Homaliodendron maolanense sp. nov. (Neckeraceae, Bryophyta), a new species from China based on molecular and morphological data

2025 ● 46 ● 5 BryologieBryologie Cryptogamie, Bryologie est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Cryptogamie, Bryologie is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Comptes Rendus Palevol, Cryptogamie sous-sections Algologie, Mycologie. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (électronique / electronic) : 1776-0992 Cryptogamie, Bryologie est indexé dans / Cryptogamie, Bryologie is indexed in: – Biological Abstracts – Current Contents – Science Citation Index – Publications bibliographiques du CNRS (Pascal) Cryptogamie, Bryologie est distribué en version électronique par / Cryptogamie, Bryologie is distributed electronically by: – BioOne® (http://www.bioone.org/loi/cryb) Directeur De la publication / Publication director: Gilles BLOCH Président du Muséum national d’Histoire naturelle réDacteur en chef / editor-in-chief: Denis LAMY assistant De réDaction / assistant editor: Violette GRUNENBERGER ([email protected]) Mise en page / Page layout: Violette GRUNENBERGER réDacteurs associés / associate editors Biologie moléculaire et phylogénie / Molecular biology and phylogeny Bernard GOFFINET Department of Ecology and Evolutionary Biology, University of Connecticut (United States) Mousses d’Europe / European mosses Isabel DRAPER Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain) Francisco LARA GARCÍA Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain) Mousses d’Afrique et d’Antarctique / African and Antarctic mosses Rysiek OCHYRA Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Krakow (Pologne) Bryophytes d’Asie / Asian bryophytes Rui-Liang ZHU School of Life Science, East China Normal University, Shanghai (China) Bioindication / Biomonitoring Franck-Olivier DENAYER Faculté des Sciences Pharmaceutiques et Biologiques de Lille, Laboratoire de Botanique et de Cryptogamie, Lille (France) Écologie des bryophytes / Ecology of bryophyte Nagore GARCÍA MEDINA Department of Biology (Botany), and Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain) couverture / cover: Extrait de la Figure 2/Extract of Figure 2 43 CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.cryptogamie.com/bryologie Homaliodendron maolanense sp. nov. (Neckeraceae, Bryophyta), a new species from China based on molecular and morphological data Jun YANG* Fumin LEI* Ruoling HUANG Yongchang ZHANG College of Life Sciences, Shanghai Normal University, no. 100, Guilin Road, 200234, Shanghai (China) *These authors contributed equally to this work and are considered co-first authors. Lin HE College of Biology and Agriculture, Zunyi Normal University, Ping’an Avenue, 563006, Guizhou (China) Shuiliang GUO Jing YU Dandan LI College of Life Sciences, Shanghai Normal University, no. 100, Guilin Road, 200234, Shanghai (China) [email protected] (corresponding author) Submitted on 14 August 2024 | Accepted on 23 May 2025 | Published on 26 September 2025 Yang J., Lei F., Huang R., Zhang Y., He L., Guo S., Yu J. & Li D. 2025. — Homaliodendron maolanense sp. nov. (Neckeraceae, Bryophyta), a new species from China based on molecular and morphological data. Cryptogamie, Bryologie 46 (5): 43-50. https://doi.org/10.5252/cryptogamie-bryologie2025v46a5. http://cryptogamie.com/bryologie/46/5 ABSTRACT The new species Homaliodendron maolanense sp. nov. is described and illustrated based on its morphological and molecular evidence. The new species can be distinguished from all congeners by the combination of the following features: 1) plants strongly flattened and frondose; 2) secondary stems with bito tripinnate branches and a few foliose pseudoparaphyllia; 3) stem leaves rather asymmetric, strongly falcate, ovate-spathulate, widest at the middle, strongly tapered to base, with a truncate or apiculate apex, as well as double and distinct costae; 4) margins of stem leaves irregularly biserrate with 6-12 sharp teeth, the terminal tooth curved, larger than the others, multicellular, coarse and blunt, entire below; and 5) cells of stem leaves strongly incrassate and smooth, strongly porose at low and basal portions. A key and digital image are presented to separate H. maolanense sp. nov. from other nine Chinese species. KEY WORDS Moss, Neckeraceae, Homaliodendron, molecular phylogeny, new species. 