Triplostegia glandulifera Wallich

Niu, Yan-Ting, Barrett, Russell L., Zhang, Zhu-Zhi, Lu, Li-Min & Chen, Zhi-Duan, 2019, Taxonomic revision of Triplostegia (Caprifoliaceae: Dipsacales), Phytotaxa 392 (1), pp. 19-32 : 24-29

Triplostegia glandulifera Wallich in Candolle (1830: 642).

Type (lectotype, designated here):— NEPAL. Himalaya, Gopam Than, August–October 1821, [1824 according to Wallich herbarium, some material at K was received in 1832], N. Wallich 436 (G barcode G00354421 [digital image!, available at https://plants.jstor. org/stable/viewer/10.5555/al.ap.specimen.g00354421]). Isolectotypes: K barcodes K000779138 [digital image!, available at https:// plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.k000779138], K000779139, [digital image!, available at https://plants.jstor. org/stable/viewer/10.5555/al.ap.specimen.k000779139], K001111025, [digital image!, available at https://plants.jstor.org/stable/ viewer/10.5555/al.ap.specimen.k001111025].

– Triplostegia glandulosa Candolle (1832: 23 . t. 5), orth. var.

= Triplostegia grandiflora Gagnepain (1900 [1901 Jan.]: 333, pl. X).

Type (lectotype, designated here):— CHINA. Environs de Yunnan Sen, on mountain, 1 September 1897, Fr. Ducloux 399 (P barcode P02284027 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02284027]). Isolectotypes: P barcodes P02284028 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02284028], P02284029 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02284029].

= Hoeckia aschersoniana Engler & Graebner (1901 [Feb.]: 598).

≡ Triplostegia aschersoniana (Engl. & Graebn.) Diels (1912b: 256) .

Type:— CHINA. [Southern Szechaun], Nan ch‘uan, Lung mo ai, Weise, September, A. von Rosthorn 873 (type: B, n.v.).

= Triplostegia repens Hemsley (1901 [post March]: 101).

Type (lectotype, designated here [or perhaps holotype]):—British [Papua] New Guinea: Wharton Range, 11100 ft a.s.l., Aug. 1897, A. Giulianetti & A.C. English s.n. (K barcode K000779137 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/ al.ap.specimen.k000779137]).

= Triplostegia delavayi Franch. ex Diels (1912a: 209) .

Type (lectotype, designated here):— China: N.W. Yunnan, eastern flank of the Lichiang [Lijiang] Range, 10000–11000 ft, Aug. 1906, G. Forrest 2755 (E barcode E00265266 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen. e00265266]).

Residual syntypes:— China: W. Yunnan, on the eastern flank of the Tali Range , 9000–10000 ft a.s.l., July 1906, G. Forrest 4646 (E barcode E00265265 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.e00265265]; P barcode P02284025 [digital image!, available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02284025]) ; China: Yunnan, Mengtze [Mengzi], eastern mountains, 6000 ft [2,000 m], A. Henry 9441 (E barcode E00265267 [digital image!, available at https:// plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.e00265267]) .

