1700 m a.s.l., twining amoung bushes on open slope, ‘‘flowers Perennial with stout subterranean rootstock. Stems 2 8m, yellow’’, 23 Apr 1954, Stainton, Sykes and Williams 5009 thin and weak, more or less terete (appearing slightly sulcate (holotype: BM!, isotypes: E!, GH!, UPS!). in the dry state). Leaves slightly reflexed on zigzag stem; petiole 0.5 3.0 cm; blade glaucous below, green above, with Margin of fruit usually with distinct rim; seeds 1.6 1.8 1 pair of alternate primary leaflets; these bi- (rarely mm, spinulose. It inhabits central Nepal and southern Tibet tri-)ternate; ultimate leaflets ovate, 10 25 7 20 mm. (Nyalam). Raceme 5 14-flowered, pendent, usually slightly obliquely so; peduncle 2 4 cm; bracts 3 10 1 2 mm, entire, sub- acute. Pedicels 10 20 mm, in fruit to 30 mm, reflexed at Additional specimen examined (selection of paratypes) base, sometimes partly fused to subtending bracts. Sepals Tibet: Nyalam, near Zhangmu, 2300 m a.s.l., 13 Aug 1972, 2 3 2.0 2.5 mm, abruptly tapering to narrow apex from unknown 1060 (PE); Zhangmu, 2700 m a.s.l., 24 Aug broad base, slightly dentate. Corolla yellow, often turning 1972, unknown 1472 (PE); Nepal: Lumle west Pokhara, reddish with age, oblong-cordate to obtusely triangular in 1500 m a.s.l., 2 Nov 1954, Stainton et al. 8279 (BM); profile, 18 20 8 12 mm; outer petals strongly keeled or central Nepal, cultivated in Uppsala Univ. Bot. Gard. Aug usually winged at base; sinus at petiole insertion 3 mm deep; 2007, Lide´n s.n. (UPS). dorsal crest of inner petals overtopping apex 1( 2) mm; This taxon is available from several garden nurseries, nectary strongly curved upwards, tapering to slender apex, often under the name ‘‘Dicentra scandens’’ (Lide´n 2008).
657 Dactylicapnos odontocarpa Lide´n sp. nov. insertion 3 4 mm deep. Inner petals: claw 10 13 mm; blade ca 5 mm; dorsal crest overtopping apex 1 mm. A D. macrocapno capsulis irregulariter grosse dentatis semi- Nectary tapering to curved apex, usually with prominent nibus multo minoribus leviter colliculatis differt. gibbosity or peg 1/3 1/2 as long as distal part of nectary (in D. macrocapnos up to 1/5). Stigma trapezoidal (in D. Type: east Nepal, Janakpur zone (Ramechhap Distr.); macrocapnos subsquarish), with small but distinct papillae in Sagarmatha zone (Solukhumbu Distr.), Bhandar (2300 m apical corners and larger papillae in basal corners. Ovary a.s.l.); Kinja (1650 m a.s.l.); Sheptom (2050 m a.s.l.); with ca 100 ovules, apparently more thick-walled than in D. Nemar (2400 m a.s.l.); Sete (2550 m a.s.l.), in shrubberies, macrocapnos. Capsule broadly elliptic-lanceolate, 22 23 ‘‘flowers yellow’’, 18 Aug 1985, H. Ohba, M. Wakabaya- 10 11 mm (in D. macrocapnos 25 35 6 7 mm); style 7 9 shi, M. Suzuki, N. Kurosaki, K. R. Rajbandhari and S. K. mm. Seeds strongly colliculate, ca 1.6 mm; elaiosome small. Wu 8571764 (holotype: E!, isotypes: KATH, TI).