44 CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) Yang J. & Lei F. et al. RÉSUMÉ Homaliodendron maolanense sp. nov. (Neckeraceae, Bryophyta), une nouvelle espèce originaire de Chine basée sur des données moléculaires et morphologiques. La nouvelle espèce Homaliodendron maolanense sp. nov. est décrite et illustrée sur la base de ses caractéristiques morphologiques et moléculaires. La nouvelle espèce se distingue de toutes les espèces apparentées par la combinaison des caractéristiques suivantes : 1) plantes fortement aplaties et frondées ; 2) tiges secondaires avec des branches bià tripennées et quelques pseudoparaphyllia foliacées ; 3) feuilles caulinaires plutôt asymétriques, fortement falciformes, ovales-spatulées, plus larges au milieu, fortement effilées à la base, avec un apex tronqué ou apiculé, ainsi que des côtes doubles et distinctes ; 4) les bords des feuilles caulinaires sont irrégulièrement biserrés avec 6 à 12 dents acérées, la dent terminale étant courbée, plus grande que les autres, multicellulaire, grossière et émoussée, entière en dessous ; 5) et les cellules des feuilles caulinaires sont fortement épaissies et lisses, fortement poreuses dans les parties basses et basales. Une clé et une image numérique sont présentées pour distinguer H. maolanense sp. nov. des neuf autres espèces chinoises. MOTS CLÉS Mousse, Neckeraceae, Homaliodendron, phylogénie moléculaire, espèce nouvelle. INTRODUCTION Homaliodendron Fleisch. sensu stricto is predomainantly reported from tropical and subtropical regions (Wijk et al. 1964; Crosby et al. 1999). Approximately twelve species of the genus are currently recognized worldwide (Fleischer 1906; Iwatsuki & Noguchi 1973; Gangulee 1976; Scott & Stone 1976; Miller et al.1978; Iwatsuki & Tan 1979; Florschütz-de Ward & Florschütz 1979; Kis 1985; Enroth 1989; Noguchi & Iwatsuki 1989; Allen 2010; Forzza 2025; Olsson et al. 2010; Enroth & Koponen 2017). Among them, nine species have been recorded in China (Ninh 1984; Redfearn et al. 1996; Su 1988; Pei et al. 2011; Wu 2011; Jia & He 2013; Enroth & Koponen 2017). Species in the genus Homaliodendron are characterized by their pleurocarpous growth habit, flattened or slightly complanate branching, and ovate to lanceolate leaves. The leaf margins are often serrate or biserrate, and the upper laminal cells tend to be elongate or linear, which distinguishes them from closely related genus Homalia Brid. Additionally, almost members of this genus exhibit strongly flattened and frondose plants with wide-spreading complanate leaves, which is a key diagnostic feature for genus-level identification. To accurately identify whether a moss specimen belongs to Homaliodendron, careful examination of the leaf arrangement, costal structure, and leaf margin dentation is required. The presence of a distinctly single costa, biserrate leaf margins with multicellular teeth, and falcate or curved leaves are key features distinguishing Homaliodendron from similar genera. During a field excursion in Guizhou province, China, we collected a moss with a Homaliodendron-like habit growing on stones. Morphologically, it is characterized by its strongly falcate branch and stem leaves with distinctly double costae. The leaf upper margins and apex are biserrate, coarse and blunt multicellular