TIBETIODES NESOM, GEN. NOV. (ASTERACEAE: ASTEREAE), INCLUDING 27 SPECIES OF HIMALAYAN ASTER

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1 Nesom, G.L Tibetiodes Nesom, gen. nov. (Asteraceae: Astereae), including 27 species of Himalayan Aster. Phytoneuron : Published 19 August ISSN X TIBETIODES NESOM, GEN. NOV. (ASTERACEAE: ASTEREAE), INCLUDING 27 SPECIES OF HIMALAYAN ASTER GUY L. NESOM Research Associate Academy of Natural Sciences of Drexel University Philadelphia, Pennsylvanica guynesom@sbcglobal.net ABSTRACT Tibetiodes Nesom, gen. nov., includes 27 species primarily Himalayan in geography T. flaccida ranges significantly more broadly and several other species occur outside of the Tibetan plateau. Molecular sequence data indicate that 7 of these species are monophyletic and distantly related to Aster sensu stricto (in the sense of Li et al 2012). The addition here of other species is based on their morphological coherence in a set of features, particularly and consistently by ray florets in 2 3 series and with short tubes and elongate styles. The following constitute the genus, all as new combinations in Tibetiodes: T. angustior, T. asteroides, T. bietii, T. brachytricha, T. brevis, T. bulleyana, T. diplostephioides, T. falconeri, T. farreri, T. flaccida, T. glandulosa, T. himalaica, T. jeffreyana, T. labrangensis, T. latibracteata, T. latisquamata, T. likiangensis, T. lipskii, T. nepalensis, T. retusa, T. salwinensis, T. setchuenensis, T. stracheyi, T. techinensis, T. tientschwanensis, T. tsarungensis, and T. yunnanensis. Formal nomenclature is provided for each species and synonyms. Lectotypes are designated for A. brachytrichus, A. delavayi, A. diplostephioides var. falconeri, A. falconeri var. nepalensis, A. farreri, A. flaccidus var. fructu-glandulosus, A. himalaicus, A. labrangensis, A. latibracteatus, A. likiangensis, A. mekongensis, A. salwinensis, A. stracheyi, A. vilmorinii, and A. yunnanensis. Molecular sequence data indicate that a group of primarily Himalayan species of Aster L. is monophyletic and evolutionarily distinct from even the broad range of species considered by Li et al. (2012) to constitute the genus. Six of the species were initially studied by Li et al.; Aster salwinensis was added to the set by Zhang et al. (2015), who updated the cladistic topology as in Figure 1. Li et al. (2012) acknowledged that this group of species "might become a new genus" and that is the course taken here. The group is part of the Asterothamnus branch of subtribe Asterinae (Nesom 2020a, 2020b). Figure 1. Relationships of seven species of Aster based on a combined data set (nuclear ribosomal DNA internal and external transcribed spacer sequences and plastid genome DNA trnl-f sequences). Modified from Zhang et al. (2015). Grierson (1964) recognized the morphological coherence of these species, placing them in subsects. Homochaeta and Heterochaeta of Aster sect. Alpigeni in his classification. His observations add 6 species to those outlined above in Figure 1: A. bietii, A. falconeri, A. himalaicus, A. latibracteatus, A. retusus, and A. stracheyi (A. tricephalus, A. neo-elegans, and A. barbellatus of Grierson's subsect. Heterochaeta are not part of the group considered here). The survey here adds another 14 species.

