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Primulina nanlingensis (Gesneriaceae), a new species from the Limestone Karst of Guangdong, China
expand article infoJin-Chu Luo‡§, Yuan-Qiu Li|, Ya-Li Li‡§, Ming-Zhao She‡§, Yang-Jin Zeng|, Fa-Guo Wang‡, Hong-Feng Chen‡
‡ South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China
§ University of Chinese Academy of Sciences, Beijing, China
| Guangdong Shimentai National Nature Reserve, Yingde, China

Corresponding author: Hong-Feng Chen ( h.f.chen@scbg.ac.cn )

Academic editor: Alan Paton
© 2025 Jin-Chu Luo, Yuan-Qiu Li, Ya-Li Li, Ming-Zhao She, Yang-Jin Zeng, Fa-Guo Wang, Hong-Feng Chen.
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: Luo J-C, Li Y-Q, Li Y-L, She M-Z, Zeng Y-J, Wang F-G, Chen H-F (2025) Primulina nanlingensis (Gesneriaceae), a new species from the Limestone Karst of Guangdong, China. PhytoKeys 254: 99-111. https://doi.org/10.3897/phytokeys.254.145138
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Abstract

Primulina nanlingensis, a new species of Gesneriaceae from the Karst of Guangdong, China, is described and illustrated. This species is morphologically similar to P. versicolor, but can be distinguished by its larger crenate-margined leaves, fewer flowers per cyme and overall cyme number, ovate-lanceolate bracts with shallow serrations, calyx lobes with 1–3 teeth per side, stamens densely glandular at base and tip and pistil densely glandular-puberulent. It also resembles P. pengii, but has significant differences in its longer corolla, pale yellow corolla colour and ovate-lanceolate bracts. Phylogenetic analyses with ITS and trnL-F sequences revealed that P. nanlingensis is sister to P. versicolor and P. pengii, but isolated from its morphological relatives. The phylogenetic and morphological relationships with similar species are discussed, including detailed descriptions, photographs and distribution information. According to the IUCN Red List Criteria, the new species is assessed as Near Threatened [NT].

Key words:

Gesneriaceae, limestone flora, morphology, phylogeny, taxonomy

Introduction

The genus Primulina Hance (1883) was initially monotypic, with its type specimen collected from the Lianjiang River Basin in northern Guangdong, China. The second species, P. guangxiensis Yan Liu & W.B.Xu, was described in 2011, based solely on morphological characteristics (Liu et al. 2011). Drawing upon molecular and morphological evidence, Primulina underwent subsequent revision and expansion, resulting in the identification of 123 species and eight varieties (Wang et al. 2011; Weber et al. 2011). The revision included numerous species from Chirita sect. Gibbosaccus, two species from Wentsaiboea D.Fang & D.H.Qin and Chiritopsis W.T.Wang which were consolidated into Primulina (Zhou et al. 2016; Xu et al. 2023). Species within the genus Primulina are perennial herbs distinguished by their rhizomatous stems, fleshy opposite leaves, corolla with an infundibuliform tube and 2-lipped lobe, two stamens with coherent anthers and a characteristic chiritoid stigma (Wang et al. 2011; Yang et al. 2023). As of February 2025, the genus Primulina has 244 accepted species, thereby constituting the largest genus in the Chinese Gesneriaceae (Chen et al. 2024; GRC 2025).

Typically residing in the sheltered and moist conditions of karst formations, such as caves and similar microhabitats (Xu et al. 2021), these species are predominantly restricted to limestone niches in southern and south-western China, as well as northern Vietnam (Kang et al. 2014; Li et al. 2019). Globally, the genus includes over 170 species that are endemic to karst regions (Xu et al. 2021; Pan et al. 2022). Guangdong Province is home to 114 species of Gesneriaceae, with a significant proportion of 48 species belonging to Primulina (Song et al. 2023). Since 2017, four new species of Primulina have been described in Guangdong Province, these being P. effusa F.Wen & B.Pan, P. anisocymosa F.Wen, Xin Hong & Z.J.Qiu, P. huangjiniana W.B.Liao, Q.Fan & C.Y.Huang and P. liangwaniae B.M.Wang & Y.H.Tong (Pan et al. 2017; Hong et al. 2019; Huang et al. 2020; Tong et al. 2023).

