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Research Article
Petrocodon wui (Gesneriaceae), a new species from Guizhou, China
expand article infoRen-Bo Zhang, Tan Deng, Nan Li, Fang Wen§|
‡ Zunyi Normal College, Zunyi, China
§ Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, Guilin, China
| National Gesneriaceae Germplasm Resources Bank of GXIB (NGGRB), Gesneriad Conservation Center of China (GCCC), Guilin, China
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Abstract

Petrocodon wui F.Wen & R.B.Zhang (Gesneriaceae), a typically lithophyte occurring in the Danxia areas of north-western Guizhou, China, is described and illustrated as new to science. The new species shows overall similarity with P. chishuiensis Z.B.Xin, F.Wen & S.B.Zhou, which is also its sister species, based on molecular evidence. The new species can be distinguished from P. chishuiensis by the elongated rhizome, the relatively long indumentum on the peduncle, the shape, size and indumentum of calyx lobes, the location of the stamens in the corolla tube and the shape, size and indumentum of the stigma. We provide a diagnosis, detailed description, photographic images and a table with taxonomic notes to distinguish several other morphologically similar Petrocodon species.

Keywords

Didymocarpoideae, flora of Guizhou, lithophyte, new taxon, Petrocodon chishuiensis, taxonomy

Introduction

The genus, Petrocodon Hance (subfamily Didymocarpoideae, family Gesneriaceae), has 47 species, including the newly-published taxon P. asterostriatus F.Wen, Y.G.Wei & W.C.Chou (Möller 2019; GRC 2022; Yang et al. 2022). All known species of this genus are lithophytes (= rock dwelling), either growing in karst topographies or in Danxia landforms. This genus is mainly distributed in karst areas from eastern and south-western China to the northern Indo-China Peninsula, especially in Guangxi of China and North Vietnam (Wei 2018; Wei et al. 2022). Undoubtedly, China is the biodiversity centre of Petrocodon because there are at least 44 species (especially in Guangxi, with 25 species) (Wei 2018). Although many new taxa have been discovered and published in recent years, there were only two new taxa discovered and confirmed from Danxia landforms, namely P. asterocalyx F.Wen, Y.G.Wei & R.L.Zhang (Zhang et al. 2018) and P. chishuiensis Z.B.Xin, F.Wen & S.B.Zhou (Xin et al. 2020). It is well known that all species of Petrocodon are typically lithophilous and usually segregated into unique habitats, for example, karst caves and Danxia gorges, so that most of the species in this genus are narrow endemics (Fu et al. 2022a).

In early August 2021, we conducted a plant diversity survey in Xishui National Nature Reserve in Guizhou Province, China. We noticed an unknown species of Gesneriaceae growing on the surface of Danxia cliff in Niuqingshan, Dabaitang of the Xishui National Reserve. Based on its lithophytic habit and taxonomical characters, we considered it might belong to the genus Petrocodon. Upon closer examination of the flowering specimens in the lab and careful observation of living plants for comparison of vegetative and reproductive organs, we soon discovered several noticeable morphological differences that do not match any known Petrocodon species. Moreover, only two known species of Petrocodon endemic to Danxia landforms were confirmed before this species was discovered. Using morphology or molecular evidence, the new taxon of Petrocodon is recovered as sister to P. chishuiensis, but remarkably different from other species in surrounding cities and counties by some obvious characters. Thus, we concluded it corresponds to a species new to science.

Materials and methods

Taxonomic revision

The studied specimens were collected from the type locality and deposited in the Guangxi Institute of Botany Herbarium (IBK) and the Botany Herbarium of Zunyi Normal College (ZY). The macromorphological features were observed on the specimen sheets and taken from field notes and reports from the conservation nurseries at the National Gesneriaceae Germplasm Resources Bank (NGGB) of the Guangxi Institute of Botany (GXIB) and the Gesneriad Conservation Center of China (GCCC). Micromorphological observations were analysed and photographed using a stereomicroscope (Olympus Optical Microscope CX23). The morphological characters were compared with the protologue and type specimens of previously described Petrocodon species (Wang et al. 1990, 1998; Wei et al. 2010; Wei 2018), in particular those involving new taxa of Petrocodon from Guangxi and adjacent provinces (see notes) and herbarium specimens deposited at relevant herbaria (e.g. HITBC, IBK, IBSC, KUN, PE and VMNM).

