Leptochilus yangjiangensis (Polypodiaceae), a New Fern Species From Guangdong, China

Leptochilus yangjiangensis (Polypodiaceae), a New Fern Species From Guangdong, China | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Ecology and Evolution This is a preprint and has not been peer reviewed. Data may be preliminary. 17 January 2026 V1 Latest version Share on Leptochilus yangjiangensis (Polypodiaceae), a New Fern Species From Guangdong, China Authors : huajing zhou 0009-0009-6542-9469 , Ting Wang , Yi Huang , Yuhan Fang , Bin Zhang 0009-0009-7029-6509 , Guodi Chen , Hong-Feng Chen , and Faguo Wang [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.176864780.00073911/v1 220 views 85 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract A new fern species, Leptochilus yangjiangensis (Polypodiaceae), was discovered during field surveys in Ziluo Mountain, Yangjiang City, Guangdong Province, China. This study provides a detailed morphological description and illustrations of the new species. Morphologically, this species resembles Leptochilus pedunculatus and L. ovatus in plant height, petiole morphology, and soral morphology, but differs from them in its coriaceous laminae and lanceolate fertile fronds. To confirm its taxonomic status, phylogenetic analyses based on three plastid genome regions (rbcL, trnL-F, and rps4 + rps4-trnS) indicated that the new species forms a distinct and well-supported monophyletic clade and is sister to L. dolichophyllus. Furthermore, the complete plastid genome of this new species is reported for the first time. Preliminarily assessed as Data Deficient (DD) according to IUCN guidelines, this new species enriches the diversity of Leptochilus. 1 Introduction Leptochilus Kaulfuss is a group of small to medium-sized ferns belonging to the family Polypodiaceae (1824, 127). It is widely distributed in tropical regions of Asia, with a few species extending to Pacific islands. In China, the genus is mainly concentrated in South China, Southwest China, and Taiwan, serving as an important component of the Asian fern flora. Owing to pronounced morphological variation among species, the taxonomic delimitation of Leptochilus has long been problematic, leading to persistent confusion and repeated revisions of species circumscription. Originally described as a small genus containing only a single species, Leptochilus was later estimated to include approximately 25 species(Zhang et al. 2015). However, species boundaries remained ambiguous, and numerous closely related taxa were independently classified rather than being assigned to this genus. Currently, based on integrative taxonomic studies including phylogenetic analyses of six plastid markers and the nuclear pgiC gene, with a sampling size of 226 accessions covering more than 70 taxa, Zhang et al.(2024) have confirmed that Leptochilus comprises 51 species. Morphologically, Leptochilus is characterized by considerable diversity, with species exhibiting terrestrial, epiphytic, or lithophytic growth habits, reflecting broad ecological adaptability.The leaves are monomorphic or dimorphic, with palmate, digitate, pinnatifid, or pinnate compound forms; sori are orbicular, elongate to linear. During recent field surveys conducted in Ziluo Mountain, Yangjiang City, Guangdong Province, China, a unique fern caught our attention. Preliminary morphological observations indicated that it could not be assigned to any known species of Leptochilus . Through comprehensive morphological and phylogenetic studies, we confirm that this species represents an undescribed new taxon, and described it here as – Leptochilus yangjiangensis . 