2 2 TIBETIODES Nesom, gen. nov. TYPE: Tibetiodes diplostephioides (DC.) Nesom See Nesom (2020c) for the taxonomic disposition of Heterochaeta DC., which might have been used as the name for this genus. Perennial herbs from a short, thickened, fibrous-rooted rhizome (stolons in T. stracheyi); cauline leaves oblong to elliptic-lanceolate, ovate, or elliptic-obovate, entire or with a few, shallow teeth or mucros, epetiolate, often subclasping; heads mostly 1; phyllaries mostly evenly herbaceous, lanceolate-triangular to lanceolate-oblong, usually longer than the disc flowers; ray flowers in 2 3 series (sometimes 1), light blueviolet to violet, tube short and styles elongate, ligules narrow, coiling; disc corollas often darkening or becoming deep violet in age; achenes narrowly obovate to oblanceolate-obovate, strongly compressed, 2- nerved; pappus usually of 2 series, the inner of relatively few, thickened bristles, the outer of short scales (all of the pappus short in T. brachytricha). Tibetiodes species mostly are restricted the Himalayan region (Tibetan Plateau). The ranges of T. asteroides, T. diplostephioides, and T. labrangensis extend to Gansu and Qinghai; T. farreri occurs only in central China, not reaching the Himalayas, and T. lipskii is restricted to Gansu and Qinghai. Unambiguously diagnostic features of the genus are in the ray flowers, which are consistently violet and usually occur in 1 or (most commonly) 2 3 series, producing short tubes and prominently long styles. In situ photos (Figs. 2-9) show the long styles. Most of the species have well-developed cauline leaves, but some have leaves mostly in a basal rosette, with cauline strongly reduced (Tibetiodes asteroides, T. flaccida, T. glandulosa, T. jeffreyana, T. likiangensis, T. retusa, T. tientschwanensis). Variation in habit and morphology in Figs Phyllaries in a few species are narrow and thickenened (Tibetiodes farreri, T. labrangensis, T. salwinensis; e.g., Fig. 12), in contrast to the broad and evenly herbaceous phyllaries characteristic of most. Variation in phyllary morphology is shown in Figs A morphological subgroup includes Tibetiodes angustior, T. brevis, T. diplostephioides, T. labrangensis, T. lipskii, and T. yunnanensis these have persistent, fibrous petiole bases (e.g., Figs. 14, 17, 18). Tibetiodes angustior and T. labrangensis (Fig. 17) consistently produce 3 5 heads per stem; T. yunnanensis usually is monocephalous but sometimes has 2 3 heads. 1. Tibetiodes angustior (Hand.-Mazz.) Nesom, comb. et stat. nov. Aster yunnanensis var. angustior Hand.-Mazz., Notizbl. Bot. Gart. Berlin-Dahlem. 13: TYPE: CHINA. Yunnan. NW Yunnan, Tung-shan in der Yangtze-Schleife, m, offene, steinige Matten, Sep, 1913, G. Forrest (holotype: E image; isotypes: BM image, K-2 sheets images). China (Sichuan, Yunnan). 2. Tibetiodes asteroides (DC.) Nesom, comb. nov. Heterochaeta asteroides DC., Prodr. 5: Aster asteroides (DC.) Kuntze, Revis. Gen. Pl. 1: TYPE: KASHMIR. Cachemire, 1833, M. Royle 114 (holotype: G-DC image). Aster heterochaeta Benth. ex C.B. Clarke, Comp. Ind., (nom. illeg. superfl.). Erigeron heterochaeta (Benth. ex C.B. Clarke) Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 16: Aster heterochaeta was a replacement name for Heterochaeta asteroides DC. Aster hedinii Ostenf. in Hedin, S. Tibet 6(3): TYPE: CHINA. Xizang (Tibet). SW Tibet, Himalayas norra fot., Från vägen mellan Läger CCIII (Dara-sumkor, 4931 m) och Läger CCIV (Buk-gyäyorap, 4870 m), Mellan båda ligger passet Tugri-la, 5270 m, 16 Jul 1907, S. Hedin s.n. (holotype, as annotated on the sheet: S image). China (Gansu, Qinghai, Sichuan, Xizang, Yunnan), Bhutan, India, Kashmir, Nepal.

