The Confusing Identity of Pternopetalum Molle (Apiaceae)

Botanical Journal of the Linnean Society, 2008, 158, 274–295. With 15 ﬁgures

The confusing identity of Pternopetalum molle (Apiaceae)

LI-SONG WANG1,2*

1State Key Laboratory of Systematics and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan, Beijing 100093, China 2Graduate School of Chinese Academy of Sciences, Beijing 100039, China

Received 25 April 2006; accepted for publication 24 June 2008

The misapplication of the name Pternopetalum molle (Franch.) Hand.-Mazz. has resulted in considerable taxonomic confusion in the genus, involving 11 names belonging to seven taxa (according to a recent treatment). A supplementary taxonomic revision of P. botrychioides (Dunn) Hand.-Mazz., P. molle and P. vulgare (Dunn) Hand.-Mazz. is presented, after a detailed examination of the morphological variation. Four new synonyms, P. longicaule var. humile R.H.Shan & F.T.Pu, P. molle var. dissectum R.H.Shan & F.T.Pu, P. radiatum (W.W.Smith) P.K.Mukherjee and P. trifoliatum R.H.Shan & F.T.Pu, are proposed. Pternopetalum delicatulum (H.Wolff) Hand.- Mazz. should be reduced to P. botrychioides rather than P. radiatum. The previous taxonomic treatments of P. cuneifolium (H. Wolff) Hand.-Mazz., P. cartilagineum C.Y.Wu ex R.H.Shan & F.T.Pu and P. molle var. crenulatum R.H.Shan & F.T.Pu are reinstated. A key and distribution maps are provided for the accepted species. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 274–295.

ADDITIONAL KEYWORDS: East Asia – morphology – taxonomic revision – Umbelliferae.

INTRODUCTION subfamily Apioideae (Pu, 1985; Pimenov & Leonov, 1993), two morphological characters, petals saccate at Pternopetalum Franch., one of the largest genera of the base and umbellules usually with only 2–3(–4) Apiaceae in Asia (Pimenov & Leonov, 2004), includes flowers (Shen et al., 1985; Pu, 2001; Pu & Phillippe, about 20–32 taxa which occur in South Korea, Japan, 2005), make it relatively easy to identify. It seems to China and the adjacent eastern Himalayan regions be a natural genus in Apiaceae (Valiejo-Roman et al., (Pu, 1985; Mukherjee & Constance, 1993; Pimenov 2002). & Leonov, 1993; Ohba, 1999; Watson, 1999; Pu & Detailed taxonomic scrutiny is hindered by inad- Phillippe, 2005), with a centre of diversity in west equate herbarium specimens and a lack of adequate Sichuan, north-west Yunnan and south-eastern Tibet field surveys, making it one of the more difficult (Shu & Sheh, 2001). As a genus endemic to the East groups in Apiaceae (M. F. Watson, Royal Botanic Asian flora (Wu, 1991; Wu et al., 2006), its phytogeog- Garden, Edinburgh, pers. comm.). Some taxa still raphy has been documented by many authors remain uncertain in the recently published Flora of (Constance, 1972; Takhtajan, 1986; Sheh & Su, 1987; China (Pu & Phillippe, 2005). The generic review Pu, 1996; Sun, 2002; Wu et al., 2003). Pternopetalum revealed that the misapplication of the name Pterno- species are mainly erect perennials with a distinct petalum molle (Franch.) Hand.-Mazz. was the great- rootstock on which the radical leaves are produced est source of confusion in this genus. each year. These plants usually grow in shady and Pternopetalum molle was first recognized by moist evergreen forest, especially in mountain areas Franchet (1894) based on Delavay, J. M. 4095 with rich streams. As part of the tribe Apieae of the (Figs 1A, 2A, B). This species is characterized by an annual habit, delicate appearance, single and fusi- *E-mail: [email protected] form root, and terminal and lateral umbels, that

Figure 1. A, Holotype of Pternopetalum molle, Delavay J. M. 4095 conserved at P. B, Pternopetalum cuneifolium, specimen referred to as P. molle in previous ﬂora treatments, Ching, R. C. 22416 conserved at PE.

Figure 2. Illustration of Pternopetalum molle: A, plant; B, fruits, illustrated from Delavay 4095 conserved at P; C, illustration of P. molle, photocopied from Shan, R. H. & Pu, F. T. 1978 with modiﬁcation; this illustration was also used in Flora Reipublicae Popularis Sinicae (1985).

Figure 3. Underground part of perennial and annual species of Pternopetalum:A,P. cuneifolium (Wang, Lisong 0039, PE); B, P. vulgare (Wang, Lisong 0037, PE); C, P. radiatum (Grierson A. J. C. & Long D. G. 1855, E). cannot be confused with other species of Pternopeta- consulted. The software packages BRAHMS 5.58 lum. However, in a later generic review (Shan & Pu, (http://herbaria.plants.ox.ac.uk/bol/home/) and DIVA- 1978), a perennial collected by R. C. Ching 22416 GIS 5.2 (http://www.diva-gis.org/) were used to (Fig. 1B) was illustrated and referred to Franchet’s P. manage specimen datasets and to produce distribu- molle (Fig. 2B). At the same time, four new taxa (P. tion maps. The characters associated with fruits and molle var. crenulatum R.H.Shan & F.T.Pu, P. molle the blade indumentum were examined, and photo- var. dissectum R.H.Shan & F.T.Pu, P. cartilagineum graphs were taken with an NDigital Camera C.Y.Wu ex R.H.Shan & F.T.Pu and P. longicaule var. DXM1200F and scanning electron microscope. humile R.H.Shan & F.T.Pu) were described in this work. The concept of P. molle in Shan & Pu (1978) was to remain in later national Chinese and local DIAGNOSTIC CHARACTERS EXAMINED ﬂora treatments (Pu, 1985, 1993; Pan, 1997). As a HABIT AND UNDERGROUND PARTS result, controversial treatments involving P. cuneifo- Plants of Pternopetalum are mainly perennial having a lium (H. Wolff) Hand.-Mazz., P. cartilagineum and P. visible short caudex on which the leaf-sheath remains molle var. crenulatum appeared. In Flora Yunnanica are preserved (for example, P. botrychioides, P. cunei- (Pan, 1997), P. cuneifolium was accepted as a distinct folium and P. vulgare) (Figs 1B, 3A, B, 4–7). Pterno- species with P. cartilagineum and P. molle var. crenu- petalum longicaule var. humile, P. radiatum, P. latum as synonyms, whereas, in the recently pub- trifoliatum and P. molle (sensu Franchet) are annuals. lished Flora of China (Pu & Phillippe, 2005), P. These plants bear a single fusiform and somewhat cartilagineum was reinstated as a distinct species, herbaceous root, usually no more than 3 cm in length and P. cuneifolium and P. molle var. crenulatum were (Fig. 3C). The root is easily broken from the stem reduced to P. molle. and is often not visible on herbarium specimens The aim of this study is to clarify the taxonomic (Figs 1A, 8B, 9). Annuals have an almost uniform identity of P. molle, and re-examine its morphological appearance to their underground parts, whereas those delimitation with P. longicaule var. humile, P. radia- of perennials vary from well branched to single and tum and P. trifoliatum. A supplementary revision is thickened rootlets (Fig. 3A, B) in response to different also provided for the other taxa confused with P. molle. habitat water availability. Pternopetalum caespitosum R.H.Shan has been recorded as an annual (Pu, 1985), MATERIAL AND METHODS but ﬁeld investigations and examination of morphological characters show that it is a typical perennial. Morphological variation was evaluated on herbarium material preserved at the following herbaria: BM, CDBI, E, GH, HGAS, IBSC, K, KUN, NAS, P, PE, STEMS AND LEAVES SM, SZ and SWFC (herbarium abbreviations follow In earlier descriptions of species (Shan & Pu, 1978; Holmgren, Holmgren & Barnett, 1990). Field investi- Pu, 1985, 2001; Pu & Phillippe, 2005), variations in gations were carried out by the author in Guizhou, the main stem were reported as ‘stem branched/ Sichuan and Yunnan Provinces, China, during several unbranched’, or ‘stems one to several’. However, as visits in the period 2003–2005. Type specimens and observed in most Apiaceae by Kljuykov (Kljuykov original descriptions were carefully examined to et al., 2004), the stems of Pternopetalum can be well assess critical diagnostic characters. Photographs of developed or not. Not all perennial species have a types from GH, L, MO, NY, UC and WU were also well-developed main stem. They usually have a

Figure 4. A, Holotype of Pternopetalum cuneifolium, Maire, E. E. 620 conserved at E. B, Holotype of P. cartilagineum, Wang, Q. W. 72593 conserved at KUN.

Figure 5. A, Lectotype of Pternopetalum molle var. crenulatum, Tsai, H. T. 55935A conserved at IBSC. B, Isolectotype of P. molle var. crenulatum, Tsai, H. T. 55935 conserved at GH.

Figure 6. A, Lectotype of Pternopetalum vulgare, Henry, A. 10675 conserved at K. B, Holotype of P. molle var. dissectum, Feng, K. M. 4800 conserved at KUN.

Figure 7. A, Lectotype of Pternopetalum botrychioides, Pratt, A. E. 839 conserved at K. B, Isotype of P. delicatulum, Limpricht 1498 conserved at WU.

Figure 8. A, Isolectotype of Pternopetalum radiatum, Gammie, G. A. 992 conserved at K. B, Pternopetalum radiatum, Wood, J.R.I. 6450 conserved at E.

