Unveiling the identity of Viburnum laterale (Viburnaceae), a mysterious species disappeared nearly 100 years, based on morphological and molecular evidence

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Unveiling the identity of Viburnum laterale (Viburnaceae), a mysterious species disappeared nearly 100 years, based on morphological and molecular evidence | 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 Search Search ADVANCED SEARCH SCROLL Ecology and Evolution This is a preprint and has not been peer reviewed.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 Unveiling the identity of Viburnum laterale (Viburnaceae), a mysterious species disappeared nearly 100 years, based on morphological and molecular evidence Authors : Liao-cheng Zhao 0000-0002-6300-7301 , Wen-jun Lyu , Hong-tao Liu , and Ming Tang 0000-0001-8419-0996 [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.176868029.91745002/v1 236 views 97 downloads Contents Abstract 2.3 Phylogenetic Analysis Supplementary Material Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Viburnum laterale Rehder is a poorly known shrub endemic to southeastern China that has remained taxonomically enigmatic since its publication more than a century ago. The species was known only from a few historical collections that lacking critical characters, and no living populations had been documented for nearly a century. Here, we report the rediscovery of this taxon and clarify its taxonomic identity using integrative approach combining morphological and molecular evidence. Observations of living plants reveal several previously undocumented reproductive traits, including inflorescences with conspicuous white sterile marginal flowers, as well as fruit and seed morphology. Comparative analyses of living material, type specimens, and protologues indicate that V. laterale is most similar to V. hanceanum Maximowicz, differing mainly in its nearly glabrous indumentum, longer peduncles, and more coarsely serrate leaf margins. Phylogenetic analyses based on chloroplast coding sequences, together with an expanded dataset comprising nuclear ITS and three plastid markers (rbcL, matK, and ndhF), consistently place V. laterale within sect. Tomentosa (Maximowicz) Nakai, where it forms a strongly supported clade with V. hanceanum. Based on morphological and molecular evidence, V. laterale is transferred to sect. Tomentosa and treated as a variety of V. hanceanum, namely V. hanceanum var. depilatum M. Tang & L.C. Zhao. Our study resolves a long-standing taxonomic problem and highlights the importance of rediscovering historically enigmatic taxa for accurate species delimitation in Viburnum. Unveiling the identity of Viburnum laterale (Viburnaceae), a mysterious species disappeared nearly 100 years, based on morphological and molecular evidence Liao-Cheng Zhao 1, 2# , Wen-jun Lyu 5, 6# , Hong-tao Liu 6-8* , Ming Tang 1, 3, 4* 1 College of Forestry, Jiangxi Agricultural University, Nanchang 330045, Jiangxi, China 2 College of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China 3 Jiangxi Provincial Key Laboratory of Conservation Biology (2023SSY02081), Jiangxi Agricultural University, Nanchang 330045, Jiangxi, China 4 Lushan National Observation and Research Station of Chinese Forest Ecosystem,Jiujiang Jiangxi 332900, China 5 National Germplasm Repository of Viburnum, Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan 430074, Hubei, China 6 CAS Key Laboratory of Aquatic Botany and Watershed Ecology, Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan 430074, Hubei, China. 7 Hubei Key Laboratory of Wetland Evolution & Ecological Restoration, Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan 430074, Hubei, China 8 University of Chinese Academy of Sciences, Beijing 100049, China. # Zhao & Lyu contributed equally to this work. *Corresponding authors: Ming Tang ( [email protected] ), Hong-tao Liu ( [email protected] ) Abstract Viburnum laterale Rehder is a poorly known shrub endemic to southeastern China that has remained taxonomically enigmatic since its publication more than a century ago. The species was known only from a few historical collections that lacking critical characters, and no living populations had been