3 3 3. Tibetiodes bietii (Franch.) Nesom, comb. nov. Aster bietii Franch., J. Bot. (Morot) 10: TYPE: CHINA. Yunnan. No other locality data, 3 Jun 1896, Prince H. d'orléans s.n. (holotype: P image). China (Xizang, Yunnan). 4. Tibetiodes brachytricha (Franch.) Nesom, comb. nov. Aster brachytrichus Franch., J. Bot. (Morot) 10: LECTOTYPE (designated here): CHINA. Yunnan. Montagne Hee-chan-men au dessus de Lankong, 11 Jul 1883, R.P. Delavay 101 (P ; isolectotypes: A image, P image). Protologue: "Yun-nan, in pascuis montis Hee-chan-men supra Lankong (R.P. Delavay n. 101 et 4466); in collibus prope collum Yen-tze-hay, 3200 m (id.)." Aster bodinieri H. Lév., Bull. Géogr. Bot. 25: LECTOTYPE (designated here): CHINA. Yunnan. Plateau du Io-chan, 3400 m, Jun 1914, E.E. Maire 706 (E ; isolectotype: E ). Protologue: "Yun-Nan: patures des plateaux et des pics de Ta-Hai, Io-Chan et de Tong-Tchouan." China (Guizhou, Sichuan, Yunnan), Myanmar. 5. Tibetiodes brevis (Hand.-Mazz.) Nesom, comb. nov. Aster brevis Hand.-Mazz., Notizbl. Bot. Gart. Berlin-Dahlem. 13: TYPE: CHINA. Yunnan. Mountains of the Chungtien plateau, SE of Chungtien, stony alpine pasture, 13,000 ft, 15 Jul 1913, G. Forrest (holotype: B?; isotype: E image, Fig. 18). China (Yunnan). 6. Tibetiodes bulleyana (Jeffrey ex Diels) Nesom, comb. nov. Aster bulleyanus Jeffrey ex Diels, Notes Roy. Bot. Gard. Edinb. 5: TYPE: CHINA. Yunnan. Western slopes of Kari pass on descent to Shu-pa valley from Pung-tzu-la to Shi-zo, moist shady places, 13,000 ft, 15 Sep 1904, G. Forrest 7661 (holotype: E image). China (Yunnan). 7. Tibetiodes diplostephioides (DC.) Nesom, comb. nov. Heterochaeta diplostephioides DC., Prodr. 5: Aster diplostephioides (DC.) Benth. ex C.B. Clarke, Comp. Ind., Erigeron diplostephioides (DC.) Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 21: TYPE: KASHMIR. Cachemire, 1833, M. Royle 147 (holotype: G-DC image). Aster delavayi Franch., J. Bot. (Morot) 10: Aster diplostephioides var. delavayi (Franch.) Onno, Biblioth. Bot. 106: Erigeron delavayi (Franch.) Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 21: LECTOTYPE (designated here): CHINA. Yunnan. Les prairies au col de Yen-tze-hay (Lan Kong), 3200 m, 18 Sep 1885, J.M. Delavay 1675 (P image; isolectotypes: P , , , , , and , images). Protologue: "Yun-nan, in pratis supra collum Yen-Tze-hay, alt m. (Delavay, n. 1675; 999); Fang-yang-tchang, in pascuis supra Mo-so-yn, alt m.)." Bhutan, China (Gansu, Qinghai, Sichuan, Xizang, Yunnan), India, Kashmir, Nepal, Pakistan. 8. Tibetiodes falconeri (C.B. Clarke) Nesom, comb. nov. Aster diplostephioides var. falconeri C.B. Clarke, Comp. Ind., Aster falconeri (C.B. Clarke) Hutch., Gard. Chron. 47: Aster diplostephioides subsp. falconeri (C.B. Clarke) Onno, Bibl. Bot. 106: Erigeron falconeri (C.B. Clarke) Botsch., Acta Inst. Bot. Acad. Sci., U.R.S.S. 21: LECTOTYPE (designated here): KASHMIR. "Kashmir," no other collection data, H. Falconer 561 (K image; isolectotypes: GOET, K , , and images, LE, M, P, S, images). Protologue: "An species? coll. Falconeri." Grierson (1964, p. 141) cited "holo.

