Phytotaxa 414 (3): 121–128 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2019 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.414.3.1

Taxonomic notes on the genus Hedyotis sensu lato (Rubiaceae) in China

RUI-JIANG WANG
Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sci-
ences, Guangzhou, Guangdong 510650, China
E-mail: wangrj@scbg.ac.cn

Abstract

The genus Hedyotis sensu lato was splitted into several genera on basis of morphological and molecular evidences. Under
this generic delimitation, seven new taxonomic treatments, including six new synonyms and one new combination, are pro-
posed for the Flora of China. The name Hedyotis tenelliflora that misapplied to Scleromitrion angustifolium is also corrected
with morphological comparison.

Keywords: Dimetia, new combination, new synonym

Introduction

Hedyotis Linnaeus (1753: 101) sensu lato includes more than 550 species worldwide and is one of the large genera
of the family Rubiaceae. Although several molecular phylogenetic analysis provided much valuable information and
robust support for the subdivision of this large genus in recent years (Terrell & Robinson 2003, Guo et al. 2011, Guo
et al. 2013, Wikström et al. 2013, Wang et al. 2014), the taxonomic nomenclature is historically intricate and still a
big obstacle for better understanding the species diversity of this complex, mainly because of not only the thousands
of validly, invalidly, legitimately, illegitimately published names, but also its worldwide distribution. The present
taxonomy of Hedyotis is therefore still regional and needs to be improved based on the comprehensive examination of
the herbarium specimens and field population observation, as well as the molecular phylogenetic analysis.
The following treatments to the Chinese Hedyotis species are mainly based on my extensive examination of the
herbarium materials, detailed observations on the field populations, and comprehensive analysis of molecular data.
The related articles of ICN (Turland et al. 2018) were followed for nomenclature changes.

1. Scleromitrion angustifolium (Chamisso & Schlechtendal) Bentham (1852: 172).—Hedyotis angustifolia Chamisso
& Schlechtendal (1829: 153); Oldenlandia angustifolia (Chamisso & Schlechtendal) Bentham (1861: 151), nom. illeg.
[later homonym of Oldenlandia angustifolia (Michaux 1803: 85) A. Gray (1853: 68). Type:—PHILIPPINES. Luzon,
Chamisso s.n., LE, n.v.).

The species Scleromitrion angustifolium is very common in South China, but it has long been misidentified as Hedyotis
tenelliflora Blume (1826: 971) in many floras, such as Ko (1974: 300, 1999: 41), Chao (1978: 278), Liu & Yang
(1998: 271), Chen (2003: 36), Ko et al. (2005: 121), Chen & Taylor (2011: 170), and Chen (2012: 476). Scleromitrion
angustifolium is morphologically different from Hedyotis tenelliflora in linear-lanceolate, 3–9 mm width (vs lanceolate
leaves in 1–1.7 cm width in H. tenelliflora) leaves, 1–4 (vs 7–9 in H. tenelliflora)-bristled stipule, loculicidally
dehiscent (vs septicidally from base to apex and then loculicidally in H. tenelliflora) capsules (Fig. 1). Geographically,
H. tenelliflora is endemic to Indonesia, especially Java Islands, according to the available collections.

Accepted by Jaideep Mazumdar: 5 Aug. 2019; published: 12 Aug. 2019 121

FIGURE 1. Morphological comparison between Hedyotis tenelliflora (A) and Scleromitrion angustifolium (B-E). A. Type of Hedyotis
tenelliflora (L0057761), B-E. Habit, inflorescence, stipule, and capsules of Scleromitrion angustifolium, respectively. Colorful images
photographed by the author.

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2. Dimetia auricularia (L.) R.J. Wang, comb. nov.—Hedyotis auricularia Linnaeus (1753: 101). Exallage auricularia
(L.) Bremekamp (1952: 142); Metabolos auricularius (L.) Blume ex Bremekamp (1939: 439); Oldenlandia auricularia
(L.) Schumann (1891: 25). Type:—SRI LANKA (holotype: LINN 123.2 [photo]!).
Hedyotis lianshanensis Ko (1995: 41). Type:—CHINA. Guangdong, Lianshan County. 23 May 1958, P.C. Tam 58017 (holotype:
IBK00191483!; isotypes: IFP, KUN331245!, PE00737198!). syn. nov.