documented for nearly a century. Here, we report the rediscovery of this taxon and clarify its taxonomic identity using integrative approach combining morphological and molecular evidence. Observations of living plants reveal several previously undocumented reproductive traits, including inflorescences with conspicuous white sterile marginal flowers, as well as fruit and seed morphology. Comparative analyses of living material, type specimens, and protologues indicate that V. laterale is most similar to V. hanceanum Maximowicz, differing mainly in its nearly glabrous indumentum, longer peduncles, and more coarsely serrate leaf margins. Phylogenetic analyses based on chloroplast coding sequences, together with an expanded dataset comprising nuclear ITS and three plastid markers ( rbc L, mat K, and ndh F), consistently place V. laterale within sect. Tomentosa (Maximowicz) Nakai, where it forms a strongly supported clade with V. hanceanum . Based on morphological and molecular evidence, V. laterale is transferred to sect. Tomentosa and treated as a variety of V. hanceanum , namely V. hanceanum var. depilatum M. Tang & L.C. Zhao. Our study resolves a long-standing taxonomic problem and highlights the importance of rediscovering historically enigmatic taxa for accurate species delimitation in Viburnum . Keywords China; taxonomy; Viburnaceae; Viburnum laterale ; rediscovery 1 Introduction Viburnum Linnaeus (Linnaeus 1753) comprises approximately 200 species of shrubs and small trees, widely distributed from subtropical regions of the Northern Hemisphere to the Andes and tropical Asia. China represents a major center of diversity for this genus, harboring 73 species, of which 45 are endemics (Yang and Malécot 2011). Originally classified within Caprifoliaceae (Cronquist 1888; Hsu 1988), subsequent phylogenetic studies based on morphological traits and molecular data have demonstrated that Viburnum is more closely related to genera currently assigned to Adoxaceae (Backlund and Bremer 1997). This treatment has since been widely accepted in recent classification systems (Yang and Malécot 2011; APG III 2009; APG IV 2016; Zhao et al. 2025). Nevertheless, Viburnum has often been treated within Adoxaceae in recent classifications, Viburnaceae is adopted here following the nomenclatural treatment of Reveal (2008) and the International Code of Nomenclature (Turland et al. 2018). Despite extensive taxonomic studies on Viburnum (Donoghue et al. 2004; Clement and Donoghue 2011, 2012; Clement et al. 2014; Landis et al. 2021), species delimitation within the genus remains challenging in China, largely due to incomplete morphological data, limited phylogenetic sampling (Zhao et al., 2025). Viburnum laterale was first collected by S.T. Dunn during an expedition to central Fujian Province, China, in 1905 (Dunn 1908), and was formally described by Alfred Rehder based on the specimen S.T. Dunn 2771 (A) ((Rehder 1911; Fig. 1A)). Since its original description, V. laterale has been rarely collected and remains poorly understood. The type material consists solely of immature fruiting specimens, lacking floral character. As a result, its taxonomic status have been unstable, and the species has been variously assigned to sect. Platyphylla and sect. Megalotinus (Maximowicz) Rehder in subsequent treatments (Hsu 1988; Yang and Malécot 2011). Notably, a fruiting specimen collected from Shaowu, Fujian Province, in 1936 ( H.C. Chou 6261 ; Fig. 1C) showed striking morphological similarity to V. hanceanum (Maximowicz 1880; Fig. 1D-F), a member of sect. Tomentosa , differing primarily in its nearly glabrous indumentum. However, the absence of flowering material and living populations has precluded a definitive assessment of their taxonomic relationship. In 2021, during a botanical survey in Jiangshi Nature Reserve, Shaowu City, Fujian Province, China (27°04′44″N, 117°13′37″E), we discovered a living population of Viburnum matching the type locality and vegetative morphology of V. laterale . These shrubs are 1–2 m tall, completely glabrous habit, and inflorescences with a prominent outer ring of white sterile marginal flowers, a feature previously unknown for the species. We collected flowering individuals and introduced them into cultivation at the National Germplasm Repository of Viburnum , Wuhan Botanical Garden, Chinese Academy of Sciences, for further observations of fruit and seed morphology. This rediscovery provides a rare opportunity to reassess the identity and taxonomic status of V. laterale using complete morphological information and molecular data. Specifically, our objectives were to (1) document previously unknown characters of V. laterale , (2) determine its phylogenetic position within Viburnum , and (3) reassess its taxonomic status. 