During a botanical survey conducted in 2024 within the Shimentai Nature Reserve, an unidentified species of Primulina was discovered on two limestone hills. The species was subsequently introduced and cultivated in the greenhouses of the South China Botanical Garden, Chinese Academy of Sciences. From April to June of 2024, the plant exhibited continuous flowering with beautiful pale yellow flowers. After rigorous comparison of this material with herbarium specimens and consultation of relevant references and monographs (Guo et al. 2015; Pan et al. 2016), we confirmed that it represents a new species of Primulina, which we describe in this study. Phylogenetic analysis, utilising ITS and trnL-F sequences, confirmed its position within the genus.

Material and methods

Morphological observation

The material of this new species was collected during a botanical survey conducted at Shimentai Nature Reserve, Yingde City, Guangdong Province. The species was cultivated for further morphological study at the South China Botanical Garden, Chinese Academy of Sciences. Morphological assessments of the new species were carried out using herbarium specimens, with measurements taken from fresh samples. Comparative morphology was conducted with morphologically similar species, utilising both living plants and specimens from institutions such as IBSC, KUN, PE and IBK, as well as digital images from JSTOR Global Plants (http://plants.jstor.org/). Indumentum characteristics were examined by an Olympus-SZ61 stereomicroscope and Olympus-BX43 optical microscope, with photographic documentation accomplished using a Nikon D810 camera.

Molecular sampling

We collected one individual from each of the two natural populations within the protected area and dried the fresh leaf samples using silica gel. Genomic DNA was extracted from the dried leaves using a modified CTAB protocol (Doyle and Doyle 1987). Based on recent phylogenetic studies (Pan et al. 2022; Yang et al. 2023; Chen et al. 2024), we retrieved ITS and trnL-F sequences for 124 Primulina species from GenBank to determine the phylogenetic position of the new species. Petrocodon ainsliifolius W.H.Chen & Y.M.Shui and Petrocodon hancei (Hemsl.) A.Weber & Mich.Möller were selected as outgroup taxa for the analysis (Chen et al. 2024). Corresponding GenBank accession numbers are presented in Table 1.

Table 1.
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Species names and GenBank accession numbers of ITS and trnL-F DNA sequences used for analysis.

