Research Article |
Corresponding author: Ren-Chao Zhou ( zhrench@mail.sysu.edu.cn ) Corresponding author: Ying Liu ( liliumrosa@163.com ) Academic editor: Ricardo Kriebel
© 2019 Qiu-Jie Zhou, Jin-Hong Dai, Che-Wei Lin, Tetsuo Denda, Ren-Chao Zhou, Ying Liu.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Zhou Q-J, Dai J-H, Lin C-W, Denda T, Zhou R-C, Liu Y (2019) Recircumscription of Bredia and resurrection of Tashiroea (Sonerileae, Melastomataceae) with description of a new species T. villosa. PhytoKeys 127: 121-150. https://doi.org/10.3897/phytokeys.127.36608
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Bredia (Melastomataceae) is an Asian genus that extends from central and southern mainland China to Taiwan and the Ryukyu islands. Molecular phylogenetic analyses reveal that the type of Bredia is nested in a clade of 20 species, while Tashiroea, a genus previously synonymized in Bredia, falls in another distantly related clade of 10 species. Our morphological survey shows that the two clades can be distinguished by several diagnostic features including leaf indumentum, texture, leaf surface sculpture under SEM, presence/absence of yellowish uniseriate trichomes, and capsule morphology. Based on molecular and morphological evidence, Bredia is recircumscribed and Tashiroea is resurrected. Description and a list of species are provided for the two genera with the description of a new species, T. villosa.
Bredia, Tashiroea, Sonerileae, Melastomataceae, morphology, taxonomy
Bredia Blume (Sonerileae, Melastomataceae) was originally described based on B. hirsuta Blume (
The generic circumscription of Bredia has long been problematic. Bredia is morphologically closely related to Phyllagathis Blume (
A recent molecular phylogenetic study, with extensive sampling of Bredia (19 species) and Phyllagathis (35 species), has shed new light on their generic delimitation (
In this paper, we revisited the morphological characters of the Bredia and Tashiroea clades in search of possible diagnostic characters. The results are presented below. Based on molecular and morphological evidence, Bredia is recircumscribed and Tashiroea is resurrected. Descriptions and a list of species are provided for the two genera with the description of a new species T. villosa. Thirteen new combinations are made.
Eight newly sequenced species were added to the combined dataset (nrITS and chloroplast trnV-trnM) published in
All species of the Bredia clade (20 species) and the Tashiroea clade (10 species) were examined, including B. changii W.Y. Zhao, X.H. Zhan & W.B. Liao, B. dulanica C.L. Yeh, S.W. Chung & T.C. Hsu, B. esquirolii (H. Lév.) Lauener, B. gibba Ohwi, B. hirsuta, B. longiloba (Hand.-Mazz.) Diels, B. microphylla H.L. Li, B. oldhamii Hook. f., B. repens R.C. Zhou, Q.J. Zhou & Ying Liu, B. rotundifolia Yan Liu & C.H. Ou, B. tuberculata (Guillaumin) Diels, B. yunnanensis, P. fordii (Hance) C.Chen, P. gracilis (Hand.-Mazz.) C. Chen, P. guidongensis K.M. Liu & J. Tian, P. latisepala C. Chen, P. longearistata C. Chen, P. longiradiosa C. Chen, P. plagiopetala C. Chen and P. velutina (Diels) C. Chen from the former clade, and B. amoena Diels, B. biglandularis C. Chen, B. okinawensis (= T. okinawensis), B. quadrangularis Cogn., B. sessilifolia H.L. Li, B. sinensis (Diels) H.L. Li (= T. sinensis Diels), B. yaeyamensis (= T. yaeyamensis), P. nudipes C. Chen, P. oligotricha Merr. and the new species T. villosa from the latter clade. Their habit, indumentum, shape and texture of the leaves, leaf surface sculpture under scanning electronic microscope (SEM), inflorescence type, stamen morphology, capsule morphology, habitat preference and geographical distribution were recorded. Data were obtained via field, herbarium and literature surveys as well as by observing living plants in the facilities of Sun Yat-sen University. Specimens of the two clades (GXMG, GXMI, HNNU, IBG, IBK, IBSC, JJF, KUN, NAS, PE, SYS) or their high-resolution photos (A, BM, CSFI, E, HAST, K, KYO, MO, NTUF, NY, PH, TAI, TAIF, TI, UC, WU) were examined. Habit, shape and texture of the leaves, and inflorescence type were obtained via visual observation. Stamen morphology and capsule morphology were determined by stereomicroscopic (Leica S8APO) examination, leaf epidermal features by desktop SEM (Phenom Pro), and indumentum by both equipment. During observation using Phenom Pro, fresh or dried tissues were directly mounted on stubs and examined without further processing. We failed to obtain the materials of two species recorded in Bredia, namely B. laisherana C.L. Yeh & C.R. Yeh and B. violacea H.L. Li, which were therefore not included in our molecular phylogenetic study. For the two species, we examined their protologue, images of herbarium specimens and color photos from the author of B. laisherana. Species circumscriptions basically follow
Diels published several names in Melastomataceae in 1924 and 1932. Although he designated type specimens, he did not specify the place of deposition for them in the protologue. Even if we assume the holotypes existed in the Berlin herbarium (where Diels worked), they were probably destroyed during the Second World War. Therefore, we follow
The combined dataset contained 1768 characters. The phylogenetic tree resulted from ML analysis is shown in Fig.
