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Research Article
Two new species and a new combination from Zhejiang, East China
expand article infoYi-Fei Lu‡, Yue-Liang Xu§, Wen-Yuan Xie|, Hong-Wei Zhang, Zi-Hong Zheng#, Xin Cai¤‡, Xiao-Feng Jin‡«
‡ Zhejiang Agricultural and Forestry University, Hangzhou, China
§ Zhejiang Museum of Natural History, Hangzhou, China
| Monitoring Center for Forest Resources in Zhejiang,, Hangzhou, China
¶ Administration of Zhejiang Qingliangfeng National Nature Reserve, Lin'an, China
# Administration of Zhejiang Jiulongshan National Nature Reserve, Suichang, China
¤ Taizhou Academy of Agricultural Sciences, Linhai, China
« Hangzhou Normal University, Hangzhou, China

Corresponding author: Xiao-Feng Jin ( docxfjin@163.com )

Academic editor: Stephen Boatwright
© 2021 Yi-Fei Lu, Yue-Liang Xu, Wen-Yuan Xie, Hong-Wei Zhang, Zi-Hong Zheng, Xin Cai, Xiao-Feng Jin.
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: Lu Y-F, Xu Y-L, Xie W-Y, Zhang H-W, Cai X, Jin X-F (2021) Two new species and a new combination from Zhejiang, East China. PhytoKeys 184: 111-126. https://doi.org/10.3897/phytokeys.184.73327
Open Access

Abstract

As the supplement of the flora of Zhejiang, East China, two new species were described with illustrations. Cerastium huadingense Y.F.Lu, W.Y.Xie & X.F.Jin (Caryophyllaceae) differs from C. qingliangfengicum in having sterile stems absent, leaves sessile, petals slightly longer than sepals, and stamens slightly shorter than sepals. Ixeridium dimorphifolium Y.L.Xu, Y.F.Lu & X.Cai (Asteraceae) differs from I. beauverdianum in having plant stoloniferous, basal leaves dimorphic, involucre 8‒10 mm long, inner phyllaries 8, and florets 7‒10. Paraphlomis setulosa C.Y.Wu & H.W.Li (Lamiaceae) was reviewed and morphological characters of the corolla and stamens of its type and the specimens collected in the field survey were critically examined. With barbate anthers and strongly divergent anther cells, Paraphlomis setulosa was transferred to Sinopogonanthera, and S. setulosa (C.Y.Wu & H.W.Li) H.W.Zhang & X.F.Jin was consequently combined.

Keywords

East China, nomenclatural novelty, seed plant, taxonomy, Zhejiang

Introduction

Zhejiang Province, between the area of 27°06'–31°11'N, 118°01'–123°10'E, is located on the southeastern coast of China, with the whole land area taking up about 1.1% of the country. It is a territory of diverse terrain, including the most mountains in the southwestern area with the highest peak Huangmaojian (1929 m a.s.l.) in Longquan County, central hills, northern alluvial plains and numerous eastern coastal islands. It has a typical subtropical monsoon climate with marked seasonal changes and good climatic conditions. According to the latest report, there are over 4800 species of vascular plants belonging to 1587 genera in 262 families in total, showing the abundance of plant species (Jin et al. unpublished).

Since the winter of 2014, we have prepared the new edition of Flora of Zhejiang. Our field surveys from Zhejiang and specimen examination and identification have resulted in the discovery of 22 new species, three subspecies and four varieties of the seed plants (Chen et al. 2017; Lu et al. 2017a, 2017b, 2019; Zhang et al. 2017; Jin et al. 2018, 2019; Xu et al. 2018; Cai et al. 2019; Xie et al. 2019a, 2019b, 2019c, 2020a, 2020b). As the supplement of the flora of Zhejiang, we herein reported the novelties below.

Results

Cerastium huadingense Y.F.Lu, W.Y.Xie & X.F.Jin, sp. nov.

urn:lsid:ipni.org:names:77221580-1
Chinese name: 华顶卷耳 Figs 1, 2

Latin diagnosis

Species nova haec C. qingliangfengico H.W.Zhang & X.F.Jin affinis est, quae caulibus sterilibus frequent praesentibus, foliis petiolatis, petiolis 5‒9 mm longis, petalis sepalis 2-plo ultra longioribus, 10‒12 mm longis, staminibus sepalis leviter longioribus differt.