Species ITS trnL-F Species ITS trnL-F
Petrocodon ainsliifolius KF202291 KF202298 Primulina lunglinensis var. amblyosepala MK747105 MK746281
Petrocodon hancei KY796057 KY796059 Primulina lungzhouensis KY394931 KY393525
Primulina alutacea KY394847 KY393441 Primulina lutea JX506921 JX506813
Primulina argentea KY394848 KY393442 Primulina lutvittata MK369978 MK369993
Primulina baishouensis KY394849 KY393443 Primulina mabaensis KY394937 KY393531
Primulina balansae MK747141 MK746274 Primulina macrodonta JX506923 JX506815
Primulina beiliuensis KY394850 KY393444 Primulina maculata KU220604 KU220609
Primulina beiliuensis var. fimbribracteata KY394851 KY393445 Primulina malipoensis MK747123 MK746240
Primulina bicolor KY394852 KY393446 Primulina medica KY394940 KY393534
Primulina bipinnatifida KY394853 KY393447 Primulina melanofilamenta MK747158 MK746277
Primulina brachytricha var. magnibracteata MK369979 MK369994 Primulina minor MK747160 MK746290
Primulina carinata KY394858 KY393452 Primulina minutimaculata KY394941 KY393535
Primulina cataractarum MW900263 MW960358 Primulina moi KF498115 KY393536
Primulina chizhouensis KY394860 KY393454 Primulina mollifolia KY394943 KY393537
Primulina confertiflora MK747101 MK746253 Primulina nandanensis KY394947.1 KY393541
Primulina cordata KC190200 KC190207 Primulina nanlingensis_YD1 PQ740297 PQ759014
Primulina cordistigma MK747118 MK746251 Primulina nanlingensis_YD2 PQ740298 PQ759015
Primulina crassirhizoma KY394864 KY393458 Primulina ningmingensis KY394949 KY393543
Primulina crassituba MK747147 MK746230 Primulina obtusidentata KF498096 KY393544
Primulina curvituba MK747137 MK746242 Primulina ophiopogoides KF498062 KY393545
Primulina danxiaensis JX506886 JX506778 Primulina orthandra MK747128 MK746286
Primulina depressa KY394869 KY393463 Primulina pengii KU220603 KU220610
Primulina diffusa KY394871 KY393465 Primulina petrocosmeoides KY394953 KY393547
Primulina dongguanica KY394872 KY393466 Primulina pinnatifida KY394954 KY393548
Primulina dryas KY394875 KY393469 Primulina polycephala KY394955 KY393549
Primulina eburnea JX506891 JX506783 Primulina porphyrea KU173793 KU173799
Primulina efusa MK369976 MK369991 Primulina pseudoeburnea KY394958 KY393552
Primulina fengkaiensis MK369975 MK369990 Primulina pseudoglandulosa KF498138 KY393482
Primulina fengshanensis MK369970 MK369985 Primulina pseudoheterotricha JX506933 JX506824
Primulina fimbrisepala JX506894 JX506786 Primulina pseudolinearifolia MK747140 MK746280
Primulina fimbrisepala var. mollis JX506895 JX506787 Primulina pseudoroseoalba KY394959 KY393553
Primulina fordii MG727881 MG727878 Primulina pungentisepala KY394962 KY393556
Primulina fordii var. dolichotricha MK747125 MK746247 Primulina purpurea KY394964 KY393558
Primulina glandaceistriata MK747114 MK746256 Primulina qingyuanensis KY394965 KY393559.1
Primulina glandulosa KY394887 KY393481 Primulina renifolia KY394966 KY393560
Primulina gongchengensis KY394889 KY393483 Primulina roseoalba KY394972 KY393566
Primulina grandibracteata MK747121 MK746266 Primulina rosulata KU528874 KU528884
Primulina guihaiensis KY394893 KY393487 Primulina rubribracteata KU173791 KU173797
Primulina halongensis KY394895 KY393489 Primulina secundiflora MK747119 MK746279
Primulina hedyotidea JX506905 JX506797 Primulina shouchengensis KY394980 KY393574
Primulina heterochroa KY394898 KY393492 Primulina sichuanensis MK747162 MK746264
Primulina hochiensis JX506903 JX506795 Primulina sinovietnamica MK369973 MK369988
Primulina huaijiensis KF498127 KY393495 Primulina spinulosa KF498063 KY393576
Primulina hunanensis KU220602 KU220608 Primulina subulata KY395020 KY393579
Primulina jiangyongensis KY394902 KY393496 Primulina subulata var. guilinensis KY394967 KY393561
Primulina jingxiensis KY394903 KY393497 Primulina subulatisepala MK747122 MK746246
Primulina jiulianshanensis OP243287 OP243283 Primulina suichuanensis KY395021 KY393580
Primulina jiuwanshanica MK747116 MK746260 Primulina tabacum KY395023 KY393582
Primulina juliae MG727889 MG727873 Primulina tenuituba KY395025 KY393584
Primulina langshanica KY394907 KY393501 Primulina tsoongii KY395029 KY393588
Primulina latinervis KY394908 KY393502 Primulina verecunda KY395031 KY393590
Primulina lechangensis KY394910 KY393504 Primulina versicolor MK747155 MK746252
Primulina leeii KY394911 KY393505 Primulina vestita MK747156 MK746282
Primulina lepingensis KY394913 KY393507.1 Primulina villosissima KY395032 KY393591
Primulina lianpingensis MH343910 MH344542 Primulina wenii MK747148 MK746284
Primulina lijiangensis KY394919 KY393513 Primulina wentsaii KY395033 KY393592
Primulina linearicalyx MH032854 MH032841 Primulina wuae MK747159 MK746265
Primulina linearifolia KY394921 KY393515 Primulina xiziae KY395038 KY393597
Primulina longgangensis JX506916 JX506808 Primulina yangchunensis KY395039 KY393598
Primulina longicalyx KY394927 KY393521 Primulina yangshanensis KY395040 KY393599
Primulina longii JX506917 JX506809 Primulina yangshuoensis KY395042 KY393601
Primulina longnanensis OP243286 OP243282 Primulina yingdeensis KU528876 KU528886
Primulina longzhouensis JX506918 JX506810 Primulina yungfuensis JX506957 JX506848
Primulina lunglinensis KY394930 KY393524 Primulina zhoui MK747104 MK746222