The description of the new species follows the terminology used by Wang et al. (1998) and Harris and Harris (2001). Assessment of the conservation status of the new species was made according to the Categories and Criteria of the IUCN (IUCN Standards and Petitions Committee 2022).

Phylogenetic analysis

Leaf material of the undescribed species was collected from the type locality in Xishui County (Guizhou, China) and immediately dried in silica gel for DNA extraction (Chase and Hills 1991). The nuclear ribosomal internal transcribed spacer (ITS) region and plastid trnL-F intron spacer region (trnL-F) were used in the study. Primers, DNA extraction, PCR amplification and sequencing followed Yang et al. (2022). To elucidate the phylogenetic affinities of the undescribed species within the genus, we incorporated 39 samples representing 25 species (Table 1), following Yang et al. (2022). The ingroup contained 37 samples from 23 species of Petrocodon. Primulina dryas (Dunn) Mich. Möller & A.Weber and P. pinnata (W.T.Wang) Yin Z. Wang were chosen as outgroups, based on previous phylogenetic analyses (Möller et al. 2009; Weber et al. 2011; Zhang et al. 2018). We performed phylogenetic analyses of the included Petrocodon species, based on the combined dataset of trnL-F and ITS sequences using Maximum Likelihood (ML). We employed IQ-TREE v.2.0.6 (Nguyen et al. 2015) with 1000 bootstrap replicates (Hoang et al. 2018) and default ModelFinder (Kalyaanamoorthy et al. 2017) and found K3Pu+F+G4 as the best fit substitution model. Tree visualisation was carried out in FigTree v.1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/). Visual comparison of optimal tree topologies of trnL-F and ITS datasets was used to compare topological inconsistencies. Conflicts between tree topologies were considered significant when the inconsistent topologies received bootstrap values ≥ 80% (Fu et al. 2022b). As visual inspection showed no significant topological contradictions for bootstrap support consistency between the trnL-F and ITS datasets (results not shown), the two regions were combined in further analyses.

Table 1.

The voucher and GenBank accession numbers used in this study.

Species name Voucher number trnL-F ITS
Primulina dryas C7a FJ501524 FJ501348
Primulina pinnata G26 FJ501526 FJ501349
Petrocodon retroflexus KX579061 KX579060
Petrocodon nivelolanosus JF697588 JF697576
Petrocodon lithophilus CWH103 KF202303 KF202296
CWH89 KF202302 KF202295
Petrocodon viridescens Y .M.Shui et al 82661 HQ632939 HQ633036
CWH41 KF202304 KF202297
Petrocodon integrifolius M.Moeller MMO 06-865 HQ632940 HQ633037
Petrocodon lui Y .G.Wei 8012 HQ632938 HQ633035
Petrocodon tiandengensis 09413 JX506850 JX506960
Petrocodon ainsliifolius Y .M.Shui et al 44071 HQ632941 HQ633038
CWH88 KF202298 KF202291
Petrocodon hispidus CWH101 KF202301 KF202294
CWH87 KF202300 KF202293
Petrocodon hunanensis WF190107-02 MK941180 MK941179
Petrocodon tongziensis Ren-Bo Zhang SBQ09383 MF872618 MF872617
Petrocodon chishuiensis FW-2014 KF680503 KF680504
Petrocodon wui WF065 OQ716553 OQ694978
Petrocodon coccineus CWH14B KF202299 KF202292
G80E FJ501516 FJ501341
Petrocodon hechiensis KR476563 KR337018
M.Moeller MMO 07-1077 HQ632942 HQ633039
Petrocodon hancei M.Moeller MMO 08-1342 HQ632944 HQ633041
KC904959 KC904956
KC904958 KC904955
GDLC05 KF498253 KF498051
Petrocodon asterocalyx FW-2013 KC904957 KC904954
Petrocodon ferrugineus M.Moeller MMO 06-784 HQ632946 HQ633043
Petrocodon multiflorus HJ01-2 KM232660 KJ475411
Petrocodon coriaceifolius M.Moeller MMO 06-913 HQ632943 HQ633040
Petrocodon scopulorus W.Fang 2010-02 HQ632947 HQ633044
GU350669 GU350637
LJM06753 KR476567 KR337023
Petrocodon dealbatus LJM1209291 KR476565 KR337020
G12B FJ501537 JF697578
LJM-2003-104 GU350668 GU350636