teeth, which further supports its placement in Homaliodendron. To confirm its genus identity, we constructed a phylogenetic tree including the newly collected moss and six other Homaliodendron species, with Neckera pennata Hedw. as an outgroup. The resulting phylogenetic tree clearly supports the placement of the moss within the genus Homaliodendron. According to its morphological features and the position on the phylogenetic tree, we confirmed that the moss clearly represented a hitherto undescribed species. MATERIAL AND METHODS Examination of spEcimEns Specimen examinations and measurements were conducted using an Olympus-BX53 light microscope, while microphotographs were obtained with a DP74 camera mounted on this microscope. For detailed analysis, leaves located at the middle of stems and branches were selected. Observations and illustrations focused on the upper, medial, and basal leaf cells. Leaf width measurements were taken at the widest part. molEcular protocols A total of 27 samples were selected for molecular analyses, representing six other species of Homaliodendron including H. flabellatum (Sm.) M.Fleisch., H. ligulifolium (Mitt.) M.Fleisch., H. montagneanum (Müll. Hal.) M.Fleisch., H. papillosum Broth., H. pulchrum L.Y.Pei & Y.Jia, H. scalpellifolium (Mitt.) M.Fleisch. These samples were collected from various regions across China, encompassing diverse ecological and geographical regions. Additionally, two species of Circulifolium S.Olsson, Enroth & D.Quandt were included in the analysis. Neckera pennata Hedw. was used as an outgroup. Taxa for the analyses and their voucher specimens are listed in Table 1. The nuclear ribosomal internal transcribed spacer region 5.8S-ITS2-26S (hereafter, ITS2), the tRNA (Gly) (UCC) (trnG), and the trnL-trnF intergenic spacer (trnL-F) have been widely used in phylogenetic analyses for mosses (Draper & Hedenäs 2009; Hedenäs 2012). These three markers were used to construct a phylogenetic tree with the above taxa. The DNA extraction method, PCR amplifications and primers for the three regions followed Li et al. (2018). 45 Homaliodendron maolanense sp. nov., a new species from China CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) phylogEnEtic trEE construction All sequences were manually aligned using PhyDE 0.9971 (Müller et al. 2010). After deletion of incomplete regions for some samples at the beginning and ends of the alignment, the total number of aligned sites from the three genes was 1701. Among them, 756 sites were variable characters and 268 were parsimony-informative. The numbers of sites in each locus based on sequence length, parsimony variable sites, parsimony informative sites, and the optimal substitution models, which were selected for Bayesian analysis, are presented in Table 2. Phylogenetic analysis of three molecular datasets were carried out using Bayesian Inferences (BI) and Maximum Likehood (ML). The best-fitting nucleotide substitution models for each gene region were determined using MrModeltest v. 2.4 (Nylander 2004) in conjunction with PAUP 4.0b10 (Swofford 2003), employing the Akaike Information Criterion (AIC). Bayesian phylogenetic tree was constructed using a Markov Chain Monte Carlo (MCMC) approach implemented in MrBayes v3.2.7 (Ronquist et al. 2012). Four independent MCMC runs, each with four chains, were performed for 1 000 000 generations, with trees sampled every 1 000 generations. Posterior probabilities were calculated after discarding the first 25% of trees as burn-in (Huelsenbeck & Ronquist 2001). Maximum Likelihood analysis was performed using RAxML v.8.2.10 (Stamatakis 2014) with 1 000 bootstrap replicates, employing a GTR + G model of nucleotide substitution for each partition. Phylogenetic trees were visualized and annotated using TreeGraph 2.0 (Stöver & Müller 2010). RESULTS The phylogenetic