Phylogenetic analyses

We constructed Maximum Likelihood (ML) and Bayesian Inference (BI) phylogenetic trees for Primulina nanlingensis and 124 Primulina species using ITS and trnL-F sequences with PhyloSuite v.1.2.2 (Zhang et al. 2020). Sequences were aligned using MAFFT v.7.471 (Katoh et al. 2019) in PhyloSuite v.1.2.3 and refined with Gblocks 0.91b (Talavera and Castresana 2007), prior to their combination. The optimal nucleotide substitution model was determined using ModelFinder (Kalyaanamoorthy et al. 2017) in PhyloSuite v.1.2.3. The ML tree was built using IQ-tree v.1.6.12 (Nguyen et al. 2015) with the TN+F+R3 model and 5000 bootstrap replicates. The BI tree was constructed using MrBayes v.3.2.6 (Ronquist et al. 2012) with the HKY+F+I+G4 model, running two chains for 3,000,000 generations and sampling every 1000 generations. Both models were selected, based on the Bayesian Information Criterion (BIC). Finally, tree visualisation was performed using ITOL v.7 (https://itol.embl.de/).

Result

Molecular phylogenetic studies

The ITS matrix, consisting of 128 sequences with an aligned length of 952 base pairs (bp), contained 52.5% variable sites and 34.5% informative sites. Similarly, the trnL-F matrix had an aligned length of 1027 bp, with 20.8% variable sites and 9.2% informative sites. Additionally, the combined ITS and trnL-F matrix had a total aligned length of 1689 bp and featured 38.8% variable sites and 24% informative sites. The two populations of the new species from Shimentai National Nature Reserve form a monophyletic group (BS = 100%, PP = 1.00), where BS stands for bootstrap support and PP stands for posterior probability. They also form a sister clade with P. versicolor and P. pengii (BS = 100%, PP = 1.00). All three form a strongly-supported clade with P. alutacea, P. suichuanensis and P. polycephala (Fig. 1).

Figure 1. 

Phylogenetic tree of Primulina generated using Maximum Likelihood (ML) analysis and Bayesian Inference (BI) of the combined ITS and trnL-F sequences. Numbers on branches indicate bootstrap support (≥ 50%) from ML and posterior probabilities (≥ 0.50, rounded to two decimal places) from Bayesian Inference (BI) analyses, while values (< 50% / 0.50) below this threshold are represented by a dash (–). * indicates the new species.

Taxonomic treatment

Primulina nanlingensis J.C.Luo & H.F.Chen, sp. nov.

urn:lsid:ipni.org:names:77359322-1
Figs 2, 3

Type.

China • Guangdong Province, Yingde City, Shimentai National Nature Reserve, 23°28′N, 113°05′E, 620 m elev., growing on top of a cliff on a limestone hill, 7 May 2024 (fl.), J.C. Luo & H.F. Chen LJC00501 (holotype: IBSC; isotypes: IBSC).

Figure 2. 

Primulina nanlingensis J.C.Luo & H.F.Chen A plants in natural habitat B habit in flowering C cyme and frontal view of corolla D the adaxial and abaxial surface of leaf blades E outside surface of bracts F frontal view of corolla G side view of corolla H opened corolla showing stamens, staminodes and colour I stamens J stigma K outside and inside surface of calyx lobes L pistil with calyx lobes and pistil without calyx lobes M infructescence.