Taxonomic treatment

Petrocodon wui F.Wen & R.B.Zhang, sp. nov.

Figs 1, 2

Diagnosis

Petrocodon wui is distinguishable by the elongated rhizome, the shape, size and indumentum of calyx lobes, the two conspicuous rows of orange glands on throat and the abaxial surface of the corolla lip. It morphologically resembles P. chishuiensis, but can be distinguished by having an elongated rhizome up to 30 cm or longer after years of growth (vs. lacking obvious rhizome in P. chishuiensis, following same order); leaf blade oval-oblong (vs. oblong or oblanceolate) and margin conspicuously undulate and densely ciliate (vs. serrate); cyme 4–10-flowered or more (vs. usually 1–3-flowered); anthers sparsely semi-transparent glands (vs. glabrous) and staminodes pale purple, club-like, glabrous (vs. absent or indistinctive).

Figure 1. 

Petrocodon wui F.Wen & R.B.Zhang, sp. nov. A habit B bracts, showing the abaxial surface C bracteoles D abaxial surfaces of calyx lobe E top view of flower F opened corolla from the dried flower G one of stamens H pistil I seed. Drawings by Tan Deng from the type specimen.

Type

China, Guizhou Province, Zunyi City, Xishui County, Xishui National Nature Reserve, Dabaitang, Niuqingshan, elev. ca. 1100 m, growing on a shaded and moist rock surface on the Danxia cliff in the gorge, Ren-Bo Zhang ZRB2401 (holotype: IBK!, isotypes: ZY!).