Maximum likelihood phylogenetic tree of Sonerileae/Dissochateae based on combined dataset of nuclear ribosomal internal transcribed spacer and chloroplast trnV-trnM sequences, showing the phylogenetic position of Tashiroea clade and Bredia clade. Numbers on the branches are bootstrap values obtained from maximum likelihood analyses (left) and Bayesian posterior probabilities (right) resulting from Bayesian inference. Boxes denote the types of the genera sampled.
Characters of the Tashiroea clade (10 species) and the Bredia clade (20 species) are summarized in Table
Comparison of the Tashiroea clade and the Bredia clade. Potential diagnostic characteristics are indicted in bold.
Tashiroea clade | Bredia clade | |
---|---|---|
Habit | Shrubs or shrublets | Shrubs, shrublets or herbs |
Indumentum of mature leaf | Glabrous, rarely pubescent and villous (T. villosa) | Sparsely to densely puberulous or strigose |
Leaf texture | Stiffly papery to leathery | Papery, rarely submembranous |
Leaf surface sculpture | Furrowed, rarely not (T. villosa) | Not furrowed |
Leaf shape | Lanceolate, ovate, elliptic or suborbicular | Lanceolate, ovate, cordate, oblong, elliptic, ovate-orbicular |
Leaf base | Cuneate, obtuse, rounded, or subcordate | Cuneate, obtuse, rounded, often cordate |
Leaf veins | 3–5 | 5–11, rarely 3 (B. guidongensis) |
Leaf margin | Entire to serrulate | Entire to serrulate |
Indumentum | Sessile gland with thin-walled head; multiseriate trichomes with glandular head or not; appressed uniseriate trichomes with glandular head; yellowish uniseriate trichomes | Sessile gland with thin-walled head; multiseriate trichomes with glandular head or not; appressed or spreading uniseriate trichomes with glandular head or not |
Inflorescence | Cymose, cymose panicle | Cymose, umbellate, or cymose panicle |
Stamens | Dimorphic and isomorphic | Dimorphic and isomorphic |
Staminal appendage of dimorphic stamens | Gibbose, tuberculate or spurred at the base of anthers in shorter stamens; connectives decurrent and prolonged, gibbose or tuberculate dorsally in longer stamens | Gibbose, tuberculate or spurred at the base of anthers in shorter stamens; connectives decurrent and prolonged, gibbose, tuberculate, or spurred ventrally in longer stamens |
Staminal appendage of isomorphic stamens | Slightly gibbose ventrally at the base of the anthers, spurred dorsally | Gibbose or tuberculate at the base of the anthers, sometimes short spurred dorsally |
Ovary top at anthesis | Slightly crowned or uncrowned | Crowned |
Capsule | Ovary crown usually evanescent, capsule uncrowned with rounded or 4-humped top; or ovary crown persistent and enlarged, enclosing an obpyramidal space (in T. nudipes and T. oligotricha) | Ovary crown persistent and enlarged, enclosing an inverted frustum-shaped depression at capsule top |
Habitat | Open or dense forests, slopes, stream banks, alt. 50–2300 m | In forest or along forest margin, stream banks, damp places, alt. 100–2500 m |
Distribution | South eastern mainland China, Taiwan, Ryukyu Islands | Central and south mainland China, Taiwan, Ryukyu Islands |
Scanning electron micrographs of lower epidermis. A–D Tashiroea clade, showing presence of furrowed surface sculpture: A T. amoena (Y. Liu 571) B T. nudipes (Y. Liu 435) C T. villosa (Y. Liu 568) D T. yaeyamensis (Y. Liu 631) E–L Bredia clade, showing absence of furrowed surface sculpture E B. dulanica (Y. Liu 565) F B. gibba (Y. Liu 566) G B. gracilis (Y. Liu 457) H B. hirsuta (Y. Liu 563) I B. longiradiosa var. pulchella (Y. Liu 485) J B. repens (Y. Liu 558) K B. tuberculata (Y. Liu 629) L B. yunnanensis (Y. Liu 627). Scale bars: 100 μm.