analysis incorporating the trnL-F, trnG, and ITS2 regions consistently identified all ten Neckeraceae Schimp. species, with high posterior probabilities (PP) and robust bootstrap support (BS) (Fig. 1). Notably, two samples collected from Maolan National Natural Reserve in Guizhou Province formed a single clade (PPBI = 1; BSML = 100), sister to a major clade consisting of H. flabellatum, H. ligulifolium, H. montagneanum, H. papillosum, H. pulchrum and H. scalpellifolium (PP BI = 1; BSML = 100). The phylogenetic analyses show that the above two samples belong to the genus Homaliodendron, and should be placed in Homaliodendron sensu stricto because of their multicellular teeth at the leaf apex and foliose pseudoparaphyllia. The two samples of Homaliodendron from Maolan are morphologically different from the other congeners by their branch and stem leaves, which are strongly falcate with widest part at the middle, and have irregularly biserrate margins with Neckera pennata CN Guizhou 1-20230430057 1/100 1/100 1/99 –/28 –/36 –/20 –/37 –/93 –/67 0.967/86 0.66/56 0.822/48 0.64/42 0.944/53 0.683/25 1/99 –/72 –/78 1/100 1/100 1/100 1/100 –/20 0.864/91 1/98 Neckera pennata CN Guizhou 1-20230504185 Circulifolium microdendron CN Guizhou 1-20181204-12 Circulifolium microdendron CN Guizhou 1-20181206-18 Circulifolium exiguum CN Guizhou 1-20181205-21 Circulifolium exiguum CN Guizhou 2-20181208-13-2 Homaliodendron maolanense sp. nov. CN Guizhou 1-20181204-005 Homaliodendron maolanense sp. nov. CN Guizhou 2-20181204-14 Homaliodendron ligulifolium CN Fujian 201508038 Homaliodendron ligulifolium CN Fujian 201508051 Homaliodendron ligulifolium CN Fujian 201508050 Homaliodendron pulchrum CN Guizhou 1-20181204-27 Homaliodendron pulchrum CN Guizhou 1-20181204-26 Homaliodendron pulchrum CN Hainan 2158029 Homaliodendron montagneanum CN Guangxi 1158 Homaliodendron montagneanum CN Guangxi 1324 Homaliodendron montagneanum CN Guangxi 1177 Homaliodendron montagneanum CN Guangxi 1341 Homaliodendron scalpellifolium CN Fujian 200911011163 Homaliodendron scalpellifolium CN Fujian 201509664 Homaliodendron scalpellifolium CN Guizhou 20170802202 Homaliodendron flabellatum CN Guangxi 1820 1 Homaliodendron flabellatum CN Guangxi 1783 Homaliodendron flabellatum CN Guangxi 214 Homaliodendron papillosum CN Guangxi 1644 Homaliodendron papillosum CN Guangxi 1938 Homaliodendron ligulifolium CN Zhejiang 20170630011 fig. 1 . — Phylogenetic tree with Bayesian posterior probabilities (PPBI) and maximum parsimony bootstrap support (BSML) on the branch, inferred from the combined datasets of trnL-F, trnG and ITS2. In violet, the clade of genus Homaliodendron Fleisch. 46 CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) Yang J. & Lei F. et al. 6-12 sharp teeth, and a large, curved, multicellular, coarse and blunt terminal tooth. Based on the results of the morphological comparisons and molecular phylogenetic analyses, we thought that these two specimens represented a hitherto undescribed species. Here we named the species as Homaliodendron maolanense J.Yang, F.M.Lei, R.L.Huang, Y.C.Zhang, L.He, S.L.Guo, J.Yu & D.D.Li, sp. nov., indicating its type locality in Maolan, Libo County, Guizhou Province, China. taxonomic trEatmEnt Family nEckEracEaE Schimp. Genus Homaliodendron Fleisch. Homaliodendron maolanense J.Yang, F.M.Lei, R.L.Huang, Y.C.Zhang, L.He, S.L.Guo, J.Yu & D.D.Li, sp. nov. (Fig. 2) table 1. — Names, origins and vouchers (herbarium) used for phylogenetic analyses in this study as well as GenBank accession numbers. Taxa Collection locality Coll. number Collector Herbarium GenBank Accession nrITS2 trnL-F trnG Circulifolium exiguum (Bosch & Sande Lac.) S. Olsson, Enroth & D. Quandt Guizhou, China 1-20181205-21 Guo S. L., Li D.D. & Zhang Y. C. SHTU PV600085 PV600058 PV603899 C. exiguum Guizhou, China 2-20181208-13-2 