Hedyotis auricularia Linnaeus (1753: 101) was designated as the generic type by Chamisso & Schlechtendal (1829:
153) because it was listed at the first by Linnaeus (1753: 101). After a century, it was also recommended to be the
generic standard-species by Hitchcock & Green (1929). But because the character of its indehiscent capsule was unfit
for the “dehiscent capsules” in the generic protologue, as noted by Blume (1826: 990), Bremekamp (1939: 438, 1952:
29, 141) proposed to choose H. fruticosa Linnaeus (1753: 101) as the generic type. This proposal was later formally
approved by the Subcommittee 3C (Javis 1992) and supported by Terrell & Robinson (2003). In addition, considering
that it was a mistake to regard Hedyotis auricularia Linnaeus as the generic type, Bremekamp (1952: 140) established
the new genus Exallage and designated E. auricularia (Linnaeus) Bremekamp (1952: 142) as the generic type after
making a new taxonomic combination based on Hedyotis auricularia Linnaeus. Molecular analysis revealed that
Exallage should be synonymized with Dimetia (Wight & Arnott 1834: 406) Meisner (1839: 160) because of their close
relationship and synapomorphic characters, such as spinous stipules, axillary or terminal inflorescences, septicidally
dehiscent or indehiscent capsules (Guo et al. 2013). Exallage therefore should be merged into Dimetia, resulting in the
new combination of Hedyotis auricularia Linnaeus.
Hedyotis lianshanensis W.C. Ko (1995: 41) has small and lacerate or bristle stipules in comparison with H.
platystipula Merrill (1922: 510) while being published (Ko 1995), but it is much similar to Dimetia auricularia in
terms of leaf, stipule and inflorescence morphology. Hedyotis lianshanensis was also accepted as a “good” species
by Ko (1999) herself and then by Ko et al. (2005: 121) and Chen & Taylor (2011: 163). Field specimen collection
and morphological observation to different populations at the type locality of H. lianshanensis shows that the stipules
usually enlarge a little at the initial developmental stage of inflorescences. This morphological variation falls into the
regular range in Dimetia auricularia populations.

3. Hedyotis baotingensis Ko (1995: 43). Type:—CHINA. Hainan, Baoting County, 11 October 1936, S. K. Lau 27972
(holotype: IBSC0017462!; isotypes: HUH-A00251057 [photo]!, KUN [No. 0777365]!). Fig. 2.
Hedyotis dinganensis Wang et al. (2019: 44). Type:—CHINA. Hainan, Dingan County, 21 April 2018, Wang Qinglong 787641 (holotype:
IBSC0831115!). syn. nov.

Wang et al. (2019) selected Hedyotis yazhouensis F. W. Xing & R. J. Wang (Wang & Xing 2003: 87) and H. paridifolia
Dunn (1912: 366), two morphologically much different species, rather than the conspecific one, H. baotingensis, for
comparison in the diagnosis while publishing H. dinganensis. Hedyotis baotingensis usually grows into a separately
rosulate herb when young (Fig. 2, A & C), but it may have a long prostrate stem from mother body to its derived
elements with aging (Fig. 2, B & D). Hedyotis baotingensis is similar to H. yazhouensis in appearance of their rosulate
figure and capitate inflorescence, but differs obviously in the leaf venation, stipule and flower morphology.

4. Hedyotis loganioides Bentham (1861: 149).—Oldenlandia loganioides (Bentham) Kuntze (1891: 292). Type:—
CHINA. Hong Kong, Hong Kong Island, on the top of Mount Gough, C. Wilford, s.n. (holotype: K000760550
[photo]!).
Hedyotis bodinieri Léveillé (1912: 64).—Oldenlandia bodinieri (Léveillé) Chun (1934: 310). Type:—CHINA. Hong Kong, New Territory,
Summit of Taimoshan, 7 May 1895, E. M. Bodinier 1158 (holotype: E00220484 [photo]!; isotype: HUH-A00233522 [photo]!). syn.
nov.

The type specimens of H. loganioides Bentham (1861: 149) and H. bodinieris Léveillé (1912: 64) were both collected
from Hong Kong. Léveillé (1912) didn’t compare with the sympatric conspecific species, H. loganioides, while
describing his new species. The difference between the two species only limits to the density of pubescence at the
young shoots and peduncles, according to the original description, that is, glabrous in H. loganioides and puberulent
in H. bodinieris. Field observation on the different populations at all known localities showed that this feature varies
a little with the changing of habitats. The young shoots are slight hairy when the habitats are opened and sunny but
glabrous when shady and humid. All stems and shoots become glabrous with aging. The species grows uncommonly
in the mountainous slope, near summit or under the forest, of the Pearl River Delta district of Guangdong province,
China.

TAXONOMIC NOTES ON THE GENUS HEDYOTIS Phytotaxa 414 (3) © 2019 Magnolia Press • 123
FIGURE 2. Hedyotis dinganensis and H. baotingensis. A-B. Holotype (IBSC0831115) and habit of Hedyotis dinganensis (photographed
by Mr. Lang-Xing Yuan), respectively, C-D. Isotype (KUN, No. 0777365) and holotype (IBSC0017462) of Hedyotis baotingensis,
respectively.