Diagnosis.

The new species is similar to Primulina versicolor F.Wen, B.Pan & B.M.Wang in terms of flower shape and corolla colour, but easily distinguished from the larger leaf blades (10–21 × 7–19 cm vs. 8–18 × 6.5–16.5 cm) with a crenate margin (vs. entire); notably lower number of flowers (3–4 cymes, 4–8 flowered vs. 4–8 cymes, 4–24 flowered or more); bracts ovate-lanceolate (vs. broadly oval or suborbicular), with shallowly serrate margins above the middle (vs. entire margins); calyx lobes densely glandular on both surfaces (vs. outside glandular-pubescent inside nearly glabrous) and with 1–3 inconspicuous teeth each side (vs. 3–5-serrate); longer pistil (3.2–3.5 cm vs. 2.5–2.8 cm) and glandular-puberulent (vs. puberulent); filaments white (vs. pale yellow) with densely glandular at base and tip, sparser mid-section (vs. only upper half sparsely glandular-puberulent). Additionally, while the leaf morphology of this new species resembles that of P. pengii W.B.Xu & K.F.Chung, it differs in having a longer corolla length (4.2–5.2 cm vs. 2.8–3.6 cm), pale yellow corollas (vs. white) and ovate-lanceolate bracts with slightly serrate edges above the middle (vs. cordate bracts with entire margins).

Figure 3. 

Primulina nanlingensis J.C.Luo & H.F.Chen A habit B side view of corolla C frontal view of corolla D opened corolla E stamens F pistil with calyx lobes G outside surface of calyx lobes H outside surface of bracts I capsule. Drawn by Mrs. Yunxiao Liu based on J.C. Luo & H.F. Chen LJC00501.

Description.

Herbs, perennial rhizome subterete, 5–8 cm × 1.5–2.5 cm, internodes indistinct. Leaves 6–8, opposite at top of rhizome; blade green, succulent to thickly chartaceous, ovate or broadly ovate to elliptic, 10–21 × 7–19 cm, apex obtuse or subacute, margin crenate, base slightly oblique or symmetrical, leaf surface and petiole densely pubescent, abaxial surface densely villous and the veins on the abaxial surface densely pubescent; lateral veins 4–6-nerved on each side; petiole cross section sub-semicircular or compressed, 2.5–9.5 cm × 0.8–1.5 cm. Cymes axillary, 3–4, 4–8 flowers per cyme; bracts 2 opposite, ovate-lanceolate, 4.1–4.4 cm × 2.2–2.4 cm, outer side is shallowly serrate, apex acuminate, outside pubescent, inside sparsely pubescent; peduncle 5–12.5 cm long, 4–6 mm across, densely pubescent; pedicel 1.2–2.8 cm long, glandular-puberulent. Calyx 5-lobed nearly to the base, 8–15 mm × 1.6–2.5 mm, lanceolate, light green, both surfaces densely glandular-puberulent, with 1–3 inconspicuous teeth on each side. Corolla 4.2–5.2 cm long, pale yellow, throat dark yellow with 2 pale purple stripes, with 3 patches at the sinuses of the 2 upper lip lobes, patches light brown outside and dark purple inside, sometimes the dark purple patches absent and these patches are glandular, outside densely glandular-pubescent; tube infundibuliform, 3.2–3.8 cm long, orifice ca. 1.3 cm in diameter; limb distinctly 2-lipped, adaxial lip 2-lobed bifid to over the middle, lobes oblong, 5–7 mm × ca. 6 mm; abaxial lip 3-lobed to one-third from the top, lobes oblong, 4–6 mm × ca. 4 mm. Stamens 2, adnate to 1.3–1.5 cm above the base of the corolla base; anthers elliptic, 2.5–3 mm long, densely glandular; filaments ca. 15 mm long, white, with the base and upper part sparsely covered with glandular-puberulent; staminodes 3, two lateral ones adnate to ca. 1.3 cm above the corolla base, ca. 7 mm long, densely glandular-pubescent, apex bent, the central one adnate to ca. 3 mm above the base of corolla base, ca. 2 mm long, glabrous; disc annular, ca. 1 mm high, yellow, glabrous. Pistil 3.2–3.5 cm long; ovary linear, ca. 1.8–2.2 cm long, densely glandular-puberulent; style 0.8–1.2 cm long, sparse glandular-puberulent. Stigmas 2-lobed, 2.8–3.6 mm long, shallowly lobed, lobes ca. 1 mm long. Capsule green, mature dark brown, 3.2–4.5 cm × 1.5–2.2 mm, with persistent calyx lobes at base, densely white-villous and pubescent.

