Nordic Journal of Botany 28: 656Á660, 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, 25Á356Á7 mm; style 5Á6 mm. 1. Margin of fruit grossly dentate; seeds faintly colliculate, Seeds colliculate to aculeate, 1.6Á1.8 mm; elaiosome small. 1.2mm ................... D. odontocarpa Flowering time: AprÁNov, fruiting time: JulÁNov. P Á Margin of fruit even; seeds sharply colliculate to Ecology: climber in forest understories at an altitudinal LANT aculeate, 1.6Á1.8mm.................... 2 range of 900Á2700 m a.s.l. 2. Capsule twice as long as broad . D. platycarpa Á Capsule 3.5Á5.0 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 Á Seeds colliculate, but not spinulose . D. macro- 1.6Á1.7 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 PECIAL ISSUE India, Garhwal, above Ghat, 2100Á2400 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 2Á8m, 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 0.5Á3.0 cm; blade glaucous below, green above, with Margin of fruit usually with distinct rim; seeds 1.6Á1.8 1 pair of alternate primary leaflets; these bi- (rarely mm, spinulose. It inhabits central Nepal and southern Tibet tri-)ternate; ultimate leaflets ovate, 10Á257Á20 mm. (Nyalam). Raceme 5Á14-flowered, pendent, usually slightly obliquely so; peduncle 2Á4 cm; bracts 3Á101Á2 mm, entire, sub- acute. Pedicels 10Á20 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, 2Á32.0Á2.5 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, 18Á208Á12 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(Á2) 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 3Á4 mm deep. Inner petals: claw 10Á13 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 1/3Á1/2 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, 22Á23 ‘‘flowers yellow’’, 18 Aug 1985, H. Ohba, M. Wakabaya- 10Á11 mm (in D. macrocapnos 25Á356Á7 mm); style 7Á9 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 2Á8 m (?), thin. Petioles 0.5Á3.0 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 5Á9 per primary leaflet, ovate, a.s.l., 9 Aug 1987, J. R. I. Wood 5675 (E); Thimphu subobtuse, 10Á207Á15 mm, glaucous below. Raceme 1Á district, J. R. I. Wood 6091 (E); Chendebi, among shrubs 3 cm, 5Á8-flowered, pendent. Peduncle 2Á4 cm. Bracts on cliff face, 2700 m a.s.l., 5 Aug 1949, Ludlow, Sherriff linear to narrowly oblanceolate, ca 51 mm, entire, and Hicks 19580 (E, BM, UPS). India: Darjeeling, Senchal subacute. Pedicels 10*20 mm, reflexed at base. Sepals 2Á WLS, way to Tung, 14 Oct 2008, A. K. Ghosh 41501 32.0Á2.5 mm, abruptly tapering to narrow apex from (CAL). PECIAL ISSUE broad base, slightly dentate. Corolla yellow, oblong-cordate S to obtusely triangular, 18Á208Á12 mm. Outer petals keeled or winged at base; sinus at petiole insertion narrow, The Dactylicapnos burmanica group (Fig.