Figure 9. A, Holotype of Pternopetalum longicaule var. humile, Liu, T. N. et al. 1310 conserved at PE. B, Holotype of P. trifoliatum, Zhou, B. K. et Pu, F. T. 211 conserved at CDBI.

Figure 10. Illustration of variable leaf segments of Pternopetalum molle from: A–C, Delavay 4095 conserved at P; D, E, Sino-British Qinghai Expedition (1997) 905 conserved at E; F, G, Cooper, R. E. 4335 conserved at E. compressed short caudex on which the basal leaves Pan, 1997; Pu, 2001; Pu & Phillippe, 2005). and an elongated peduncle are produced, as in P. However, this character is unstable (Table 1), and botrychioides, P. cuneifolium and P. vulgare (Figs 3A, shows continuous variation in closely related taxa B, 4–7). The elongated peduncle occasionally bifur- (for example, P. longicaule var. humile, P. radiatum, cates in its upper part, where one or several slim P. trifoliatum and P. molle) (Fig. 10), even in a single leaves appear (cauline or upper leaves), but some- plant (for example, Pan, Z. H. et al. 84-100 NAS!). times cauline leaves are totally absent (Fig. 5). Pter- Therefore, this character is not suitable for interspe- nopetalum delavayi, P. longicaule var. humile, P. cific delimitation. molle, P. radiatum and P. trifoliatum have a continu- The blades of P. vulgare vary from entirely glabrous ously developed main stem on which the leaves are or sparsely strigose (Fig. 11A) to densely strigose more or less uniformly attached. They often bear one (Fig. 11B). Pternopetalum botrychioides and P. molle or several slim radical leaves (Figs 1A, 2A). have entirely glabrous blades (Fig. 11C, D, G, H). The Leaf division is diagnostically important in Pter- blades of P. cuneifolium are slightly strigose (Fig. 11E, nopetalum, although it is variable in several species. F). Its hairs are more or less cartilaginous, but not It is ternate with three distinct leaflets in P. uniformly throughout the whole blades in a single vulgare, with leaflets that vary from undivided, two- plant. For hairy species, the indumentum is usually to three-lobed to irregularly parted (Fig. 6). Ternate visible on the abaxial rather than the adaxial surface, or biternate leaves are found in P. longicaule var. and usually occurs along veins and margins of blades humile, P. radiatum, P. trifoliatum, P. molle and P. (Fig. 11). The blade indumentum has proven to be of cuneifolium (Figs 1A, 2A, 4, 8, 9). Pternopetalum little taxonomic value in Pternopetalum, as it is easily botrychioides is characterized by ternate-pinnate induced by habitat water availability. leaves, and its terminal ultimate leaf segments are caudate (Fig. 7). According to Franchet’s observations (Franchet, UMBELS 1885, 1894), the homomorphic or heteromorphic ulti- The umbels of Pternopetalum are only terminal or mate leaf segments of Pternopetalum are diagnosti- terminal and lateral. Terminal umbels are attached cally important. This opinion was followed by later on an elongated peduncle or pseudo-scape, which authors (Wolff, 1927; Shan & Pu, 1978; Pu, 1985; sometimes bifurcates in the upper part (for example,

Table 1. Variation in the patterns of the attached position of the leaves and shape of the ultimate leaf segments in Pternopetalum taxa accepted in the Flora of China (2005), Flora of Bhutan (1999), Umbelliferae (Apiaceae) of India (1993) and Chinese local ﬂora treatments

Basal Cauline Ultimate leaf Species Taxon leaves leaves segments complex

P. bipinnatum*PA– P. botrychioides var. botrychioides P A/P HO P. botrychioides var. latipinnulatum P A/P HO P. caespitosum P P HO/HE P. tanakae P. cardiocarpum P P HO/HE P. delavayi P. cartilagineum P A/P HO P. cuneifolium P A/P HO P. davidii P A/P HO P. delavayi P P HO/HE P. delavayi P. delicatulum P A/P HO P. ﬁlicinum P P HO/HE P. tanakae P. gracillimum PA – P. heterophyllum PP HE P. tanakae P. leptophyllum PP HO P. longicaule var. longicaule AP HO P. longicaule var. humile P P HO/HE P. molle P. molle var. molle P P HO/HE P. molle P. molle var. dissectum P A/P HO P. nudicaule PA – P. radiatum P P HO/HE P. molle P. rosthornii PP HO P. senii PA – P. sinense PP HO P. delavayi P. subalpinum PA – P. tanakae var. tanakae P A/P HE P. tanakae P. tanakae var. fulcratum PP HE P. tanakae P. trichomanifolium PA – P. trifoliatum PP HO P. molle P. vulgare var. vulgare P A/P HO P. vulgare var. acuminatum P A/P HO P. vulgare var. strigosum P A/P HO P. wolffianum P A/P HO P. yiliangense P A/P HO

A, absent; HE, heteromorphic ultimate leaf segments; HO, homomorphic ultimate leaf segments; P, present. Species complexes were identiﬁed by phenetic analyses (Wang, 2007). *New species described by Wang (2005).

P. botrychioides, P. cuneifolium and P. vulgare) CALYX TEETH, STYLOPODIUM AND STYLES (Figs 3, 4). Lateral umbels have short peduncles, usually shorter than the subtending petioles The characters calyx teeth, stylopodium and styles (Mukherjee & Constance, 1993) (for example, P. dela- have been well documented in earlier research (Pu, vayi, P. longicaule var. humile, P. trifoliatum, P. radia- 1985, 2001; Pu & Phillippe, 2005), but their taxo- tum and P. molle) (Figs 1A, 2A, 8B, 9). This character nomic value was not fully recognized. The robust was documented for P. radiatum (Mukherjee & Con- perennials (usually more than 30 cm high) (for stance, 1993), but not clearly indicated in P. caespito- example, P. botrychioides, P. davidii, P. delavayi and sum, P. cardiocarpum, P. longicaule var. humile, P. P. vulgare) have distinct and triangular or subulate sinense and P. trifoliatum (Pu & Phillippe, 2005); they calyx teeth, with erect styles which are approximately usually bear both terminal and lateral umbels. twice the length of the stylopodium. The stylopodia

Figure 11. Blade indumentum on adaxial (left) and abaxial (right) side of Pternopetalum vulgare, P. botrychioides, P. cuneifolium and P. molle:A,B,P. vulgare (Wang Lisong 0092, PE); C, D, P. botrychioides (Wang Lisong 0072, PE); E, F, P. cuneifolium (Wang, C. W. 72420, PE); G, H, P. molle (Delavay 4095, P).

Figure 12. Fruit shape and size, stylopodium, styles, calyx teeth and ribs: A, Pternopetalum vulgare (Wang, Lisong 0092); B, P. botrychioides (Wang, Lisong 0042); C, P. cuneifolium (Wang, Lisong 0037); D, P. molle (Delavay 4095); E, P. longicaule var. humile (Liou, T. N. & Tsoong, P. C. 1310); F, P. radiatum (John, R. I. Wood 7022). are conical in shape (Fig. 12A–C). In contrast, the lum have five more or less prominent and filiform ribs calyx teeth of delicate members (usually no more than on one mericarp (Fig. 12). These are more evident 30 cm high) (for example, P. leptophyllum, P. longi- in species having a robust appearance (for example, caule var. humile, P. trifoliatum, P. radiatum, P. molle P. botrychioides, P. cuneifolium and P. vulgare) than and P. tanakae) are minute or absent. The styles are in species having a delicate appearance (for example, recurved at the apex, equal or subequal to the sty- P. longicaule var. humile, P. trifoliatum, P. radiatum, lopodium. The delicate members have low-conical sty- P. molle and P. tanakae). Therefore, this character is lopodia (Figs 12D–F, 13A). Phenetic analyses (Wang, not clear-cut for the identification of species of Pter- 2007) have indicated that the characters calyx teeth, nopetalum, as it is variable in some species but stable stylopodium and styles are diagnostically important in others. The ribs of P. vulgare vary from smooth, in the identification of the two phenetic groups, minutely scaberulous (Mukherjee & Constance, 1993) and can be used to determine sectional divisions in to denticulate (Fig. 12A). Pternopetalum botrychioides Pternopetalum. has smooth or scabrid ribs (Fig. 12B), whereas the ribs of P. cuneifolium are denticulate or denticulate- fringed (Fig. 12C). The ribs of P. longicaule var. FRUITS AND RIBS humile, P. trifoliatum, P. radiatum and P. molle are Variations of the rib surface were originally empha- entirely smooth (Fig. 12B, D–F). sized by Wolff (1927), who used them to key out P. Variations in the pericarp surface have not been davidii with seven other species in section Dentari- investigated in previous treatments of Pternopetalum. oideae H.Wolff. According to Wolff (1927), the surface Through examination by scanning electron micros- of the fruit ribs was ‘serrulato-fimbriatus’inP. davidii copy, the surface is scale-like in P. longicaule var. and ‘glaber laevisque’ in the seven other species of humile, P. trifoliatum, P. radiatum and P. molle Pternopetalum. However, later authors (Pu, 1985; (Fig. 13B, C), and smooth and ridged in other taxa of Pan, 1997; Pu & Phillippe, 2005) expanded the Pternopetalum (Fig. 13D). Considering the common diagnostic value of this character to the whole genus, habitat preference (along margins of Abies/Tsuga and provided descriptions of ‘fruit ribs denticulate or woodland) of P. longicaule var. humile, P. trifoliatum, finely scabrid’ versus ‘fruit ribs filiform’ (Pu, 1985; Pu P. radiatum and P. molle, the scale-like structure may & Phillippe, 2005). However, all taxa of Pternopeta- facilitate the capture of heat and water better than