Morphology:—Herbaceous perennial, erect, 15–45 cm tall, sometimes clonal. Taproot simple or bifurcate, brown, nearly fusiform, 30–50 mm long, 2–5 mm diam., slightly fleshy. Rhizomes thin, horizontal, 4-angular, short–elongate with 2–6 internodes, each 5–20 mm long, nodes rooted. Stems solitary, simple, slightly to distinctly 4-angular, glabrous or sparsely to densely white villous, often glandular hairy above (usually above the first inflorescence bracts). Basal leaves dense, rosulate; leaf blade serrate or pinnatifid; cauline leaves decussate, otherwise similar to the basal ones. Leaves opposite, sessile to petiolate; petiole 0–50(–80) mm long; leaf blade slightly amplexicaul, obovate to obovatelanceolate, 7–119 mm long and 3–47 mm wide, with base cuneate, apex rounded, pinnatifid or pinnatilobate, more deeply so at base; terminal lobe largest, all lobes serrate; abaxially villous on veins, adaxially sparsely villous or both surfaces villous, ultimately glabrescent; margin irregularly lobed or serrate; upper leaves gradually smaller and becoming bractlike, lobed, sessile. Inflorescences thyrsoidal, branches compact to elongate, flowers in remote, terminal, simple or compound dichasia, all parts (except corolla) densely white villous and glandular hairy; bracts leaf-like to linear, 2–20 mm long, midvein prominent, lamina sometimes white villous and glandular hairy, margin shallowly serrate or entire, sparsely ciliate, apex obtuse; pedicels 1–3 mm long; involucels (epicalyx whorls) 2, urceolate, outer involucel 4-lobed; lobes lanceolate, 1.5–3 mm long, abaxially densely covered with purple (to black) gland-tipped hairs; inner involucel urceolate to ovoid, c. 1.5 mm long, hirsute, 8-ribbed, apex acute or constricted into a short limb; limb (5–)8-toothed or (5–)8-dentate, teeth acute at base. Flowers nearly actinomorphic. Calyx reduced, 4 (possibly rarely 5)-lobed. Corolla white or rose, tubular-funnelform, (4)5-lobed, 3–12 mm long, outside sparsely villous; lobes subequal, c. 1/3 as long as total length of corolla, apex obtuse, imbricate in bud. Stamens 4, equal in length, alternate with corolla lobes, slightly to prominently exserted; filaments 3–9 mm long; anthers introrse, versatile, white to yellow; pollen all tricolpate, oblate spheroidal with a tectate, rather thick exine with prominent, acute or obtuse spines and micro-spines. Ovary enveloped by urceolate involucel. Style solitary, slightly shorter than to longer than stamens; stigma capitate. Achenes 8-ribbed, enveloped by urceolate involucels; involucels 3–4 mm long in fruit, glandular hairy, 4-lobed; lobes long acuminate, hooked or not. Seeds subglobose, acute at both ends, smooth, with 2 obscure angles; endosperm copious; embryo small.

Typification:—We have chosen the sheet G00354421 as the lectotype of the name T. glandulifera as this is the material examined by Candolle. Several plants are mounted on the sheet and there is a flowering inflorescence in a packet. The three sheets at K (K000779138, K000779139 and K001111025) are clearly duplicate specimens, so they are considered isolectotypes.

The sheet chosen as lectotype of T. grandiflora (P02284027) bears two whole plants, both in flower, and a detailed illustration of floral parts drawn by Gagnepain. The two sheets recognised as isolectotypes (P02284028 and P02284029) are clearly duplicates, noting that P02284028 bears an erroneous collection number, 339 rather than 399.

When describing Hoeckia aschersoniana, Engler & Graebner (1901: 598) only listed a single collection by A. von Rosthorn 873 (B), which was also cited by Diels (1912b: 256). We have been unable to locate this specimen, or any duplicates, and it has possibly been destroyed. While designation of a lectotype may seem desirable, until original material can be located, we do not consider this to be appropriate.

Hemsley (1901: 101) simply cited ‘Wharton Range, 11100 ft. ’ The specimen chosen as lectotype of T. repens (K000779137) is the only one found from this location. It has a copy of the original description attached to the sheet. The sheet chosen as lectotype of T. delavayi (E00265266) has three whole plants, all in flower. A designation as ‘lectotype’ has been added by C.G.C.J. van Steenis in 1957, however we have found no evidence that this was ever published. We agree that this sheet is a good choice as lectotype due to the quality of the specimen and formally designate it as such here.

While there are two collections at P (mounted on four sheets) by J.M. Delavay, to whom the species is dedicated, they were not cited by Diels (1912), so they are not considered syntypes as there is no direct evidence that they were seen by Diels or included in his concept of the species. The specimens do bear the name T. delavayi , but this was probably added by Franchet, not Diels.

Notes:—The precise publication date of three synonyms is important for establishing the priority of names. The cover page for the Bulletin de la Société Botanique de France vol. 47, fascicle 8, states that it was printed on 31 January, 1901. Stafleu & Mennega (1998) list Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie vol. 29 part 5 as published in February 1901. The volume Bulletin of Miscellaneous Information (Royal Botanic Gardens, Kew) for 1899 simply states that it was printed in 1901; however on page 81 of the volume for 1901, it is noted that ‘the volume for 1899 will shortly be issued’ [apparently read to the Linnaean Society in March 1901]. This would suggest it was published after March 1901.