Guo S. L., Li D.D. & Zhang Y. C. SHTU PV600086 PV600059 PV603900 C. microdendron (Mont.) S. Olsson, Enroth & D. Quandt Guizhou, China 1-20181204-12 Guo S. L., Li D.D. & Zhang Y. C. SHTU PV600087 PV600060 PV603901 C. microdendron Guizhou, China 1-20181206-18 Guo S. L., Li D.D. & Zhang Y. C. SHTU PV600088 PV600061 PV603902 Homaliodendron flabellatum (Sm.) M.Fleisch. Guangxi, China 1783 Wang Y. F. & Zuo Q. HSNU PV600089 PV600062 PV603903 H. flabellatum Guangxi, China 1820-1 Wang Y. F. & Zuo Q. HSNU PV600090 PV600063 PV603904 H. flabellatum Guangxi, China 214 Wang Y. F. & Zuo Q. HSNU PV600091 PV600064 PV603905 Homaliodendron ligulifolium (Mitt.) M.Fleisch. Zhejiang, China 20170630011 Guo S. L. SHTU PV600092 PV600065 PV603906 H. ligulifolium Fujian, China 201508038 Wang Y. F., Wei Q. Q. & Zhang B. Y. HSNU PV600093 PV600066 PV603907 H. ligulifolium Fujian, China 201508050 Wang Y. F., Wei Q. Q. & Zhang B. Y. HSNU PV600094 PV600067 PV603908 H. ligulifolium Fujian, China 201508051 Wang Y. F., Wei Q. Q. & Zhang B. Y. HSNU PV600095 PV600068 PV603909 Homaliodendron maolanense sp. nov. Guizhou, China 1-20181204-005 Yang J. et al. SHTU PV600096 PV600069 PV603910 H. maolanense sp. nov. Guizhou, China 2-20181204-14 Yang J. et al. SHTU PV600097 PV600070 PV603911 Homaliodendron montagneanum (Müll. Hal.) M. Fleisch. Guangxi, China 1158 Wang Y. F. & Zuo Q. HSNU PV600098 PV600071 PV603912 H. montagneanum Guangxi, China 1177 Wang Y. F. & Zuo Q. HSNU PV600099 PV600072 PV603913 H. montagneanum Guangxi, China 1324 Wang Y. F. & Zuo Q. HSNU PV600100 PV600073 PV603914 H. montagneanum Guangxi, China 1341 Wang Y. F. & Zuo Q. HSNU PV600101 PV600074 PV603915 Homaliodendron papillosum Broth. Guangxi, China 1644 Wang Y. F. & Zuo Q. HSNU PV600102 PV600075 PV603916 H. papillosum Guangxi, China 1938 Wang Y. F. & Zuo Q. HSNU PV600103 PV600076 PV603917 Homaliodendron pulchrum L.Y. Pei & Y. Jia Guizhou, China 1-20181204-27 Guo S. L., Li, D.D. & Zhang Y. C. SHTU PV600104 PV600077 PV603918 H. pulchrum Guizhou, China 1-20181204-26 Guo S. L., Li, D.D. & Zhang Y. C. SHTU PV600105 PV600078 PV603919 H. pulchrum Hainan, China 2158029 Wang Q. H. & Yu N. N. PE PV600106 PV600079 PV603920 Homaliodendron scalpellifolium (Mitt.) M.Fleisch. Guizhou, China 20170802202 Guo S. L., Li, D.D. & Zhang Y. C. SHTU PV600107 PV600080 PV603921 H. scalpellifolium Fujian, China 200911011163 Wang Y. F., Wei Q. Q. & Zhang B. Y. HSNU PV600108 PV600081 PV603922 H. scalpellifolium Fujian, China 201509664 Wang Y. F., Wu W. Y. & Liu Q. HSNU PV600109 PV600082 PV603923 Neckera pennata Hedwig. Guizhou, China 1-20230430057 Guo S. L., Yu J. & Huang R. L. SHTU PV600083 PV600056 PV603897 N. pennata Guizhou, China 1-20230504185 Guo S. L., Yu J. & Huang R. L. SHTU PV600084 PV600057 PV603898 47 Homaliodendron maolanense sp. nov., a new species from China CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) kEy to Homaliodendron spEciEs in china 1. Capsules immersed ........................................................................................................ H. neckeroides Broth. — Capsules exserted ........................................................................................................................................ 2 2. Stipe leaf apices rounded-obtuse ....................................................H. montagneanum (Müll. Hal.) M.Fleisch. — Stipe leaf apices acute or narrowly obtuse .................................................................................................... 3 3. Upper cells of stem leaves long rhomboid .................................................................................................... 4 — Upper cells of stem leaves rounded, quadrate to hexagonal . ......................................................................... 