Figure 13. Scanning electron micrographs: A, stylopodium, calyx teeth, styles; B–D, surface structure of pericarps; A–C, Pternopetalum molle, sample from Expedition of Chinese Medical Plants on Tibet 4098 (PE); D, P. cuneifolium, sample from Ching, R. C. 22416 (PE). a smooth and ridged surface. This is possibly an var. humile, P. radiatum, P. trifoliatum and P. molle. indicator of ecological adaptation to an alpine Furthermore, annuals usually occur above 3000 m, environment. and are found on the margins of alpine needle-leaf forest (Abies/Tsuga forest) or in alpine meadows. By contrast, perennials often occur below 2800 m, and DISCUSSION AND CONCLUSIONS prefer shady, wet and less disturbed evergreen forest In summary, the morphology of the calyx teeth, sty- vegetation. lopodium and styles is important in the identiﬁcation The lateral umbels are particularly distinct in P. of the two sections of Pternopetalum, and may be caespitosum, P. cardiocarpum, P. delavayi, P. longi- responsible for its earlier phenotypic divergence caule, P. longicaule var. humile, P. radiatum, P. sinense, (Wang, 2007). The annual habit and associated mor- P. trifoliatum and P. molle, and are occasionally found phological variations, such as plant appearance, in P. tanakae. As observed in Anthriscus Pers. (Spalik, absence/presence of rhizome and attached position of 1996), the lateral formation of umbels allows continu- leaf, are useful in the identiﬁcation of P. longicaule ous growth of the main stem and the production of

KEY TO SPECIES 1. Plant annual, small and fusiform root slightly lignified, without well-developed caudex and distinct leaf-sheath remains at the base, leaves mainly cauline, basal leaves scanty or occasionally 1–2, umbels terminal and lateral, stylopodium low-conical, calyx teeth minute or absent, ribs smooth...... P. molle 1. Plant perennial, root intensely lignified with well-developed caudex and distinct leaf-sheath remains at the base, leaves mainly basal, cauline leaves 1–2 or sometimes absent, umbels only terminal, stylopodium conical, calyx teeth distinctly triangular or subulate, ribs smooth or denticulate ...... 2 2. Leaves ternate-pinnate or pinnate, caudate at the apex of terminal segments, blades entirely glabrous ...... P. botrychioides 2. Leaves ternate or biternate, acute at the apex of terminal segments, blades glabrous, strigose or densely strigose along veins and margins...... 3 3. Leaves ternate, leaflets 3, ultimate leaf segments larger, ovate or oblong ovate, usually 4–6.5 ¥ 2.5–4 cm, umbellules two- to five-flowered, filiform ribs smooth, scabrid or denticulate...... P. vulgare 3. Leaves biternate, ultimate leaf segments smaller, rhomboidal, ovate or oblong-ovate, often 1–3 ¥ 1–2 cm, umbellules two-flowered, ribs denticulate-fringed...... P. cuneifolium additional umbels if more resources become available. Spec. Nov. Regni Veg. 27: 181. 1929. Type: China, This character has been shown to be an ecological ‘Yünnan, Vallées-rives des torrents à Tsien-sin, preadaptation to ruderal habitats in Anthriscus 3000 m.s.m.’ [Now Yunnan Province], Maire, E. E. 620 (Spalik, 1996), and is correlated with the degree of (holotype: E! Barcode: E00000547) (Fig. 4A). selfing (Jury, 1986). In Pternopetalum, species having = Pternopetalum cartilagineum C.Y.Wu ex R.H.Shan lateral umbels usually occur in alpine habitats (above & F.T.Pu, in Acta Phytotax. Sin., 16(3): 70. 1978. Type: 2800 m) with one aborted mericarp in the schizocarp. China, Yunnan, Zhengkang Xian, iii.1936, 2500 m, C. This suggests that Pternopetalum may also experience W. Wang 72593 (holotype: KUN!; isotype: NAS! PE!) similar evolutionary changes to Anthriscus under high (Fig. 4B). selection pressure. = Pternopetalum molle var. crenulatum R.H.Shan & In contrast to reproductive characters, vegetative F.T.Pu, in Acta Phytotax. Sin., 16(3): 72. 1978. Type: characters, such as leaf division, blade indumentum China, Yunnan, Tsai, H. T. 55935A [lectotype here and the size and shape of ultimate leaf segments, designated: IBSC!; isolectotypes: (Tsai, H. T. 55935) exhibit much more plasticity in response to the NAS! PE! SZ! GH photograph seen] (Fig. 5). environment. They are usually polymorphic in widely distributed species, such as P. botrychioides, P. molle Distribution and habitat: North-west and north-east and P. vulgare, and are relatively stable in species Yunnan, China (Fig. 14), streamsides under ever- having a restricted distribution, such as P. cuneifo- green forest, 2400–3100 m. lium. Any abnormal variations are usually found in peripheral populations under harsh environmental conditions. This facilitates phenotypic plasticity, espe- Remarks: The specimen R. C. Ching 22416 (Figs 1B, cially variation in the size and shape of vegetative 2C) belongs to P. cuneifolium, a species recognized by characters. Therefore, it is best to group variants Wolff (1929) and characterized by the perennial habit, into a single polymorphic species, such as P. molle distinct rhizome and terminal umbels only (Figs 1, 2, or P. vulgare, rather than to treat them as separate 4). Shan and Pu (Shan & Pu, 1978; Pu, 1985, 1993) taxonomic units. misidentified this plant as Franchet’s P. molle Denticulate or scabrid ribs are known only in (Figs 1A, 2A), which is characterized by an annual P. botrychioides, P. cuneifolium, P. davidii and P. habit, delicate appearance, single fusiform root, with vulgare, and vary in widely distributed species. terminal and lateral umbels. As a result, a perennial Therefore, the diagnostic value of this character could having characters of leaves mainly basal, blade margin not be expanded to the whole genus for the identifi- crenulate on upper part and fruit ribs denticulate was cation of major species groups. described as a new variant: P. molle var. crenulatum (Shan & Pu, 1978). In this work, a perennial with ternate leaves, umbellules two-flowered, leaf segments TAXONOMIC TREATMENT somewhat coriaceous, veins and margins cartilaginous Pternopetalum cuneifolium (H.Wolff) Hand.-Mazz., and ribs denticulate was also described as a new in Umbelliferae. Symbolae Sinicae, 7: 718. 1933 species: P. cartilagineum. Being unaware of confusion Basionym: Cryptotaeniopsis cuneifolia H.Wolff, with P. molle, P. cuneifolium and P. molle were merged Umbelliferae Asiaticae Novae Relictae II. Repert. as a single species (Pu & Phillippe, 2005).

Figure 14. Distribution of Pternopetalum botrychioides (᭹), P. vulgare (), P. cuneifolium ( ) and type locality of P. cartilagineum ( ).

Indeed, P. cartilagineum, P. cuneifolium and P. identified as P. cuneifolium by Shan R. H. and Pan Z. molle var. crenulatum are conspecific, as determined H., whereas the specimen at GH was identified as P. by their common diagnostic characters, such as leaves vulgare by C. Norman (Fig. 5B), and the specimen at mainly basal and one- to two-ternate, cauline leaves SZ was identified as P. vulgare and P. molle by Hu, W. absent or occasionally one, ultimate leaf segments K. and K. T. Fu, respectively. It seems that, in order ovate or rhomboidal, c. 1–3.5 ¥ 1–1.5 cm, umbellules to qualify the new variant P. molle var. crenulatum, uniformly two-flowered and ribs denticulate (Figs 1B, Shan & Pu (1978) renumbered the specimens at IBSC 4A, 12C). Therefore, the treatment of Pan (1997) and KUN as Tsai, H. T. 55935A, and designated the should be reinstated because it is consistent with the latter as the holotype of the name P. molle var. crenu- morphological variation displayed in these plants. latum. As the specimen at KUN could not be found Detailed examination of the type materials indi- after an extensive search, the specimen at IBSC cated that the original collection Tsai, H. T. 55935 (Fig. 5A) was chosen as the lectotype of P. molle var. consisted of seven specimens, which are widely dis- crenulatum. tributed at GH, IBSC (two sheets), KUN, NAS, PE Pternopetalum cuneifolium is closely related to P. and SZ. The seven specimens were proven to have vulgare, but has weak morphological differences and been collected from one population, and belong to one geographical isolation from the latter. In addition, species, as determined by their common characters. there are a few populations that could be referable to However, the identification of this collection was this species, so that further field investigations are problematic. For example, the specimen at NAS was needed in the future.

