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Phytotaxa 414 (6): 269–279
https://www.mapress.com/j/pt/
Copyright © 2019 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Renato Goldenberg: 25 Jul. 2019; published: 21 Aug. 2019
https://doi.org/10.11646/phytotaxa.414.6.1
269
A new species of Tashiroea (Melastomataceae, Sonerileae) from Guangxi, China
QIU-JIE ZHOU1, JIN-HONG DAI1, REN-CHAO ZHOU1 & YING LIU1*
1State Key Laboratory of Biocontrol and Guangdong Key Laboratory of Plant Resources, School of Life Sciences, Sun Yat-sen Univer-
sity, No. 135, Xin-Gang-Xi Road, Guangzhou 510275, China
*Corresponding author: liuyng73@mail.sysu.edu.cn
Abstract
Tashiroea dayaoshanensis (Melastomataceae, Sonerileae), a species from northeastern Guangxi, China and previously
often identified as Phyllagathis nudipes or P. oligotricha in Chinese herbaria, is described as new based on morphological
and molecular data. Indumentum, leaf texture and surface sculpture, capsule morphology and molecular phylogenetic data
strongly support the placement of T. dayaoshanensis within the Tashiroea clade which should be accommodated in Tashiroea.
Morphology as well as sequence divergence of the nrITS region indicate that T. dayaoshanensis is well differentiated from
other species in the same clade, and thus should be treated as a distinct species. Tashiroea dayaoshanensis is phylogenetically
closest to Bredia sessilifolia but can be easily distinguished by its petiolate leaves (vs. sessile or subsessile), and crowned
ovary (vs. uncrowned). It resembles P. nudipes and P. oligotricha in habit and stamen morphology but differs from both in
its leaf apex acuminate (vs. acute to obtuse), connective ventrally tuberculate (vs. not tuberculate) and ovary crown exerted
from calyx tube during young fruit stage (vs. not exerted).
Keywords: Bredia, Tashiroea, taxonomy
Introduction
Tashiroea Matsumura (1899: 489) was established based on two species from Ryukyu Islands, namely T. yaeyamensis
Matsumura (1899: 489) and T. okinawensis Matsumura (1899: 490). Diels (1924, 1932) recognized Tashiroea and
added one Chinese species T. sinensis Diels (1924: 198) to the genus, but Li (1944) considered Tashiroea as congeneric
with another Asian genus Bredia Blume (1849: 24) and placed it in synonym under Bredia. Li’s treatment was thereafter
adopted (Chen 1984a, b, Hansen 1992, Chen & Renner 2007). Recent phylogenetic studies, with extensive sampling
of Bredia and related genera, indicated that Bredia as currently circumscribed is not monophyletic, encompassing
two distantly related clades, with the type of Bredia being grouped in one clade and Tashiroea in the other (Zhou
et al. 2019, Zhou et al. unpublished data). The latter clade (hereafter referred to as the Tashiroea clade), therefore,
should be excluded from Bredia and accommodated in Tashiroea. Kokubugata et al. (2019) also suggested the same
taxonomic treatment, but only for T. okinawensis, T. yaeyamensis, and T. sinensis, because of limited taxa sampling.
Morphologically, the Tashiroea clade can be easily distinguished from Bredia by the glabrescent, stiffly papery to
leathery leaves with furrowed surface sculpture under scanning electronic microscope (SEM) (vs. puberulous and
papery leaves and smooth surface), presence of yellowish uniseriate hairs (vs. absence) and uncrowned capsules or
and crowned capsules with an obpyramidal depression on the top (vs. crowned capsule and inverted frustum-shaped
depression) (Zhou et al. 2019, Zhou et al. unpublished data).
During a field trip for a project on the systematics of Bredia and Phyllagathis Blume (1831: 507), an unknown
plant was discovered along the forest margin of Dong-wang-shan in Pingnan County, eastern Guangxi. Subsequent
herbarium survey revealed additional specimens of the same species from Jinxiu County, northeastern Guangxi (Fig.
1). Detailed examination of these plants revealed a character combination of yellowish uniseriate hairs covering the
apical and axillary buds, stiffly papery leaves glabrescent with furrowed surface sculpture under SEM, and ovary crown
enlarged in young fruit (Figs. 2–4) but evanescent in old fruit (S.S. Sin 23662, IBSC0246945). The above characters
are consistent with those of the Tashiroea clade. Herbarium specimens of the same plant were previously identified
as Bredia amoena Diels (1924: 197), B. quadrangularis Cogniaux (1891: 473), B. sinensis (Diels) H.L. Li (1944:
22) (= T. sinensis), Phyllagathis anisophylla Diels (1932: 115) [= P. oligotricha Merrill (1930: 74)], or most often as
ZHOU ET AL.