5 4. Branch leaves nearly similar in width throughout, acute or narrowly obtuse at the apex ................................. ................................................................................................................... H. ligulifolium (Mitt.) M.Fleisch. — Branch leaves usually wider at the base, acute at the apex ....................... H. scalpellifolium (Mitt.) M.Fleisch. 5. Branch leaf cells more or less distinctly mammillose . ..................................................... H. papillosum Broth. — Branch leaf cells smooth .............................................................................................................................. 6 6. Costae single, strong or slender ................................................................................................................... 7 — Costae absent or short and double ............................................................................................................... 9 7. Branch leaves with the same width from base to upper, costae slender ......................................................... 8 — Branch leaves with the base broader than the apex, costa strong .................................. H. crassinervium Thér. 8. Stipe leaves appressed, seta 2.5-4 mm ............................................................ H. flabellatum (Sm.) M.Fleisch. — Stipe leaves spreading, seta longer than 1.5 cm ............. H. fruticosum (Mitt.) S.Olsson, Enroth & D.Quandt 9. Branch leaf margins irregularly dentate at the apex, and costae double, short, sometimes indistinct or absent ........................................................................................................................... H. pulchrum L.Y.Pei & Y.Jia — Branch leaf margins strongly, irregularly biserrate teeth at the apex and costae distinctly double ..................... ................................. H. maolanense J.Yang, F.M.Lei, R.L.Huang, Y.C.Zhang, L.He, J.Yu & D.D.Li, sp. nov. t ypE matErial . — China • Guizhou Province, Libo County, Maolan National Natural Reserve, Forest of Stone Scenic Spot; on rocks; 874 m alt.; 25°17’31.41”N, 107°56’16.75”E; 04.XII.2018; leg. J. Yang et al. 1-20181204-005 (holotype: SHTU!; isotype: KUN!). E tymology . — The specific epithet indicates the type locality: Maolan, Libo County, Guizhou Province, China. habitat. — Growing on rocks under broadleaved forest, along roads. Diagnosis. — Similar to Homaliodendron pulchrum, but differing in having strongly falcate, asymmetrial leaves widest in midleaf, apices with irregularly biserrate teeth, and costae distinctly double. DEscription Plants medium to large-sized, to 10 cm long, strongly flattened, frondose, mostly with glossy, green or yellowish green, becoming tawny with age. Primary stems prostrate, with few reddish rhizoids, secondary stems erect, bipinnately or tripinnately branched, with few foliose pseudoparaphyllia. Central strand absent, outer cortex of 6-8 layers of thickwalled collenchymatous cells, inner cortex of thin-walled parenchymatous cells. Stipe leaves oblong-spathulate to obovate, obtuse at apices, 2.9-3.3 mm long, decurrent; costae double, short, distinct. Stem leaves, 2.3-3.3 mm, horizontally spreading, very asymmetric, strongly falcate, ovate-spathulate, with widest part at the middle, 0.851.45 mm wide, strongly tapered to base, 0.35-0.55 mm at base; apex truncate or apiculate, irregularly biserrate with 6-12 sharp teeth, the terminal tooth curved, larger than the others, 0.2-0.28 mm long, multicellular, coarse and blunt, entire below; often inflexed on one side at base; costae double, disitinct; cells strongly incrassate, smooth, round to longly rhomboidal, 4-8 × 10-18 μm at tip and margin, gradually longer below, 5-10 × 16-38 μm in mid-leaf, 5-9 × 35-95 μm at base near costa, basal leaf cells distinctly porose, at margin becoming smaller, shortly rhomboid