Representative specimens examined: CHINA:YUNNAN: well developed and leaf-sheath remains invisible at Dali Xian: in shady thickets in side valleys on the the base. Stem single. Leaves basically cauline, eastern flank of the Tali Range, lat. 25°40′N, 7000– occasionally with one or two basal leaves, one- to 9000 ft, vi.1906, Forrest, 4623 (E); Jingdong Xian: Mt. two-ternate, ultimate segments ovate, broad-ovate or Wu Liang, Luo Shui Dong, 2490 m, iv.1959, X. G. Xu rhomboidal, 1–3.5 ¥ 1–1.5 cm, sometimes with lan- 4674 (PE); Shunning Xian: snow mountain, 3100 m, ceolate segments on upper leaves, blades smooth, 28.v.1938, T. T. Yu 16045 (NAS, PE); Yangbi Xian: margins serrate or incised-serrate; mucronate at Shang Yi County, Li Zi Ping, 27.iv.1929, R. C. Ching apex; cuneate at base. Loosely arranged umbels 22416 (PE); Yunlong Xian: Cao Jian County, Zi Ben terminal and lateral, the lateral umbels usually with Shan, 2417 m, 26.v.2005, Wang, Li-Song 0039 (PE); peduncle shorter than subtending petiole and its rays. Zhenkang Xian: possible Zheng Kang snow mountain, Bract absent. Rays (4–)6–15(–20), 1–3.5 cm long. vii.1936, C. W. Wang 72420 (KUN, PE); ibid., 2400 m, Bracteoles 1–2, 0.4–0.8 mm long, or occasionally vii.1936, C. W. Wang 72426 (PE). scanty. Umbellules (1–)2–3(–4)-flowered; pedicels 0.3– 2.5 mm long. Calyx teeth minute or absent. Petals Pternopetalum molle (Franch.) Hand.-Mazz., in white, c.2¥ 0.9 mm. Stylopodium low-conical; styles Hand.-Mazz., Umbelliferae. Symbolae Sinicae, VII. equal or subequal to stylopodium, reflexed at apex. 718. 1933 Fruit ovoid or oblong-ovoid, c. 1–2 ¥ 0.5–1 mm; later- Basionym: Carum molle Franch., in Notes sur ally compressed, five filiform ribs obscure, smooth; quelques Ombellifères du Yunnan. Bull. Soc. Philom. commissural plane to concave, vittae 1–3 in each Paris, Sér. 8, 6: 120: 1894. Type: China, ‘Yunnan, in furrow, 2–4 on commissure. Flowers April to June, silvis ad San-tcha-ho, supra Mo-so-yn; fruct. Vix mat. fruits July to September. 10 Aug. 1889’ [now Yunnan Province, Erh-Yuan Xian], Delavay, J. M. 4095 (holotype:P!;isotypes:K!NY Illustration: Figure 2A. photography seen) (Fig. 1A). pro. parte. Wolff, H. 1927. Umbelliferae-Apioideae-Ammineae-Carinae, Distribution and habitat: China: south Gansu, south- Ammineae novemjugat. et gen. (Cryptotaeniopsis). in west Shaanxi, Sichuan, south-east Xizang (Tibet), Engler, A. (ed.) Pflanzenr. pp. 178. auct. non. R. H. north-west Yunnan. Adjacent Himalayan regions: Shan & F. T. Pu 1978. in Acta Phytotaxon Sinica Bhutan, Sikkim (Fig. 15). It is usually found in alpine 16(3): 71–72.; auct. non. F. T. Pu 1985. in Shan, R., H., meadows or coniferous forest at 2800–4200 m; one of Sheh, M. & L. (eds.) Flora Reipublicae Popularis the few typical delicate alpine herbs found in this Sinicae. Pp. 50–51. auct. non. F. T. Pu 1993. in Wang, genus. W. T. et al. (eds.) Vascular Plants of Hengduan Moun- tains. pp. 1315. auct. non. Z. H. Pan, 1997. in C. Y. Wu Remarks: According to Shan & Pu (1978), P. longi- et al. (eds.) Flora Yunnanica. pp. 530. caule var. humile (Fig. 9A) is similar to P. longicaule = Pternopetalum radiatum (W.W.Smith.) P.K.Mukher- Shan, but differs from the latter in its dwarf appear- jee, in Indian Forester 97(1):55. 1971. ance, 4–30 cm high (usually 20 cm), occasionally with Basionym: Pimpinella radiata W.W.Smith., in Rec. one basal leaf, the upper ultimate leaf segments Bot. Surv. India, Iv. 266. 1911. Syn. Nov. Type: elongate-lanceolate. Pternopetalum longicaule is a Sikkim, Yeumtong, Gammie, G. A. 992 (lectotype: species recognized by its well-developed main stem, CAL; isolectotype:K!pro parte, the plant at the left and homomorphic and uniformly attached cauline part) (Fig. 8A). leaves (Shan, 1940). Shan (1940) also suggested that = Pternopetalum longicaule var. humile R.H.Shan & P. longicaule was closely related to P. filicinum F.T.Pu, in Acta Phytotax. Sin., 16(3): 76 1978. Syn. (Franch.) Hand.-Mazz. However, this is not the case Nov. Type: China. Shaanxi: Meixian, Mt. Taibai, Dou- as P. longicaule is very similar to P. delavayi.They Mu-Gong, east gorge, Liou, T. N. & P. C. Tsoong 1310 share many morphological characters, such as a (holotype: PE!) (Fig. 9A). conical stylopodium, distinct and triangular calyx = Pternopetalum trifoliatum R.H.Shan & F.T.Pu, in teeth, terminal and lateral umbels, and characters Acta Phytotax. Sin., 27(1): 64. 1989. Syn. Nov. Type: associated with the leaf. The similarity of P. longi- China, Sichuan, Ma-erh-k’ang, Mt. Tihu, 3500 m, caule and P. delavayi resulted in Hiroe (1979) grow on moss under Abies forests, 1.ix.1984, B. K. merging them as a single species, although he erro- Zhou, & F. T. Pu 211 (holotype: CDBI!; isotype: NAS!) neously used P. longicaule as the legitimate species (Fig. 9B). name. Although P. longicaule var. humile shares several Description: Delicate annual herb, glabrous, 10–35 cm characters, such as terminal and lateral umbel and high. Small and single fusiform root, 1–1.5 cm long, well-developed main stem, with P. longicaule and P. 2–3 mm in diameter, slightly lignified, caudex not delavayi, they have different characters with regard

Figure 15. Distribution of Pternopetalum molle (), P. longicaule var. humile (᭿), P. radiatum (᭜) and type locality of P. trifoliatum ( ).

to the stylopodium, calyx teeth, styles and habit (see low conical, styles usually reflexed at apex and ovate above). The distinctness of P. longicaule var. humile fruits usually with one aborted mericarp. in comparison with P. longicaule resulted in Fu The confusion with P. molle inevitably complicated elevating it as a species (Fu, 1981), a treatment efforts to examine its taxonomic relationships with P. that has not been accepted by Pu (Pu, 1985; Pu & radiatum (Fig. 8), a Himalayan species (Mukherjee, Phillippe, 2005). 1971, 1977; Mukherjee & Constance, 1993; Watson, This confused taxon relationship evidently derived 1999) which has been proposed as a threatened plant from confusion regarding the identity of P. molle. in India (Nayar & Sastry, 1987; Walter & Gillett, 1998). Indeed, P. longicaule var. humile and the more With the help of M. F. Watson (E), in addition to the recently described new species P. trifoliatum (known type specimens, the materials represented by three (or only from the type specimen) by Shan & Pu (1989) are four) populations of P. radiatum from Bhutan (Upper conspecific with P. molle. These plants share nearly kuru Chu: Cooper, R. E. 4335; Bumthang: Grierson A. the same morphological variation, such as delicate J. C. & Long D. G. 1855, Thimphu: Wood, J. R. I. 7022, appearance, usually not more than 30 cm high, mainly 6450) (Fig. 8B) were examined in this study. The cauline leaves, ternate or biternate, terminal and results indicate that P. molle and P. radiatum are lateral umbels, loosely arranged rays numbered from conspecific, as evidenced by their common delicate six to 15 in each umbel, umbellules two- to three- appearance, lateral umbels, mainly cauline and flowered, calyx teeth minute or obsolete, stylopodium ternate or biternate leaves etc. (Figs 8, 9).