Trichomes in the Tashiroea clade and the Bredia clade. A–D Micrographs of trichomes in the Tashiroea clade: A sessile gland with thin-walled head in T. nudipes (Y. Liu 435) B multiseriate trichomes in T. amoena (Y. Liu 571) C appressed uniseriate trichomes with glandular heads in T. nudipes (Y. Liu 435) D yellowish uniseriate trichomes (branched or unbranched) in T. quadrangularis (Y. Liu 585). E–H Micrographs of trichomes in the Bredia clade: E sessile gland with thin-walled head in B. longiradiosa var. pulchella (Y. Liu 485) F multiseriate trichomes in B. repens (Y. Liu 558) G appressed uniseriate trichomes with glandular heads in B. gibba (Y. Liu 566) H spreading uniseriate trichomes in B. hirsuta (Y. Liu 563). I–L Stereoscopic images of the Tashiroea clade, showing buds covered in yellowish uniseriate trichomes I T. amoena (Y. Liu 571) J T. nudipes (Y. Liu 435) K T. quadrangularis (Y. Liu 585) L T. yaeyamensis (Y. Liu 631). M–P Stereoscopic images of the Bredia clade, showing buds without yellowish uniseriate trichomes: M B. dulanica (Y. Liu 565) N B. gibba (Y. Liu 566) O B. gracilis (Y. Liu 457) P B. yunnanensis (Y. Liu 627). Scale bars: 100 μm (A–H); 1 mm (I–P).
Stamen morphology in the Bredia clade. A–E Dimorphic stamens: A B. esquirolii (Y. Liu 587) B B. hirsuta (Y. Liu 563) C B. hirsuta var. scandens (Y. Liu 539) D B. rotundifolia (Y. Liu 538) E B. tuberculata (Y. Liu 629). F–O Isomorphic stamens: F B. fordii (Y. Liu 444) G B. fordii var. micrantha (Y. Liu 580) H B. gracilis (Y. Liu 457) I B. guidongensis (Y. Liu 472) J B. longearistata (Y. Liu 496) K B. longiradiosa var. pulchella (Y. Liu 485) L B. microphylla (Y. Liu 551) M B. plagiopetala (Y. Liu 460) N B. repens (Y. Liu 558) O B. yunnanensis (Y. Liu 627). Scale bar: 5 mm.
Longitudinal section of ovary (A–H) and old fruit (I–P). A–D Tashiroea clade, crowned or uncrowned ovaries at anthesis: A T. amoena (Y. Liu 571) B T. quadrangularis (Y. Liu 585) C T. biglandularis (Y. Liu 553) D T. sinensis (Y. Liu 569). E–H Bredia clade, crowned ovaries at anthesis: E B. hirsuta (Y. Liu 563) F B. fordii var. micrantha (Y. Liu 580) G B. plagiopetala (Y. Liu 459) H B. longiradiosa var. pulchella (Y. Liu 485). I–L Tashiroea clade, uncrowned, rarely crowned capsules: I T. amoena (Y. Liu 571) J T. biglandularis (Y. Liu 553) K T. sinensis (Y. Liu 569) L T. oligotricha (Y. Liu 468). M–P Bredia clade, crowned capsules: M B. dulanica (Y. Liu 565) N B. fordii (Y. Liu 444) O B. latisepala (Y. Liu 557) P B. repens (Y. Liu 558). Scale bars: 2 mm.
Field and literature surveys revealed that species of both Tashiroea clade and Bredia clade prefer shaded or moist habitats in forests or along forest margin, often along stream banks, from 50 m to 2500 m alt. The Tashiroea clade occurs in southeastern mainland China and the Ryukyu islands, while the Bredia clade, aside from the above areas, also extends to the central and southwestern Chinese provinces of Guizhou, Hubei, Chongqing, Sichuan and Yunnan.