270 • Phytotaxa 414 (6) © 2019 Magnolia Press
Phyllagathis nudipes C. Chen (1984a: 47), all of which are members of the Tashiroea clade. The plant in question is
10–30 cm tall with adventitious roots at lower nodes (Fig. 1A), and the stamens are subequal and subisomorphic (Figs.
3–4), readily distinguished from B. amoena, B. quadrangularis, and B. sinensis (vs. 30–120 cm, without adventitious
roots, unequal and dimorphic stamens) (Chen 1984b). It resembles P. nudipes (Fig. 5A) and P. oligotricha (Fig. 5B) in
the stoloniferus habit and subequal stamens, but differs from the latter in its glabrous mature leaf (vs. sparsely setose
with multiseriate hairs), and from both in its acuminate leaf apex (vs. acute to obtuse), ventrally tuberculate connective
of stamen (vs. not tuberculate) and ovary crown exerted from calyx tube in young fruit (vs. not exerted) (Figs. 2–4).
Based on the above evidence, we suspected that it represented an undescribed species in the Tashiroea clade.
In this study, we performed phylogenetic analyses based on sequence data of the nuclear ribosomal internal
transcribed spacer (nrITS) to evaluate the phylogenetic position of the unknown plant. Our results confirmed that
this plant represented a previously unrecognized species in the Tashiroea clade. We name it as T. dayaoshanensis and
describe it in detail below. A key to separate it from other species of the Tashiroea clade is also provided.
FIGURE 1. Herbarium specimens of Tashiroea dayaoshanensis. A. Jinxiu Exped. 5-1-196 (GXMI) collected from Liu-xiang, Jinxiu
County, Guangxi, China, previously misidentified as Bredia sinensis and B. amoena. B. Da-yao-shan Exped. 13748 (IBK, IBSC) collected
from Mt. Luo-xiang, Jinxiu County, Guangxi, China, previously misidentified as Phyllagathis nudipes.
Materials and methods
Morphological data were obtained through field, herbarium and literature surveys as well as by observing living plants
in the facilities of Sun Yat-sen University. Specimens of the species concerned (GXMG, GXMI, HNNU, IBG, IBK,
IBSC, JJF, KUN, NAS, PE, SYS) or their high-resolution photos (A, K, MO, NY, TAIF, UC) were examined.
For phylogenetic analyses, outgroup and ingroup taxa were selected based on previous studies (Zhou et al.
2019, Zhou et al. unpublished data). To infer the phylogenetic position of T. dayaoshanensis, the type of Bredia
and Phyllagathis, one representative each from several related genera, and all known species of the Tashiroea clade
except B. laisherana C.L. Yeh & C.R. Yeh (2008: 400) were included in the analyses. The nrITS dataset contained
21 accessions representing 17 species from eight genera, with Blakea P. Browne (1756: 323) (Blakeeae) chosen as an
A NEW SPECIES OF TASHIROEA (MELASTOMATACEAE) Phytotaxa 414 (6) © 2019 Magnolia Press • 271
outgroup. Total DNA was extracted from silica-gel dried leaves using the CTAB procedure (Doyle & Doyle 1987).
Nuclear ribosomal ITS region of T. dayaoshanensis and T. okinawensis was amplified and sequenced using universal
primers ITS4 and ITS5 (White et al. 1990), following the same procedure described in Boufford et al. (2014). The
remaining sequences were downloaded from GenBank. The source of the materials and GenBank accession numbers
are given in Table 1.
TABLE 1. Source of materials studied and GenBank accession numbers for nrITS. Newly generated sequences are indicated
in bold.