In addition to the common morphological charac- 24.x.1982, F. T. Pu 492 (CDBI, NAS); Ya Zhui forestry ters of P. longicaule var. humile, P. trifoliatum, P. centre, 3500 m, 16.x.1982, Y. B. Yang et al. 217 radiatum and P. molle, these plants actually occur in (CDBI, NAS); Pingwu Xian: Zhuo Ca river, 2600 m, a continuous distribution range from south Gansu 2.viii.1985, s.n. 198 (CDBI); Ruo-er-gai Xian: Lang and Shaanxi Provinces, China, extending to eastern Mu Shi, 3200 m, 4.viii.1978, s.n. 0225 (SM); Songpan Himalayan regions (Fig. 15). Within this area, the Xian: Gong Ga Ling, 3400 m, 25.viii.1984, Cao, Y. L. diversiﬁed ultimate leaf segments of P. molle (Fig. 10) 070 (CDBI); Kang Xi Gorge, 2700 m, 19.x.1983, F. T. show a geographical divergence to some extent. Some- Pu 198 (CDBI); Xiaojin Xian: Liang He Kou, San Song what heteromorphic upper leaves (Fig. 10C, E, G) are Ping, 3600 m, 29.viii.1979, C. Q. Yuan 0037 (NAS); usually found from southward or northward periph- TIBET (XIZANG): without detailed locality, 3048 m, eral populations under harsh environments, such as 9.vii.1935, Kingdon-Ward, Francis s.n. (E); without Wood, J. R. I. 6450 (Fig. 8B) (Thimphu, Bhutan, c. detailed locality, 13.vii.1965, Y. T. Zhang, et al. 621 27.5°N, 89.7°E) and Liou,T. N. & P. C. Tsoong 1310 (PE); Bomi Xian: 3.iv.1982, B. S. Li, et al. 559 (PE); (Fig. 9A) (Mt. T’ai-pai, Meixian, Shaanix, c. 33.95°N, Guo Gu county, 4000 m, 6.viii.1965, J. S. Ying, et al. 107.75°E). This may indicate an ecological adaptation 0889 (PE); Ch’a-yu Xian: west slope of de la mu, of P. molle in response to suboptimal habitats. 4200 m, 20.viii.1973, s.n. 888 (PE); upper Ch’a-yu, Su Ku, 2880 m, 7.viii.1981, Tibet Group 73992 (PE); Representative specimens examined: CHINA:GANSU: Milin Xian: Zhe Mi Gang, 3800 m, 6.vii.1972, Expe- Kangle Xian: Mt. Lian Hua, 2500 m, 11.vii.1994, Y. F. dition of Chinese Medical Plants on Tibet 3898 (KUN, Wang 94236 (PE); Yuzhong Xian: Mt. Xing Long, PE); valley of Baga, 3400 m, 20.vii.1972, Expedition of 2150 m, 12.vi.1990, C. Y. Wu et al. 9205 (KUN); Zhuo Chinese Medical Plants on Tibet 4098 (PE); Ba-ga- Ni Xian: 26.vii.1936, 2800 m, Z. B. Wang 5416 (NAS); nan-jiang, 3400 m, 28.vii.1975, Tibet Group Supple- SHAANXI: Fopin Xian: Hei Tao Ping, Shan Ren Du, mentary Subgroup 750951 (PE); YUNNAN: Deqin Xian: 1900 m, 25.vi.1952, K. T. Fu4759 (IBSC, PE); without Yong Zhong snow mountain, 3200–3400 m, 6.vii.1981, detailed locality, 12.viii.1938, T.N.Liou&P.C. Tibet Group 2187 (CDBI); Atuntze, 3200 m, Tsoong 3298 (PE); Lueh-Yang Xian: Mt. Han Feng, 16.vi.1937, T. T. Yu 8542 (NAS, PE); Lijiang Xian: Mt. 2050 m, 13.iv.1952, K. T. Fu5868 (PE); Mei Xian: s.d., Yu Long, Gan Hai Zi, 3400 m, Z. H. Pan 84249 (NAS); P. C. Tsoong 2653 (PE); Kwanshan, Iungte, 2500 m, Ju Dian, a-shi-zhu, 2200 m, 13.vi.1964, Zhou, X. 1141 15.vii.1942, Z. B. Wang 13198 (NAS); SICHUAN: (KUN); Adjacent Himalayan Regions: Sikkim Hima- Baoxing Xian: near Qiao Qi,? Skang snow mountain, layan at Yeumtong, 13 000 ft., s.d., Gammie s.n. (K); 300 m, 30.vii.1978, Collection Team of Baopu County, Bhutan: Bumthang: Byakar, 3100 m, 12.vi.1979, Chongqing Municipal Academy of China (1978) Grierson A. J. C. & Long D. G. 1855 (E); Thimphu: 780802 (SM); Butuo Xian: near Wu Tuo rangeland, Doden to Barshong, 3 h above Dodena, 3200 m, 3600 m, 21.viii.1979, s.n. 1788 (SM); Dao’u Xian: Luo 23.vii.1989, Wood, J. R. I. 7022 (E); Hongtsu: to the Ka County, 2900 m, 8.iv.1960, s.n. 620158 (NAS); west of the town, 3100 m, 10.vii.1988, Wood,J.R.I. Jiulong Xian: Wu Xu Hai, 3400 m, 7.viii.1979, Hu, X. 6450 (E); Upper kuru Chu: gulu, 3210 m, 3.viii.1915, H. 20568 (CDBI); Under Tang Gu Community to Ran Cooper, R. E. 4335 (E). Da Valley, 3500–3900 m, 27.vii.1980, Q. Q. Wang 22714 (CDBI); Kangding Xian: Da Jian Lu, 3100 m, Pternopetalum botrychioides (Dunn) Hand.-Mazz., 2.viii.1934, Smith Harry 11076 (PE); Ma-erh-k’ang Umbelliferae. Symbolae Sinicae, 7: 718. 1933 Xian: Zhou Ke Ji to Hong Jun Ping, 3300 m, F. T. Pu Basionym: Cryptotaeniopsis botrychioides Dunn, J. 90019 (CDBI); ibid., 2900 m, 20.iv.1990, F. T. Pu Linn. Soc., Bot. 35: 494. 1903. Type: China, ‘West 90023 (CDBI); ibid., 21.iv.1990, F. T. Pu 90033 Szechuen at 9000–13 500 ft’ [type sheet label: ‘West (CDBI); 32 km N to Ma-erh-K’ang, 3000 m, Szechuen and Tibetan Frontier: Chieﬂy near 21.iv.1990, F. T. Pu 90029 (CDBI); 34 km near Mt. Tachienlu at 9000–13 500 feet’. Now Sichuan Prov- Meng Bi, 3600 m, 3.iv.1984, Yao, G. 232 (CDBI, NAS); ince, Kangding Xian], Pratt, A. E. 839 (lectotype:K! A-Mu-Jiao, 3000 m, 18.vii.1957, Z. Y. Zhang, et al. here designated; isolectotype: P! Barcode: P00432442) 23053 (IBSC, NAS, SZ); near rd of Mt. Zhe Gu, (Fig. 7A). 3500 m, 1.iv.1984, B. K. Zhou, 212 (CDBI, NAS); = Pternopetalum delicatulum (H.Wolff) Hand.-Mazz., Maowen Xian: Lu Shun Community, Wu Peng Zi, Umbelliferae. Symbolae Sinicae, 7: 718. 1933. 2750 m, 17.viii.1975, s.n. 11045 (CDBI, NAS, PE); Nan Xin community rangeland, 2000 m, 16.iv.1979, C. Basionym: Carum delicatulum H.Wolff, Botanische Q. Yuan 0081 (NAS); Meigu Xian: Hong Xi District, Yi Reisen in der Hochgebirgen Chinas und Ost-Tibet von Zi Yakou, 2700 m, 20.vii.1976, s.n. 13064 (CDBI, NAS, Dr. W. Limpricht: Umbelliferae 1922. Type: ‘China: PE); Hong Xi District, Mao Hong, 23.vii.1976, s.n. Sze-tchuan; Kiating-fu, Omi-schan, an Felsen, c. 13151 (CDBI, NAS, PE); Muli Xian: 3400 m, 3000 m.ü.M’ [now Sichuan Province, Emei Xian, Mt.