2 Material and Methods 2.1 Morphological Observations Living plants of Viburnum laterale were observed and photographed during the flowering stage in Jiangshi Nature Reserve, Fujian Province, China, and during the fruiting stage from individuals cultivated at the National Germplasm Repository of Viburnum , Wuhan Botanical Garden, Chinese Academy of Sciences. Detailed observations focused on inflorescence architecture, the presence and morphology of sterile marginal flowers, and fruit characters. All major morphological features of V. laterale and V. hanceanum were documented on living plants using a digital camera (Olympus TG-6, Tokyo, Japan). 2.2 Taxon Sampling, DNA Extraction, and Data Collection Taxon sampling followed previous phylogenetic studies of Viburnum (Donoghue et al. 2004; Clement and Donoghue 2011, 2012; Clement et al. 2014; Landis et al. 2021; Zhao et al. 2025) and the taxonomic framework proposed by Clement and Donoghue (2011). For analyses based on chloroplast coding sequences (CDS), 29 accessions representing 26 species and one variety from eight sections of Viburnum were sampled. To further ensure phylogenetic robustness, an expanded dataset was assembled using four commonly employed markers-three plastid markers ( mat K, ndh F, rbc L) and the nuclear ribosomal internal transcribed spacer (nrITS), including 62 accessions representing 57 species and three varieties across 10 sections within this genus . Sambucus canadensis Linnaeus, Adoxa moschatellina Linnaeus, Tetradoxa omeiensis (Hara) C.Y. Wu, and Sinadoxa corydalifolia C.Y. Wu, Z.L. Wu & R.F. Huang were selected as outgroups. Newly generated sequences were obtained from three accessions of V. laterale , whereas all remaining sequences were downloaded from GenBank (https://www.ncbi.nlm.nih.gov/). Voucher information and GenBank accession numbers for all samples are provided in Tables S1 and S2. Total genomic DNA was extracted from silica gel–dried leaf material of V. laterale using a modified CTAB protocol (Doyle and Doyle 1987). DNA integrity was assessed by 1% (w/v) agarose gel electrophoresis, and DNA quality and concentration were measured using a NanoDrop 2,000 spectrophotometer (Thermo Scientific, Waltham, MA, USA). Library preparation and sequencing were performed by Novogene Co., Ltd. (Beijing, China). Paired-end libraries (2 × 150 bp) were constructed following the Nova-PE150 strategy, generating 2 Gb of raw sequencing data per sample. Raw reads were filtered using Trimmomatic v. 0.39 (Bolger et al. 2014) to remove low-quality bases and unpaired reads. Filtered reads were assembled with GetOrganelle v. 1.7.7 (Jin et al. 2020), yielding complete chloroplast (cp) genomes and nuclear ribosomal DNA (nrDNA) sequences. Assembly graphs were visualized and manually inspected using Bandage v. 0.8.1 (Wick et al. 2015). Chloroplast genome annotation was conducted in CPGAVAS2 (Shi et al. 2019) using V. japonicum (Thunberg) C.K. Sprengel (GenBank accession OP644292) as a reference. Plastid markers ( mat K, ndh F, rbc L) were extracted from annotated cp genomes using Geneious Prime 2020 (Kearse et al. 2021), and nrITS sequences were extracted from nrDNA assemblies with ITSx (Bengtsson et al. 2013). 2.3 Phylogenetic Analysis Sequence alignments were generated using MAFFT v. 7.450 (Katoh and Standley 2013) with default parameters. Maximum likelihood (ML) analyses were conducted in IQ-TREE (Nguyen et al. 2015), with nodal support assessed using 100,000 ultrafast bootstrap replicates. The best-fit nucleotide substitution models were selected under the Bayesian Information Criterion (BIC) using ModelFinder (Kalyaanamoorthy et al. 2017). Bootstrap support values (MLBS) ≥ 70% were considered strongly supported (Huelsenbeck and Hillis 1993). Resulting phylogenetic trees were visualized and edited in FigTree v. 