Of the two species without molecular data, B. laisherana is endemic to Taiwan and B. violacea occurs in northern Vietnam. The former species has glabrous stems and leaves, and uncrowned ovaries and capsules (
Species of the Tashiroea clade and Bredia clade are similar in habit, habitat preference, most indumentum types, and in having lanceolate, ovate to suborbicular leaves, a cuneate to cordate leaf base, cymose to cymose panicles, dimorphic or isomorphic stamens, and gibbose, tuberculate, or spurred staminal appendages (Figs
However, our survey of morphological characters has shown that the two clades can be distinguished by several diagnostic features, including leaf indumentum, texture, leaf surface sculpture under SEM, presence/absence of yellowish uniseriate trichomes, and capsule morphology. Species of the Tashiroea clade differ from those of the Bredia clade by the glabrous, stiffly papery to leathery mature leaves with furrowed surface sculpture under SEM (vs. puberulous and papery, without furrowed sculpture) (Figs
The two clades are also morphologically distinguishable from their close relatives. Phylogenetic analyses showed that the Tashiroea clade is most closely related to Scorpiothyrsus and Driessenia axantha while Bredia clade is closest to Blastus, Fordiophyton, and Plagiopetalum (Fig.
Two species previously placed in Bredia were not sampled in the molecular phylogenetic studies: B. laisherana from Taiwan and B. violacea from north Vietnam. Bredia laisherana was initially identified as B. quadrangularis by
Morphological evidence and molecular phylogenetic data confirmed that the Tashiroea clade and the Bredia clade represent two distantly related lineages morphologically well differentiated from each other and from their possible relatives. We therefore resurrect the generic name Tashiroea Matsum for the former clade and redefine Bredia Blume to include the latter. For the species lack of molecular data, we place B. laisherana and B. violacea in Tashiroea and Bredia respectively based on morphology. Species circumscriptions basically follow
Tashiroea yaeyamensis Matsum., J. Coll. Sci. Imp. Univ. Tokyo 12: 489. 1899. (Designated here)
Shrubs or shrublets, erect, rarely creeping in the lower parts. Stems terete or slightly 4-sided, glabrous or glabrescent, rarely densely hairy (in T. villosa), terminal and axillary buds pubescent with yellowish uniseriate branched trichomes. Leaves petiolate; leaf blade lanceolate, ovate, elliptic, rarely suborbicular, stiffly papery to leathery, glabrescent when mature, rarely hairy (in T. villosa), secondary veins 1 or 2 on each side of midvein, margin remotely serrulate, or almost entire. Inflorescences terminal, few-flowered cymes to cymose panicles; bract minute, rarely to 1–2 cm long (in T. villosa), usually caducous. Flowers 4-merous. Hypanthium campanulate, rarely funnel shaped. Calyx lobes repand, crenate or triangular. Petals pink or purplish red, ovate, oblong to suborbicular, more or less oblique, apex acute or acuminate. Stamens 8, unequal or subequal; filaments filiform; anthers dimorphic or isomorphic, subulate to oblong-linear, gibbose, tuberculate or spurred at base, sometime unappendaged adaxially. Ovary half inferior, slightly crowned or uncrowned, ovoid-globular or turbinate, 4-celled. Style filiform; stigma apiculate. Capsule cup-shaped or subglobular, more or less 4-sided, woody, uncrowned, apex rounded or 4-humped, or crown persistent and enlarged, enclosing an obpyramidal space (in T. nudipes and T. oligotricha). Seeds numerous, minute, cuneate, densely granulate. (Figs
Eleven species, eight in southeastern mainland China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Zhejiang), one in Taiwan, and two in the Ryukyus (Fig.
Bredia amoena Diels, Notizbl. Bot. Gart. Berlin-Dahlem 9(83): 197–198. 1924 (Basionym). Type: China. Zhejiang: Yentang, 16 Aug 1920, H.H. Hu 30 (lectotype, designated here: A! [A00071979]; isolectotype: UC! [UC231837]).
Bredia chinensis Merr., J. Arnold Arbor. 8(1): 11–12. 1927. Type: China. Zhejiang: Yentang, 16 Aug 1920, H.H. Hu 30 (holotype: UC! [UC231837]; isotype: A! [A00071979]).
Bredia pricei F.P. Metcalf, Lingnan Sci. J. 12: 153–154. 1933. Type: China. W.R. Price 1200A (K! [K001325176]).
Bredia biglandularis C. Chen, Bull. Bot. Res., Harbin 4(3): 39. 1984 (Basionym). Type: China. Guangxi: Dongxing, Lu-bao-shan, 2 Oct 1976, T. Fang 1519 (holotype: GXMI! [GXMI050232]).
Bredia laisherana C.L. Yeh & C.R. Yeh, Edinburgh J. Bot. 65(3): 400 (–403; figs 3, 4C–D). 2008 (Basionym). Type: Taiwan, Pingtung, Mt Laisher, on the ridge of a mountain and steep valley and in a dense cloudy forest, 1600–1800 m, 15 Sept 2005, C.L. Yeh & C.R. Yeh 33 (holotype: PPI 065514).