4 4 K" is the only one of the K sheets with an institutional label of "Type Specimen" but otherwise it is not distinguished from the others. Kashmir. 9. Tibetiodes farreri (W.W. Smith & Jeffrey) Nesom, comb. nov. Aster farreri W.W. Smith & Jeffrey, Notes Roy. Bot. Gard. Edinb. 9: Aster diplostephioides subsp. farreri (W.W. Smith & Jeffrey) Onno, Biblioth. Bot. 106: Erigeron farreri (W.W. Smith & Jeffrey) Botsch., Botsch., Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 21: LECTOTYPE (designated here): CHINA. Xizang. Near Gansu frontier, sporadic in the higher valley fields and alps, in hay grass, 12 Aug 1914, R. Farrer & W. Purdom 174 (E image; isolectotype: E image). The protologue cited only "Farrer and Purdom. No. 174." Aster nigrotinctus Y. Ling, Contr. Inst. Bot. Natl. Acad. Peiping 2: Aster vilmorinii var. nigrotinctus (Y. Ling) Y. Ling, Contr. Inst. Bot. Natl. Acad. Peiping 2: LECTOTYPE (designated here): CHINA. Xizang. Hsiaowutaishan, 2400 m, 27 Jul 1931, T.-P. Wang 760 (PE image; isolectotypes: PE and images). China (Gansu, Hubei, Qinghai, Shanxi, Sichuan). 10. Tibetiodes flaccida (Bunge) Nesom, comb. nov. Aster flaccidus Bunge, Mém. Acad. Imp. Sci. St.-Pétersbourg Divers Savans 2: Erigeron flaccidus (Bunge) Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 16: TYPE: RUSSIA. Protologue: "Hab. in alpinis ad Tschujam et in subalpinis Sailughem; floret Julio mense [perennial]" (Probable type material: P image, "alp. Tschujae "; possible type material: GH image). Aster argunensis DC., Prodr. 5: TYPE: RUSSIA. Ad fl. Argunum in Dahurie, 1820, Razgildeev s.n. (holotype: G-DC image). Aster glarearum W.W. Smith & Farrer, Notes Roy. Bot. Gard. Edinb. 9: TYPE: CHINA. Gansu. Kansu, high alpine shingles above Siku, 11,000 12,000 ft, 21 Jun 1914, R. Farrer and W Purdom 131 (holotype: E image; isotype: E image). f Bhutan, China (Gansu, Hebei, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan), India, Iran, Kashmir, Kazakhstan, Mongolia, Nepal, Pakistan, Russia, Uzbekistan. 11. Tibetiodes glandulosa (Keissl.) Nesom, comb. nov. Aster flaccidus var. glandulosus Keissl., Ann. Naturhist. Hofmus. Wien 22: Aster glandulosus (Keissl.) Hand.-Mazz., Ost. Bot. Zeit. 79: (not Aster glandulosus Labill. 1806). Aster flaccidus subsp. glandulosus (Keissler) Onno, Biblioth. Bot. 26(Heft 106): TYPE: CHINA. Xizang. Tibet, Sudufer des Mangzaka, 5370 m, Umgebung von Thermen, 1907, E. Zugmayer s.n. (holotype: W, photo- E image). Aster flaccidus var. fructu-glandulosus Ostenf. in Hedin, Southern Tibet 6(3): Aster flaccidus subsp. fructu-glandulosus (Ostenf.) Onno, Bibl. Bot. 106: LECTOTYPE (designated here): CHINA. Xizang. Eastern or Inner Tibet, Mandarlik, near Camp XLIV, 5127 m, 18 Aug 1901, S. Hedin s.n. (C image; isolectotype: S [as cited by Onno 1932 and by Grierson 1964]). The lectotype comprises the 3 small plants at the bottom right of the sheet; another Hedin collection (2 plants) of the same species is above, another (1 plant) to the left. Protologue: "Northern Tibet, Camp XVII, at a river, 4024 m, 31 Jul 1900 (flow.); Kar-yakaksai, Camp X, Chimen-tagh, 3984 m, 21 Jul 1900 (flow.); Mandarlik, 3437 m, medio July 1900 (flow.); some spec. very large); Eastern or Inner Tibet, near Camp XLIV, 5127 m, 18 Aug 1901 (flow.); S.W. Tibet, Camp CCXI, East of the lake Manasarovar, 4654 m, 24 Jul 1907 (flow.)." Ostenfeld (1922) was aware of Keissler's earlier var. glandulosus from Tibet but noted that "As nothing is said [in Keissler's protologue] about the achenes I cannot refer it to its proper place and do not know if it is a variety of A. flaccidus or of A. Hedinii described below."