2 Materials and Methods 2.1 Morphological Study This study first adopted the Leptochilus species identification key constructed by Zhang et al.(2024) for preliminary taxonomic screening, and then retrieved and collated specimen information of related taxa from digital databases (CVH, GBIF, Global Plants on JSTOR). Based on these, field-collected specimens of L. yangjiangensis were used for comparative morphological analyses with its similar species ( L. pedunculatus : isotypes E00507825, E00507826; L. ovatus : holotype MICH 1190696, isotype BM 000036785) and the closely related species L. dolichophyllus (Fu et al. 2025). The overall morphology and habitat of L. yangjiangensis were photographed using a digital camera (Canon EOS R). Morphological data of sterile and fertile fronds of L. yangjiangensis (IBSC 1094808) were measured using MATO (Liu et al. 2023). Scale color and other key traits were observed using a stereomicroscope (OLYMPUS-SZ61) and a biological microscope (OLYMPUS-BX43), and spore ornamentation characteristics were obtained using a Scanning Electron Microscope (JSM-IT210LV). 2.2 Taxon Sampling and Sequencing Leaves from two different individuals of the new species were collected and dried in silica gel. Total DNA was extracted using the cetyltrimethylammonium bromide (CTAB) method(Allen et al. 2006). Genomic DNA was fragmented to approximately 350 bp, followed by end repair, 3’ adenylation, adapter ligation, fragment selection, and PCR amplification to construct a sequencing library. After quality control, the library was sequenced on the Illumina Novaseq platform with PE150, and raw data were filtered for low-quality sequences using fastp before subsequent analyses(Yan et al. 2013). 2.3 Plastome Assembly and Annotation The cleaned reads were de novo assembled into plastome contigs using the GetOrganelle pipeline (v1.7.5+, https://github.com/Kinggerm/GetOrganelle; Jin et al. 2020). To ensure accuracy, the assembled plastome was visually inspected and edited with Bandage v.0.8.1 (Wick et al. 2015) to resolve ambiguities, yielding a complete circular genome. Annotation was performed in Geneious v.11.1.5 (Kearse et al. 2012) with reference to the plastome sequence of the closely related species Leptochilus hemionitideus (NC_040177.1). 2.4 Phylogenetic Analyses Maximum Likelihood (ML) and Bayesian Inference (BI) methods were used for molecular phylogenetic analyses. According to Zhang et al.(2020), target genes were extracted from chloroplast genomes, including 43 rbcL , 42 trnL-F , and 44 rps4 + rps4- trnS, which were downloaded from GenBank for 46 taxa to construct phylogenetic trees (Table 1). Sequence alignment was performed using MAFFT v.7.475(Katoh & Standley, 2013), and regions with low alignment quality were trimmed using trimAl (Capella-Gutiérrez et al. 2009) to optimize the dataset. Phylogenetic trees were constructed using two methods: Maximum Likelihood (ML) and Bayesian Inference (BI). The optimal evolutionary models for ML and BI were selected using ModelTest-NG (Darriba et al. 2020) under the Bayesian Information Criterion (BIC), with the parameter ”-T raxm1” specified for ML and ”-T mrbayes” for BI. ML analysis was implemented in RAxML v.8.2.10 (Stamatakis, 2014), and node support values were evaluated via the Rapid Bootstrap Support (RBS) method with 1000 pseudoreplicates. BI analysis was conducted using MrBayes v.3.2 (Ronquist et al. 2012) with 1,000,000 generations, and trees were sampled every 100 generations. Table 1 Voucher and GenBank accession information of species used for plastid genome assembly. A dash (—) indicates missing data. Leptochilus bolikhamsaiensis Liang Zhang, Khamphanh Thepkaysone & Zhuo Zhou Zhuo Z. et al. LZ156 (KUN) PQ316915 PQ317022 PQ316962 Leptochilus cantoniensis (Baker) Ching Dong S.