Taxon Locality Voucher ITS
Blakea schlimii (Naudin) Triana - - AY460441
Tr. Dissochaeteae
Dissochaeta gracilis Blume Java, Indonesia Liu 601 (SYS) MG644479
Tr. Sonerileae
Blastus cochinchinensis Lour. Yilan, Taiwan Liu 528 (SYS) MG644466
Bredia hirsuta Blume Komi, Iriomote, Japan Alison Wee & Akiyo Naiki 8032 (SYS) MG644416
Bredia hirsuta Blume Taidong, Taiwan Liu 563 (SYS) MG644417
Fordiophyton faberi Stapf Linchuan, Guangxi, China Liu 552 (SYS) MG644456
Phyllagathis rotundifolia (Jack) Blume Kuala Lumpur, Malasysia Zhou et al. M50 (SYS) MG644436
Scorpiothyrsus oligotrichus H.L. Li Ledong, Hainan, China Liu 454 (SYS) MG644440
MG644441
Tashiroea clade
Tashiroea dayaoshanensis Pingnan, Guangxi, China Liu 476 (SYS) MK860817
Tashiroea okinawensis Matsumura Nago, Okinawa, Japan Liu 636 (SYS) MK860816
Tashiroea sinensis Diels Zixi, Jiangxi, China Liu 546 (SYS) MG644397
Tashiroea sinensis Diels Pingnan, Fujian, China Liu 569 (SYS) MG644398
Tashiroea yaeyamensis Matsumura Komi, Iriomote, Japan Alison Wee & Akiyo Naiki 8024 (SYS) MG644396
Bredia amoena Diels Zixi, Jiangxi, China Liu 545 (SYS) MG644389
Bredia amoena Diels Pingnan, Fujian, China Liu 571 (SYS) MG644390
Bredia biglandularis C. Chen Fangcheng, Guangxi, China Liu 553 (SYS) MG644392
Bredia quadrangularis Cogn. Guidong, Hunan, China Liu 473 (SYS) MG644391
Bredia sessilifolia H.L. Li Huaiji, Guangdong, China Liu 540 (SYS) MG644393
Phyllagathis nudipes C. Chen Ruyuan, Guangdong, China Liu 435 (SYS) MG644394
Phyllagathis oligotricha Merr. Rucheng, Hunan, China Liu 468 (SYS) MG644395
Sequences obtained were aligned using SeqMan v.7.1.0 (DNASTAR Inc., Madison, WI). Prior to phylogenetic
analyses, GTR+G was selected as the best-fitting model using the Akaike information criterion in Modeltest version
3.7 (Posada & Crandall 1998). Bayesian inference (BI), Maximum likelihood (ML) and Maximum parsimony (MP)
analyses were performed using MrBayes 3.2.6 (Huelsenbeck & Ronquist 2001), RAxML version 8.2.10 (Stamatakis
2014) and PAUP* 4a165 (Swofford 2003) respectively. For BI analyses, two independent Markov Chain Monte-Carlo
analyses (MCMC) were performed each with four simultaneous chains. A total of 10,000 trees were sampled evenly.
Convergence was considered as reached when average deviation of split frequencies fell below 0.01 at the end of
MCMC analyses. We also assessed the effective sample sizes (ESS) for all parameters and statistics using Tracer
version 1.7.1(Rambaut et al. 2018). All ESS were obtained with values higher than 3000, indicating that all parameters
were sufficiently sampled for all chains to converge. Finally, the first 25% of the trees (2,500 trees) were discarded as
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272 • Phytotaxa 414 (6) © 2019 Magnolia Press
burn-in, and the remaining trees were used to construct a 50% majority-rule consensus tree with Bayesian posterior
probabilities (PP). For MP analyses, a heuristic search strategy was conducted of 1000 random addition replicates, with
the tree-bisection-reconnection (TBR) branch swapping algorithm and MultTrees on. Maxtree was set to 500. Node
support (BSMP) was evaluated by 1000 bootstrap replicates of 1000 random additions. ML analyses were performed
under GTR+G model. Bootstrap support (BS) was calculated with 1,000 bootstrap replicates using a fast bootstrapping
algorithm (Stamatakis et al. 2008).
Results
The aligned sequence matrix contained 666 characters. Statistics of sampled sequences were summarized in Table 2.