Gagnepain (1901) described T. grandiflora in Dipsaceae (as “Dipsacées”), differentiating the species from T. glandulifera on account of the leaf morphology and the length of the corolla. Gagnepain (1901) commented on the bizarre appearance of the genus, allying it to Scabiosa L. based on pollen morphology.

While it may be possible that larger flowers are produced by polyploid plants, there is currently no evidence for this in Triplostegia . If this was the case, there would need to be multiple cases of polyploid events to explain the current distribution of large flowered populations.

It has generally been overlooked that Diels (1912b) recombined Hoeckia aschersoniana into Triplostegia , differentiating it from T. delavayi based on its smaller flowers and inconspicuous inner calyx. There is no direct reference to the basionym, however it is considered that an indirect reference is made through the inclusion of ‘Engl. & Grabn.’, validating the new combination ( Turland et al. 2018; Art 41.3).

Triplostegia repens was described by Hemsley (1901) based on a small specimen without mature flowers, distinguished from Chinese plants based on the small habit, a feature also noted for New Guinea specimens by van Steenis (1951). Merrill & Perry (1949) noted that the species is very variable in habit depending on the environmental conditions. A representative set of collections from New Guinea was examined at CANB, but no morphological characters were identified that suggest it should be recognised as a distinct taxon ( Fig. 2 & Table 2).

Triplostegia delavayi was described in Dipsacaceae , separated from T. glandulifera by larger flowers and more deeply incised leaves ( Diels 1912a: 209).

Phenology:—Flowers and fruits mostly from July–October, but also record for April and December, so possibly all year round under favourable conditions.

Cytology:—2 n = 18 ( Goldblatt 1984).

Habitat:—Grows in a variety of habitats including forests, forest margins, grassy slopes, meadows and along streams; 1500–4000 m a.s.l.

Conservation:—While not threatened as a species, and not listed under IUCN criteria, some populations of Triplostegia do require protection from over-collection for medicinal use ( Niu et al. 2018a).

Ethnobotany:— Triplostegia is utilised as medicinal plants, with no distinction in use between the morphotypes previously recognised as two different species ( Ma et al. 1992, Liu et al. 2012).

Distribution:—From the eastern Himalaya ( India and Nepal) northward to Gansu ( China), southward to Malaysia, with disjunct populations eastward to Taiwan and further south in Indonesia and Papua New Guinea ( Fig. 3 ). The disjunctions are likely to be the result of recent range contractions ( Niu et al. 2018a).

Selected specimens examined:— BHUTAN (1 of 6). Thimphu: summit of Dochong La, 27°29’N, 89°45’E, 3080 m a.s.l., I.W.J. Sinclair & D.G. Long 4752 (E).

NEPAL (8 of 28). Gandaki: Kaski District, Ghodepani , 2760 m a.s.l., 28 August 1994, M. Mikage et al. 9485559 (L, TI, n.v.) ; Kaski District, Banthanti, Deurali & Ghodepani Deurali , 29°20’N, 83°40’E, 2700 m a.s.l., 24 August 1988, M. Suzuki et al. 8881333 (L, TI, n.v.) ; Gorkha District, Lho to Sama Gaon , 28°34’27”N, 84°40’13”E, 3510 m a.s.l., October 2012, H. Ikeda et al. 20811134 (E) ; Gorkha District, Lho to Sama Gaon , 28°34’19”N, 84°42’23”E, 3100 m a.s.l., October 2012, H. Ikeda et al. 20812155 (E) ; Koshi: Sankhuwasabha District, Camp site, 27°49’N, 87°22’E, 3140 m a.s.l., 22 August 1998, S. Noshiro et al. 9830117 (L, TI, n.v.) ; Sagarmatha: Eastern Development Region, Solu Khumbu, Namche Bazar , Hillside above Namche on the route Tengboche , 27°48’22”N, 86°42’54”E, 3500 m a.s.l., 13 September 2005, M.F. Watson et al. DNEP3 AY26 (E) ; Bagmati: Sindhupalchok District, Langmara Kharka , 27°55’42”N, 85°50’22”E, 3043 m a.s.l., 11 September 2011, M.F. Watson et al. EKSIN19 (E) ; Rasuwa District, Sunchet Kharka (2530 m a.s.l.)— Gopchet Kharka (3390 m a.s.l.)— Lipchet Kharka (3580 m a.s.l.), 28°07’N, 85°07’E, 2640 m a.s.l., 15 August 1994, F. Miyamoto et al. 9440174 (E) .