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/). 3 Results 3.1 Morphological Study Observations of living plants reveal several reproductive features of Viburnum laterale that were previously undocumented (Fig. 2). The inflorescences are compound corymbs with a conspicuous outer side of white sterile marginal flowers, surrounding numerous small yellow fertile flowers. Each fertile flower bears five stamens, a short style, and a capitate stigma. Fruits are ovoid drupes that change form green to red and turn black at maturity, and seeds show two longitudinal grooves on the dorsal surface and a single groove on the ventral surface. Comparisons based on specimens and living plants (Figs. 1-3) indicate that V. laterale and V. hanceanum share a highly similar inflorescence architecture, including the presence of white sterile marginal flowers, typically five primary rays, and fertile flowers borne on the third-order branches. The two taxa differ mainly in vegetative characters. V. laterale is nearly glabrous throughout, with longer peduncles and more coarsely serrate leaf margins than those observed in V. hanceanum . 3.2 Phylogenetic Study Maximum likelihood analysis based on chloroplast coding sequences (CDS) (best-fit BIC model = TVM+F+I+R3) strongly supports the monophyly of Viburnum and its eight major clades (MLBS = 100; Fig. 4). Within this framework, the three accessions of V. laterale form a well-supported clade with V. hanceanum (MLBS = 100), together constituting sect. Tomentosa . Analysis based on an expanded dataset combining ndh F, ma tK, rbc L, and nrITS (best-fit BIC model = TPM3u+F+R2) yield a congruent topology (Fig. 5). Viburnum laterale neither clusters with V. amplifolium Rehder, despite its placed in sect. Platyphylla by P.S. Hsu (1988), nor associates with the four sections segregated from sect. Megalotinus by Clement and Donoghue (2011): sect. Coriacea Kern, sect. Sambucina Kern, sect. Lutescentia Kern, and sect. Punctata Kern. Instead, V. laterale consistently forms a strongly supported clade with V. hanceanum (MLBS = 100; Fig. 5). 3.3 Taxonomic Treatment Viburnum hanceanum var. depilatum M. Tang & L.C. Zhao, var. nov. (Figs. 1 A-C, 2) Type. CHINA. Fujian Province. Without precise locality, collection date unknown, S.T. Dunn 2771 (holotype:A00031569; isotype: IBSC0006026). (Fig. 1A, B) Diagnosis. Viburnum hanceanum var. depilatum is most closely allied to V. hanceanum in inflorescence architecture, particularly in having corymbose inflorescences with conspicuous white sterile marginal flowers, but differs by its nearly glabrous habit (vs. branchlets, petioles, peduncles, and leaves densely stellate-pubescent), longer peduncles (6–9 cm vs. 2–5 cm), and more coarsely serrate leaf margins. Description. Shrubs, 1–2 m tall. Branchlets slender, terete, green to grayish brown, glabrous or very sparsely pubescent when young. Winter buds naked. Leaves opposite, petioles 1–3 cm long, glabrous; leaf blades ovate to elliptic, 6–12 cm long, 3–6 cm broad, papery to thinly coriaceous, both surfaces glabrous, lateral veins 6–8 pairs, impressed adaxially, prominent abaxially, base broadly cuneate to rounded, margin coarsely serrate, apex acuminate. Compound corymbs terminal, 5–9 cm in diameter, usually with five primary rays; peduncles slender, 6–9 cm long, glabrous. Marginal flowers sterile: calyx green, glabrous; corolla conspicuously enlarged, white, irregularly 4- or 5-lobed, lobes broadly ovate, corolla 2-3 cm in diam.; stamens and pistils not developed. Inner flowers fertile, bisexual; calyx like fertile flowers; corolla white, rotate, ca. 3 mm in diam., glabrous. Stamens 5, style short, stigma capitate. Fruits, drupaceous, ovoid, 6–8 mm long, ca. 4 mm broad, initially red, turning black at maturity, endocarp compressed. Seeds with two longitudinal grooves on the dorsal surface and one groove on the ventral surface. Phenology. Flowering from April to May; fruiting from July to September. Distribution and habitat. Endemic to southeastern China, currently known only from the type locality in Fujian Province. Grows in valleys, streams and