Trees generated by ML, MP and BI analyses had identical topology. Only the phylogenetic tree obtained from ML
analysis is shown here, with Bayesian posterior probabilities (PP) and MP bootstrap support values (BSMP) indicated
near the nodes. As shown in Fig. 6, T. dayaoshanensis clustered within the well supported Tashiroea clade (PP = 1.0,
BS = 100%, BSMP = 100%) which was sister to Scorpiothyrsus H.L. Li (1944: 33), whereas the type of Bredia and
Phyllgathis formed a clade with Blastus Loureiro (1790: 526) and Fordiophyton Stapf (1892: 314) (PP = 1.0, BS =
81%, BSMP = 86%). Two subclades were recovered within the Tashiroea clade with strong support. One subclade
contained the species originally described in Tashiroea, T. sinensis, T. yaeyamensis, and T. okinawensis (PP = 1.0, BS
= 99%, BSMP = 99%); the other comprised B. amoena, B. quadrangularis, B. glandularis C. Chen (1984a: 39), B.
sessilifolia H.L. Li (1944: 22), P. nudipes, P. oligotricha, and T. dayaoshanensis (PP = 1.0, BS = 98%, BSMP = 96%),
with T. dayaoshanensis sister to B. sessilifolia (PP = 1.0, BS = 97%, BSMP = 94%).
TABLE 2. Summary statistics of sequences used for phylogenetic analyses. PIS, parsimony-informative sites.
Marker Accessions sampled Aligned base pairs Variable sites % Variable sites Conserved sites PIS % PIS Model
nrITS 21 666 243 36% 414 155 23% GTR+G
Discussion
Tashiroea dayaoshanensis was recovered within the Tashiroea clade. It has yellowish uniseriate hairs, glabrescent,
stiffly papery leaves with furrowed surface sculpture, and evanescent ovary crown in the old fruit (Figs. 2–4), all of
which are typical of the Tashiroea clade (Zhou et al. 2019; Zhou et al. unpublished). The generic placement of T.
dayaoshanensis in Tashiroea is therefore strongly corroborated.
Tashiroea dayaoshanensis differs markedly from B. laisherana, the only potential member of the Tashiroea clade not
included in phylogenetic analyses, in height (10–30 vs. 70–100 cm), stem and inflorescence glabrescent (vs. pubescent
with spreading hairs), calyx tube shortly campanulate (vs. narrowly elliptic to long cup-shaped), and connective of
longer stamen dorsally spurred (vs. not spurred). Between T. dayaoshanensis and other species of the Tashiroea clade
sampled in the present analyses, we detected 16 to 37 nucleotide substitutions at the nrITS region, showing that it is
well diverged from other species in the same clade. Tashiroea dayaoshanensis is phylogenetically most closely related
to B. sessilifolia (Fig. 6), however, it can be easily distinguished from the latter by its petiolate leaves (vs. sessile or
subsessile) and crowned ovary (vs. uncrowned) (Fig. 5C–D). Morphologically, T. dayaoshanensis is similar to P.
nudipes and P. oligotricha in the stoloniferus habit. Therefore, it is understandable that herbarium specimens of this
species were often identified as P. nudipes or P. anisophylla (= P. oligotricha). Tashiroea dayaoshanensis differs from
P. oligotricha in its glabrous mature leaf (vs. sparsely setose), and from both P. oligotricha and P. nudipes in its leaf
apex acuminate (vs. acute to obtuse), connective ventrally tuberculate (vs. not tuberculate) and ovary crown exerted
from calyx tube during young fruit stage (vs. not exerted) (Figs. 2–4). In addition, their flowering seasons are different.
Phyllagathis nudipes and P. oligotricha flower from April to June, while T. dayaoshanensis flowers from July to
August.
Therefore T. dayaoshanensis is quite distinct in molecular constitution, morphology, and phenology from related
species, leading us to recognize it as a new species. A key is provided below to separate it from other species of the
Tashiroea clade.
A NEW SPECIES OF TASHIROEA (MELASTOMATACEAE) Phytotaxa 414 (6) © 2019 Magnolia Press • 273
FIGURE 2. Tashiroea dayaoshanensis, all from Y. Liu 476 (A, SYS). A. Habitat. B. Yellowish hairs covering axillary buds. C. Adaxial
and abaxial leaf surface. D. Furrowed leaf surface sculpture under SEM. E. Flowering branch. F. Inflorescence. Scale bars: 2 mm (B); 50
μm (D).
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274 • Phytotaxa 414 (6) © 2019 Magnolia Press
FIGURE 3. Tashiroea dayaoshanensis, all from Y. Liu 476 (A, SYS). A. Top view of a flower. B. Side view of a flower. C. Longitudinal
section of a flower showing subequal stamens. D. Anther morphology in detail. E. Longitudinal section of ovary at anthesis showing the
ovary crown. F. Longitudinal section of young fruit showing the enlarged ovary crown. Scale bars: 2 mm (A–F).