5 5 China (Xizang), India. 12. Tibetiodes himalaica (C.B. Clarke) Nesom, comb. nov. Aster himalaicus C.B. Clarke, Comp. Ind., LECTOTYPE (designated here): INDIA. Sikkim. Jongri [Dzongri], 13,000 ft, 15 Oct 1875, C.B. Clarke (K image; isolectotype: K image). No collections were cited in the protologue. Clarke is among sheets with a handwritten label by Clarke as A. himalaicus. Aster mekongensis Onno, Bibl. Bot. 106: LECTOTYPE (designated here): CHINA. Yunnan. Lichiang Range, 27 24' N, stony alpine pasture, 14,000 ft, Sep 1913, G. Forrest (E image; isolectotype: PE image-fig. 25). Without designating a type, Onno cited these collections by Forrest: 11137, 14241, 17230, 18892, 20710, 21984, and 22276, all from herb. E. Forrest (E) has Onno's annotation as "Aster mekongensis nova. sp." and matches his description of stems as ca. 15 cm tall. Grierson (1964), followed by Ling (1985) and Chen et al. (2011), cited "Aster mekongensis p.p." as a synonym of both A. himalaicus and A. bietii. Bhutan, China (Sichuan, Xizang, Yunnan), India, Myanmar, Nepal. 13. Tibetiodes jeffreyana (Diels) Nesom, comb. nov. Aster jeffreyanus Diels, Notes Roy. Bot. Gard. Edinb. 5: TYPE: CHINA. Yunnan. Base of Lichiang peak, Lichiang valley, marshy hillsides, 10,000 ft, Sep 1904, G. Forrest 669 (holotype: E image-fig. 25; isotypes: BM image, P image). China (Guizhou, Sichuan, Yunnan). 14. Tibetiodes labrangensis (Hand.-Mazz.) Nesom, comb. nov. Aster labrangensis Hand.-Mazz., Notizbl. Bot. Gart. Berlin-Dahlem 13: Aster yunnanensis var. labrangensis (Hand.- Mazz.) Y. Ling, Fl. Reipubl. Popularis Sin. 74: TYPE: CHINA. Xizang (Tibet). Eastern Tibet, grasslands between Labrang and Yellow River, in meadow of Wago (Waro) valley beyond Kadja, 11,500 ft, Jul 1926, J.F. Rock (holotype: B not seen; isotypes: E- image, GH image-fig. 17, W not seen). Protologue: "Rock 14447: Herb. Berlin, Mus. Wien." Aster kawaguchii Kitamura, Acta Phytotax. Geobot. 15: TYPE: CHINA. Xizang (Tibet). Circa Lhasa, Mt. Cho-mo-sese, 6 Sep 1914, E. Kawaguchi (holotype: SM [National Science Museum, Tokyo] 96879, Fig. 4 in the protologue). China (Gansu, Qinghai, Sichuan, Xizang) 15. Tibetiodes latibracteata (Franch.) Nesom, comb. nov. Aster latibracteatus Franch., J. Bot. (Morot) 10: LECTOTYPE (designated here): CHINA. Yunnan. Lieux frais un peu ombragés, au sommet du Tsang-chan, au dessus de Ta-li, 4000 m, 5 Aug 1884, R.P. Delavay 1207 (P image; isolectotypes: K image, P 11697, 11698, and 11699, images). Protologue: "Yun-nan, ad cacumina montis Tsang-chan, alt m. (R.P. Delavay, n. 1207, 2650)." China (Yunnan), Myanmar. 16. Tibetiodes latisquamata (C.E.C. Fischer) Nesom, comb. nov. Doronicum latisquamatum C.E.C. Fischer, Bull. Misc. Inform. Kew 1937: Aster platylepis Y.L. Chen [nom. nov.], Kew Bull. 39: (not Aster latisquamatus (Maxim.) Hand.-Mazz. 1938). TYPE: INDIA. Sikkim. Tsamgo Lake, common all around lake on rocky shale, 12,000 ft, 3 Aug 1935, C.S. Cutting and A.S. Vernay 20 (holotype: K image, photo-e!). Aster heliopsis Grierson, Notes Roy. Bot. Gard. Edinb. 26: TYPE: NEPAL. Lihkoo- Shailoong, 14,000-15,000 ft, 1930, Capt. L. Dhwoj 403 (holotype: E; isotypes: BM-2 images). China (Xizang), India.