-Y. 1034 (IBSC) EU363245 — EU363258 Leptochilus cf . ellipticus Chen C.-C. 1065 (H) MH665038 MH665169 MH665102 Leptochilus cf. flexilobus Zhang L.-B. et al. 6710 (CDBI, MO, VNMN) MH768417 MH768545 MH768479 Leptochilus cf. hemitomus Zhang L.-B. et al. 6484 (CDBI, MO, VNMN) MH768427 MH768555 MH768489 Leptochilus cf. pothifolius Cicuzza D. 1998 (HITBC) PQ316873 PQ317004 PQ316943 Leptochilus chingii Li Bing Zhang & Liang Zhang Zhang L.-B. et al. 7453 (CDBI, MO, VNMN) MH768437 MH768565 MH768502 Leptochilus daklakensis Liang Zhang, Li Bing Zhang, X.M.Zhou & Thien Tam Luong Zhang L.-B. et al. 8944 (CCDBI) PQ316871 PQ317002 PQ316941 Leptochilus digitatus (Baker) Noot. A.R. Smith 00–036 (UC) EU482948 EU483044 EU482998 Leptochilus dissimilialatum (Bonap.) Li Bing Zhang & Liang Zhang Zhang L.-B. et al. 6362 (CDBI, MO, VNMN) MH768419 MH768547 MH768481 Leptochilus dolichophyllus H.H.Fu & H.J.Wei She-Lang Jin, Hou-Hua Fu JSL9400 PV442128 PV442128 PV442128 Leptochilus flexilobus (Christ) Li Bing Zhang & Liang Zhang Zhang L. et al. 3050 (KUN) PQ316887 PQ317020 PQ316960 Leptochilus gracilis Z.L.Liang, Liang Zhang & Li Bing Zhang Liang Z.-L. et al. 607 (KUN, CDBI) — MW142228 MW142229 Leptochilus hemionitideus (C.Presl) Noot. Wu S.-K. et al. WS-2437 (KUN) JX103694 JX103778 JX103736 Leptochilus hemitomus (Hance) Noot. Zhang X.-C. 3302 (PE) EU482951 EU483047 EU483001 Leptochilus henryi (Baker) X.C.Zhang DJY04047 (CDBI) MH768428 MH768556 MH768490 Leptochilus heterophyllus (S.K.Wu & K.L.Phan) Christenh. Wu S.-K. et al. WP-136 (KUN) JX520933 JX520937 JX520935 Leptochilus kachinensis Liang Zhang & Li Bing Zhang Deng Y.-F. et al.3200 (CDBI) PQ316865 PQ316996 PQ316936 Leptochilus khammouanensis Liang Zhang, Thepkaysone & Z.Zhou Shui Y.M. et al. LK136 (KUN) PQ316872 PQ317003 PQ316942 Leptochilus leveillei (Christ) X.C.Zhang & Noot. Zhang L.-B. et al. 475 (CDBI) PQ316867 PQ316998 PQ316938 Leptochilus leveillei (Christ) X.C.Zhang & Noot. Zhang X.-C. 4312 (PE) MH665047 MH665179 MH665112 Leptochilus leveillei (Christ) X.C.Zhang & Noot. Zhang L.-B. et al. 6623 (CDBI) — PQ316990 PQ316929 Leptochilus leveillei (Christ) X.C.Zhang & Noot. Zhang L.-B. et al. 7069 (CDBI) PQ316861 PQ316991 PQ316930 Leptochilus luangprabangensis Liang Zhang & Zhuo Zhou Zhuo Z. et al. LZ415 (KUN) PQ316898 PQ317034 PQ316974 Leptochilus macrophyllus (Blume) Noot. Wade 1772 (TAIF) MH768447 MH768573 MH768511 Leptochilus morsei (Ching) Fraser-Jenk. Jin S.-L. et al. JSL7322 (CSH) — PQ317014 PQ316954 Leptochilus multilobus Liang Zhang & X.M.Zhou YLZB2004 (CDBI) PQ316868 PQ316999 — Leptochilus oblongus Li Bing Zhang, Liang Zhang & N.T.Lu Zhang L.-B. et al. 6299 (CDBI, MO, VNMN) MH768429 MH768557 MH768491 Leptochilus ovatifolius Zhe Zhang, S.W.Yao & Yi Huang Zhang L. NAS20180929_4 (KUN) PQ316880 — PQ316951 Leptochilus ovatus Copel. Wade 1583 (TAIF) PQ316863 PQ316994 PQ316934 Leptochilus pedunculatus (Hook. & Grev.) Fraser-Jenk. Zhang L. et al. 2758 (KUN) PQ316917 PQ317024 PQ316964 Leptochilus pedunculatus (Hook. & Grev.) Fraser-Jenk. Zhang L. et al. 4470 (KUN) PQ316907 PQ317043 PQ316981 Leptochilus pedunculatus (Hook. & Grev.) Fraser-Jenk. Zhang L. et al. 2855 (KUN) PQ316882 PQ317012 PQ316952 Leptochilus pentaphyllus (Baker) Li Bing Zhang & Liang Zhang Xu C.-D. A0357 (PE) MH665043 MH665175 MH665108 Leptochilus pteropus (Blume) Fraser-Jenk. Zhang L.-B. et al. 8014 (CDBI, MO, VNMN) MH768412 MH768540 — Leptochilus sanjiangensis Liang Zhang & Li Bing Zhang Jin S.