Taxonomic treatment
Tashiroea dayaoshanensis Q.J. Zhou & Ying Liu, sp. nov. (Figs. 2–4)
Type:—CHINA. Guangxi: Pingnan County, Si-wang town, Dong-wang-shan, forest margin, 800–1000 m elev., 12 Aug 2016, Y. Liu 476
(holotype A!, isotypes SYS!).
Diagnosis:—Resembles P. nudipes and P. oligotricha in the stoloniferus habit and isomorphic stamens, while differs from both in having
acuminate leaf apex (vs. acute to obtuse), ventrally tuberculate connective (vs. not tuberculate) and ovary crown exerted from calyx
tube during young fruit stage (vs. not exerted).
A NEW SPECIES OF TASHIROEA (MELASTOMATACEAE) Phytotaxa 414 (6) © 2019 Magnolia Press • 275
FIGURE 4. Drawing of Tashiroea dayaoshanensis, all from Y. Liu 476 (A, SYS). A. Habit. B. Bud of a flower. C. Top view of a flower. D.
Longitudinal section of a flower (part). E. Anther morphology. F. Longitudinal section of young fruit. G. Old fruit. Drawn by Ms. Rong-
Mei Zhang (KUN).
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276 • Phytotaxa 414 (6) © 2019 Magnolia Press
FIGURE 5. Species morphologically or phylogenetically close to Tashiroea dayaoshanensis. A. Phyllagathis nudipes, from Y. Liu 435
(SYS). B. P. oligotricha, from Y. Liu 468 (SYS). C–D. Bredia sessilifolia, from Y. Liu 540 (SYS).
FIGURE 6. Maximum likelihood (ML) phylogenetic tree based on nrITS sequence data showing the phylogenetic position of Tashiroea
dayaoshanensis. Numbers near the nodes are bootstrap values obtained from maximum likelihood analyses (BS, left), Bayesian posterior
probabilities (PP, middle) and bootstrap values resulting from maximum parsimony analyses (BSMP, right). Tashiroea dayaoshanensis is
indicated in bold. Asterisks denote the three species originally described in Tashiroea, while boxes denote the type species of Bredia and
Phyllagathis.
A NEW SPECIES OF TASHIROEA (MELASTOMATACEAE) Phytotaxa 414 (6) © 2019 Magnolia Press • 277
Shrublets, 10–30 cm tall, stoloniferus, branched or not, with adventitious roots at lower nods. Stems terete, apical
and axillary buds covered with yellowish uniseriate hairs, glabrescent at maturity. Leaves opposite, puberulous when
young, glabrous at maturity; petiole 0.7–3 cm long; leaf blade ovate, 4.2–12 × 1.8–7.4 cm, papery to stiffly papery,
abaxial surface pale green, adaxial surface green to dark green, secondary veins 2 on each side of midvein, base obtuse
to rounded, seldom cuneate, margin inconspicuous serrulate, apex acuminate to caudate. Inflorescences terminal and
axillary, cymose, 3.5–9 × 2–5 cm. Peduncles, pedicels and calyces sparsely puberulous or glabrescent. Pedicels 3–8
mm long. Hypanthium shortly campanulate, obtusely 4-sided, 3–4 mm long. Calyx lobes broadly triangular, 1 mm
long, apex acute. Petals purple or pink, ovate to oblong, 7 × 4–5 mm, slightly oblique, apex acute or apiculate. Stamens
subisomorphic. Alternipetalous stamens ca. 12 mm long; anthers lanceolate, ca. 6 mm long; connective decurrent,
slightly prolonged, tuberculate ventrally, forming a short spur dorsally. Antepetalous stamens ca. 9 mm long, anthers
lanceolate, ca. 4 mm long; connective decurrent, tuberculate ventrally, forming a short spur dorsally. Ovary half
inferior, locules 4, ovary crowned, apex slightly 4-lobed, margin ciliate with glandular trichomes. Style 1 cm long,
puberulous in lower part or glabrous. Ovary crown enlarged in young fruit, but evanescent in the old fruit. Capsule
cup-shaped; hypanthium ca. 3–4 × 5 mm, glabrescent; placental column distally entire, placentas non-thready. Seed
numerous, very minute, cuneate.
Phenology:—Flowering July–August, fruiting August–September.
Etymology:—The specific epithet is derived from Dayaoshan Mountain, the distribution range of T.
dayaoshanensis.