Type

China. Zhejiang (浙江): Tiantai (天台), Mount Huading (华顶山), near Huading Temple (华顶讲寺), roadside at forest margin, 29°15'06.31"N, 121°05'24.52"E, alt. 1000 m, 25 April 2020, Xiao-Feng Jin & Yi-Fei Lu 4583 (holotype: ZM; isotypes: HTC, PE).

Description

Herb perennial, 12‒25 cm tall. Roots slender. Stems caespitose, unbranched, erect, green, sparsely to densely white-pubescent. Leaves opposite; basal and lower leaf blades spatulate or narrowly spatulate, 1.2‒3.2 cm long, 0.4‒0.8 cm wide, entire and sparsely ciliate at margin, apex acute, base attenuate, sparsely pubescent on both surfaces; upper leaf blades ovate, ovate-elliptic or oblong, 1.2‒4 cm long, 0.5‒1.5 cm wide, entire and sparsely ciliate at margin, apex acute, base slightly cordate, sparsely pubescent on both surfaces, sessile. Cyme terminal or axillary in upper part of stems, 6‒15-flowered; bracts leaf-like but smaller, narrowly ovate or lanceolate, herbaceous, sparsely pubescent on both surfaces; pedicels slender, 3‒12 mm long, white-pubescent; flowers white, 1.1‒1.3 cm in diam. Sepals 5, lanceolate, green, 5‒6 mm long, apex acuminate, margin membranous, abaxially pilose and glandular-pubescent. Petals 5, narrowly obovate, white, 6‒7 mm long, slightly longer than sepals, 2-lobed for ± 1/4 of length, lobes broadly ovate-oblong, apex acute, base glabrous. Stamens 10, filaments glabrous, slightly shorter than sepals. Styles 5, linear. Capsules cylindric, ± 2 as long as sepals, 10-toothed. Seeds numerous, subtriangular-globose, slightly compressed, brown, ca. 1 mm in diam., tuberculate. Flowering and fruiting April-May.

Figure 1. 

Cerastium huadingense sp. nov. A habit B indumentum on stem C basal leaf D upper leaf E indumentum on peduncle F vertical section of flower (showing sepals, petals, stamens and pistil) G petal H stamen I capsule with sepals and petals J seed (Illustrated by Hong Wang; based on Xiao-Feng Jin & Yi-Fei Lu 4583, ZM).

Figure 2. 

Comparison of Cerastium huadingense and C. qingliangfengicum A–E Cerastium huadingense sp. nov. A habitat B habit C indumentum on stem D flower E capsule F–I Cerastium qingliangfengicum F habitat G indumentum on stem H flower I capsule.

Distribution and habitat

This new species is known from Mount Huading of Tiantai County and Mount Siming of Yuyao County, eastern Zhejiang. It grows under forest, in wet places or roadside at forest margin at the elevation of 900‒1000 m.

Phenology

Flowering and fruiting from mid-April to mid-May.

Etymology

The specific epithet ‘huadingense’ refers to the type locality of the new species.

Conservation status

Vulnerable, VU B2ab(iii)C1 (IUCN 2019). The new species is only known from two localities, Mount Huading in Tiantai and Mount Siming in Yuyao, and occupied less than 400 km2 with about 5000 mature individuals. This species is considered as Vulnerable (VU) according to IUCN Red List Categories and Criteria (IUCN 2019) based on the current data.

Specimen examined

Zhejiang (浙江): Tiantai (天台), Mount Huading (华顶山), Mochi (墨池), in wet place, alt. 900 m, 14 April 2018, Wen-Yuan Xie TT18041411 (PE, ZJFC, ZM), the same locality, under forest, 12 May 1978, Chao-Fang Zhang 3881 (ZJFC, ZM), roadside, alt. 1000 m, 25 April. 2020, Xiao-Feng Jin & Yi-Fei Lu 4584 (HTC, PE, ZM), 11 May 2020, Yi-Fei Lu 235 (HTC, PE, ZJFC, ZM); Mount Huading (华顶山), Ximaopeng (西茅蓬), roadside, alt. 900 m, 26 April 1976, Chao-Zong Zheng 8160 (HTC, ZM). Yuyao (余姚), Mount Siming (四明山), 28 April. 2018, Zheng-Hai Chen & Jian-Sheng Wang YY18042801 (HTC, ZJFC, ZM).