bushes at elevations of ca. 200–800 m. Etymology. The varietal epithet depilatum refers to the nearly glabrous indumentum of this taxon, in contrast to the densely pubescent typical variety of Viburnum hanceanum . Hence we propose the Chinese name ‘无毛蝶花荚蒾 (wu mao die hua jia mi)’. IUCN Red List Category :— Viburnum hanceanum var. depilatum seems currently known only from its type locality. Only one small population of this species, with less than 50 individuals, has been discovered there. The habitat of this species is now well preserved. Further populations of this species is expected to be discovered as botanical exploration of northwestern Fujian proceeds. According to the IUCN Red List Categories and Criteria (IUCN 2022), the new taxon is best categorized as Data Deficient (DD). Additional specimens examined: CHINA. Fujian Province, Shaowu City, Jiangshi Nature Reserve, 11 May 2024, L. C. Zhao 56 (JXAU, paratype), L. C. Zhao 57 (JXAU, paratype), L. C. Zhao 58 (JXAU, paratype). Notes. Morphological and molecular evidence support the treatment of Viburnum laterale as an infraspecific taxon of V. hanceanum. The diagnostic characters are stable in both wild and cultivated populations and correspond to minor vegetative differentiation rather than species-level divergence. 4 Discussion 4.1 Resolving the Identity of a Long-overlooked Taxon The rediscovery of Viburnum laterale after nearly a century allows a reassessment of a taxon whose identity has long been obscured by incomplete original material. The type specimen lacks flowers, which precluded the recognition of key reproductive traits and contributed to inconsistent sectional placements in earlier treatments (Hsu 1988; Yang and Malécot 2011). Our observations of living plants reveal that V. laterale possesses inflorescences with conspicuous white sterile marginal flowers, together with taxonomic fruit and seed features that were previously unavailable for evaluation. Sterile marginal flowers are relatively uncommon in Chinese Viburnum and occur in only a limited number of taxa (e.g. V. keteleeri Carrière, V. macrocephalum Fortune, V. opulus Linnaeus, V. plicatum, V. hanceanum , V. sympodiale Graebner, V. opulus var. sargentii Koehne, V. hanceanum var. Depilatum , V. plicatum var. formosanum Y.C. Liu & C.H. Ou and V. plicatum f. tomentosum (Miq.) Rehder) among four sections: sect. Viburnum , sect. Pseudotinus C.B. Clarke, sect. Opulus (Miller) Candolle, and sect. Tomentosa . The scattered phylogenetic distribution of sterile marginal flowers suggests that this character has evolved independently multiple times within Viburnum (Park and Donoghue 2021). Although of limited value for diagnosing sections, this character remains taxonomically informative at the species and infraspecific levels when evaluated together with vegetative morphology and molecular evidence. 4.2 Phylogenetic Position Both morphological and molecular evidence place V. laterale within sect. Tomentosa (MLBS = 100; Figs. 4-5), where it forms a strongly supported clade with V. hanceanum (MLBS = 100). This placement contrasts with earlier assignments to sect. Platyphylla or sect. Megalotinus (Hsu 1988; Yang and Malécot 2011). The molecular results indicate that the absence of complete floral characters in the type specimen (Fig. 1A-C), rather than genuine phylogenetic affinity, likely underlies these earlier interpretations. 4.3 Taxonomic Status Morphologically, V. laterale closely resembles V. hanceanum , particularly in inflorescence architecture and organization of fertile flowers. The difference between the two taxa are mainly in vegetative traits, notably in indumentum, peduncle length, and leaf margin serration. Phylogenetic analyses consistently recover V. laterale as sister to V. hanceanum with strong support (MLBS = 100; Figs. 4-5), and the known distribution of V. laterale lies entirely within the range of V. hanceanum (Hsu 1988; Yang and Malécot 2011). Taken together, these evidence suggest that the observed differences represent infraspecific variation rather than species-level divergence. We therefore treat V. laterale as a variety of V. hanceanum and recognize it as V. hanceanum var. depilatum M. Tang & L.C. Zhao. Author contributions Liao-Cheng