-L. et al. JSL7568 (CSH) PQ316889 PQ317025 PQ316965 Leptochilus saxicola (H.G.Zhou & H. Li) Li Bing Zhang & Liang Zhang Zhang L.-B. et al. 6772 (CDBI, MO, VNMN) MH768441 MH768569 MH768505 Leptochilus scandens H.J.Wei & Yi Huang bis Jin S.-L. et al. JSL8000 (CSH) PQ316911 — PQ316985 Leptochilus shintenensis (Hayata) X.C.Zhang & Noot. Knapp R. 3874 (P) MH768454 — MH768518 Leptochilus sp . Knapp R. 3849 (P) MH768443 MH768571 MH768507 Leptochilus wrightii (Hook. & Baker) X.C.Zhang Li 47 (PE) MH665064 MH665197 MH665130 Leptochilus wusugongii Liang Zhang & Li Bing Zhang Wu S.-K. et al. WS-2591 (KUN) PQ316874 PQ317005 PQ316944 Leptochilus yangjiangensis F. G. Wang, Y. Huang & H.J. Zhou Yi Huang, Guo-Di Chen HY2868 (IBSC) Outgroup Microsorum commutatum Copel. Wade 3768 (TAIF) MH051171 MH113503 MH113470 Microsorum insigne (Blume) Copel. Liu 204 (PE) EU482957 EU483054 EU483008 Microsorum punctatum (L.) Copel. Wade 1390 (TAIF) MH051177 MH113509 MH113476 Phymatosorus longissimus (Blume) Pic.Serm. Cheng X. et al. FB042 (KUN) MT130640 MT130640 MT130640 3 Results 3.1 Characteristics of the Plastid Genome We successfully assembled the complete plastid genome of Leptochilus yangjiangensis , with a total length of 153,177 bp and a GC content of 43.9% (Figure 1). The plastid genome exhibits a typical quadripartite structure, consisting of a pair of inverted repeat (IR) regions (24,880 bp), a large single-copy (LSC) region (81,525 bp), and a small single-copy (SSC) region (21,892 bp). The plastid genome of L. yangjiangensis encodes 115 genes, including 82 protein-coding genes, 29 transfer RNA (tRNA) genes, and four ribosomal RNA (rRNA) genes. Figure 1 Plastome annotation map of Leptochilus yangjiangensis . The darker gray in the inner circle corresponds to GC content. Outside the GC content region, IRA and IRB (two inverted repeat regions); LSC (large single-copy region); and SSC (small single-copy region) are indicated. 3.2 Phylogenetic Relationships Using Microsorum commutatum , M. insigne , M. punctatum , and Phymatosorus longissimus as the outgroup, phylogenetic analyses based on three molecular markers showed that the newly discovered L. yangjiangensis forms a distinct, strongly supported clade, and is sister to L. dolichophyllus (Figure 2). In addition, although L. yangjiangensis is morphologically similar to L. pedunculatus and L. ovatus , the phylogenetic results indicate that it is only distantly related to these two species. Figure 2 Maximum Likelihood (ML) and Bayesian Inference (BI) phylogenetic tree constructed based on plastid genome regions ( rbcL , trnL-F , and rps4 + rps4-trnS ). Numbers adjacent to nodes represent ML bootstrap support values (BS) and BI posterior probabilities (PP) (presented as “BS/PP”). The asterisk (*) at nodes indicates 100% bootstrap support (for ML) and 1.00 posterior probability (for BI). Red text denotes the new species, whereas blue text denotes the species morphologically similar to the new species. 4 Taxonomic Treatment Leptochilus yangjiangensis F. G. Wang, Y. Huang & H.J. Zhou, sp. nov. (Figures 3 and 4) Figure 3 Line drawings of Leptochilus yangjiangensis . (A) Habit. (B) Portion of fertile frond showing sori. (C) Sori and paraphyses. (D) Scale. These illustrations were drawn by Yun-Xiao Liu. Figure 4 Habitat and morphology of Leptochilus yangjiangensis . (A) Habitat. (B) Whole plant. (C) Habit. (D) Close-up of fronds with sori. (E) Scale structure under a biological microscope. (F) Spore morphology under a scanning electron microscope. Photographs by Guo-Di Chen (A–D). Type: CHINA. Guangdong Province: Ziluo