Notes

The genus Cerastium contains ca. 100 species and is almost cosmopolitan, but is mainly distributed in north temperate regions. Twenty-five species were recorded in China (Lu and Morton 2001; Zhang et al. 2008; Yao et al. 2021). A recent study revealed that the monotypic genus Pseudocerastium was merged in Cerastium (Yao et al. 2021).

Cerastium huadingense X.F.Jin, W.Y.Xie & Y.F.Lu is most similar to C. qingliangfengicum in pubescent stems, 5 styles and petals longer than sepals, but differs in having sterile stems absent, leaves sessile, petals slightly longer than sepals, and stamens slightly shorter than sepals. With loosely flowered cymes, pubescent pedicels, and petals 2-lobed at the apex, the new species, C. qingliangfengicum, C. jiuhuashanense and C. wilsonii are similar to each another and form a complex. The morphological characters distinguished from these species are shown in Table 1.

Table 1.
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Comparisons of Cerastium huadingense, C. qingliangfengicum, C. jiuhuashanense and C. wilsonii.

Traits Cerastium huadingense C. qingliangfengicum C. jiuhuashanense C. wilsonii
Leaf
Petiole sessile on sterile stems 5‒9 mm long basal leaves: 4‒5 mm long; upper ones: sessile basal leaves: 3‒6 mm long; upper ones: sessile
Indumentum sparsely pubescent on both surfaces densely pubescent on both surfaces sparsely pubescent on both surfaces sparsely pubescent on both surfaces
Flower
Sepal 5‒6 mm long 6‒8 mm long 5‒7 mm long ca. 6 mm long
Petal 6‒7 mm long, slightly longer than sepals, 2-lobed for ± 1/4 of length 10–12 mm long, ca. twice as long as sepals, 2-lobed for ± 1/2 of length 10–12 mm long, ca. twice as long as sepals, 2-lobed for ± 1/2 of length ca. 12 mm long, ca. twice as long as sepals, 2-lobed for ± 1/2 of length
Filament glabrous glabrous villous on lower part glabrous
Seed shape subtriangular-globose subtriangular-globose subtriangular-globose compressed globose

Sinopogonanthera setulosa (C.Y.Wu & H.W. Li) H.W.Zhang & X.F.Jin, comb. nov.

urn:lsid:ipni.org:names:77221582-1
Chinese name: 小刺毛髯药草 Figs 3, 4

≡ Sinopogonanthera zhejiangensis H.W.Zhang & X.F.Jin, Fl. Zhejiang (New Ed.) 7: 230. 2020, nom. nud.

Basionym

Paraphlomis setulosa C.Y.Wu & H.W.Li, Fl. Reipubl. Popularis Sin. 65(2): 602. 1977. Type: China. Anhui (安徽), Xiuning (休宁), 20 June 1959, Ren-Hua Shan, Xiu Wu 2226 (holotype: NAS 00072429!).

Note

Based on the type specimen and the other related specimen examination, we emended the morphology of the species and described it below

Description emended

Herb perennial, rhizomatous. Stems erect, unbranched, 30‒80 cm tall, 2‒3.5 mm in diam., obtusely 4-angled, shallowly canaliculate, densely retrorsely white-setulose. Leaves opposite; blades elliptic, oblong-ovate, elliptic-ovate or ovate, 7‒19 cm long, 3‒9.5 cm wide, apex acuminate, base broadly cuneate, margin serrate, adaxially green, abaxially pale green, sparsely pubescent on both sides, densely pubescent on mid-ribs, lateral veins 5‒8 pairs; petioles 0.5‒4.5 cm long, adaxially flat, densely white-setulose. Verticillasters axillary, (3‒)7‒14-flowered, 1.5‒3 cm in diam.; pedicels 1‒2 mm long or subsessile; bracteoles subulate, 1.5‒3 mm long, densely setuloae. Calyx tubular-obconical, 7‒9 mm long, outside densely setulose, 5-veined, 5-lobed; teeth oblong-lanceolate or triangular, 2.5‒3 mm long, apex acuminate, white-setulose. Corolla pale purple or pink, 10‒15 mm long, outside densely appressed-pilose, inside sparsely pilose, hairy-annulate on lower part; tube 6‒8 mm long, nearly straight, upper part slightly curved, amplified, with limb 2-lipped; upper lip straight, obovate-oblong, 5‒5.5 mm long, apex emarginate; lower lip 3-lobed, middle lobe broadly obovate, ca. 3.5 mm long, 2.5‒3 mm wide, apex emarginate, lateral lobes ovate, ca. 2.5 mm long, ca. 1.5 mm wide. Stamens 4, didynamous, nearly included, posterior 2 inserted in lower lip of corolla larynx, anterior 2 inserted in upper lip of corolla larynx, filaments sparsely pilose, anthers barbate, cells strongly divergent. Ovary glabrous; style slender, slightly exserted, apex 2-cleft. Disc persistent, cup-shaped. Nutlets obovoid, obtusely trigonous, gray-brown, ca. 2 mm long, apex truncate. Flowering and fruiting June-August.