Phyllagathis nudipes C. Chen, Bull. Bot. Res., Harbin 4(3): 47. 1984 (Basionym). Type: China. Guangdong: Ruyuan, Wuzhi Shan, 21 May 1973, Guangdong73-101 (holotype: IBSC! [IBSC0003997]).
Bredia okinawensis (Matsum.) H.L. Li, J. Arnold Arbor. 25: 21. 1944.
Japan. Okinawa: in montosis tractus Kunchan, Apr 1887, Y. Tashiro (5) (lectotype, designated here: TI! [TI00002346]). Additional syntypes: Japan. Okinawa: S. Tanaka 216 (TI! [TI00002347]), Matsumura (TI! [TI00002348]).
Phyllagathis oligotricha Merr. Sunyatsenia 1: 74. 1930 (Basionym). Type: China. Guangdong: Lok Chang, 8 Jun 1929, Tso 21016 (holotype: NY! [NY00273007]; isotypes: IBSC! [IBSC0003939, IBSC0003940]).
Phyllagathis anisophylla Diels, Bot. Jahrb. Syst. 65(2–3): 115. 1932. Type: China. Hunan, no precise location, 1926, Hunan Museum 60 (lectotype, designated here: IBSC! [IBSC0003938]; isolectotypes: IBSC! [IBSC0003936, IBSC0003937]).
Bredia quadrangularis Cogn. Monogr. Phan. 7: 473–474. 1891 (Basionym). Type: South China. Seemann s.n. (LE).
Bredia sessilifolia H.L. Li, J. Arnold Arbor. 25: 22. 1944 (Basionym). Type: China. Guangxi: Shang-sze District, Shih Wan Tai Shan, Tang Lung Village, 25 Sept 1934, W. T. Tsang 24346 (holotype: A! [A00071991]; isotype: IBSC! [IBSC0003956]).
Bredia sinensis (Diels) H.L. Li, J. Arnold Arbor. 25: 22. 1944.
Bredia glabra Merr., J. Arnold Arbor. 8: 12. 1927. Type: China. Zhejiang: Pinyong Xian, 11 Jul 1924, Ling Kan 7333 (holotype UC! [UC252284]).
China. Fujian: Chung-an District, 27 Jul 1921, H.H. Hu 1343 (lectotype, designated here: A! [A00073233]).
Bredia yaeyamensis (Matsum.) H.L. Li, J. Arnold Arbor. 25: 21. 1944.
Tashiroea yaeyamensis var. tanakaea Matsum., J. Coll. Sci. Imp. Univ. Tokyo 12: 490. 1899. Type: Okinawa: Yaeyama archipelago, Jul 1890, S. Tanaka 344 (holotype: TI! [TI00002345]).
Japan. Okinawa: Iriomote 1890, S. Tanaka 345 (lectotype, designated here: TI! [TI00002344]). Additional syntype: Japan. Okinawa: in Yaeyama, Aug 1887, Tashiro (TI! [TI00002343]).
China. Fujian: Pingnan County, Lingxia Town, 1000 m, 16 Jul 2017, Y. Liu 568 (holotype: A!; isotype: SYS!).
Resembles T. amoena in height, leaf size and shape, inflorescence and stamen morphology, while differing from the latter in the dense indumentum covering the whole plant and much larger bracts.
Shrubs or shrublets, 20–60 (–90) cm tall. Stem, leaves, peduncles, bracts, pedicels and hypanthium densely pubescent and villous with multiseriate or sometimes uniseriate glandular or non-glandular trichomes. Stems cylindrical, branchlets slightly 4-sided, sometimes rubescent. Leaves opposite; petiole 1.2–4.5 cm long; leaf blade ovate to ovate-elliptic, 4.2–12 × 1.8–6 cm, papery, abaxial surface pale green, adaxial surface green, secondary veins 3 on each side of midvein, base cordate to rounded, margin ciliate and inconspicuously serrulate, apex acuminate or short acuminate. Inflorescences terminal, cymose, 7–14 × 3.5–6.5 cm, bracts 9–19 × 5–8 mm, deciduous or sometimes persist till anthesis. Pedicels 1–3 mm. Hypanthium short campanulate, 4-sided, 3–5 mm long, calyx lobes broadly triangular, 1 mm long, apex acute. Petals purplish pink or pink, ovate to ovate-oblong, 7–10 × 3.5–5 mm, slightly oblique, apex acute. Stamens 8, dimorphic, unequal. Longer stamens antesepalous, ca. 15 mm long; anthers lanceolate, ca. 8 mm long, geniculate; connective decurrent, slightly prolonged, forming a short spur dorsally. Shorter stamens antepetalous, ca. 8 mm long, anthers lanceolate, ca. 4 mm long, base gibbose ventrally and forming a short spur dorsally. Ovary half inferior, locules 4, apex slightly 4-lobed, margin ciliate with glandular trichomes. Style ca. 0.6 cm long, puberulous with glandular trichomes basally. Capsule cup-shaped; hypanthium ca. 4 × 3.5 mm long; placental column distally entire, placentas non-thready. Seeds numerous, minute, cuneate, granulate. Flowering July-August, fruiting August-October.