Mountain, Xinzhou Town, Yangdong District, Yangjiang City, 21°49′34″ N, 112°22′41″ E, elev. 600 m, 10 October 2025, Yi Huang, Guo-Di Chen HY2868 (Holotype: IBSC!). Diagnosis: Leptochilus yangjiangensis is similar to both L. pedunculatus and L. ovatus , but can be distinguished by the following diagnostic characters (Table 2) : (1) lamina coriaceous (vs. herbaceous in both L. pedunculatus and L. ovatus ); (2) fertile fronds lanceolate (vs. ovate-lanceolate in both L. pedunculatus and L. ovatus ); (3) life form lithophytic (vs. lithophytic in L. pedunculatus and hemiepiphytic in L. ovatus ). Additionally, it can be distinguished from its sister species L. dolichophyllus by the smaller plant size (fertile fronds 26.0–43.0 cm long vs. 43.0–60.0 cm long in L. dolichophyllus ) and distinct life form (lithophytic vs. hemiepiphytic in L. dolichophyllus ). Table 2 Morphological comparison with similar and closely related species. Life form lithophytic lithophytic hemiepiphytic hemiepiphytic Frond dimorphism dimorphic subdimorphic subdimorphic subdimorphic Lamina texture coriaceous herbaceous herbaceous herbaceous Fertile frond shape lanceolate ovate-lanceolate ovate-lanceolate Linear to linear lanceolate Description: Plants perennial, evergreen, lithophytic. Rhizome creeping, densely rooted, uniformly covered with scales throughout; scales brown, ovate-lanceolate, 2.3–3.1 × 0.8–1.0 mm, apex acuminate, margins sparsely serrate. Fronds distant, distinctly dimorphic, coriaceous, glabrous on both surfaces; sterile fronds: 8.0–20.0 cm long; stipe 3.0–79.0 mm long; lamina lanceolate, 8.0–26.0 × 0.7–2.5 cm, apex acuminate to shortly acuminate. Fertile fronds: 14.0–22.0 cm long, reaching 9.0–25.0 cm; lamina lanceolate, 26.0–43.0 × 1.2–2.1 cm, apex long acuminate; adaxially dark green, abaxially pale green. fertile fronds are approximately twice as long as sterile fronds. Sori linear, borne on anastomosing veins, 1 regular row between each pair of lateral veins, obliquely emerging from midrib, most extending to leaf margin. Spores reniform, 21.7–36.7 × 13.5–21.3 μm. Additional specimens examined: CHINA. Guangdong: Xinhui, Gudoushan Nature Reserve, ca. 450 m elev., 3 January 2002, Yue-Hong Yan (HUST 00001462). Guangdong: Jiangmen City, Taishan City, Beidou Town, Mount Ziluo, ca. 420 m elev., 14 December 2025, Guo-Di Chen (IBSC 1095498). Guangdong: Jiangmen City, Taishan City, Beidou Town, Mount Ziluo, ca. 420 m elev., 14 December 2025, Guo-Di Chen (IBSC 1095499). Guangdong: Jiangmen City, Taishan City, Beidou Town, Mount Ziluo, ca. 420 m elev., 14 December 2025, Guo-Di Chen (IBSC 1095500). Guangdong: Jiangmen City, Taishan City, Beidou Town, Mount Ziluo, ca. 420 m elev., 14 December 2025, Guo-Di Chen (IBSC 1095501). Distribution and Habitat: Leptochilus yangjiangensis occurs on mixed forest slopes along ravines in the coastal regions from Yangjiang City to Taishan City, Guangdong Province, China. Etymology: The specific epithet yangjiangensis refers to Yangjiang City, Guangdong Province, China, the type locality of this species. Vernacular Name: 阳江线蕨 (yang jiang xian jue). Conservation status： Approximately 200 mature individuals of Leptochilus yangjiangensis have been observed in Mount Ziluo, Yangjiang City, Guangdong Province, China. This species has also been found in the coastal mountain forests of Taishan City, Guangdong Province, yet the population size at this locality remains unconfirmed. In view of the existing gaps in population data and insufficient distribution information for this species, it is recommended to be categorized as Data Deficient (DD) following the IUCN Red List Categories and Criteria (IUCN, 2024). 