Specimen examined

Anhui (安徽), Xiuning (休宁), 29 June 1959, Ren-Hua Shan, Xiu Wu 2674 (paratype: NAS). Zhejiang (浙江): Kaihua (开化), Mount Gutian (古田山), roadside under forest, alt. 400 m, 28 June 2018, Xiao-Feng Jin 4230 (HTC, ZM). Lin’an (临安), Tuankou Town (湍口), Tangli Village (塘里村), in grasses under Torreya forest, alt. 550 m, 4 August 2014, Xiao-Feng Jin 3316, 3317, 3318 (ZJFC, ZM), the same locality, under forest, 5 June 2013, Hong-Wei Zhang 0001 (HTC, ZM), cultivated in Changhua (昌化), introduced from Tangli Village (塘里村) of Tuankou Town (湍口镇), 11 June 2021, Yi-Fei Lu & Xiao-Feng Jin 242 (holotype: ZM; isotypes: HTC, KUN for Sinopogonanthera zhejiangensis). Qujiang (衢江), Qianligang (千里岗), 22 July 2017, Zheng-Hai Chen s.n. (HTC).

Figure 3. 

Sinopogonanthera setulosa A lower part of habit B upper part of habit C indumentum on stem D flower with bracteole E opened corolla F anther G stigma H dorsal section of nutlet I ventral section of nutlet (Illustrated by Xiao-Feng Jin; based on Yi-Fei Lu & Xiao-Feng Jin 242, ZM).

Figure 4. 

Comparison of Sinopogonanthera setulosa and S. intermedia A–F Sinopogonanthera setulosa A habitat B rhizome C calyx D inflorescence E corolla F nutlet G–K Sinopogonanthera intermedia G habit H inflorescence I nutlet J calyx K corolla.

Notes

Twenty-eight species with nine varieties within the genus Paraphlomis are currently recognized, which are mainly distributed in tropical and subtropical evergreen and mixed forests (Chen et al. 2021). Li (1965) revised Paraphlomis from China and 21 species with eight varieties were recognized. Wu & Li (1977) published Paraphlomis setulosa and P. reflexa as two new species, and they described that Paraphlomis setulosa had anther cells strongly divergent. The taxonomic treatment of the genus Paraphlomis in Flora of China was just the same as Wu & Li in Flora Reipublicae Popularis Sinicae (Li & Hedge 1994). Guo (1993) established a new genus Pogonanthera close to Paraphlomis but mainly differs from barbate anthers and filament appendages at the apex. Unfortunately, Pogonanthera H.W.Li & X.H.Guo is a later homonym and Li (1993) proposed Sinopogonanthera H.W.Li as the replacement name, which were accepted and adopted (Guo 1995; Zheng 2005).

The genus Sinopogonanthera contains three species, and the key to these three species is shown as follows. Sinopogonanthera caulopteris H.W.Li has verticillasters up to 30-flowered, leaf blade long-elliptic, base cuneate and decurrent, petioles very short or subsessile, and stems narrowly alate. Sinopogonanthera intermedia (C.Y.Wu & H.W.Li) H.W.Li has verticillasters 10‒14-flowered, leaf blade ovate to ovate-rounded, base broadly cuneate, petioles 1‒6 cm long. Sinopogonanthera setulosa is somewhat similar to S. intermedia in attenuate but not decurrent leaf blades at the base and leaf blade petiolate, but differs in having calyx lobes oblong-lanceolate, 2.5‒3 mm long, corolla outside densely appressed-white-pilose, and leaves without glands. The species Sinopogonanthera zhejiangensis (nom. nud.) is consistent to S. setulosa and here reduced to synonym.

