Ainsliaea Polystachya (Asteraceae), a New Species from Fujian, China Based on Morphological and Molecular Evidence

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Ainsliaea polystachya (Asteraceae), a new species from Fujian, China based on morphological and molecular evidence

Article in Phytotaxa · April 2021 DOI: 10.11646/phytotaxa.497.3.6

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Ainsliaea polystachya (Asteraceae), a new species from Fujian, China based on morphological and molecular evidence

MEI-JIAO ZHANG1, XIANG-XIU SU2, CHANG AN3, HONG-QING LI1 & ZHEN ZHANG4* 1School of Life Sciences, East China Normal University, 500 Dongchuan Road, Shanghai, 200241, China �[email protected]; https://orcid.org/0000-0003-4378-3538 �[email protected]; https://orcid.org/0000-0003-0658-6295 2Fengyang Nursery, 137 East Wenhua Road, Pingnan County, Fujian,352300 China �[email protected]; https://orcid.org/0000-0001-5117-5407 3College of Pharmacy, Fujian University of Traditional Chinese Medicine, 1 Qiuyang road, Fuzhou, 350122 China �[email protected]; https://orcid.org/0000-0002-7657-7936 4College of Architecture and Urban Planning, Tongji University, 1239 Siping Road, Shanghai 200092, China �[email protected]; https://orcid.org/0000-0003-0271-0973 * Correspondence author

Abstract

A new species, Ainsliaea polystachya X. X. Su & M. J. Zhang is described from Fujian, China, based on morphological traits and molecular markers. The new species is distinct by open panicles, capitula bearing only one flower, and actinomorphic corolla. Keys to the morphologically similar species and phylogenetic allies are proposed.

Keywords: Ainsliaea, new species, Compositae, morphology, phylogeny

Introduction

Ainsliaea Candolle (1838: 13) belongs to the tribe Mutisieae Cassini (1819: 199) in Asteraceae. The monophyly of this genus was established by Mitsui et al. (2008). Morphologically, Ainsliaea is relatively easily recognizable in the field by its unbranched stem, rosulate leaves, capitula with a few flowers (often 3), and plumose pappi. This genus is mainly distributed in southeastern Asia, mostly in south China but with some species in Afghanistan, Bangladesh, Bhutan, India, Indonesia, Japan, Korea, Myanmar, Nepal, Pakistan, Philippines, Thailand and Vietnam (Freire 2007; Gao et al. 2011). Over 50 species have been recorded in Ainsliaea, of which about 40 species (30 endemic) can be found in China (Gao et al. 2011). Undoubtedly, China is the center of species diversity for Ainsliaea. In the recent years, new species of Ainsliaea were described in China (Wang et al. 2010; Zhang et al. 2019; Peng et al. 2020), indicating that some further unrecorded species can be found there. In this study, we described a novel species in Ainsliaea from northeast Fujian, China, which occurs in the Wuyi Mountains covered by the well-developed subtropical evergreen broad-leaf forest. For the added reliability of the new species, both morphological examination and molecular phylogenetic analysis were accomplished.

Material and methods

Morphological observations:—A well-developed population has been surveyed in Jiaocheng District, Fujian Province, southeast China. Some morphological characteristics, such as life form, indumentum, color of leaves and flowers, and inflorescence were recorded in the field or based on cultivated living plants. Size of leaves, flowers, and fruits were measured on the type specimens (and cultivated living plants). Specimens were stored in the Herbarium of the East China Normal University (HSNU) and the duplications are available in the herbaria of the Shanghai Chenshan Botanical Garden (CSH) and the Institute of Botany, Chinese Academy of Sciences (PE). The morphological comparisons between the new species and allied or sympatric species in Ainsliaea were carried out.