5 Discussion As an important group of Polypodiaceae, Leptochilus has long suffered from ambiguous taxonomic delimitation and specimen identification errors due to strong morphological plasticity and overlapping diagnostic characters among some species(Dong et al. 2008; Zhang et al. 2019). The new species described herein has long been misidentified as Leptochilus pedunculatus in southern China. However, recent studies by Zhang et al. (2024) have revealed that the natural distribution of L. pedunculatus may be restricted to low-altitude areas of the Himalayas, while the discovery site of this new species is the coastal mountains of Yangjiang City, Guangdong Province, China. The geographical distributions of the two are completely isolated with no overlap, directly negating the rationality of them being conspecific from a biogeographic perspective. Integrating evidence from molecular phylogenetics, plastid genome data, and morphology, this study confirms that the species is an independent new species of Leptochilus , which not only corrects long-standing taxonomic misplacement but also provides key supplements for understanding the species diversity and distribution pattern of the genus in southern China. Molecular evidence has become an important basis for modern species delimitation and confirmation of the evolutionary independence of new species. Phylogenetic analyses based on molecular markers show that the new species forms a highly supported monophyletic clade (BS=100), and its evolutionary branch is clearly separated from other congeneric species, confirming its independent evolutionary status. The phylogenetic tree also reveals that Leptochilus yangjiangensis forms a sister group relationship with Leptochilus dolichophyllus . The complete plastid genome of L. yangjiangensis successfully obtained in this study further strengthens the molecular evidence chain. Its genomic structure is consistent with the conserved characteristics of previously published Leptochilus plastid genomes(Fu et al. 2025; Min et al. 2018), verifying its correct affiliation at the generic level. This is consistent with the conclusion of Zhang et al. (2019) that plastid genome sequences can effectively resolve the relationships among species within Leptochilus , highlighting the important value of genomic data in addressing the taxonomic ambiguity of this genus. Although L. yangjiangensis is closely related to L. dolichophyllus phylogenetically, nevertheless they exhibit significant morphological differentiation and clear biogeographic isolation. The taxonomic clarification of L. yangjiangensis is of great significance for the revision of Leptochilus . The long-term misidentification of L. yangjiangensis as L. pedunculatus indicates that morphological similarity within Leptochilus is prone to classification errors, and it is urgent to re-evaluate herbarium collections and regional floristic records. Meanwhile, the discovery of this new species expands the known diversity of Leptochilus in southern China, suggesting that there may still be undiscovered cryptic species in subtropical and tropical Asia. In conclusion, based on integrative evidence from phylogenetics, plastid genomics, morphology, and biogeography, this study confirms that Leptochilus yangjiangensis is an independent taxon. This research not only corrects long-standing taxonomic misplacement but also provides new perspectives for understanding the evolutionary differentiation and biogeographic pattern of Leptochilus , and offers a more comprehensive scientific basis for the systematic revision