Emei], H. W. Limpricht 1498 (isotype WU, photograph without specific locality, s.d., R. C. Ching et al. 49 seen) (Fig. 7B). (PE); s.d., L. Y. Dai 41 (NAS); s.d., M. Y. Fang 117711 (SZ); s.d., W. R. He 257 (KUN); 740 m, 5.xi.1956, X. Li Distribution and habitat: China: South Chongqing, 47274 (KUN); s.d., T. H. Tu 278 (PE); 1930, J. Tang, Guizhou, Sichuan and Yunnan (Fig. 14), shady and 49 (PE); s.d., Y. H. Tao, 53967 (SZ); E. H. Wilson 3668, moist forest, streamside, occurring from 740 to 4925 (IBSC, K, P); 900 m, 11.v.1978, s.n. 0135 (SM); 2900 m. 28.iv.1979, s.n. 0246 (SM); 980 m, 24.v.1979, s.n. 487 (SM); Baoxing Xian: 1925, K. L. Chu 9931 (PE); Butuo Remarks: Pternopetalum botrychioides was first Xian: Jiao Ji He District, Lian He Community, recognized by Dunn (1903). Two specimens, Faber 30.v.1979, Collection Team of Butuo County, Chong- 629 and Pratt 839 (Fig. 7A), were cited but without qing Municipal Academy of China (1978) 0243 unambiguous typification in the original description. (SM); Chengdu: near medical factory of Chendu, Therefore, the two specimens are syntypes of 16.xii.1932, Z. R. Wang 7574 (SZ); Ebian Xian: s.d., Z. P. botrychioides under Art. 9.4 of the ICBN (McNeill X. Zhao 067 (KUN, NAS); ibid. 199 (KUN, NAS); et al., 2006). In the protologue, Dunn suggested that Emei Xian: Mt. O-mei: 4.v.1925, C. P. Qian, 5483 (SZ); the leaf characters of P. botrychioides are: ‘. . . Folia Wan Nian Shi, 25.vi.1976, 236 Army 1511 (PE); near radicalia minora simpliciter ternate foliolis Xi Xiang Chi, 29.vi.1976, 236 Army 1760 (PE); near ovatis . . .’. He also indicated the flower characters Jiu Lao Dong, 1.vii.1976, ibid. 1819 (PE); between the are: ‘. . . Petala late obovata, biloba, acumine rd of Guan Xi slope to Chu Dian, 23.vii.1962, S. Y. inflexo . . .’. It is clear that these characters are Chen 7504 (SM); Shi Shun Gorge, 9.v.1957, S. Y. Chen abstracted from Pratt 839 rather than Faber 629,as et al. 3111 (NAS); Jiu Lao Dong, 7.vii.1957, ibid. Faber 629 does not have basal leaves and flowers 3480 (NAS, SZ); 25.iv.1946, L. Y. Dai 1820 (NAS); because of the poor collection during fruiting time. 10.vi.1935, D. H. Du 228 (NAS, PE, SZ); 1300 m, Therefore, Pratt 839 is designated here as the lecto- viii.1928, W. P. Fang 2571 (NAS); 4500 m, type of P. botrychioides. 18.viii.1926, ibid. 3156 (NAS, PE); 15.vii.1930, ibid. Wolff did not see any specimens of P. botrychioides 7584 (NAS, SZ); 1100 m, 28.vii.1927, ibid. 12609 (SZ); (‘Specimina mihi non visa; . . .’) (Wolff, 1927), and 1600 m, 19.v.1940, ibid. 14707 (KUN, NAS, SZ); therefore he recognized Limpricht 1498, collected 2.xi.1940, ibid. 15088 (KUN, NAS); 12.iv.1930, ibid. from Mt. Emei, Sichuan of China, as a new species: P. 16143 (KUN, NAS, SZ); Zhang Lao Ping, 19.iv.1930, delicatulum. However, the type material of both P. ibid. 16273 (IBSC, NAS, SZ); 1800 m, 30.vi.1930, ibid. botrychioides and P. delicatulum was collected from 17097 (KUN); 20.iv.1931, ibid. 18483 (NAS, SZ); the same area, and they show nearly the same mor- 900 m, 20.iv.1931, ibid. 18496 (NAS, SZ); 1800 m, phological variation, especially the ternate-pinnate 6.v.1931, ibid. 18690 (PE, SZ); 9.viii.1951, ibid. 20624 leaves and terminal ultimate leaf segments usually (NAS, PE, SZ); 2900 m, 9.v.1964, K. J. Guan, et al. somewhat caudate. The seeming distinctness of Lim- 431 (CDBI); 1400 m, 17.viii.1964, ibid. 1503 (CDBI, pricht 1498, as indicated by the epithet ‘delicatulum’, PE); 1950 m, 3.viii.1964, ibid. 1565 (PE); 1950 m, s.d., is only because it is a young flowering individual of P. ibid. 1656 (CDBI, PE); 1000 m, 2.xi.1964, ibid. 2475 botrychioides. This situation was also found for other (PE); 30.vii.1935, X. Y. He 5548 (NAS, NY); 1200 m, herbarium materials of P. delicatulum, e.g. Zhao, Z. 5.viii.1956, L. C. Hu 51292 (SZ); 9.v.1959, Y. Y. Hua X. 199 and 067 (KUN and NAS). Therefore, P. deli- 3111 (SM); 1200 m, 7.x.1940, T. C. Lee 3709 (NAS); catulum should be reduced to P. botrychioides rather 2700 m, 11.vi.1983, M. L. Sheh 83617 (NAS); 1100 m, than to P. radiatum (Watson, 1999), or even as a 12.vi.1983, ibid. 83627 (NAS); 22.vi.1955, Sino- separate species (Pu & Phillippe, 2005), as its mor- Russian Investigation Group 2061 (PE, SZ); 5.v.1939, phological variation and geographical distribution do S. L. Sun, 0078 (NAS, SZ); 21.iv.1940, ibid. 1630 not justify this taxonomic conclusion. (NAS, SZ); 21.v.1940, ibid. 2120 (NAS, SZ); 1200 m, 14.vi.1940, ibid. 2360 (KUN, NAS); 1.x.1939, T. N. Representative specimens examined: CHINA: without Liou et al. 1113 (PE); 1700 m, 17.x.1976, D. D. Tao, specific locality, 2000 m, 28.v.1922, Smith Harry 2170 11457 (KUN); 1200 m, 5.viii.1956, Y. H. Tao 51292 (K); CHONGQING: Pengshui Xian: Cha Ban District, (NAS, SZ); 31.viii.1956, ibid. 51852 (SZ); 1950 m, 780 m, 24.vi.1979, s.n. 719 (SM); GUIZHOU: Xi-An 29.iv.1957, ibid. 54269 (SZ); 1200 m, 26.vii.2005, Jiu Community(?), 1100 m, 23.ix.1959, Libo Team 2122 Long Yan, Wang, Li-Song 0144 (PE); 28.v.1975, S. X. (PE); Anlong Xian: Long Shan District, Guizhou Expe- Wang 392 (CDBI, PE); 1600 m, 24.v.1980, C. Y. Wu dition (1960) 4411B (KUN); Leishan Xian: Mt. Lei 050 (KUN); 24.viii.1980, C. Y. Wu et al. 50 (PE); Gong, under slope of rangeland, 1450 m, 8.vi.1974, 1100 m, 2.iv.1956, Y. X. Xiao 48380 (SZ); 21.iv.1956, s.n. 74456 (HGAS); HUBEI: Shennongjia: 1700 m, ibid. 48703 (SZ); 3.v.1952, J. H. Xiong et al. 30343 24.vi.1934, Smith Harry 10079 (PE); SICHUAN: (NAS, PE); 14.viii.1938, G. J. Xu, 185 (NAS);

7.ix.1938, ibid. 543 (NAS); 1.x.1938, ibid. 784 (NAS); Xian: 27.v.1979, Collection Team of Meipu Chongqing 800 m, 17.v.1995, ibid. 3372 (PE); 850 m, 17.v.1995, Municipal Academy of China (1978) 0196 (SM); ibid. 3827 (PE); 1730 m, s.n., H. G. Xu 5392 (PE); 7.vii.1979, Collection Team of Meipu Chongqing 1200 m, 8.v.1957, G. H. Yang 54541 (NAS, PE, SZ); Municipal Academy of China (1978) 0369 (SM); 3060 m, 15.vi.1957, ibid. 55361 (PE, SZ); 1800 m, Muchuan Xian: 9.vii.1979, Collection Team of 16.viii.1957, ibid. 56676 (NAS, PE, SZ); 2000 m, Muchuan County, Chongqing Municipal Academy of 10.iv.1987, H. Z. You, et al. 380 (KUN); 1400 m, China (1979) 0353 (SM); 8.vii.1979, s.n. 0343 (SM); 31.v.1956, S. Z. Yu 49566 (SZ); 1100 m, 22.vi.1956, Pingshan Xian: 1500 m, 12.vii.1976, Collection Team ibid. 50062 (SZ); 950 m, 16.vi.1932, T. T. Yu 319 (NAS, of Chongqing Municipal Academy of China 358 (SM); PE); 3.vi.1939, G. Zhang 198 (NAS); 1.vii.1935, X. B. Shimian Xian: 1600 m, 24.vi.1978, Collection Team of Zhang 249 (NAS); 14.v.1940, W. J. Zheng 10362 Shimian County, Chongqing Municipal Academy of (IBSC, SZ); 27.v.1940, ibid. 10434 (IBSC, SZ); Guan China (1978) 780495 (SM); Shizhu Xian: 1500 m, Xian: Mt. Qingcheng, 9.iii.1928, W. P. Fang 13546 10.viii.1919, Collection Team of Chongqing Municipal (IBSC, NAS, SZ); Mt. Qingcheng, Tian Shi Dong, Academy of China 1203 (SM); Tianquan Xian: 1925, 740 m, 5.xi.1954, D. P. He 47274 (IBSC, SZ); 900 m, D. H. Du 2568 (KUN); 2900 m, 18.v.1956, D. P. He 6.v.1987, Z. L. Zhao 870396 (PE); Hongxi Xian: 43686 (SZ); Mt. Erh-lang, 26.vi.1951, W. G. Hu et al. 2200 m, 18.vii.1959, Chuan-Jing-Liang 1341 (PE); 10154 (PE, SZ); 1700 m, 26.iv.1986, T. Naito et al. 159 2100 m, 3.viii.1959, Z. T. Guan 9129 (NAS, PE); (PE); 980 m, 20.vi.1982, D. Y. Peng 45565 (CDBI, Hongya Xian: 2400 m, 8.vi.1994, W. K. Bao 2041 KUN); 1700 m, 25.iv.1980, Y. B. Yang et al. 21672 (CDBI); 1850 m, 26.vi.1994, ibid. 2423 (CDBI); (CDBI, KUN); 1840 m, 21.iv.1930, s.n. 00075 (NAS); 20.vii.1931, D. H. Du et al. 353 (NAS, PE); 1992, Z. W. Xiangcheng Xian: 3500 m, 18.vii.1979, Investigation Wang A00152 (CDBI); 19.vi.1939, C. W. Yao 3698 Team of Medical Plants from District of Ganzhi, (NAS); 1957, X. B. Zhang 03021 (SZ); 1200 m, Sichuan (1979) 0428 (SM); Xingwen Xian: 6.viii.1977, 19.vi.1979, s.n. 308 (SM); JiuLong Xian: 3300 m, Collection Team of Xingwen County, Chongqing 15.viii.1980, Z. A. Liu 22890 (CDBI); Leibo Xian: Municipal Academy of China (1977) 77325 (SM); 2200 m, 18.ix.1960, Chinese Medicine Investigation Xinjin Xian: 4.vii.1938, C. W. Yao 2087 (NAS, PE); Team (1960) 27974 (PE); 1800 m, 12.v.1959, Chuan- Ya-an Xian: 1800 m, 24.v.1978, Ya-an Team of Jing-Liang 0043 (KUN); 2170 m, 13.v.1983, M. Y. Sichuan (1978) 0283 (SM); Yibing Xian: 1000 m, Fang 117704 (SZ); 2175 m, 21.v.1983, ibid. 117760 5.vii.1977, s.n. 509 (SM); YUNNAN: without speciﬁc (CDBI, SZ); 2176 m, 21.v.1983, ibid. 117765 (SZ); locality, 10.vii.1913, Kingdon-Ward, Francis 748 (E); 2203 m, 30.v.1983, ibid. 117851 (SZ); 1350 m, 22.vi.1955, Sino-Russia Team 2061 (KUN); Dali Xian: 30.v.1959, Z. T. Guan 8261 (NAS, PE); 2100 m, Lugu Lake, 3658 m, vii.1910, Forrest 7143 (E); Cang 9.vi.1959, ibid. 8856 (NAS); 1500 m, 14.vii.1959, ibid. Shan, Hua Dian Ba, 3100 m, 20.v.1981, Sino-British 9328 (NAS); 1250 m, 27.iv.1983, M. Y. He et al. 117017 Expedition to Cangshan (1981) 0911 (E, KUN); (SZ); 2000 m, 12.v.1983, ibid. 116733 (CDBI, SZ); 2870 m, 21.v.1981, Sino-British Expedition to Cangs- 2200 m, Investigation Team of Medical Plant of han (1981) 1003 (E, KUN); 2700 m, v.1935, C. W. Sichuan 27994 (SM); 1200 m, 4.vi.1973, X. Li 110315 Wang 63347 (NAS, PE); Jingdong Xian: Mt. Wu (SZ); 800 m, 22.iv.1964, K. P. Yin et al. 0043 (CDBI); Liang, 2400 m, 5.iv.1959, X. G. Xu 3566 (KUN); 2100 m, 24.vii.1983, Q. S. Zhao 121371 (CDBI, SZ); 1980 m, 5.v.1959, ibid. 4855 (KUN); 2100 m, iv.1959, 2200 m, 18.vii.1983, ibid. 121457 (CDBI, SZ); 1250 m, ibid. 4915 (KUN); 2100 m, iv.1959, ibid. 5018 (KUN); 18.v.1983, Q. S. Zhao et al. 117134 (CDBI, SZ); Lijiang Xian: 2.vii.1939, R. C. Ching 20845 (PE); 2170 m, 27.iv.1983, ibid. 117638 (SZ); 2180 m, 10.vii.1959, K. M. Feng 22343 (KUN); 21.vii.1939, 3.v.1983, ibid. 117687 (SZ); 2180 m, 3.v.1983, ibid. Zhao, Y. K. 20845 (KUN); Malipo Xian: Mt. Lao Jun, 117689 (CDBI); 2100 m, 12.v.2005, ibid. 118704 26.v.1962, K. M. Feng 22824 (KUN); Suijiang Xian: (CDBI, SZ); 2000 m, 10.vii.1983, ibid. 118954 (CDBI, Luo Han Ping, 1450 m, 10.v.1973, B. X. Sun et al. SZ); Ma Liu Wan, 1800 m, 12.v.1959, s.n. 0043 (KUN, 0188 (KUN, PE); Luo Han Ping, 1260 m, 3.vi.2004, PE, SZ); 1800 m, 29.v.1964, s.n. 0258 (SZ); Luding Wang, Li-Song 0042 (PE); 1500 m, 5.vi.2004, ibid. Xian: 2200 m, 25.vi.1994, Y. Tang et al. 687 (CDBI); 0072 (PE); Tengchong Xian: 2600 m, 18.v. 1964, S. G. 2500 m, 28.vi.1994, s.n. 1256 (CDBI); Lushan Xian: Wu 6623 (KUN); Weixi Xian: 2600 m, 19.iv.1940, K. 1230 m, 10.v.1983, D. Y. Peng 47744 (CDBI); Mabian M. Feng 3542 (PE); Li Di Ping, 3300 m, 13.vi.1940, Xian: 940 m, 1.vi.1978, Collection Team of Mabian ibid. 4520 (NAS, PE); 3000 m, vi.1935, C. W. Wang County, Chongqing Municipal Academy of China 63719 (IBSC, KUN, NAS, PE); 3500 m, vi.1935, ibid. (1978) 325 (SM); 3400 m, 1.vii.1978, Collection Team 63923 (NAS, PE); 3500 m, vi.1935, ibid. 63990 (NAS, of Mabian County, Chongqing Municipal Academy of PE); Li Di Ping, 3136 m, 22.v.2004, Wang, Li-Song China (1978) 797 (SM); 1934, T. T. Yu 4280 (PE); 0037 (PE); 3092 m, 22.v.2004, Wang, Li-Song 0038 1650 m, 1.ix.1962, Q. L. Zhang 121359 (NAS); Meigu (PE); Yangbi Xian: 2480 m, 23.v.1963, Northwest