Accepted by Alexander Sennikov: 5 Apr. 2021; published: 22 Apr. 2021 277 1 Ainsliaea acerifolia var. acerifolia 1 100/100 Ainsliaea cordifolia 1 100/100 Ainsliaea acerifolia var. subapoda 96/88 Ainsliaea dissecta Ainsliaea apiculata Ainsliaea apiculata var. acerifolia 1 Ainsliaea linearis 84/81 1 Ainsliaea macroclinidioides var. okinawensis 98/93 Ainsliaea oblonga 1 100/99 Ainsliaea macroclinidioides var. okinawensis 1 Ainsliaea fragrans 1 93/82 Ainsliaea fragrans var. integrifolia 85/76 1 1 Ainsliaea simplicissima 81/66 100/100 Ainsliaea simplicissima Ainsliaea polystachya SXX20201101-1 0.81 1 A 1 B 74/57 100/100 Ainsliaea polystachya SXX20201101-2 100/100 Ainsliaea polystachya SXX20201101-3 sp. nov. Ainsliaea macroclinidioides 1 Ainsliaea gracilis 89/81 Ainsliaea walkeri 0.98 0.97 83/68 Ainsliaea grossedentata 88/95 Ainsliaea trinervis 1 Ainsliaea aptera 0.78 96/94 Ainsliaea apteroides 39/- Ainsliaea foliosa 0.57 32/- 1 Ainsliaea fulvipes 73/71 0.99 Ainsliaea macrocephala 0.97 54/60 0.006 40/- 1 Ainsliaea heterantha 100/100 0.77 Ainsliaea hypoleuca 47/- Ainsliaea gongshanensis 1 1 Ainsliaea elegans var. strigosa 100/100 93/91 Ainsliaea pertyoides 1 Ainsliaea henryi 91/80 0.91 45/- Ainsliaea lijiangensis 0.6 Ainsliaea yunnanensis -/55 Ainsliaea henryi 1 Ainsliaea henryi 97/95 Ainsliaea latifolia 97/97 Ainsliaea spicata 1 Ainsliaea glabra 1 100/98 Ainsliaea sutchuenensis 100/100 Ainsliaea lancifolia

1 Pertya robusta Pertya rigidula 100/100 0.99 69/58 0.99 Pertya scandens 75/55 Pertya glabrescens

FIGURE 1. Bayesian phylogenetic tree based on three concatenated loci (ITS+ETS+ndhF). The posterior probability (PP) values are listed above the branches, the maximum likelihood bootstrap supports (MLBS) and the maximum parsimony bootstrap supports (MPBS) below the branches.

Phylogenetic inferences:—Two nuclear (ITS, ETS) and one chloroplast (ndhF) loci were used to infer the phylogenetic position of the novel species. Nine new sequences from three individuals of the new species were acquired (Genbank accessions: MW829545-MW829547 for ITS, MW836815-MW836817 for ETS and MW836818- MW836820 for ndhF). The sequences of other species of Ainsliaea were obtained from the works of Mitsui et al.

278 • Phytotaxa 497 (3) © 2021 Magnolia Press Zhang ET AL. (2008), Funk et al. (2014), and Zhang et al. (2019) (detailed information on the sequences refers to Table 1 in Zhang et al. 2019). The sister genus Pertya Schultz Bipontinus (1862: 109) was set as outgroup according to Mitsui et al. (2008) and Funk et al. (2014). In this paper, three complementary approaches (maximum parsimony, maximum likelihood, and Bayesian inference) were implemented to construct a phylogenetic tree. The methods of alignment and phylogenetic reconstruction were referred to Zhang et al. (2019).

Results

Morphological observations:—Morphologically, the vegetative organs of the new species are similar to Ainsliaea gracilis Franchet (1894: 297) except its less-developed indumentum on leaves and cordiform base of non-decurrent lamina. However, its capitulum resembles A. simplicissima Zhang & Li (2019: 243) in a single flower of the same size and actinomorphic corolla, though its open inflorescence rachis and slim peduncle make it distinguishable. The new species is obviously distinct from the other species with one flower in capitulum, including A. uniflora Schultz Bipontinus (1861: 187), A. lancangensis Qian (2000: 161), and A. simplicissima, in the shape and arrangement of leaves. The sympatric species, such as A. fragrans Champ. ex Bentham (1852: 236) and A. macroclinidioides Hayata (1908: 141), can be easily distinguished from the new species by their shape and size of leaves. Molecular inference:—Three individuals of the new species form a well-supported monophyletic group (PP=1, MLBS=1, MPBS=1) in the phylogenetic tree based on three concatenated loci (Fig. 1, Clade A). It is phylogenetically allied to A. simplicissima and A. fragrans (PP=1, MLBS=81, MPBS=66). The close relationship between the new species and A. simplicissima is supported by their shared character of capitulum, which comprises only one flower. However, the new species and A. fragrans seem to have no obvious morphological similarity to confirm their phylogenetic relationship. Another morphologically related species, A. gracilis is far away from the new species on the phylogenetic tree. The other species in the Clade B are also easily distinguishable from the new species.