Climbing perennial herbs, with stout subterranean root- Additional specimens examined (paratypes) stock. Stems 2 8 m (?), thin. Petioles 0.5 3.0 cm. Leaves Bhutan: Thimphu district, above end of road along with 1 pair of alternate primary leaflets; primary leaflets Thimphu Chu just below Changri monastery, 2500 m biternate; ultimate leaflets 5 9 per primary leaflet, ovate, a.s.l., 9 Aug 1987, J. R. I. Wood 5675 (E); Thimphu subobtuse, 10 20 7 15 mm, glaucous below. Raceme 1 district, J. R. I. Wood 6091 (E); Chendebi, among shrubs 3 cm, 5 8-flowered, pendent. Peduncle 2 4 cm. Bracts on cliff face, 2700 m a.s.l., 5 Aug 1949, Ludlow, Sherriff linear to narrowly oblanceolate, ca 5 1 mm, entire, and Hicks 19580 (E, BM, UPS). India: Darjeeling, Senchal subacute. Pedicels 10*20 mm, reflexed at base. Sepals 2 WLS, way to Tung, 14 Oct 2008, A. K. Ghosh 41501 3 2.0 2.5 mm, abruptly tapering to narrow apex from (CAL). PECIAL ISSUE broad base, slightly dentate. Corolla yellow, oblong-cordate S to obtusely triangular, 18 20 8 12 mm. Outer petals keeled or winged at base; sinus at petiole insertion narrow, The Dactylicapnos burmanica group (Fig. 2) 3 mm deep. Inner petals claw 11 mm, blade 5.5 mm; dorsal crest overtopping apex 1( 2) mm. Nectary 4 5 mm, The D. burmanica group is characterised by very short- slightly curved upwards, tapering to slender apex, with a stalked rather thin leaves with two long-stalked opposite swelling 1/3 1/2 from base. Capsule lanceolate, flattened, twice ternate primary leaflets, long narrow thin-walled 30 35 6 8 mm, with thin valves; style 5 6 mm. Seeds fruits, dorsal median crests of inner petals protruding 3 4
AXONOMY faintly colliculate, 1.2 mm, with small elaiosome. mm beyond the apical junction of the petals (1 2mmin T Similar to D. macrocapnos, but capsule with distinct other species) and bifid nectaries. Further, it has a swollen grossly and irregularly dentate rim, seeds small, 1.1 1.2 mm, carrot-like root-stock (only known for D. burmanica, only faintly colliculate, and nectary with inconspicuous though), like the D. scandens group. It has a disjunct lateral wart. distribution with one species in eastern Nepal and Darjeel- LANT ing, the other in western Yunnan and northern Burma. For P differences between the species see Table 1. Additional specimen examined (paratype) India: Sikkim, Lachung, 8500 feet, 31 Aug 1892, G. A. Gammie 1077 (CAL). Dactylicapnos cordata Lide´n sp. nov. A D. burmanica floribus brevioribus latioribus, nectariis non
ASCULAR Dactylicapnos platycarpa Lide´n sp. nov. reflexis inaequaliter bifidis, stylo breviore, seminibus multo minoribus differt. V A D. macrocapno capsulis brevioribus subduplo latioribus differt. Type: Nepal: Tamur river near Chirwa, 27829’N, 87846’E, 1350 m a.s.l., among rocks in broad-leaved forest, Type: Bhutan, Thimphu district, Dotena, Thimphu Chu, scrambling herb, flowers pale orange, sometimes with
SIATIC 27835’N, 89838’E, amongst shrubs on river bank, 2550 m brown tips to petals, 3 Sep 1989, Kew Edinburgh a.s.l., 5 Sep 1984, I. W. J. Sinclair and D. G. Long 4833 Kathmandu expedition to eastern Nepal (KEKE) 189 A (holotype: E!). (holotype: E!, isotypes: K, KATH).
Herb [perennial], climber. Stems 3 4 m, thin. Leaves9 Root-stock unknown (in D. burmanica carrot-shaped). reflexed on zigzag stem; petiole 5 15 mm; blade glaucous Stem climbing on shrubs, thin (1 2 mm), leafy and below, dark green above, with two alternate primary pinnae. branched throughout. Petioles 5 10 mm; petiolules of Each primary leaflet with petiolule 15 25 mm; blade bi- primary leaflets 2 4( 5) cm, thin; primary leaflets twice (rarely tri-)ternate; ultimate leaflets ovate, 10 25 5 15 ternately compound, each with 6 to 11 very unequally sized mm. Raceme 5 10-flowered, pendent; peduncle 2 5 cm; ultimate leaflets; these ovate, 10 40 7 30 mm, soft and bracts 4 7 1 mm, entire, acuminate. Pedicels 10 25 mm, thin, sub-obtuse with very small recurved mucro. Racemes in fruit to 30 mm. Sepals 3 2 mm, abruptly narrowed nutant, ca 1 cm, 4 7-flowered; peduncle 2 5 cm. Bracts from broad base to narrow apex, slightly dentate. Corolla linear-lanceolate, 3 4 mm. Pedicels thin, 10 25 mm, 20 yellow, oblong-cordate, 20 23 9 12 mm; outer petals 30 mm in fruit. Sepals triangular, subentire, 2 3 1.0 1.5 with prominent dorsal wing at base; sinus at petiole mm. Corolla yellow to orange-yellow, ovate-cordate in
658 A SIATIC V ASCULAR
Figure 2. Flowers, nectaries and seeds of the Dactylicapnos burmanica group (note that flowers and fruits are actually pendent, but here illustrated with the pedicel attachment at page bottom). A) Dac. burmanica (also fruit); B) Dac. cordata. profile, 16 23 9 16 mm; outer petals broadly saccate, (basal part of the flower very narrow), and petiolules are usually with a dorsal wing at base; sinus at petiole insertion shorter than in Nepalese specimens). P
3 5 mm deep; dorsal crest of inner petals conspicuous, LANT overtopping petal apex 3 mm; nectaries directed out-wards, rather thick, unequally bifid, outer (longer) branch slightly Dactylicapnos burmanica (K. R. Stern) Lide´n (2007, upcurved. Stigma longer than broad, with rather small p. 35) papillae. Capsule thin-walled with ca 30 ovules, lanceolate, T AXONOMY 25 33 5 6 mm excl. persistent style 3 5 mm. Seeds Basionym: Dicentra burmanica K. R. Stern (1967, p. 280). (immature) 0.9 1.1 1.2 mm, aculeate with broad smooth Type: Burma: Kachin, N’Changyang, 25850?N, 97848?E, beak, with small elaiosome. Flowering: May Sep, fruiting: 2 000 feet (610 m a.s.l.), 12 Jul 1939, R. Kaulback 267 Sep. (holotype: BM!, isotype: E!).