and species diversity conservation of the genus. Future studies should expand the sampling scope of Leptochilus and integrate nuclear gene and plastid genome data to more comprehensively reveal the evolutionary history of Leptochilus species. Author Contributions Hua-Jing Zhou: data curation (lead), formal analysis (equal), investigation (equal), visualization (lead), writing-original draft (lead). Ting Wang: methodology (lead), data curation (equal), formal analysis (lead), visualization (equal), writing-original draft (supporting). Yi Huang: investigation (equal), data curation (equal). Yu-Han Fang: data curation (equal), investigation (equal), supervision (equal). Bin Zhang: data curation (equal), formal analysis (equal). Guo-Di Chen: investigation (lead). Hong-Feng Chen: conceptualization (equal), methodology (equal), supervision (equal). Fa-Guo Wang: conceptualization (lead), investigation (equal), funding acquisition (lead), supervision (lead), writing-review and editing (equal). Acknowledgments. We thank Senior Engineer Yun-Xiao Liu for assisting in the illustration of figures, and Chao-Qi Wang for conducting spore ultrastructure studies via scanning electron microscopy. We acknowledge the financial support of Guangdong Flagship Project of Basic and Applied Basic Research (Grant number: 2023B0303050001). Data Availability Statement Type specimens of the new species described in this study are deposited at IBSC(1094808). Conflicts of Interest The authors declare no conflicts of interest. Ethical statement No ethical statement was reported. Author ORCIDs Hua-Jing Zhou https://orcid.org/0009-0009-6542-9469 Ting Wang https://orcid.org/0000-0002-8622-5459 Yi Huang https://orcid.org/0000-0002-3651-2898 Yu-Han Fang https://orcid.org/0000-0002-1604-1988 Bin Zhang https://orcid.org/0009-0009-7029-6509 Guo-Di Chen https://orcid.org/0009-0003-6907-4569 Hong-Feng Chen https://orcid.org/0000-0002-8415-3260 Fa-Guo Wang https://orcid.org/0000-0002-9326-8000 References Allen, G. C., Flores-Vergara, M. A., Krasynanski, S., Kumar, S., & Thompson, W. F. (2006). A modified protocol for rapid DNA isolation from plant tissues using cetyltrimethylammonium bromide. Nature Protocols , 1 , 2320-2325.Capella-Gutiérrez, S., Silla-Martínez, J. M., & Gabaldón, T. (2009). trimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics (Oxford, England) , 25 , 1972-1973.Darriba, D., Posada, D., Kozlov, A. M., Stamatakis, A., Morel, B., & Flouri, T. (2020). 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Collection Ecology and Evolution Keywords description molecular evolution molecular genetics plants terrestrial Authors Affiliations huajing zhou 0009-0009-6542-9469 Chinese Academy of Sciences South China Botanical Garden View all articles by this author Ting Wang South China Botanical Garden View all articles by this author Yi Huang Chinese Academy of Sciences South China Botanical Garden View all articles by this author Yuhan Fang Chinese Academy of Sciences South China Botanical Garden View all articles by this author Bin Zhang 0009-0009-7029-6509 Chinese Academy of Sciences South China Botanical Garden View all articles by this author Guodi Chen Yangjiang Municipal People's Government View all articles by this author Hong-Feng Chen South China Botanical Garden View all articles by this author Faguo Wang [email protected] South China Botanical Garden View all articles by this author Metrics & Citations Metrics Article Usage 220 views 85 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation huajing zhou, Ting Wang, Yi Huang, et al. 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