Taxonomic Treatment

Ainsliaea polystachya X. X. Su & M. J. Zhang, sp. nov. (多穗兔儿风) (Fig. 2–4)

Type:—CHINA. Fujian: Ningde, Jiaocheng, Hubei, Shitang, Qiduxi, 1st November 2020, Xiang-Xiu Su SXX20201101-1 (holotype HSNU00079855!, Fig. 2); isotypes SXX20201101-2 (HSNU00079856!); SXX20201101-3 (CSH0183101!), SXX20201101-4 (CSH0183102!), SXX20201101-5 (CSH0183103!); SXX20201101-6 (PE02347182!), SXX20201101-7 (PE02347183!).

Diagnosis:—The new species resembles Ainsliaea gracilis in the vegetative organs, differing from the latter by its open panicles, capitulum bearing only one flower, and actinomophic corolla. It is also similar to A. simplicissima in the capitulum, but can be distinguished by its open panicles, longer and slimmer peduncle, and white corolla lobes. Description:—Perennial herbs, 25–75(–90) cm tall when flowering. Roots numerous, lathy, fibrous. Rhizome short or extended, sometimes to 10 cm, cylindrical, 1–4 mm in diameter, with 1–4 scale leaves. Rosette leaves aggregated near the base or the lower part of the stem; petioles unequal, (2–)4–11 cm, sparsely pubescent, subglabrous when mature except the base; leaf blade thickly chartaceous to thinly leathery, cordiform, ovate-cordiform to ovate- lanceolate, 2–5(–11)×1.5–4.5(–8) cm; apex acuminate, acute or obtuse, base shallowly cordiform, margin with sparse acicular denticles, adaxial surface pale green, abaxial surface dark red to purple, sparsely pubescent on the veins and margin to glabrous when mature; lateral veins 2–3 pairs, all veins slightly raised on both sides. Inflorescence in open panicles, arising from the center of the rosette leaves, 20–70 cm high, rachis glabrous, ebracteate. Capitula pedunculate, peduncle 3–7 mm, slim, longest at the middle of the rachis, only 1-flowered; involucre cylindric, ca. 1 mm in diameter; phyllaries 6–8-seriate, herbaceous, glabrous, with purple sides, outer phyllaries ovate, 1–2 mm, inner ones oblong, ca. 7 mm. Florets hermaphrodite; corolla actinomorphic, white, tubular, deeply 5-lobed, lobes linear, 6–8 mm, twisted, 1.5 to 2 times longer than corolla tube; anthers 5, coalescent into a tube, 4–5 mm long, higher than corolla lobes, apical appendages truncate, white, base tails ca. 1.5 mm long; filaments free, beard on corolla throat, ca. 5 mm; style extended out of the anther tube, ca. 11 mm long, glabrous; stigmas 2, apex tuberculate; ovary cylindrical, densely pubescent. Achenes subfusiform, 4–5 mm long, ca. 1 mm in diameter, brown, densely hirsute; pappi dark yellow to brown, plumose, 4–5 mm long. Fl. Oct–Nov; fr. Nov–Jan.

280 • Phytotaxa 497 (3) © 2021 Magnolia Press Zhang ET AL. FIGURE 3. Illustration of Ainsliaea polystachya. A, Habitat. B, Living plants in the field. C, Rosulate leaves. D, Fruiting plant. E, Inflorescence. F, Part of the raceme. G, Leaves. H, Capitulum. I, Flower. J, Style and stigma with synantherous stamen. K, Juvenile fruit. L, Mature fruit.

Ainsliaea polystachya, a new species from Fujian Phytotaxa 497 (3) © 2021 Magnolia Press • 281 FIGURE 4. Line drawing of Ainsliaea polystachya. A, Plant with panicle. B, Leaves. C, Capitulum. D, Phyllaries. E, Flower. F, Pistil. G, Stamen. H, Achene with pappus.