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FIGURE 3. The variation pattern of indument and flowers in different populations of Hedyotis matthewii in South China. Photographed
by the author.

TAXONOMIC NOTES ON THE GENUS HEDYOTIS Phytotaxa 414 (3) © 2019 Magnolia Press • 125
5. Hedyotis matthewii Dunn (1909: 376).—Oldenlandia mathewii (Dunn) Chun (1934: 312). Type:—CHINA.
Guangdong Province, Lianzhou River, December 1907, C. G. Matthew, Hongk. Herb. No. 5042 (holotype: HK13684!,
isotypes: IBSC0021613!, K000174523[photo]!). Fig. 3.
Hedyotis assimilis Tutcher (1915: 32).—Oldenlandia assimilis (Tutcher) Chun (1934: 310). Type:—CHINA. Guangdong Province, North
River, Kou Wang Shek, 12 April 1914, W. J. Tutcher, Hongk. Herb. No. 10883A. (lectotype: HK13622!, isolectotype: K000760561
[photo]!, designated by Institute of Botany, Chinese Academy of Sciences (1959: 1615)). syn. nov.
Hedyotis mellii Tutcher (1915: 32).—Oldenlandia mellii (Tutcher) Chun (1934: 313). Type:—CHINA. Guangdong Province, Fungman,
12 April 1914, R. Mell, Hongk. Herb. No. 10942. (holotype: HK13687!; isotype: IBSC0285854!). syn. nov.

These species are very common in South China, especially in Guangdong Province, China. The morphological
differences to delimit them include the stipule shape (entire, glandular-serrate or -serrulate, lobbed, or setose),
inflorescence morphology (e.g., flower number, pedicel length), ratio of corolla lobes to tube, indument of calyxes,
corolla, filaments and styles (glabrous vs hairy). Field observation on the abundant populations growing from the north
to central Guangdong and adjacent provinces revealed that the stipule and leaf shape, pedicel length, the hair density
covering the stems, leaves and inflorescences, inflorescence structure, and the length of corolla lobes and tube vary
continuously and have no much distinct morphological difference. The plants growing in north Guangdong are usually
dense hairy probably because of the high altitude and latitude and low temperature of the habitats. For the populations
in central Guangdong, they often have sparse hairs or are glabrous probably due to the low altitude and latitude and
high temperature. Molecular phylogenetic analysis sampling from its distribution region also indicated that these
species always cluster in the same clade (Guo et al. 2011, Guo et al. 2013). These available evidence indicate that these
three species are actually conspecific.

6. Hedyotis shenzhenensis Chen (2007: 331). Type:—CHINA. Guangdong, Shenzhen, Longgang, Paiyashan Mountain,
16 May 2007, Tao Chen 8043 (holotype: SZG0029990!; isotypes: HUH-A00291304 [photo]!, SZG0029991!,
SZG0029992!, SZG0029993!, SZG0029994!).
Hedyotis shiuyingiae Chen (2008: 283). Type:—CHINA. Hong Kong, Tai Po, 12 July 2007, Tao Chen, Y. W. Lam & K. Y. Lam 2007004
(holotype: HK, n.v.; isotypes: E00268786 [photo]!; HUH-A00291305 [photo]!). syn. nov.

According to the original description, H. shiuyingiae Chen (2008: 283) differs from H. shenzhenensis Chen (2007:
331) by its conspicuous internodes, not rosette leaves, subtetragonous inflorescence axis and branches, narrowly
infundibuliform corolla and stamen position in the flower (Chen 2008, 2012). My field observation and examination
on the populations in the type localities of both species showed that the internode length varies from very short when
growing in dry and open habitats to long in humid and shady places. The stamen position in long- and short-styled
flowers is not so clearly distinguished because of the very small size of the flowers. The previous molecular analyses
also indicated that the two species had very close relationship all the time (Guo et al. 2011, Wang et al. 2015, Wang
et al. 2018). In addition, the type localities of the two species are not so far away from each other. They are therefore
recognized as conspecific.

Acknowledgements

The work was financially supported by National Science Foundation of China (31770217). I am also grateful to Dr.
Kwok-Leung Yip, Mr. Ying-Wai Lam and Mr. Kai-Yip Tam from Agriculture, Fisheries and Conservation Department,
the Government of the Hong Kong Special Administrative Region and Mr. Guo-Bin Jiang, Mr. Lang-Xing Yuan and
Miss Dan Liang for their field assistance. Much thanks should also be extended to the curators of herbaria CUHK, E,
HK, HUH, IBK, IBSC, IFP, K, KUN, L, LE, LINN, PE, SZG for their kind help. I am indebted to Ms. Jian-Rong Li
for preparing the digital images of Hedyotis dinganensis and H. baotingensis.

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