Corresponding author: Ren-Chao Zhou (
Academic editor: Ricardo Kriebel
Zhou Q-J, Dai J-H, Lin C-W, Denda T, Zhou R-C, Liu Y (2019) Recircumscription of
The generic circumscription of
A recent molecular phylogenetic study, with extensive sampling of
In this paper, we revisited the morphological characters of the
Eight newly sequenced species were added to the combined dataset (nrITS and chloroplast
All species of the
Diels published several names in
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
Characters of the
Comparison of the
Habit | Shrubs or shrublets | Shrubs, shrublets or herbs |
Indumentum of mature leaf |
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Leaf texture |
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Leaf surface sculpture |
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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 ( |
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; |
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 |
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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 |
Leaf morphology of the
Leaf morphology of the
Scanning electron micrographs of lower epidermis.
Trichomes in the
Stamen morphology in the
Stamen morphology in the
Longitudinal section of ovary (
Field and literature surveys revealed that species of both
Of the two species without molecular data,
Species of the
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
The two clades are also morphologically distinguishable from their close relatives. Phylogenetic analyses showed that the
Two species previously placed in
Morphological evidence and molecular phylogenetic data confirmed that the
Shrubs or shrublets, erect, rarely creeping in the lower parts. Stems terete or slightly 4-sided, glabrous or glabrescent, rarely densely hairy (in
Eleven species, eight in southeastern mainland China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Zhejiang), one in Taiwan, and two in the Ryukyus (Fig.
Distribution of
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]).
China. Fujian: Chung-an District, 27 Jul 1921, H.H. Hu 1343 (lectotype, designated here: A! [A00073233]).
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
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.
Line illustration of
Y. Liu 568 (A) collected from Lingxia, Pingnan County, Fujian, China, holotype of
The specific epithet refers to the dense pubescent and villous indumentum.
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
Twenty-one species: 15 in central and southern mainland China (Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Chongqing, Yunnan, Zhejiang), one in north Vietnam, five in Taiwan and one extending to the Ryukyu islands (Fig.
Distribution of
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]).
Taiwan. Pingtung, Sungshan, J. Ohwi s.n. (holotype: KYO! [KYO00022400]).
Japan. K. Ito s.n. (lectotype, designated here: L! [L0170980]).
In the protologue,
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]).
Three gatherings were cited in the protologue of
China. Guangxi: Guilin District, Chi-fen Shan, Xichang Cun and vicinity, W.T. Tsang 28432 (holotype: A! [A00071988]; isotype: IBSC! [IBSC0003950]).
Taiwan, near Tamsuy, Jan 1864, R. Oldham 118 (holotype: K! [K000978944]; isotype: GH! [GH00071989] P! [P02274733] US! [US00120439]).
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).
Three gatherings were cited in the protologue of
Vietnam. Tonkin: Tian-yen, Ho Yung Shan & vicinity, 13 Oct–22 Nov 1940, W.T. Tsang 30751 (holotype: A! [A00071992]).
China. Yunnan: Mengzi, 1000–2300 m, A. Henry 13479 (lectotype, designated here: K! [K000867582]; isolectotypes: A! [A00055334] NY! [NY00221472]).
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
Table S1. Source of materials studied and GenBank accession numbers for nrITS and chloroplast intergenic spacer
molecular data