Description

Perennial herb, strictly lithophytic. Rhizome brown, abundant fibrous roots, especially at the nodes, rhizome becoming very long and up to 30 cm or longer after years of growth, the lower half of long rhizome usually growing downwards along the surface of rock with lots of fibrous roots, apex of rhizome usually curved and forming a hooked shape, some persistent base of petioles spirally arranged on the surface of rhizome; upper rhizome densely covered with villous multicellular hairs ca. 2 mm long with 4‒6 cells. Leaves in whorls of three, 6–15 crowded in a basal rosette or clustered at the top of elongated rhizome after years of growth, but usually some dried leaves persistent below foliage; petiole green, up to ca. 4 cm long, cylindrical, densely white pubescent; leaf blade chartaceous and thinly coriaceous when dried, oval-oblong, 6‒10 × 1‒3 cm, apex obtuse to acute or subacute, base cuneate, margin entire to inconspicuously or conspicuously undulate and densely ciliate, both surfaces densely white pubescent, lateral veins 4‒5-paired; Inflorescences 1‒4 or more, axillary, cymose, 4–10-flowered or more; peduncle pale green, 1‒4 cm long, ca. 1.5 mm in diameter, densely white villous; bracts 2, opposite, pale green, lanceolate, ca. 10 × 0.5 mm, apex acute, margin entire, both surfaces densely covered with villous multicellular hairs, ca. 1.5 mm long with ca. 3 cells; bracteoles 2, pale green, opposite, narrowly lanceolate, ca. 3 × 0.25 mm, indumentum same as bracts, but hairs on only ca. 2 cells; pedicels pale green, 0.8‒2 cm long, indumentum same as peduncle. Calyx 5-sected to near the base, but base slightly united forming calyx tube ca. 1 mm long; lobes equal, pale green to whitish-green, nearly linear, 6‒8 mm long, 5‒6 mm wide at the base, apex obtuse to rounded, margin entire, outside densely covered with white villous hairs, inside sparsely covered with white villous hairs. Corolla tubular, white, zygomorphic, ca. 2.5 cm long, outside densely white pubescent, inside nearly glabrous, upper part of corolla close to mouth puberulent; corolla tube 1.7–2.2 cm long, ca. 2.5 mm wide at the base of corolla tube and ca. 4.5 mm at the widest part of corolla tube; limb 2-lipped, adaxial lip shorter, 2-lobed to the middle, lobes broadly triangular, ca. 1.5 mm long, ca. 2.5 mm at the bottom of lobe, abaxial lip longer, 3-lobed to the middle or slightly exceeding the middle, lobes ovate, central one longer than lateral ones, ca. 3.5 mm long, lateral ones ca. 2.8 mm long, with two conspicuous rows of orange glands on abaxial lip and corolla throat. Stamens 4, two longer ones adnate to corolla tube ca. 9.5 mm from the base, filaments ca. 4.5 mm long, two shorter ones adnate to corolla tube ca. 8.5 mm from the base, filaments ca. 4 mm long, all filaments linear, straight, but slightly arched at the base and turning into a sheet at the base, white to semi-transparent, densely with brownish-black glands, especially from the middle to the base and glandular-puberulent close to the upper of filament; anthers brownish-purple to dark purple, dorsi-fixed, elliptic to nearly rounded, ca. 1 mm long, ca. 0.9 mm wide, coherent in pairs, thecae confluent at middle, sparsely semi-transparent glands, dehiscing longitudinally. Staminode 1, pale purple, club-like, glabrous, adnate to corolla tube ca. 8 mm from the base. Disc annular, ca. 1 mm high, margin entire. Pistil ca. 2.5 cm long, densely erectly glandular-pubescent; ovary linear-cylindrical, ca. 2 cm long, ca. 1.5 mm wide, 1-loculed, placentas 2, parietal; style ca. 6 mm long, ca. 0.8 mm wide; stigma 2, lobes lamellar, rounded to shallowly spatulate, glabrous, ca. 1 mm long, 0.9‒1 mm wide. Fruit a capsule, ca. 5.5 cm long, linear-cylindrical, 4-valved, pubescent. Seeds appendaged, grain shortly cylindrical, rough, ca. 0.5 mm long, ca. 0.3 mm wide, covered densely verrucate.

Phenology

Flowering occurs in August in the wild; fruiting should occur in October, based on current flowering patterns.

Etymology

We dedicate this new species of Petrocodon to Wu Zheng-Yi (Wu Chengyih) (1916–2013), who devoted over 70 years to the flora of China. The scientific name, “wui”, is the latinisation of Wu Zheng-Yi’s family name. Coincidentally, a plant enthusiast, Lady Xiang-Hong Wu, took this species’ flowering photos in 2017 and sent them to one of the authors (Fang Wen) and her surname is also Wu.

Vernacular name

The Chinese name proposed here is “吴氏石山苣苔.” Phonetically, it is “Wú Shì Shí Shān Jù Tái”.

Distribution and ecology

The new species is endemic to Guizhou Province and known only from the type locality, Xishui National Nature Reserve in Xishui County. It grows on the steep Danxia cliff in an evergreen, broad-leaved forest in a valley of the Danxia landform, at an altitude of 1100‒1600 m. The cliff slope faces northwest at an angle of up to 60 to 80 degrees. The tree cover is up to 12 m tall, the canopy cover is 75%, the shrub layer cover is 85% and the herb layer cover is 35%.

Conservation status

Petrocodon wui is known only from the type locality, which is protected by national and local laws and regulations. However, it is clearly scarce, being known from only one very small area of occupancy, estimated at 20 m2 on a rock surface in a valley of the Danxia landform. Obviously, this area of occupancy of P. wui we found so far is significantly lower than the smallest AOO unit of IUCN which is 4 km2 (2 × 2 km2 grid) for Critically Endangered B2. According to the detailed information from our careful field observations on the surroundings of the type area, the known population has about 50 individuals, half of those being mature individuals and half being seedlings. According to the Guidelines for using the IUCN Red List Categories and Criteria (IUCN Standards and Petitions Committee 2022), P. wui is provisionally assessed as “Critically Endangered, CR B2ab(ii) + C2b” because of its limited distribution and vulnerable habitat.