6 6 17. Tibetiodes likiangensis (Franch.) Nesom, comb. nov. Aster likiangensis Franch., J. Bot. (Morot) 10: LECTOTYPE (designated here): CHINA. Yunnan. Les hauts côteaux du Li Kiang Suee Chan (Li Kiang), 4000 m, 14 Aug 1886, R.P. Delavay 2098 (P image; isolectotypes: A image, P and images). Aster costei H. Lév., Bull. Géogr. Bot. 25: Aster likiangensis subsp. costei (H. Lév.) Onno, Biblioth. Bot. 26 (Heft 106): Aster asteroides subsp. costei (H. Lév.) Grierson, Notes Roy. Bot. Gard. Edinb. 26: TYPE: CHINA. Yunnan. Pâtures du Io-Chan, 3300 m, 15 Aug 1912, E.E. Maire s.n. (holotype: E image). Bhutan, China (Sichuan, Xizang, Yunnan). 18. Tibetiodes lipskii (Komarov) Nesom, comb. nov. Aster lipskii Komarov, Bot. Mater. Gerb. Glavn. Bot. Sada R.S.F.S.R. 2: Not seen. According to Chen et al. (2011), the original description was based on seeds collected from near Qinghai Hu (Qinghai Province) and cultivated at the Komarov Botanical Gardens. Treated as a good species by Ling (1985) and by Chen et al. (2011). Two collections from the region of the type locality are morphologically distinctive and have been identified as Aster lipskii: Gansu. Lapuleng, weedy slope, 3800 m (12,500 ft), 19 Jul 1937, T.-P. Wang 7256 (PE, Fig. 14). Qinghai. Halihatu Mountain, Xireg, Ulan County, Chaidamu Basin, 26 Jul 1956, Qinghai Collecting Team 642 (PE, Fig. 15). China (Gansu, Qinghai). 19. Tibetiodes nepalensis (Grierson) Nesom, comb. et stat. nov. Aster falconeri subsp. nepalensis Grierson, Notes Roy. Bot. Gard. Edinb. 26: LECTOTYPE (designated here): NEPAL. Bhurchula Lekh, near Jumla, open slopes among dwarf rhododendrons, 12,500 ft, 11 Jul 1952, O. Polunin, W.R. Sykes, and L.H.J. Williams 4554 (E image; isolectotype: BM, as cited by Grierson). Grierson did not designate a type but cited many collections from Kumaun (India) and Nepal. India, Nepal. 20. Tibetiodes retusa (Ludlow) Nesom, comb. nov. Aster retusus Ludlow, Bull. Brit. Mus. (Nat. Hist.), Bot. 2: TYPE: CHINA. Xizang. SE Tibet, Kungbo, Nambu La, on rocks, 14,000 ft, 18 Jul 1947, F. Ludlow & G. Sherriff (holotype: BM image). China (Xizang). 21. Tibetiodes salwinensis (Onno) Nesom, comb. nov. Aster salwinensis Onno, Biblioth. Bot. 26 (Heft 106): 74, pl. IB LECTOTYPE (designated here): CHINA. Yunnan. No other locality data, , G. Forrest (K image, Fig. 22). There are 7 other K collections by Forrest with the same minimal label data, each with a different collection number, each with a handwritten annotation by Onno in 1930 as "Aster salwinensis Onno nova sp." China (Sichuan, Xizang, Yunnan), Myanmar. Phyllary morphology of T. salwinensis (Figs. 12A, 22) is unusual, as are the relatively broad, few ray flowers in a single series. But morphology otherwise seems characteristic of the genus and molecular data place it there. 22. Tibetiodes setchuenensis (Franch.) Nesom, comb. nov. Aster setchuenensis Franch., J. Bot. (Morot) 10: TYPE: CHINA. Sichuan. "Thibet Oriental, Ta-tsien-lou (Principauté de Kiala)," 1893, J.-A. Soulié 472 (holotype: P image). China (Sichuan).