282 • Phytotaxa 497 (3) © 2021 Magnolia Press Zhang ET AL. Distribution:—Two allopatric populations from Chenfeng village (E119.391432, N27.045481), Shicheng town, Zhouning County, and Shitang (E119.300752, N26.739047), Hubei country, Jiaocheng District in Fujian, China. At altitudes of 300–850 m. More than 2,000 individuals exist in the two well-developed populations. Ecology:—The new species grows in forb communities along forest margins, often in moist sites or along the creeks in the subtropical evergreen broad-leaf secondary forest. Etymology:—The species epithet describes the open panicles of this species, which is rather unique in Ainsliaea. Its Chinese name should be spelled as “duo sui tu er feng”. Note:—The new species seem to be a plausible morphological chimera, in which the vegetative organs are similar to A. gracilis while the capitulum resembles A. simplicissima. The new species and A. simplicissima are sympatric, and the latter can be found around the type population of the new species. Additionally, they are closely related on the phylogenetic tree (Fig. 1, Clade A). It seems to indicate that the new species originated from natural hybridization between A. simplicissima and A. gracilis. However, A. gracilis is allopatric and shows a distinct phylogenetic relationship (Fig. 1, Clade A), and they seem to have no chance to hybridize. This phenomenon also appeared in A. simplicissima, whose morphological traits resemble the sympatric species A. trinervis Tseng (1993: 367) except for the number of florets, and its phylogenetic relationships are obviously distinct from A. trinervis (Zhang et al. 2019; Fig. 1). So, further explorations of the genetic background and floral organogenesis of these species are necessary to understand the evolutionary processes in Ainsliaea.

Key of species in the Clade B in the phylogenetic tree of Ainsliaea (Fig. 1)

1. Leaf blades linear to elliptic, at least two times longer than wide, base cuneate to decurrent...... 2 - Leaf blades ovate, cordiform to palmatifid, at most two times longer than wide, base ± cordate...... 5 2. Leaf blades linear to elliptic, apex acute, 1-nerved...... 3 - Leaf blades lanceolate, apex acuminate, 3-nerved...... 4 3. Leaf margins with 4 to 8 teeth on each side...... Ainsliaea linearis - Leaf margins with 1 or 2 subterminal teeth on each side...... Ainsliaea walkeri 4. Capitula 1-flowered...... Ainsliaea simplicissima - Capitula 3-flowered...... Ainsliaea trinervis 5. Leaves lobed...... 6 - Leaves not lobed...... 7 6. Blades palmately lobed; achenes hirsute...... Ainsliaea apiculata - Blades shallowly pinnately lobed; achenes glabrous...... Ainsliaea grossedentata 7. Indumentum well developed on rachis, petiole, and leaf margins...... Ainsliaea fragrans - Indumentum sparse or absent on rachis, petiole, and leaf margins...... 8 8. Abaxial leaf surface green...... Ainsliaea macroclinidioides - Abaxial leaf surface red...... 9 9. Capitula 3-flowered, arranged in spikes...... Ainsliaea gracilis - Capitula 1-flowered, arranged in open panicles...... Ainsliaea polystachya (sp. nov.)

Key of species with only one flower in capitulum in Ainsliaea

1. Leaves partite; pappus biseriate; anther appendages pilose...... Ainsliaea uniflora - Leaves not lobed; pappus uniseriate; anther appendages glabrous...... 2 2. Abaxial leaf surface red...... Ainsliaea polystachya (sp. nov.) - Abaxial leaf surface green...... 3 3. Leaf blades narrowly elliptic or lanceolate; capitula arranged in unilateral racemes...... Ainsliaea simplicissima - Leaf blades 7- to 11-palmatifid, lobes triangular; capitula arranged in open panicles...... Ainsliaea lancangensis

Acknowledgements

This work was funded by the Biodiversity Investigation, Observation and Assessment Program (2019–2023) of the Ministry of Ecology and Environment of China. The authors are grateful to Jian-Hang Zhang and Xin-Jie Jin for their help in molecular experiments, and to Shuai Liao for his help in literature retrieval and specimen examination.

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