Distinguished from D. cordata by longer and narrower Additional specimens examined (paratypes) flowers, longer style and larger seeds (Table 1). S 8 ? 8 ?
Nepal: Chyangtha[ng], 27 17 N, 87 56 E, 4500 feet (1370 ISSUE PECIAL m a.s.l.). On shady banks among shrubs, scandent, flowers yellow, 12 Jun 1969, Williams 586 (E). Arun valley, Sedua Additional specimens examined northwest of Num, 27835?N, 87817?E, 6000 feet (1800 m Burma: Kachin, Pooritong, flowers yellow, Sep 1899, Shaik a.s.l.), climbing on shrubs, corolla orange yellow, 16 Jun Mokim 721 (CAL); China: Yunnan, Wanding county, 1956, Stainton 689 (E). sine loco: (probably east Nepal) shady place in forest, 19 Aug 1976, Pei Shengji 14056 V. Puri 326 (CAL). India: Darjeeling, Kurseong, May (KUN); India: Nagaland, Kohima, cultivated in gardens 1915, E. A. C. Modder 82 (CAL) (in this specimen, the one (rare), 2 290 m a.s.l., flowers yellow, tuber carrot-like, 27 Jul complete flower seems abnormal or stunted in development 2005, A. A. Mao 109227 (CAL) (according to information
Table 1. Comparison between Dactylicapnos cordata and D. burmanica.
D. burmanica D. cordata
Corolla outline oblong-cordate; more than twice as long as broad ovate-cordate; less than twice as long as broad Corolla total length (mm) 25 30 8 11 16 23 9 16 Dorsal crest of outer petals absent usually obvious at base of flower Dorsal crest of inner petals overtopping apex by 4 mm overtopping apex by 3 mm Style length (mm) 6 73 5 Seed number apparently few (ca 10?) ca 30 (?) Seed size (mm) ca 1.8 2.2 ca 0.9 1.1 (?immature) Nectary bent down-wards, equally bifid bent out-wards, unequally bifid Distribution west Yunnan, north Burma; (Nagaland) east Nepal; Darjeeling
659 on the label, the rhizome of this species is used in local Stern (Fumariaceae). Rep. Spec. Nov. Reg. Veg. 83: 511 medicine for women having ‘‘warmb’’ [?]-problems). 575. Lide´n, M. et al. 1997. Phylogeny and classification of Fumar- iaceae, with emphasis on Dicentra s.l. based on the rps16 Acknowledgements My thanks go to the herbaria listed above, intron. Plant Syst. Evol. 206: 411 420. especially to Colin Pendry and Mark Watson in Edinburgh for Lide´n, M. 2007. New species, combinations, and records of generous help and for inviting me to join the ‘Flora of Nepal’ Hypecoum, Dactylicapnos and Corydalis (Fumariaceae) in team, which instigated this revision. China. Nord. J. Bot. 25: 33 37. Lide´n, M. 2008. Climbing hearts identified. Plantsman 7: 118 121. References Prain, D. 1896. Noviclae Indicae X: Some Additional Fumar- iaceae. J. Asiat. Soc. Bengal, pt. 2, Nat. Hist. 65: 10 41. Hutchinson, J. 1921. The genera of Fumariaceae and their Stern, K. R. 1967. A new species of Dicentra from Burma. distribution. Kew Bull. Misc. Inform. 35: 97 115. Brittonia 19: 280 282. Khanh, T. C. 1972. Beitrag zur Kenntnis der Sippenstruktur des Genus Dicentra Bernh. subgenus Dactylicapnos (Wall.) K. R. PECIAL ISSUE S AXONOMY T LANT P ASCULAR V SIATIC A
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