to irregular rectangular. Branch leaves similar to stem leaves in shape, but smaller in size, 2.0-2.5 mm long, strongly falcated, costae double, dentate at the apex, with widest part at the middle, 0.75-0.90 mm wide, 0.32-0.34 mm at base; cells similar to those of stem leaves but smaller. Sporophytes not seen. DISCUSSION Fleischer (1905-1906) established the genus Homaliodendron, dividing it into two sections: Homaliodendron and Circulifolia Fleisch. Species in section Circulifolia differ from these in section Homaliodendron by having smaller, complanate leaves, minute, crenulate leaf apices and filiform pseudoparaphyllia (Ninh 1984; Buck 1998). Olsson et al. (2010) elevated Circulifolia to the generic level, naming it Circulifolium S.Olsson, Enroth & D.Quandt. In our phylogenetic analyses, Circulifolium exiguum and C. microdendron are confirmed as a distinct genus, differing from Homaliodendron stricto sensu by their strongly complanate leaves with minute, crenulate leaf dentation and filiform pseudoparaphyllia, consistent with the findings on the phylogenetic 48 CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) Yang J. & Lei F. et al. AD E FGH IJ K LM NOPQ RST UVW XYZ AA AB AC AD C B A, E, I B, F, J C, D, G, H L-Z AA-AB AC-AD fig. 2. — Homaliodendron maolanense sp. nov.: A, B, apical cells of branch leaves; C, upper cells of branch leaf; D, basal cells of branch leaf; E, F, apical cells of stem leaves; G, upper cells of stem leaf; H, basal cells of stem leaf; I, J, apical cells of stipe leaves; K, cross-section of stem; L-Q, branch leaves; R-W, stem leaves; X-Z, stipe leaves; AA, AB, pseudoparaphyllia; AC, plant when wet; AD, plant when dry (all from SHTU 2-20181204-14). Scale bars: A, C-E, G-I, AA, AB, 50 µm; B, F, J, 200 µm; K, 100 µm; L-Z, 500 µm; AC, AD, 500 mm. 49 Homaliodendron maolanense sp. nov., a new species from China CRYPTOGAMIE, BRYOLOGIE • 2025 • 46 (5) relationships within the Pinnatella clade of the Neckeraceae (Olsson et al. 2010). Additionally, the newly identified species H. maolanense sp. nov., along with six other species of Homaliodendron stricto sensu, form a well-supported monophyletic clade. In the study of Indochinese Homaliodendron, Ninh (1984) highlighted that the shape and arrangement of stem leaves and branch leaves were the most important characteristics for species identification. In the clade of seven Homaliodendron species on the phylogenetic tree, H. ligulifolium can be easily distinguished from other species by their lingulate, oblonglingulate or broadly lingulate stem leaves. Homaliodendron pulchrum, first described by Pei et al. (2011), is characterized by its oblong-spathulate to ovate leaves with indistinct or short double costae. Our phylogenetic analysis places three specimens of H. pulchrum, collected from Guizhou and Hainan Provinces, within a distinct clade. This clade is sister to H. montagneanum, H. scalpellifolium, H. flabellatum and H. papillosum. These findings reinforce the recognition of H. pulchrum as a unique species within the genus Homaliodendron. Among the other four species on the phylogenetic tree, H. montagneanum was also distinct and mostly easily identified by its broadly triangular to lanceolate stem leaves, and acute or narrowly obtuse stipe leaf apices. Leaf characters such as the margin, cell structure and costa are also valuable for distinguishing Homaliodendron species (Ninh 1984; Olsson et al. 2010). Homaliodendron papillosum is unique within the genus for its medial cells of branch leaves and stem leaves more or less distinctly mammillose, a feature that clearly differentiates it from other species(Wu 2011; Enroth & Koponen 2017). Homaliodendron maolanense sp. nov. and H. pulchrum exhibit morphological similarities, notably in their leaf apices, which are characterized by multicellular dentate structures, as