7 7 23. Tibetiodes stracheyi (Hook. f.) Nesom, comb. nov. Aster stracheyi Hook. f., Fl. Brit. India 3(8): LECTOTYPE (designated here): INDIA. Uttarakhand. Kumaon, Pindari, 12,000 ft, 1848, R. Strachey & J.E. Winterbottom 2 (K image; isolectotypes: BM, K, P, images). Protologue: "WESTERN ALPINE HIMALAYA; Kumaon, Wallich; Pindari, alt. 12,000 ft., Str. & Wint., Stewart." Bhutan, China (Xizang), India, Nepal. 24. Tibetiodes techinensis (Y. Ling) Nesom, comb. nov. Aster techinensis Y. Ling, Fl. Reipubl. Pop. Sin. 74: TYPE: CHINA. Yunnan. Y. Atuntze, Yangtsatung, 3900 m, 24 Aug 1938, T.T. Yü 9828 (holotype: PE image; isotypes: A image, KUN image, PE-5 sheets images). China (Yunnan). 25. Tibetiodes tientschwanensis (Hand.-Mazz.) Nesom, comb. nov. Aster tientschwanensis Hand.- Mazz., Oesterr. Bot. Zeit. 89: TYPE: CHINA. Sichuan. Tien-chuan-hsien, 3340 m, 13 Jun 1936, K.L. Chu 2780 (holotype: B?; isotypes: E image, PE image). China (Sichuan). 26. Tibetiodes tsarungensis (Grierson) Nesom, comb. nov. Aster flaccidus subsp. tsarungensis Grierson, Notes Roy. Bot. Gard. Edinburgh 26: Aster tsarungensis (Grierson) Y. Ling, Fl. Reipubl. Popularis Sin. 74: TYPE: CHINA. Xizang (Tibet). Doker La, 28 20'N, 3960 m Aug 1918, G. Forrest (holotype: E image-fig. 19). China (Sichuan, Xizang, Yunnan). 27. Tibetiodes yunnanensis (Franch.) Nesom, comb. nov. Aster yunnanensis Franch., J. Bot. (Morot) 10: Aster diplostephioides var. yunnanensis (Franch.) Onno, Biblioth. Bot. 106: LECTOTYPE (designated here): CHINA. Yunnan. Les bois de Fang-yang-tchang, 3000 m, 19 Jul 1887, R.P. Delavay 3107 (P image; isolectotypes: K image, P and images). Protologue: "Likiang in pratis et silvis haud procul a nivibus perpetuis, alt m. (Delavay, n et 2482);" in silvis ad Kan-ho, prope Fang-yang-tchang, alt m. (id., n et 3701; ad collum Lo-pin-chan (id.)." Aster vilmorinii Franch., J. Bot. (Morot) 10: Erigeron vilmorinii (Franch.) Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 21: LECTOTYPE (designated here): CHINA. Sichuan. Ta-tsien-lou, 17 Jun 1891, M. Pratt 631 (P image; isolectotype: BM image). Protologue: "Se-tchuen occid., in silvis prope Ta-tsien-lou (R.P. Soulié, n. 418; Pratt, n. 631; Tongolo ad limites tibetana, in quercetis et betuletis (id., n. 505)." China (Sichuan, Yunnan). ACKNOWLEDGEMENTS Thanks to Dave Boufford (Harvard University Herbaria) for help in interpreting Chinese names and geography. Photos by David Bouffourd and Susan Kelley are from the "Biodiversity of the Hengduan Mountains and adjacent areas of south-central China" website. LITERATURE CITED Chen, Y.-L., L. Brouillet, and J.C. Semple Aster (Astereae). Pp , in Flora of China, Vol Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis). Grierson, A A revision of the asters of the Himalayan area. Notes Roy. Bot. Gard. Edinb. 26: Nesom, G.L. 2020a. Revised subtribal classification of Astereae (Asteraceae). Phytoneuron : 1 39.