INDIA (4 of 4). Sikkim: Zema Valley , 12000 ft a.s.l., 20 July 1909, Smith & Cave 2757 (L) ; below Jauga , 10000 ft a.s.l., 16 August 1909, Smith & Cave 2527 (L) ; 9–12000 ft a.s.l., n.d., J.D. Hooker s.n. (U.1749651); 10–12000 ft a.s.l., 1857, T. Thomson s.n. (L).

CHINA (71 of 410): Xizang: Chimo Dzong, Cyamda Chu , 29°38’N, 94°16’E, 10000 ft a.s.l., 19 August 1938, 6098 (E) ; Bomi, south bank of Guxiang lake , 29°55’N, 95°30’E, 3100 m a.s.l., 27 August 1965, J.S. Ying & D.Y. Hong 651248 ( PE) ; Chayu, Shangshiyu, Suku , 28°39’36”N, 97°28’12”E, 2800 m a.s.l., 7 August 1973, Qingzang team 73- 1003 ( KUN) ; Chi-na-tung, Tsa-wa-rung, 28°39’36”N, 97°28’12”E, 2800 m a.s.l., August 1935, C.W. Wang 65229 ( PE) ; Cuona, Mama, Chafang , 27°59’24”N, 91°57’36”E, 2800 m a.s.l., 26, August 1975, Qingzang team 751688 ( KUN) ; Gengzhang , 29°43’48”N, 94°5’24”E, 3300 m a.s.l., 31 July 1974, Anon. 2060 ( PE) ; Gongbo Gyamda, 3 km from Bahe ( Beba ) Zeng (town) on road to Basong Hu ( Basum Tso ) ( Lake ), 29°53’26”N, 93 39’23”E, 3275 m a.s.l., 14 August 2000, D.E. Boufford et al. 30037 (E) ; Linzhi, below Nixi , 29°38’24”N, 94°21’36”E, 3200 m a.s.l., 29 July 1965, Anon. 2339 ( KUN) ; Linzhi, near the Western medicine factory, 29°38’24”N, 94°21’36”E, 3200 m a.s.l., 3 August 1965, 2420 ( KUN) ; Linzhi, in the Bayi College of agriculture and animal husbandry, 29°38’24”N, 94°21’36”E, 2970 m a.s.l., 29 July 1989, K. Yao 1950 ( NAS) ; Linzhi, western slope of Mt. Shejila , 29°38’24”N, 94°21’36”E, 3700 m a.s.l., 2 August 1975, Qingzang team 751130 ( PE) ; Milin, station of Wolong 159 regiment, 29°13’12”N, 94°12’36”E, 3100 m a.s.l., 9 August 1982, Z.C. Ni et al. 2857 ( PE) ; Milin, Jiage river , 29°13’12”N, 94°12’36”E, 3450 m a.s.l., 13 July 1972, Xizang Chinese herbal medicine census team 3758 ( PE) ; Milin, Hongwei forestry station, 29°13’12”N, 94°12’36”E, 3100 m a.s.l., 29 July 1975, Qingzang team 750982 ( KUN) ; Motuo, Gedangxing , 29°19’48”N, 95°19’48”E, 2400 m a.s.l., Qingzang team 74-4934 ( PE) ; Nielamu, Zhangmu , 27°59’24”N, 85°58’48”E, 2810 m a.s.l., 14 August 1972, Xizang Chinese herbal medicine census team 171 ( PE) ; Yadong, Renqinggang , 27°28’48”N, 88°54’36”E, 2900 m a.s.l., 24 July 1960, Xizang investigation team 438 ( PE) ; Yunnan: Environs de Yunnan Sen , on mountain, 10 April 1904, Fr. Ducloux 2575 (L, P) ; Vicinity of Yunnan Sen , [rec. Nov. 1906], E.E. Maire 75 (L) ; N.W. Yunnan, 27°20’N August 1906, A.K. Bulley et al. 2683 (E); Chuxiong Yi Aut. Pref., Xiang Yun, Kunming to Dali Road , Tiang Shen Tung Village , 25°19’5”N, 101°1’56”E, 2480 m a.s.l., 4 September 1997, Gaoligong Shan Expedition 8502 (E) ; Dali, stony pastureland on the eastern flank of the Tali Range , 25°40’N, 9–10000 ft a.s.l., July 1906, A.K. Bulley et al. 4646 (E) ; Deqin , A-tun-tze, 27°49’N, 99°42’E, 2700 m a.s.l., September 1935, C.W. Wang 69884 ( KUN) ; Dongchuan, Guomindazhu , 26°4’48”N, 103°11’24”E, 2800 m a.s.l., 12 August 1984, S.B. Lan 95 ( PE) ; Eryuan, colliery in Sanyingdongshan , 23 September 1975, B.Y. Qiu 60962 ( KUN) ; Pi-lo-shan, Che-tse-lo, 4000 m a.s.l., 14 September 1934, H.T. Tsai 58589 ( KUN) ; Heqing, Lianping, Niaoershan , 26°33’36”N, 100°10’48”E, 3100 m a.s.l., R.C. Qin 23976 ( PE) ; Huize, Dahaihengshan , 26°13’48”N, 103°15’36”E, 3400 m a.s.l., N.E. Dian team 308 ( KUN) ; Kunming Municipality, Qiongzhusi ( Bamboo Temple ), 25°04’N, 102°37’E, 2300 m a.s.l., 30 July 1984, 1984 Sino-Amer. Bot. Exped. 1469 (E) ; Lijiang, Xuesong Village , 2800 m a.s.l., 13 August 1937, Yu 15428 ( KUN) ; Lijiang, western slope of Mt. Yulong, Shangmoguxi Village , 2100 m a.s.l., 24 September 2007, Peng Hua et al. 6675 (E) ; Lushui, Pianma, way from Yakou to Erdao Yakou , 3500 m a.s.l., 17 August 1964, S.G. Wu 8461 ( KUN) ; Zhongdian Xian, Dongwang, Sichuan-Yunnan border area, Village of Dongwang on N side of pass over Daxue Shan , 28°37’8”N, 99°49’17”E, 3700 m a.s.l., 26 July 1998, D.E. Boufford et al. 29245 (E) ; Sichuan: Dujiangyan Municipality (formerly Guan Xian) ; between Guanmengzi and Guihuashu, upstream from Longxi along the Longxi River and on adjacent slopes, 1300 m a.s.l., 3 September 1988, D.E. Boufford & B. Bartholemew 24461, G. Li & G.H. Zhu (A, n.v., CAS, L, PE) ; Baiyu Xian, Gan-Bai Lu ( Road from Baiyu to Ganzi ) then S on narrow side road off Ou Qu ( Ou River ) at Zhandu Xiang along branch of the Ou Qu in Ase Gou ( Ase Gorge ), 31°5’50”N, 98°58’45”E, 3250 m a.s.l., 23 August 2006, D.E. Boufford 37198 ( PE) ; Dajin, Wanlixiang, Wanlichenggou , 3200 m a.s.l., 24 July 1958, A. Li 78190 ( WUK) ; Tah-Liang-shan, 2500 m a.s.l., 31 August 1934, T.T. Yv 3930 ( PE) ; Daocheng, Julongxiang, 3600 m a.s.l., 17 July 1973, Plant investigation team of Sichuan province 2136 ( CDBI) ; Omei-hsien, Mt. Omei , 4500–6000ft a.s.l., 19 August 1928, W.P. Fang 3185 (E) ; Guan, Guangguangshan , 2100 m a.s.l., 25 August 1987, D.Z. Fu & Z.L. Zhao 87-2072 ( HX) ; Huidong , 2500 m a.s.l., 12 September 1961, T.L. Dai & L.P. Tang 11918 ( NAS) ; Huei-li Hsien , 2300 m a.s.l., 6 September 1932, T.T. Yv 1434 ( PE) ; Dajin , 3000 m a.s.l., 31 July 1958, A. Li 78237 ( PE) ; Kang-ting-hsien, 2600–3500 m a.s.l., 3 September 1934, C.S. Liu 1438 ( PE) ; Kangding, Zheduotang , 3300 m a.s.l., 3 August 1963, K.J. Guan & W.C. Wang 1349 ( PE) ; Luding Xian, Moxi, Gongga Shan , east side, downstream from Hailuo Guo Glacier , vicinity of Camp Number 2, 29°35’11”N, 102°1’40”E, 2430–2620 m a.s.l., 19 August 1997, D.E. Boufford et al. 27338 (E) ; Miyaluo, Qiangou , 2750 m a.s.l., 6 August 1956, D.P. He 45588 ( QTPMB) ; Tianyang Xian, Muyepeng, East side of Erlang Shan , 29°54’47”N, 102°12’45”E, 2580 m a.s.l., 13 August 1997, D.E. Boufford et al. 27156 (E) ; Wuxi, Gaolouxiang, Fenghuangtou , 2000 m a.s.l., 21 August 1958, G.H. Xiang 65063 ( IBSC) ; Xiangcheng Xian, Reda Lincha upstream on Mayi River from the town of Reda , 29°8’31”N, 99°38’59”E, 3450–3500 m a.s.l., 17 July 1998, D.E. Boufford et al. 28851 (E) ; Xiangcheng Xian, Maxiong Gou , 29°8’28”N, 99°57’52”E, 3700 m a.s.l., 9 July 1998, D.E. Boufford et al. 28440 (E) ; Xiangcheng Xian, Reda, Vicinity of the town of Reda , 29°6’11”N, 99°37’55”E, 3450–3650 m a.s.l., 16 July 1998, D.E. Boufford et al. 28729 (E) ; Yajiang Xian, Road (highway 318) from Yajiang to Litang , 29°59’20”N, 100°56’7”E, 3480–3640 m a.s.l., 6 August 2006, D.E. Boufford et al. 35871 ( PE) ; Chongqing: Chengkou, Qigan Mountain, Fangshengping , 2050 m a.s.l., 3 September 1958, T.L. Dai 102351 ( PE) ; Chengkou, Houping Xiang, Longpanshui , 1250 m a.s.l., 30 July 1958, T.L. Dai 10578 ( PE) ; Chengkou, Caimengqu , 1850 m a.s.l., 27 September 1958, T.L. Dai 106151 ( PE) ; Fengjie, Xinglong Maocao Zhen , 1740 m a.s.l., 27 September 1964, H.C. Zhou & H.Y. Su 111208 ( PE) ; Wushan, Zhuxiangongshe , 1280 m a.s.l., 22 August 1964, H.C. Zhou & H.Y. Su 110153 ( PE) ; southern Gansu: Dangchang, Xigu, Chaganglidongshan , 2200 m a.s.l., 13 September 1939, W.Y. Hsia 6740 ( PE) ; Wen, Tielougongshe, Yanggashan , Dahaigou , 2250 m a.s.l., 9 August 1967, C.L. Tang 1698 ( WUK) ; Xigu, Gongbasihe, Kazizhuang , 2200 m a.s.l., 22 August 1951, J.B. Wang 14881 ( WUK) ; Zhouqu, near Chagangxiang , 33°35’93”N, 104°19’30”E, 2000 m a.s.l., 20 July 1998, Bailongjiang investigation team 447 ( PE) ; southern Shanxi: Pingli, Qianjiaping forestry station, 31°0.76’N, 109°16.828’E, 1730 m a.s.l., 7 July 2004, Y.S. Chen et al. 1035 ( WUK) ; western Hubei: Shennongjia Forest District , 31°30’N, 110°30’E, 2000 m a.s.l., 15 September 1980, 1980 Sino-Amer. Exped. 1349 (E) ; Shennongjia, Hongping forestry station, 2150 m a.s.l., 11 August 1976, Hubei Shennongjia plant investigation team 32351 ( PE) ; Shennongjia, Tianjia Mountain pharmaceutical factory, 1680 m a.s.l., 29 August 1976, Hubei Shennongjia plant investigation team 32743 ( PE) ; Shennongjia, Xiaojiu Lake , 1850 m a.s.l., 23 August 1976, Hubei Shennongjia plant investigation team 11487 ( PE) ; Guizhou: Jiangkou Xian, Fanjing Shan , vicinity of summit, 2000–2300 m a.s.l., 29 August 1986, Sino-American Guizhou Botanical Expedition 665 ( PE) ; Taiwan: Central Formosa, from Ton Pou [?Tongluo] to Rossison , 2600 m a.s.l., 14 August 1966, C.G.G.J. van Steenis 20906 (L) ; Nantou Hsien, Jenai Hsiang, Tienchih-Boundary stone table, 2850 m a.s.l., 7 September, T.Y.A. Yang et al. 13946 ( PE) .