Phenology.

Flowering from late April to early June, fruiting from June to August.

Distribution and ecology.

Primulina nanlingensis is known only from two separate limestone hills in the Shimentai National Nature Reserve, Yingde City, Guangdong Province, China. Companion species were calcareous herbs such as Selaginella effusa Alston, S. delicatula (Desv.) Alston., Pilea peltata Hance and Ficus sarmentosa var. henryi (King ex Oliv.) Corner etc.

Etymology.

The species epithet refers to the type locality, the Nanling Mountains.

Vernacular name.

南岭报春苣苔 (Chinese name); Nán Lǐng Bào Chūn Jù Tái (Chinese pronunciation).

Provisional conservation status.

At present, only two populations of Primulina nanlingensis have been discovered on limestone hills in the Shimentai National Nature Reserve, where a substantial area of 35 km2 has been identified as suitable habitat for the species. The two naturally distributed populations are no more than 10 km apart and each population consists of no more than 100 mature individuals. Currently, the two populations are stable, as the habitat is under protection by the administrators of the scenic area. However, considering the overall low number of individuals across the species populations and the conservation measures in place, it could be provisionally classified as Near Threatened [NT] according to the IUCN Red List Categories and Criteria (IUCN Standards and Petitions Committee 2024).

Additional specimens examined.

China • Guangdong Province, Yingde City, Shimentai National Nature Reserve, 23°28′N, 113°05′E, 620 m elev., growing on top of a cliff on a limestone hill, 25 May 2024 (fl.), J.C. Luo & H.F. Chen LJC00502 (IBSC).

Discussion

The karst regions of southern and south-western China, as well as northern Vietnam, are hotspots for diversity of Primulina species, predominantly consisting of endemic species with limited populations confined to isolated sites (Kang et al. 2014; Tong et al. 2020; Wei et al. 2022). The Nanling Mountain Range serves as a habitat for Primulina, where the complex topography and soil heterogeneity foster a high level of species diversity and endemism (Wang et al. 2017). The type localities for both P. nanlingensis and P. versicolor are situated within Yingde City, with a distance of over 50 km between them. Furthermore, P. pengii is found in Yangshan County, Guangdong Province, which is more than 120 km away from P. nanlingensis. Notably, P. nanlingensis closely resembles P. versicolor in both flower shape and colour, but differs in several internal floral structures, such as the pistil with glandular-pubescent and stamens densely glandular at the base and tip, sparser in the middle, fewer flowers per cyme and overall cyme number. Phylogenetic analysis shows that they are closely related, yet their morphological differences suggest that P. nanlingensis represents a new species. Detailed comparisons of the three species are provided in Table 2 and Fig. 4. Finally, given the small population size and restricted distribution to just two locations, conservation efforts for P. nanlingensis are of utmost importance. In all, P. nanlingensis, as a newly-discovered species in Primulina, not only enhances the plant diversity in Naning Mountain, but also provides valuable insights for further study on the local adaptation in karst regions.

Table 2.
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Comparisons of Primulina nanlingensis to P. versicolor and P. pengii, respectively.