The specific epithet refers to the dense pubescent and villous indumentum.
Tashiroea villosa is currently known from Pingnan, Jianou and Jianyang, northern Fujian, China (Fig.
Tashiroea villosa is discovered by Mr. Xiang-xiu Su. He is an amateur collector in Fujian who had made an important contribution to the description of this new species. We therefore include him as the author of this name. Tashiroea villosa is the sole species currently known in Tashiroea with densely puberulous and villous leaves (vs. glabrous) and smooth leaf surface sculpture (vs. furrowed). It is morphologically and phylogenetically closest to T. amoena. The two species are similar in height, leaf size and shape, inflorescence and stamen morphology. Tashiroea villosa is distinct from T. amoena in the dense indumentum covering the whole plant (vs. petioles and inflorescences pubescent or sometimes glabrescent) and much larger bracts (9–19 × 5–8 mm vs. 1–2 × 1 mm) in the florescence (Fig.
Bredia hirsuta Blume, Mus. Bot. 1(2): 25. f. 4. 1849.
Shrubs, shrublets or herbs, erect, ascending or creeping. Stems terete or more or less 4-sided, sparsely to densely puberulous, rarely glabrescent. Leaves petiolate; leaf blade ovate, cordate, oblong, elliptic, ovate-orbicular, rarely lanceolate, papery, rarely submembranous, sparsely to densely puberulous or strigose, secondary veins 2–5 on each side of midvein, margin serrulate or entire. Inflorescences terminal, umbellate, cymes or cymose panicles. Flowers 4-merous. Hypanthium funnel shaped to campanulate. Calyx lobes conspicuous, linear-lanceolate to triangular. Petals pink or purplish red, ovate to oblong, more or less oblique, apex acute or acuminate. Stamens 8, unequal or subequal; filaments filiform; anthers dimorphic or isomorphic, subulate to oblong-linear, gibbose, tuberculate or spurred at base, rarely unappendaged abaxially. Ovary half inferior, crowned, ovoid, 4-celled. Style filiform; stigma apiculate. Capsule turbinate to cup-shaped, more or less 4-sided, crown persistent and enlarged, enclosing an inverted frustum-shaped depression at capsule apex. Seed numerous, minute, cuneate, densely granulate. (Figs
China. Jiangxi: Chongyi County, Niedu Town, 579 m, 1 Aug 2016, W. Y. Zhao et al. LXP-13-22114 (holotype: SYS!; isotype: IBSC!).
Taiwan, Taitung, Mt Dulan, on the ridge of a mountain, 1000–1200 m, 14 Oct 2007, S.W. Chung, T.C. Hsu & C.R. Yeh 16 (holotype: TAIF! [TAIF348619]; isotypes: TAIF! [TAIF348620, TAIF348621, TAIF348622]).
Barthea esquirolii H. Lév., Repert. Spec. Nov. Regni Veg. 11(301–303): 494. 1913 (Basionym). Type: China. Guizhou: Tchai-choui-ho, July 1909, Esquirol 1581 (holotype: E! [E00090793]).
Bredia cordata H.L. Li, J. Arnold Arbor. 25(1): 24–25. 1944. Type: China. Sichuan: Ya-an, dense forest shade, 686 m, 30 Jul 1939. C. Y. Chiao 1205 (holotype: A! [A00071982]).
Bredia esquirolii var. cordata (H.L. Li) C. Chen, Bull. Bot. Res., Harbin 4(3): 40. 1984.
Otanthera fordii Hance, J. Bot. 19: 47. 1881 (Basionym). Type: China. Hong Kong, Jul 1880, C. Ford. herb no. 21099 (lectotype, designated here: BM! [BM000629024]; isolectotype BM! [BM000629025]).