Distribution:—Tashiroea dayaoshanensis is currently known from Dayaoshan Mountain, Pingnan and Jinxiu,
northeastern Guangxi, China (Fig. 7). It occurs in dense or open forest, often on slopes, along forest margin, or in
bushes along trails, at 700–1100 m.
Conservation status:—This species is narrowly endemic to Jinxiu and Pingnan. The populations in Jinxiu are
under protection in the Dayaoshan Nature Reserve, whereas the population discovered in Pingnan is threatened by
habitat destruction because of human activities. Tashiroea dayaoshanensis should be considered endangered (EN)
according to IUCN (2012).
FIGURE 7. Distribution of Tashiroea dayaoshanensis (triangle) and three close species Bredia sessilifolia (solid triangle), Phyllagathis
nudipes (solid circle) and P. oligotricha (solid square).
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Additional specimen examined:—CHINA. Guangxi: Jinxiu County, Liu-xiang, 25 Aug 1951, Anonymous 328
(HUUN [00058872]); Jinxiu County, Liu-xiang, Yao-shan, 22 Aug 1951, Anonymous 293 (HUUN [00022855]); Jinxiu
County, Liu-xiang, 4 Jul 1977, Jinxiu Exped. 5-1-196 (GXMI [010646]); Jinxiu County, Mt. Luo-xiang, 700 m, 30 Mar
1982, Da-yao-shan Exped. 13748 (IBK [00127578], IBSC [0246652]); Jinxiu County, Liu-xiang, Sheng-tang-wei,
1090 m, 10 Nov 1981, Da-yao-shan Exped. 12452 (IBK [00127579], IBSC [0246651]); Yao-shan, Ku-chun, 21 Jul
1934, S.S. Sin 23662 (IBSC [0246945, 0246946]); Yao-shan, 28 May 1928, S.S. Sin 208 (IBSC [0246944]).
Key to species of Tashiroea
1. Shrublets 10–30 cm tall, ascending, with adventitious roots at lower nodes .....................................................................................2
- Shrubs or shrublets to 40–200 cm tall, usually without adventitious roots at lower nodes .............................................................. 4
2. Petiole and leaf blade of mature leaves sparsely setose with multiseriate hairs ............................................................P. oligotricha
- Petiole and leaf blade of mature leaves glabrous ...............................................................................................................................3
3. Leaf apex acute to obtuse; connective ventrally not tuberculate; ovary crown not exerted from calyx tube in young fruit ...............
............................................................................................................................................................................................. P. nudipes
- Leaf apex acuminate; connective ventrally tuberculate; ovary crown exerted from calyx tube in young fruit ..... T. dayaoshanensis
4. Young stem pubescent with spreading multiseriate hairs or inflorescence puberulous with spreading glandular hairs ....................5
- Stem and inflorescence glabrous or glabrescent without spreading hairs ..........................................................................................6
5. Connective of longer stamen decurrent, not spurred dorsally; calyx tube narrowly elliptic to long cup-shaped ..........B. laisherana
- Connective of longer stamen decurrent, spurred dorsally; calyx tube shortly campanulate-funnel-shaped .......................B. amoena
6. Peduncles slender, inflorescence pendent............................................................................................................... B. quadrangularis
- Peduncles not slender, inflorescence erect .........................................................................................................................................7
7. Leaves sessile or subsessile ............................................................................................................................................B. sessilifolia
- Leaves petiolate ..................................................................................................................................................................................8
8. Leaf blade basally 2-glandular ...................................................................................................................................B. biglandularis
- Leaf blade basally not glandular .........................................................................................................................................................9
9. Leaf blade 3–7 × 1.2–2.8 cm; inflorescence 1–3-flowered .......................................................................................... T. okinawensis
- Leaf blade 5–22.3 × 2–6 cm; inflorescence 5–35-flowered .............................................................................................................10
10. Leaf blade 5–13 × 2–6 cm, secondary veins obscure .......................................................................................................... T. sinensis
- Leaf blade 9.5–22.3 × 3.3–5.5 cm, secondary veins conspicuous................................................................................T. yaeyamensis
Acknowledgements
We thank Ms. Xiao-Lan Wang and Miss Chun-Yu Zou for their help during field work, and Ms. Rong-Mei Zhang for
preparing the illustration. This work was supported by the National Natural Science Foundation of China (31770214)
and the Science and Technology Planning Project of Guangdong Province (2015A030302011).
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