Taxonomic and phylogenetic notes

The aligned matrix of trnL-F and ITS sequences comprised 1594 characters. Of the 370 (23.21%) variable characters, 222 (13.93%) were parsimony informative. The phylogenetic trees revealed that all sampled Petrocodon taxa clustered together as a monophyletic group (BP = 100%), which is consistent with previous studies (Yang et al. 2022). Three strongly-supported clades are attributed to Petrocodon. Of these, the new species belonged in a moderately-supported subclade (BP = 75%) that also includes P. hunanensis X.L.Yu & Ming Li (Weber et al. 2011), P. tongziensis R.B.Zhang & F.Wen and P. chishuiensis (Petrocodon_sp_FW2014) (Fig. 3). This clade, denoted in Zhang et al. (2019), has four fertile stamens as a synapomorphy and our morphological observation of the new species supported this (Fig. 2). Within this clade, the new species is most closely related to P. chishuiensis (BP = 100%) (Fig. 3), whereas it can be easily distinguished from the latter by its rhizome, leaf blade, flowers number per cyme, bracts, bracteoles, calyx, filaments, anthers and staminodes, all of which are presented in Table 2.

Table 2.

Morphological comparison of Petrocodon wui and P. chishuiensis.

Characters P. wui P. chishuiensis
Rhizome present, becoming very long and up to 30 cm or longer after years of growth lacking rhizome
Leaf blade
Shape oval-oblong oblong or oblanceolate
Margin entire to inconspicuously or conspicuously undulate and densely ciliate serrate
lateral veins 4‒5-paired 5‒6-paired
Flowers number per cyme 4–10-flowered or more usually 1–3-flowered
Bracts
Shape lanceolate oblong
Width ca. 0.5 mm wide ca. 3 mm wide
Indumentum both surfaces densely covered multicellular nodose villous and hairs ca. 1.5 mm long with ca. 3 cells outside whitish pubescent, inside sparsely pubescent
Bracteoles Size ca. 3 × 0.25 mm 6–7 × ca. 1.5 mm
Calyx 5-sected to near the base, but base slightly united forming calyx tube and tube ca. 1 mm long 5-sected from base
Filaments
Length two longer ones 4.5 mm long, two shorter ones ca. 4 mm long two longer ones ca. 9 mm long, two shorter ones ca. 8 mm long
Indumentum densely with brownish-black glands especially from the middle to the base and glandular-puberulent close to the upper of filament densely with glandular-puberulent hairs especially at the base
Anthers
Length ca. 1 mm long ca. 1.8 mm long
Indumentum sparsely semi-transparent glands glabrous
Staminodes pale purple, club-like, glabrous absent or extremely indistinctive
Figure 2. 

Petrocodon wui F.Wen & R.B.Zhang, sp. nov. A plants in bloom in natural habitat B plant in flower C upward view of plant showing the abaxial surfaces of leaf blade and petiole D flowering cyme E cymes, calyx and immature capsule F frontal view of corolla G lateral view of corolla and extended pistil H opened corolla I four fertile stamens J calyx and pistil K mature and dehiscent capsules (Photographed by F. Wen and R.B. Zhang).

Figure 3. 

Phylogenetic tree of Petrocodon generated from Maximum Likelihood (ML) of trnL-F and ITS datasets. Numbers on the branches indicate ML bootstrap values (≥ 50%).

Acknowledgements

We thank Michael LoFurno, Adjunct Professor, Temple University, Philadelphia, the USA, for his editorial assistance and Dr Long-Fei Fu for his help in molecular analysis. This study was financially supported by the National Natural Science Foundation (31860047), the Key Science & Technology Research and Development Project of Guangxi (Guike AD20159091 & ZY21195050), the capacity-building project of SBR of CAS (KFJ-BRP-017-68), the Basic Research Fund of Guangxi Academy of Sciences (CQZ-C-1901), the Foundation of Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain (22-035-26) and the Comprehensive Scientific Investigation by Xishui National Nature Reserve Administration.

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