Phyllagathis fordii (Hance) C. Chen, Bull. Bot. Res., Harbin 4(3): 50. 1984.
Bredia sepalosa Diels, 65(2–3): 109–110. 1932. Type: China. Guangxi: Yao shan, 1928, S.S. Sin & K.K. Whang 648 (lectotype, designated here: IBSC! [IBSC0003942]).
Phyllagathis fordii var. micrantha C. Chen, Bull. Bot. Res., Harbin 4(3): 50–51. 1984 (Basionym). Type: China. Guizhou: Dushan, in convallibus montanis, 600 m, 22 Aug 1930, Y. Tsiang 6563 (holotype: IBSC! [IBSC0003995]; isotypes: NAS! [NAS00052126] PE! [PE00782806, PE00782810]).
Bredia penduliflora S.S. Ying, Quart. J. Chin. Forest. 6(1): 167. 1972. Type: Taiwan, Sakanyalan-Tawu, 10 Oct 1972, S.S. Ying 1517 (lectotype, designated here: NTUF! [F00004987]; isolectotypes: NTUF! [F00004986, F00004989, F00004990, F00004991, F00004992, F00004993]).
Taiwan. Pingtung, Sungshan, J. Ohwi s.n. (holotype: KYO! [KYO00022400]).
Fordiophyton gracile Hand.-Mazz., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Anz. 63: 3, 10. 1926 (Basionym). Type: China. Hunan: Heng Shan, Wukang, 1150–1300 m, 4–8 Aug 1918, Hand.-Mazz. 12380 (holotype: WU! [WU0059491]; isotypes: A! [A00055337] E! [E00090795]).
Phyllagathis gracilis (Hand.-Mazz.) C. Chen, Bull. Bot. Res., Harbin 4(3): 51. 1984.
Phyllagathis guidongensis K.M. Liu & J. Tian, Phytotaxa 263(1): 58–62 (Basionym). Type: China. Hunan: Guidong County, Pule Town, 970 m, 3 Jul 2013, K.M. Liu, R.Y. Yi & L. Peng 24147 (holotype: HNNU; isotypes: HNNU CSFI).
Japan. K. Ito s.n. (lectotype, designated here: L! [L0170980]).
In the protologue,
Bredia scandens (Ito & Matsum.) Hayata. J. Coll. Sci. Imp. Univ. Tokyo 30(1): 114. 1911.
Taiwan, inter Suiteiryō et Niki, C. Owatari, Jan 1898 (the date “1896” cited in the protologue is probably erroneous) (lectotype, designated here: TI! [TI00002337]; isolectotype: TI!, [TI00002339]).
Phyllagathis latisepala C. Chen, Bull. Bot. Res., Harbin 4(3): 53–54. 1984 (Basionym). Type: China. Hubei: Hefeng, ad pedes montis calcareo, 18 Sept 1958, H.J. Li 6451 (holotype: IBSC! [IBSC0003996]; isotype: PE! [PE00025692]).
Phyllagathis longearistata C. Chen, Bull. Bot. Res., Harbin 4(3): 52–53. 1984 (Basionym). Type: China. Guangxi: Hechi, prope rivulos in convallibus montanis, 19 May 1928, L.H. Chun 91861 (holotype: IBK! [IBK00190677]; isotype: IBK! [IBK00190678]).
Fordiophyton gracile var. longilobum Hand.-Mazz., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Anz. 63: 3, 10. 1926 (Basionym). Type: China. Jiangxi: between Ningdu and Ki-an, T.H. Wang 493 (holotype: WU! [WU0059490]).
Phyllagathis longiradiosa C. Chen, Bull. Bot. Res., Harbin 4(3): 51. 1984 (Basionym). Type: Based on Barthea cavaleriei H. Lév.
Barthea cavaleriei
H. Lév., Repert. Spec. Nov. Regni Veg. 8(160–162): 61. 1910. Type: China. Guizhou: near Mou-you-sé, J. Cavalerie 1552 (lectotype, designated by
Fordiophyton cavaleriei (H. Lév.) Guillaumin, Bull. Soc. Bot. France 60: 275. 1913.
Bredia cavaleriei (H. Lév.) Diels, Bot. Jahrb. Syst. 65(2–3): 110. 1932.
Bredia longiradiosa C. Chen, Fl. Yunnan 2: 105, f. 27, 1–5. 1979. nom. inval. (reference to place of publication of basionym not provided).