Key to the species of Sinopogonanthera

1 Stems narrowly alate; leaf base cuneate and decurrent, petioles very short or subsessile; verticillasters 12‒30-flowered S. caulopteris
– Stems obtusely tetragonal; leaf base broadly cuneate or attanuate, but not decurrent, petioles 0.5‒6 cm long; verticillasters 3‒14-flowered 2
2 Calyx lobes broadly triangular, ca. 1 mm long, outside densely pubescent; corolla outside sparsely puberulent and glabular S. intermedia
– Calyx lobes oblong-lanceolate, 2.5‒3 mm long, outside densely setulose; corolla outside densely pilose S. setulosa

Ixeridium dimorphifolium Y.L.Xu, Y.F.Lu & X.Cai, sp. nov.

urn:lsid:ipni.org:names:77221583-1
Chinese name: 二型叶小苦荬 Figs 5, 6

Latin diagnosis

Species nova I. beauverdiano (H. Lév.) Spring. affinis est, sed planta stolonibus, foliis basilaribus dimorphis, involucris 8‒10 mm longis, phyllis intimis 8, flosculis 7‒10 differt.

Type

China. Zhejiang (浙江): Suichang (遂昌), Jiulongshan National Nature Reserve (九龙山国家级自然保护区), Yangmei Keng (杨梅坑), in grasses along streams, alt. 550 m, 17 April 2020, Xiao-Feng Jin 4568 (holotype: ZM; isotypes: HTC, ZM).

Description

Herb perennial, glabrous totally. Stolons elongate, creeping. Stems erect, 20‒35 cm tall, slender, with sparse branches from base. Basal leaves crowed, present at anthesis, dimorphic; blades of early ones nearly orbicular to long-elliptic, 0.9‒2.5 cm long, 0.7‒1.3 cm wide, apex obtuse and mucronate, base broadly cuneate to rounded, margin entire and with sparsely slender ciliate-teeth, with petioles 1‒3 cm long; blades of later ones narrowly elliptic to lanceolate, 3‒6 cm long, 0.6‒1.1 cm wide, apex acute, base attenuate to a 2‒3 cm long petiole, margin entire or with sparsely slender ciliate-teeth below. Cauline leaves 1‒3; blades linear-lanceolate, 2.5‒7.5 cm long, 0.4‒0.7 cm wide, apex acuminate, base attenuate, margin entire and with/without sparsely slender ciliate-teeth below. Synflorescence corymbiform, with 6‒16 capitulae; capitula with 7‒10 florets, base with slender, long peduncle. Bracts small, broadly ovate-triangular, ca. 1 mm long, margin entire or with 1-pair ciliate-teeth. Involucre narrowly cylindrical, 8‒10 mm long. Phyllaries 2 series, abaxially glabrous; outer phyllaries broadly ovate, 0.5‒0.7 mm long, apex obtuse; inner phyllaries 8, linear-lanceolate, 7‒9 mm long, apex obtuse, short-ciliate. Receptacle flattened, glabrous, alveolate. Florets 7‒10, yellow, corolla 10‒11 mm long, tube 3‒3.5 mm long, ligule linear, apex 5-lobed. Achenes brown or pale brown, narrowly fusiform or linear-fusiform, slightly compressed, 4.5‒5 mm, finely spiculate, with ribs finely spiculate, apex attenuate to a 1‒1.5 mm long slender beak. Pappi pale brown, ca. 5 mm long, scabrid. Flowering and fruiting April‒ July.

Figure 5. 

Ixeridium dimorphifolium sp. nov. A habit B bract C outer phyllary D inner phyllary E opened involucre (showing receptacle) F floret G achene (Illustrated by Xiao-Feng Jin; based on Xiao-Feng Jin 4568, ZM).

Figure 6. 

Comparison of Ixeridium dimorphifolium and I. beauverdianum A–E Ixeridium dimorphifolium sp. nov. A habit B basal part (showing dimorphic leaves and stolon) C cauline leaves D capitula (showing involucre) E capitula (showing florets) F–J Ixeridium beauverdianum F habit G basal part (showing monomorphic leaves) H cauline leaves I capitula (showing involucre) J capitula (showing florets).

Distribution and habitat

Ixeridium dimorphifolium is currently known from only two localities in Suichang and Qingyuan counties, southwestern Zhejiang Province, East China. It grows in grassy roadside along stream or wetland at the elevation of 500‒1450 m.

Phenology

Flowering and fruiting from mid-April to mid-July.

Etymology

The specific epithet ‘dimorphifolium’ refers to two different shapes of basal leaves.