well as in their thick-walled laminal cells and distinctly porose basal laminal cells. However, the leaves of H. maolanense sp. nov. are strongly falcate with widest part at the middle, distinctly asymmetric, strongly and irregularly biserrate teeth at the apex and costae distinctly double, while the leaves of H. pulchrum are oblong-spatulate to obovate, slightly asymmetric, irregularly dentate at the apex, and costae double, short, sometimes indistinct or absent (Pei et al. 2011). The new species, Homaliodendron maolanense sp. nov., could be identified by the combination of the following features: 1) primary stems prostrate secondary stems erect, bior tri-pinnately branched, with few foliose pseudoparaphyllia; 2) branch and stem leaves strongly falcate with widest part at the middle; 3) costae double, short, distinct; 4) the teeth of upper margins and apex of leaves multicellular, biserrate, coarse and blunt; and 5) basal leaf cells distinctly porose. Our results indicate that the new species, H. maolanense sp. nov., belongs to Homaliodendron stricto sensu. This placement is supported by both molecular data and morphylogical characteristics: notably presence of multicellular teeth at the leaf apex and foliose pseudoparaphyllia. So far, a total of ten species of Homaliodendron have been reported from China. The following key facilitates the identification of H. maolanense sp. nov. among the nine other Chinese species in the genus. Acknowledgements We are grateful to the financial support from the National Nature Science Foundation of China (No. 32100171, 31860162) and the Shanghai Sailing Program (No. 20YF1435500). Thanks are also due to the curators of PE and HSNU for their help in sending specimens on loan, and Prof. Yu Jia, Chinese Academy of Sciences, for his valuable suggestions. We also thank the anonymous referees for their valuable work. REFERENCES allEn b. 2010. — Anomodontaceae-Symphyodontaceae, in allEn b. (ed.), Moss flora of Central America. Part 3. Missouri Botanical Garden press, St. Louis, 731 p. buck W. r. 1998. — Pleurocarpous Mosses of the West Indies. Memoirs of the New York Botanical Garden, 82: 1-400. c rosby m. r., m agill r. E., a llEn b. & h E s. 1999. — A checklist of the mosses. Missouri Botanical Garden Press, St. Louis, 325 p. DrapEr i. & hEDEnäs l. 2009. — Circumscription of European taxa within the Sciuro-hypnum reflexum complex (Brachytheciaceae, Bryophyta), based on molecular and morphological data. Taxon 58: 572-584. https://doi.org/10.1002/tax.582021 Enroth J. 1989 — Bryophyte Flora of the Huon Peninsula, Papua New Guinea. XXVII. Neckeraceae (Musci). Acta Botanica Gennica 137: 41-80. Enroth J. & koponEn t. J. 2017. — Bryophyte flora of Hunan Province, China. 24. Neckeraceae and Miyabeaceae (Musci). Acta Bryolichenologica Asiatica 7: 147-176. flEischEr m. 1906. — Neue Familien, Gattungen und Arten der Laubmoose. Hedwigia 45: 65-87. florschütz-DE WaarD J. & florschütz p. a. 1979. — Estudios sobre criptógamas Colombianas III. Lista comentada de los musgos de Colombia. Bryologist 82: 215-259. forzza r. c. 2025. — Lista de espécies Flora do Brasil. Jardim Botânico do Rio de Janeiro, Rio de Janeiro. https://floradobrasil. jbrj.gov.br/reflora/herbarioVirtual/ [Accessed on August 28, 2025]. g angulEE h. c. 1976. — Mosses of Eastern India and Adjacent Regions, Part 5. Privately published, Calcutta: 1135-1462. hEDEnäs l. 2012. — Molecular differentiation within European Cratoneuron filicinum, and differences from Asiatic and American table 2 . — Numbers of sites in each locus based on sequence length (bp), parsimony variable (p.v.) sites, parsimony informative (p.i.) sites, and the models selected for Bayesian analysis. Gene Length (bp) p.v. sites p.i. sites Model trnL-F 497 137 34 HKY+I trnG632 345 148 GTR+G its2 572 274 86 GTR+G Total 1701 756 268 —