INDONESIA (4 of 5). Sulawesi Selatan: Latimojong Ranges, Pos 7 on track from Karangan to Gunung Rantemario , 3132 m a.s.l., 8 July 2002, G. Brown, L. Craven & L. Juswara 69 (BO, n.v., CANB) ; G. Rantemario, Latimojong Range , 3250 m a.s.l., 10 February 1981, J.M.B. Smith 683 (L) ; Ertsberg: 200 m W of the Ertsberg, Carstensz Mountains, West Irian , 3480 m a.s.l., 11 December 1971, G.S. Hope ANU 10863 ( CANB) ; Mt Trikora: Second Valley W of Somalak Valley, 3550 m a.s.l., 13 August 1984, J.-M. Mangen 1048 (L) .

PAPUA NEW GUINEA (5 of 69). Isuani Basin: SE flank of Mount Victoria, Central District , 2800 m a.s.l., 16 July 1974, L.A. Craven 3116 (CANB, LAE, n.v.) ; Wabag: Northern slopes of Sugarloaf complex (along Wapu River ), Wabag subdistrict, Western Highlands, c. 9500 ft a.s.l., 12 July 1960, R.D. Hoogland & R. Schodde 7028 (A, n.v., BISH, n.v., BH, n.v., CANB, L, LAE, n.v.) ; Laiagam: Yobobos grassland area, source of Lagaip River, Laiagam subdistrict, Western Highlands, c. 8500 ft a.s.l., 19 August 1960, R.D. Hoogland & R. Schodde 7507 (CANB, L, LAE, n.v.) ; Monarauwe: Salawaket Range, Huon Peninsula , Morobe District, c. 9500 ft a.s.l., 31 August 1964, R.D. Hoogland 9706 (CANB, L, LAE, n.v., US, n.v.) ; West Sepik: Star Mountains, West Sepik District, Camp 1, Tel Basin , 2960 m a.s.l., 1 April 1975, J.F. Veldkamp 6230 (CANB, L) .

MYANMAR (2 of 3). Mt. Victoria: 9000 ft a.s.l., 28 October 1956, F. Kingdon Ward 22799 (E); Mindat: 5000 ft a.s.l., 16 July 1956, F. Kingdon Ward 22488 (E).

Figures

FIGURE 2. Triplostegia glandulifera. Representative specimens from Papua New Guinea. A, B: Leaves. C: Stem indumentum. D–F: Inflorescence, floral bracts, epicalyx and flowers. Vouchers: A, C, D: R.D. Hoogland & R. Schodde 7507; B: R.D. Hoogland 9706; E, F: R.D. Hoogland 7902 (all CANB).

FIGURE 3. Distribution range of Triplostegia. Red circles represent locations of specimens with detailed records.