Phyllagathis longiradiosa var. pulchella C. Chen, Bull. Bot. Res., Harbin 4(3): 52. 1984 (Basionym). Type: China. Guangxi: Daxin, Longjin, 4 May 1959, F.F. Huang 3596 (holotype: GXMI! [GXMI050237]).
Three gatherings were cited in the protologue of Barthea cavaleriei, viz. Cavalerie 1552, Esquirol 1581 and Esquirol 215, without designation of a type. Two of the syntypes, Esquirol 1581 and 215 were later cited as the types of Barthea esquirolii H. Lév. (1913) and Barthea blinii H. Lév. (1913), respectively.
China. Guangxi: Guilin District, Chi-fen Shan, Xichang Cun and vicinity, W.T. Tsang 28432 (holotype: A! [A00071988]; isotype: IBSC! [IBSC0003950]).
Bredia oldhamii var. ovata Ohwi, J. Jap. Bot. 12(9): 661–662. 1936. Type: Taiwan, Taitung, in open forest, forest margin, 100–1200 m, J. Ohwi 425 (holotype: KYO).
Taiwan, near Tamsuy, Jan 1864, R. Oldham 118 (holotype: K! [K000978944]; isotype: GH! [GH00071989] P! [P02274733] US! [US00120439]).
Phyllagathis plagiopetala C. Chen, Bull. Bot. Res., Harbin 4(3): 44–45. 1984 (Basionym). Type: China. Hunan: Xinning, Ziyun Shan, in dense silvis apice montium, 800 m, 11 Jul 1959, P.C. Tam 63423 (holotype: IBK! [IBK00127587]).
China. Hunan: Sangzhi County, from Shayuan to Nanmuping village, 430–470 m, 11 Nov 2016, Y. Liu 558 (holotype: SYS!; isotypes: A! SYS!).
Taiwan, Chiayi, Juili, Ou 2869 (holotype: NCUF).
Fordiophyton tuberculatum Guillaumin, Notul. Syst. (Paris) 2(11): 326. 1913 (Basionym). Type: China. Yunnan: Tchen fong chan, Delavay 5053 (lectotype, designated here: E! [E00285959]; isolectotypes: P! [P02274731], P! [P02274729] individual on the right side of the sheet).
Bredia omeiensis H.L. Li, J. Arnold Arbor. 25(1): 24. 1944.Type: China. Sichuan: Emei Shan, 1100 m, 21 Aug 1937, Y.S. Liu 1080 (holotype: A! [A00071990]; isotype: LBG! [LBG00089612]).
Three gatherings were cited in the protologue of Fordiophyton tuberculatum, viz. Delavay 5053, Ducloux 2192 and Wilson 4906.
Vietnam. Tonkin: Tian-yen, Ho Yung Shan & vicinity, 13 Oct–22 Nov 1940, W.T. Tsang 30751 (holotype: A! [A00071992]).
Phyllagathis velutina (Diels) C. Chen, Bull. Bot. Res., Harbin 4(3): 51. 1984.
China. Yunnan: Mengzi, 1000–2300 m, A. Henry 13479 (lectotype, designated here: K! [K000867582]; isolectotypes: A! [A00055334] NY! [NY00221472]).
Blastus yunnanensis H. Lév., Repert. Spec. Nov. Regni Veg. 11(286–290): 300–301. 1912 (Basionym). Type: China. Yunnan: Vallée de Long-Ky, pied des rochers humides, 700 m, Aug 1911, Maire s.n. (holotype: E! [E00285956]).
Blastus mairei H. Lév., Repert. Spec. Nov. Regni Veg. 11(286–290): 300. 1912. Type: China. Yunnan: Bord des eaux, Vallée de Long-Ky, 700 m, Jul 1911, Maire s.n. (holotype: E! [00285957]).
We thank curators and staff of herbaria cited in the text for permission to examine the specimens or high-resolution photos of them, especially Akiko Shimizu (TI) who kindly searched for original material to facilitate lectotypification. We are deeply grateful to Dr. Hideaki Ohba and Dr. Shinobu Akiyama for providing important information regarding the type materials of B. hirsuta; Dr. David E. Boufford, Dr. Fabian A. Michelangeli and Dr. Susanne S. Renner for their valuable comments on the manuscript; Dr. Bing-lan Zhang for her help with leaf surface scanning using Phenom Pro. This work was supported by the National Natural Science Foundation of China (31770214) and the Science, Technology Planning Project of Guangdong Province (2015A030302011) and Chang Hungda Science Foundation of Sun Yat-sen University.
Table S1. Source of materials studied and GenBank accession numbers for nrITS and chloroplast intergenic spacer trnV-trnM
Data type: molecular data