Conservation status

Endangered, EN B2ab(iii) (IUCN 2019). The new species is only known from two localities, and occupied less than 100 km2 with about 1000 mature individuals. This species is considered as Endangered (EN) according to IUCN Red List Categories and Criteria (IUCN 2019) based on the current data.

Specimen examined

Same locality as type, alt. 500 m, 5 May 2017, Yue-Liang Xu 234, 239, 274 (ZM), alt. 550 m, 17 April 2020, Xiao-Feng Jin 4569 (HTC, ZM). Qingyuan (庆元), Huangpi (黄陂), in wetland, alt. 1450 m, 12 July 2020, Xiao-Feng Jin, Yi-Fei Lu & Dan-Qi Liu 4600 (HTC, ZM).

Notes

Ixeridium (A. Gray) Tzvel. is a moderatedly sized genus within the tribe Cichorieae. It contains 15 species and distributes in E and SE Asia, with eight species occurring to China (Shi & Kilian, 2011). The genus Ixeridium is most morphologically similar to Ixeris (Cass.) Cass., but is distinguished in having achenes with 9‒12 prominent but not wing-like ribs (vs. 10 prominent and wing-like ribs) (Shih, 1997). Ixeridium dimorphifolium is similar to I. beauverdianum in having the cauline leaf blades linear to linear-elliptic, margin nearly entire, and pappi brown or pale brown, but differs from the latter in its dimorphic basal leaves, involucres longer, 8‒10 mm long, inner phyllaries 8, florets 7‒10, and plant with stolons.

Morphology of pollens and achenes of the species in Asteraceae has taxonomic significance and sometimes was used to identify similar species (Biyiklioğlu et al. 2018; Joujeh et al. 2020; Nurgül et al. 2021). The micromorphology of achenes and pollen grains of Ixeridium dimorphifolium and I. beauverdianum was shown in Figure 7 and Table 2. The new species, Ixeridium dimorphifolium, has the achenes narrowly fusiform or linear fusiform, with surfaces and ribs both finely similar spiculate, apex gradually narrow to a beak. While those of I. beauverdianum are fusiform, with spicules on surfaces shorter than those on ribs, apex abruptly narrow to a beak. The pollen grains of Ixeridium dimorphifolium and I. beauverdianum are similar to each other.

Figure 7. 

Micromorphology of achenes and pollen grains of Ixeridium dimorphifolium and I. beauverdianum A–E Ixeridium dimorphifolium sp. nov. (from Suichang) F–J Ixeridium dimorphifolium sp. nov. (from Qingyuan) A and F achene shape (scale bar: 1mm) B and G middle part (scale bar: 200μm) C and H spicules on surfaces and ribs (scale bar: 50μm) D and I pollen grains (scale bar: 20μm) E and J surface sculpture (scale bar: 5μm) K–O Ixeridium beauverdianum K achene shape (scale bar: 1mm) L middle part (scale bar: 200μm) M spicules on surfaces and ribs (scale bar: 50μm) N pollen grains (scale bar: 20μm) O surface sculpture (scale bar: 5μm).

Table 2.
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Comparisons of the achenes and pollen grains among Ixeridium dimorphifolium (two samples) and I. beauverdianum.

Traits/species Ixeridium dimorphifolium (Suichang) I. dimorphifolium (Qingyuan) I. beauverdianum
Achene
shape narrowly fusiform, slightly compressed linear-fusiform, slightly compressed fusiform, slightly compressed
length 4.98±0.42 mm 5.54±0.60 mm 4.72±0.34 mm
surface finely spiculate, with ribs finely spiculate finely spiculate, with ribs finely spiculate finely spiculate, with ribs finely spiculate
rib with spicule similar to surface with spicule similar to surface with spicule longer than those on surface
apex gradually narrow to a beak gradually narrow to a beak abruptly narrow to a beak
Pollen
shape tricolporate, spherical tricolporate, spherical tricolporate, spherical
size 27.55±3.60 μm 26.70±2.84 μm 24.62±1.95 μm
surface echinophate echinophate echinophate

Acknowledgements

This work was supported by the National Natural Science Foundation of China (Nos. 32070218, 31770212), the provincial project of the new edition of Flora of Zhejiang (No. 335010-2015-0005), ‘Biodiversity Survey, Monitoring and Evaluation of Ministry of Ecology and Environment’ (Jingning 2020-2021) and the project of Science & Technology Bureau of Hangzhou (20160432B07).

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