Zhao: formal analysis (lead), methodology (lead), software (lead), visualization (lead), conceptualization (supporting), writing–original draft (lead), writing–review and editing (supporting). Wen-jun Lyu: data curation (lead), investigation (lead), resources (supporting), conceptualization (supporting). Hong-tao Liu: data curation (supporting), investigation (supporting), resources (supporting), conceptualization (supporting). Ming Tang: formal analysis (supporting), methodology (supporting), writing–original draft (supporting), writing–review and editing (lead), resources (lead), conceptualization (lead), funding acquisition (lead), project administration (lead), supervision (lead) Acknowledgements We are grateful to the anonymous reviewers for their helpful comments on how to improve our manuscript. We thank Mr. G.J. Yan for his permission of using some images of V. hanceanum, and also thank A, AU, HIB, K, P, GH, NAS, IBK, LBG, JJF, GNNU, CSH, CSFI, SN, IBSC, GZAC, JXAU, KUN and PE for allowing us to examine and use their scanned images of specimens. Figure 1. Viburnum laterale and V. hanceanum . A Fujian, China, S.T. Dunn 2771 (A00031569, holotype of V. laterale ) B Fujian, China, S.T. Dunn 2771 (IBSC0006026, isotype of V. laterale ) C Shaowu city, Fujian, China, H.C. Chou 6261 (IBSC0492224, V. laterale ) D Baiyun Mountain, Guangzhou City, Guangdong, China, G.T. Sampson 10838 (K000797923, holotype of V. hanceanum ) E Baiyun Mountain, Guangzhou City, Guangdong, China, G.T. Sampson 10838 (P00644640, isotype of V. hanceanum ) F Baiyun Mountain, Guangzhou City, Guangdong, China, G.T. Sampson 10838 (GH00031565, isotype of V. hanceanum ). Figure 2. Morphology of Viburnum laterale. A habit B inflorescence C leaves (glabrous on both adaxial ( a ) and abaxial ( b ) sides) D inflorescence axis E white sterile flowers on the outer edge of the inflorescence F fertile flowers on the inner edge of the inflorescence G fruit (the left one) and seed (the right three). Figure 3. Morphology of Viburnum hanceanum . A habit B inflorescence C leaves (yellow-brown fasciculate stellate-pubescent on both adaxial ( a ) and abaxial ( b ) sides) D inflorescence axis E white sterile flowers on the outer edge of the inflorescence F fertile flowers on the inner edge of the inflorescence. Figure 4. The maximum likelihood tree of Viburnum based on chloroplast coding sequence (CDS), with three V. laterale samples highlighted in red font. Bootstrap values (BS) are shown above the branches. Dashes (-) indicate MLBS < 70. Figure 5. The maximum likelihood tree of Viburnum based on nrITS, rbc L, mat K and ndh F, with three V. laterale samples highlighted in red font. Bootstrap values (BS) are shown above the branches. Dashes (-) indicate MLBS < 70. Conflict of Interest The authors declare no conflicts of interests. Ethical statement No ethical statement was reported. 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PLoS One 20(4): e0312920. https://doi.org/10.1371/journal.pone.0312920 Supplementary Material File (table s1.xlsx) Download 12.62 KB File (table s2.xlsx) Download 19.97 KB Information & Authors Information Version history V1 Version 1 17 January 2026 Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Ecology and Evolution Keywords comparative description ecosystem evolutionary ecology plants Authors Affiliations Liao-cheng Zhao 0000-0002-6300-7301 Jiangxi Agricultural University College of Forestry/School of Landscape and Art View all articles by this author Wen-jun Lyu Chinese Academy of Sciences Wuhan Botanical Garden View all articles by this author Hong-tao Liu Chinese Academy of Sciences Wuhan Botanical Garden View all articles by this author Ming Tang 0000-0001-8419-0996 [email protected] Jiangxi Agricultural University College of Forestry/School of Landscape and Art View all articles by this author Metrics & Citations Metrics Article Usage 236 views 97 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Liao-cheng Zhao, Wen-jun Lyu, Hong-tao Liu, et al. Unveiling the identity of Viburnum laterale (Viburnaceae), a mysterious species disappeared nearly 100 years, based on morphological and molecular evidence. Authorea . 17 January 2026. 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