Part P. nanlingensis P. versicolor P. pengii
Leaf blade ovate or broadly ovate to elliptic, 10–21 cm × 7–19 cm, margin crenate broadly oval or nearly cordate, 8–18 cm × 6.5–16.5 cm, margin entire ovate to broadly ovate, 14–25 cm × 9.5–15 cm, the margin shallowly repand to crenate
Cyme 3–4, 4–8-flowered 4–8, 4–24-flowered or more 3–4, 4–12-flowered
Bracts ovate-lanceolate, 4.1–4.4 cm × 2.2–2.4 cm, shallowly serrate above the middle, apex acuminate, outside pubescent, inside sparsely pubescent broadly oval or suborbicular, 5–5.5 cm × 4.4–5 cm, apex acute, outside densely appressed pubescent, inside nearly glabrous, margin entire cordate, 2.6–3.2 cm × 2.5–3 cm, the margin entire to shallowly repand, the apex acute, outside pubescent, inside sparsely pubescent
Corolla pale yellow, 4.2–5.2 cm, outside densely glandular-pubescent, inside nearly glabrous; throat dark yellow with 2 pale purple stripes canary yellow, 3.5–4.2 cm, outside densely glandular-pubescent, inside nearly glabrous; throat dark yellow with 2 brownish-purple stripes white, 2.8–3.6 cm long, outside glandular pubescent, inside sparsely puberulent, with 2 pale purple stripes
Calyx lobes 8–15 mm × 1.6–2.5 mm, both surfaces densely glandular, with 1–3 inconspicuous teeth each side 8.5 mm × 2 mm, outside densely glandular-pubescent, inside nearly glabrous, margin 3–5-serrate 8–10 mm × ca. 2 mm, outside glandular pubescent, inside sparsely pubescent, margin serrulate
Stamens filaments ca. 15 mm long, white, densely glandular at base and tip, sparser mid-section, anthers elliptic, 2.5–3 mm long, densely glandular filaments ca. 12.5 mm long, pale yellow, glabrous, but the upper half of filament sparsely glandular puberulent, anthers semicircular, 5–6 mm long, glabrous filaments ca. 14 mm long, white, sparsely puberulent, anthers reniform, ca.4 mm long, puberulent
Pistul 3.2–3.5 cm long, densely glandular-puberulent 2.5–2.8 cm long, densely puberulent 2.4–3.1 cm long, densely puberulent
Figure 4. 

Morphological comparison between Primulina nanlingensis (A1–A4), P. versicolor (B1–B4, photos by Fang Wen.) and P. pengii (C1–C4, photos by Weibin Xu). Legends: Leaf blade (A1, B1, C1); Cyme (A2, B2, C2); Corolla and Calyx lobes (A3, B3, C3); Bracts (A4, B4, C4).

Acknowledgements

We would like to express our gratitude to Yunxiao Liu (IBSC) for preparing the line drawing. We are grateful to Dr. Weibin Xu (Guangxi Institute of Botany, Chinese Academy of Sciences) for providing photos of Primulina pengii and to Dr. Fang Wen (same institute) for photos of P. versicolor and his guidance on plant identification.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was financially supported by the Science and Technology Projects in Guangzhou (E33309) and Guangdong Flagship Project of Basic and Applied Basic Research (2023B0303050001).

Author contributions

Hong-Feng Chen, Yuan-Qiu Li, Jin-Chu Luo: Conceptualisation, Methodology; Hong-Feng Chen, Yuan-Qiu Li, Yang-Jin Zeng, Jin-Chu Luo, Ming-Zhao She: Field investigation, Materials collection; Jin-Chu Luo: Data analyses and visualisation; Jin-Chu Luo: Manuscript writing; Hong-feng Chen, Yuan-Qiu Li, Ya-Li Li, Fa-Guo Wang: Manuscript revision. All authors have read and approved the manuscript.

Author ORCIDs

Jin-Chu Luo https://orcid.org/0009-0003-0526-8962

Yuan-Qiu Li https://orcid.org/0009-0005-8135-500X

Ya-Li Li https://orcid.org/0000-0003-4667-5241

Ming-Zhao She https://orcid.org/0009-0003-1223-6037

Fa-Guo Wang https://orcid.org/0000-0002-9326-8000

Hong-Feng Chen https://orcid.org/0000-0002-8415-3260

Data availability

All of the data that support the findings of this study are available in the main text.

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Jin-Chu Luo and Yuan-Qiu Li contributed equally to this work.
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