Yunnan Jinsha River Investigation Group 4129 (PE); Huo Jia Wan, 1250 m, 20.viii.1957, ibid. 63571 (NAS, Cang Shan, Hua Dian Ba, 2700 m, 24.vi.1984, Sino- PE, SZ); south of Jinfo Shi, 2250 m, s.n., Scientific American Botanical Expedition of Yunnan Province, Institute of West China 3139 (PE); Qiao Tou He Ba, China 470 (KUN); 2700 m, 25.vi.1984, ibid. 494 16.vii.2005, Wang, Li-Song 0097 (PE); Qiang Dao (KUN); Yongshan Xian: 2500 m, 20.vi.1932, X. G. Xu Ping, 17.vii.2005, ibid. 0123 (PE); Xiao He, Bao Jia 51056 (KUN). Ping, 1700 m, 19.vii.1957, J. H. Xiong et al. 95247 (NAS, SM, SZ); 1450 m, 3.v.1957, ibid. 90660 (PE, SZ); Pternopetalum vulgare (Dunn) Hand.-Mazz., 1900 m, 15.viii.1957, ibid. 92743 (PE, SZ); GANSU: Umbelliferae. Symbolae Sinicae, VII. 719. 1933 Wen Xian: Bikou, Shanwang Miao, 1400 m, Basionym: Cryptotaeniopsis vulgaris Dunn, Hook. 19.vii.1991, X. L. Chen et al. 911109 (PE); Bikou Dong, Ic. Pl. t. 2737. 1902. p. p. Type: China, ‘Yunnan, 920 m, 17.iv.2005, X. Wang, 10 (PE); GUANGXI: Yung- Fengchenlin, in forests at 7000 feet’ [Now Yunnan Ning: Jian Tong Ping, Yun Mu Xi, 800 m, 17.vi.1988, Province, Yuanyang Xian, Fengchunling], Henry, A. J. C. Zhang et al. 409 (PE); GUIZHOU: 22.vi.1935, S. W. 10675 (lectotype:K!;isolectotypes: E! MO photography Deng 0642 (PE); 26.iv.1936, ibid. 90167 (PE); Anlong seen) (Fig. 6A). Xian: Xinlong District, Xianhe Ping, 1700 m, 23.viii. = Pternopetalum molle var. dissectum R.H.Shan & 1981, C. Z. Deng 1188 (HGAS); 1280 m, Guizhou F.T.Pu, Acta Phytotax. Sin., 16(3): 72. 1978. Syn. Nov. Expedition (1960) 2875 (PE); Xianhe Ping, 1300 m, Type: China, Yunnan, Weixi Xian, 3000–3200 m, 14.vii.2004, Wang, Li-Song 0092 (PE); Bijie Xian: streamside under forest, 17.vi.1940. K. M. Feng 4800 1450 m, 6.ix.1957, P. H. Yu 649 (PE); Duyun Shi: (holotype: KUN!; isotype: PE!) (Fig. 6B). Yunfoushan, 730 m, 21.vii.1936, Y. Tsiang, 5982 (NAS); Jiangkou Xian: Da He Yan Gorge, 1100 m, Distribution and habitat: China: Gansu, Guizhou, 23.v.1964, Z. S. Zhang et al. 400781 (PE); 1700 m, Guangdong, Guangxi, Hubei, Hunan, Jiangxi, 3.vi.1964, ibid. 402230 (PE); Leishan Xian: Niao Dong Sichuan, Chongqing, Yunnan; adjacent Himalayan rangeland, 1500 m, 10.v.1985, Q. H. Chen, 3462 regions: north-east India, north Myanmar, Nepal (HGAS); Taoliang, Lengzhu Shan, 1450 m, Q. H. Chen (Fig. 14). This species has variable ultimate leaf seg- et al. 3032 (HGAS); Mt. Lei Gong, rangeland, 1450 m, ments in terms of size, shape and division, with a s.n. 74412 (PE); 1450 m, 8.vi.1974, s.n. 74412 (PE); broader distribution range and ecological adaptation rangeland, 1450 m, s.n. 74418 (HGAS, PE); 1610 m, than other members of the genus; it occurs from 700 s.n. 74455 (PE); east of rangeland, 1450–1550 m, s.n. to 3000 m. 74475 (HGAS); Libo Xian: Wengxi Community, Mogan Zhai, Z. R. Xu L1094 (PE); s.n., ibid. 1135 (PE); Remarks: As the epithet suggests, P. molle var. dis- Nayong Xian: Juren district, 1900 m, 25.vii.1929, sectum was mainly established on the basis of the Bi-Jie Expedition (1959) 397 (PE); Rongjiang Xian: irregular lobed or pinnatisected ultimate leaf seg- Xiaodan River, 680 m, s.n., Q. H. Chen, 3268 (HGAS); ments, which are larger than those of P. molle at Tongzi Xian: Guochang Gorge, 1200 m, 30.iv.1963, 4–7 cm long and 2–3 cm wide, according to Shan & Pu Normal University of Southwest of China 575 (PE); (1978). Examination of the type collection, K. M. Feng Yinjiang Xian: 1300 m, 2.ix.1963, Z. P. Jian et al. 4800, showed that P. molle var. dissectum has the 31116 (PE); Mt. Fanjing, at mountain peak, 18.v.1959, same typical ternate and three leaflet leaves as P. North Guizhou Investigation Group 01036 (NAS, PE); vulgare. The irregularly lobed or dissected leaflets 980 m, 7.vi.1959, ibid. 1050 (NAS, PE); 1800–2000 m, also found in Indian materials of P. vulgare was s.n., S. Z. Xin, 51132 (PE); s.n., ibid. 51233 (PE); indicated by Mukherjee, P. K. and Constance, L. Xiaojia River, Shai Shi Ban, 1160 m, 21.iv.1964, Z. S. (Mukherjee & Constance, 1993) as ‘leaves...ternate Zhang et al. 401295 (PE); Huguo Shi, 1200 m, or biternate..., the larger often few lobed or 13.v.1964, ibid. 401369 (PE); Huguo Shi, 1350 m, parted...’. 15.v.1964, ibid. 401861 (PE); 1800 m, 14.vii.1931, s.n. 51132 (PE); Zhenfeng Xian: Longtou Mountain, Leng- Representative specimens examined: CHINA: without shui River, 1600 m, 19 vii. 1986, Beijing Yong Expedi- specific locality, s.n., Z. W. Wang, A00434 (CDBI); tion (1986) 0325 (PE); HUBEI: without specific locality, 3000 m, 23.iv. 1882, Watt G. 6724 (E); 2438 m, s.n., A. Henry 5419 (IBSC, K); s.n., ibid. 5444 (K, P); 14.vi.1981, Stainton 8285 (K); E. H. Wilson 617 (K); s.n., ibid. 5444A (IBSC, K, P); Hefeng Xian: 11.v.1990, CHONGQING: Jiangjin Xian: 1500 m, 20.iv.1959, s.n. R. C. Wang HF90051 (NAS); Xuan’en Xian: Ba Da 0168 (SM); Nanchuan Xian: Mt. Jinfo: 30.v.1935, K. L. Gong Mountain, 1400 m, 17.vii.1958, H. J. Li 3963 Chu 1010 (NAS, PE, SZ); south of Jin Fo Shi, 1932, D. (PE); 1600 m, 12.vii.1958, H. J. Li 4103 (PE, SZ); H. Du 03139 (PE); 2439–2743 m, 31.v.1928, W. P. Fang HUNAN: without specific locality, Ho & Li 51058 (K); 1166 (NAS, PE); Yang Yu Ping and Lei Jiao Ping, Qianyang Xian: 13.viii.1953, D. H. Du 757 (PE); 1450 m, 28.v.1957, G. F. Li 61357 (NAS, PE, SZ); near Sangzhi Xian: Yang Hong Ping, 1200 m, 4.x.1976,