8 8 Nesom, G.L. 2020b. The genus Aster (Asteraceae) in the strictest sense. Phytoneuron : Nesom, G.L. 2020c. Lectotypification of Heterochaeta DC. (Asteraceae). Phytoneuron : 1. Ostenfeld, C.H. and O. Paulsen A list of plants from Inner Asia collected by Dr. Sven Hedin (Asteraceae by Ostenfeld). Pp. 1 98, plates 1-8, in S. Hedin, Southern Tibet, Vol. 6. Onno, M Geographisch-morphologische Studien über Aster alpinus L. und verwandte Arten. Biblioth. Bot. 106: Zhang G.-J., H.-H. Hu, C.-F. Zhang, X.-J. Tian, H. Peng, and T.-G. Gao Inaccessible biodiversity on limestone cliffs: Aster tianmenshanensis (Asteraceae), a new critically endangered species from China. PLoS ONE 10(8): e Figure 2. Tibetiodes diplostephioides. Sichuan. Photo by Dave Boufford, 2005.

9 9 Figure 3. Tibetiodes diplostephioides. St. Andrews Botanic Garden. Photo by Kirsty Wilson, Figure 4. Tibetiodes yunnanensis. Sichuan. Photo by Susan Kelley, 2006.

10 10 Figure 5. Tibetiodes yunnanensis. Sichuan. Photo by Susan Kelley, 2006.

11 11 Figure 6. Tibetiodes labrangensis. Tibet. Photo by Dave Boufford, 2004.

12 12 Figure 7. Tibetiodes flaccidus. Sikkim. Photo by Harry Jans, 2018, inaturalist.

13 13 Figure 8. Tibetiodes flaccidus. Yunnan. Photo by Dave Boufford, Figure 9. Tibetiodes himalaicus. Nepal. Photo by T. Abe Lloyd, 2017.

14 14 Figure 10. Tibetiodes involucres. A. T. stracheyi. B. T. latisquamatum. C. T. brachytricha. D. T. himalaica. E. T. jeffreyana. F. T. yunnanensis.

15 15 Figure 11. Tibetiodes involucres. A. T. likiangensis B. T. tientschwanensis. C. T. retusa. D. T. asteroides. E. T. flaccidus. F. T. brevis.

16 16 Figure 12. Tibetiodes involucres. A. T. salwinensis. B. T. labrangensis. C. T. farreri.

17 17 Figure 13. Tibetiodes diplostephioides. Yunnan. Forrest (E).

18 18 Figure 14. Tibetiodes lipskii. Gansu. Wang 7256 (PE).

19 19 Figure 15. Tibetiodes lipskii. Qinghai. Qinghai Collecting Team 642 (PE).

20 20 Figure 16. Tibetiodes bietii. Yunnan. Rock 9728 (E).

21 21 Figure 17. Tibetiodes labrangensis. Tibet. Rock 14447, isotype (GH).

22 22 Figure 18. Tibetiodes brevis. Yunnan. Forrest 10516, asdfafd, Aster brevis (E).

23 23 Figure 19. Tibetiodes tsarungensis. Yunnan. Forrest 16873, Aster flaccidus subsp. tsarungensis, holotype (E).

24 24 Figure 20. Tibetiodes asteroides. Tibet. Lepcha 226 (K).

25 25 Figure 21. Tibetiodes likiangensis. Yunnan. Forrest 6001 (E).

26 26 Figure 22. Tibetiodes salwinensis. Yunnan. Forrest 19088, lectotype (K).

27 27 Figure 23. Tibetiodes stracheyi. Nepal. Hoshino et al (E).

28 28 Figure 24. Tibetiodes brachytricha. Yunnan. Rock 9728 (US).

29 29 Figure 25. Tibetiodes himalaicus. Yunnan. Forrest 11134, isolectotype of Aster mekongensis (PE).

30 30 Figure 26. Tibetiodes jeffreyana. Yunnan. Forrest 669, holotype of Aster jeffreyanus (E).