Hunan Team 802 (PE); 1650 m, 28.vi.1975, B. G. Li 22.v.1978, ibid. 379 (CDBI, SZ); Tianquan Xian: et al. 750058 (PE); 650 m, 19.vi. 1991, Q. Lin 604 (PE); 1640 m, 17.v.1983, D. Y. Peng 47869 (CDBI); Wen- Sha Di Ping, 1250 m, 29.iv.1958, L. H. Liu 9210 (NAS, chuan Xian: 1120 m, Training Group of Graduate PE); JIANGXI: Dayu Xian: 750 m, 10.vi.1962, J. S. Yue, Student (1978) 79033 (SZ); Xingwen Xian: 1340 m, 1216 (NAS, PE); Suichuan Xian: 14.v.1970, 236 13.v.1959, Chuan-Jing-Yi 0407 (PE, SZ); Xiushan Working Group 725 (PE); 900 m, 5.x.1963, J. S. Yue Xian: 1200 m, 3.vii.1978, Collection Team of Fuling 4599 (KUN); SICHUAN: without speciﬁc locality, Region, Chongqing Municipal Academy of China 3.viii.1959, Chuan-Jing-Liang 4391 (PE); 25.vi.1955, (1979) 0819 (SM); 1500 m, 7.viii.1979, Collection Team Xuedong ping, Sino-Russian Investigation Group 2310 of Fuling Region, Chongqing Municipal Academy of (PE, SZ); Emei Xian: Mt. O-mei, 27.vi.1976, 236 Army China (1979) 1027 (SM); Xuyong Xian: 1500 m, 1650 (PE); 1938, C. Y. Chiao et al. 737 (E); 2300 m, 15.v.1964, Collection Team of Xuyong County, Chong- 18.vi.1935, D. H. Du 328 (NAS, PE, SZ); Jie Ying Dian, qing Municipal Academy of China (1978) 0048 (SM); 2300 m, Z. X. Qu 328 (PE); Gulin Xian: 950 m, Ya-an Xian: 1000 m, 7.v.1978, Collection Team of 23.viii.1976, Collection Team of Guling County, Ya-an County, Chongqing Municipal Academy of Chongqing Municipal Academy of China (1976) 922 China (1978) 0032 (SM); 2000 m, 18.vii.1978, Collec- (SM); 1200 m, 20.vii.1976, s.n. 0751 (SM); Hejiang tion Team of Ya-an County, Chongqing Municipal Xian: 500 m, 25.vi.1977, 32 Working Group (1977) Academy of China (1978) 0720 (SM); 2000 m, 0213 (SM); Hongya Xian: Mt. Wa Wu, 1850 m, 18.vii.1978, Collection Team of Ya-an County, Chong- 2.vi.1993, W. K. Bao 799 (CDBI); 1950 m, 4.vi.1993, qing Municipal Academy of China (1978) 0723 (SM); ibid. 893 (CDBI); 1000 m, 15.vi.1979, Collection Team YUNNAN: without speciﬁc locality, iv.1917, Forrest of Hongya County, Chongqing Municipal Academy of 13747 (E); v.1919, ibid. 17870 (E); 2439 m, 18.iv.1882, China (1979) 0169 (SM); Jinyang Xian: 2300 m, G. Watt 6556 (CAL, E, K); Binchuan Xian: Huang 7.viii.1978, Collection Team of Jinyang County, Ping, H. C. Wang 1939 (PE); Dali Xian: Cang Shan, Chongqing Municipal Academy of China (1978) 432 2134 m, vi.1906, Forrest 4623 (E, K); Pingbian Xian: (SM); 2200 m, 5.vi.1979, Collection Team of Liangshan 1500 m, P. Y. Mao 02378B (KUN, PE); 1400 m, Region, Chongqing Municipal Academy of China 12.vii.1934, H. T. Tsai 62563 (KUN, NAS, PE, SZ); (1979) 000121 (SM); Junlian Xian: 1500 m, Suijiang Xian: 24 rd, Niu Lu Kou, 1500 m, 27.v.1973, 28.vii.1977, Collection Team of Junlian County, B. X. Sun et al. 0461 (KUN, PE); Tengchong Xian: Chongqing Municipal Academy of China (1978) 1135 1900 m, Hou Qiao Community, 19.v.1964, S. G. Wu (SM); Leibo Xian: 13.v.1959, Chuan-Jing-Liang 0104 6702 (NAS); Weixi Xian: along Lancing River and Nu (PE); 2600 m, 21.vi.1959, ibid. 0887 (CDBI, PE); River, 2100 m, 6.v.1940, K. M. Feng 3660 (PE); 2100 m, 19.v.1959, Chuan-Jing-Pei 0192 (PE); Weideng county, 2400 m, 19.v.1982, Tibet Group 6566 25.v.1979, Collection Team of Leibo County, Chongqing (PE); Wenshan Xian: 1900 m, 20.viii.1947, K. M. Feng Municipal Academy of China (1978) 0402 (SM); at 11363 (PE); Xichou Xian: 1420 m, 25.v.1964, S. Z. 257 km rd, 2250 m, 29.v.1983, M. Y. Fang 117839 Wang 433 (KUN); Yiliang Xian: 1972, C. Y. Wu 77 (CDBI); 1050 m, 25.vi.1976, Training Group of Plant (KUN). Taxonomy 76195 (SZ); Mabian Xian: 2000 m, 9.vi.1978, Collection Team of Mabian County, Chong- ACKNOWLEDGEMENTS qing Municipal Academy of China (1978) 475 (SM); 1500 m, 7.vi.1978, Collection Team of Mabian County, The author would like to thank the curators of CDBI, Chongqing Municipal Academy of China (1978) 0507 HGAS, IBSC, KUN, NAS, PE, SM, SZ and SWFC for (SM); 2400 m, 10.vii.1978, Collection Team of Mabian allowing the study of their Pternopetalum specimens, County, Chongqing Municipal Academy of China and of B, BM, E, GH, K, MO, NY, P, UC and WU for (1978) 921 (SM); 2500 m, v.1931, J. Tang et al. 22913 providing valuable type specimens and information (PE); Pingshan Xian: 1000 m, 22.vi.1959, s.n. 1168 for this research. Sincere thanks go to M. F. Watson (PE); Pingwu Xian: 1820 m, 16.v.1958, X. Q. Jiang (E) and David E. Boufford (GH) for sending additional 10233 (NAS, PE, SZ); Puge Xian: 2700 m, 13.vi.1979, Pternopetalum specimens for generic review, and for Collection Team of Puge County, Chongqing Municipal comments on earlier versions of the manuscript. Academy of China (1979) 688 (SM); Qingchuan Xian: Thanks are due to Christina Flann for improving the 1400 m, 4.viii.1965, s.n. 2715 (SM); Shimian Xian: English and to A. L. Li (PE) for drawing the illustra- 2300 m, 20.ix.1978, Collection Team of Shimian tions. This work was supported by the National County, Chongqing Municipal Academy of China Natural Science Foundation of China (30600035), a (1978) 781263 (SM); Hai Zi Shan, s.n., C. J. Xie 42618 project from the Chinese Academy of Sciences (PE); Shizhu Xian: 1600 m, vi.1978, Y. Chen 2472 (KSCX2-YW-Z-067) and Digital Herbarium Project (CDBI, SZ); 1050 m, 25.vii.1978, ibid. 2972 (CDBI); from the Ministry of Science and Technology of China 1300 m, 20.v.1978, W. H. Wang 360 (CDBI); 1300 m, (2005DKA21401).

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