Abstract
This study describes a new species of Sedum (Crassulaceae), Sedum qingyuanensesp. nov. discovered in Qingyuan City, Guangdong Province, China. Morphologically, S. qingyuanense resembles S. alfredii and S. emarginatum. The new species may have previously been misidentified as S. alfredii. However, the new species can be readily distinguished from S. alfredii by its creeping sterile stems, opposite leaves, emarginate leaf apices, and smaller sepals. Compared to S. emarginatum, it is differentiated by its creeping sterile stems, smaller sepals, and conspicuous pseudopetioles. Phylogenetic analysis based on the nuclear ribosomal DNA internal transcribed spacer (ITS) region revealed that the new species forms a sister clade to S. emarginatum within Sedum sect. Sedum, supported by moderate to strong bootstrap support (> 80%). Integrating both morphological evidence and the phylogenetic tree, we establish it as an independent lineage.
Key words: New species, nrITS, phylogeny, Sedum
Introduction
The genus Sedum L. (Crassulaceae), comprising approximately 474 accepted species (WFO 2025), represents the most species-rich genus within the family. It has a broad distribution across temperate and subtropical regions of the Northern Hemisphere, with biodiversity hotspots concentrated in the Mediterranean Basin, Central America, the Himalayas, and East Asia (Stephenson 1994; Thiede and Eggli 2007). A few species extend into the Southern Hemisphere, particularly in Africa and South America. Sedum species typically exhibit herbaceous to subshrubby growth forms with succulent leaves and stems, an adaptation to xeric habitats such as deserts, cliffs, rock crevices, and sandy slopes (Thiede and Eggli 2007). Certain taxa, including Sedum lineare Thunb. and S. sarmentosum Bunge, are widely used in urban green roofing due to their compact growth habit, shallow root systems, stress tolerance, low maintenance requirements, and ornamental value (Schindler et al. 2019).
East Asia constitutes one of the major centers of Sedum diversity (Thiede and Eggli 2007; Ito et al. 2017a). According to the Flora of China (Fu and Ohba 2001), 121 Sedum species occur in China, of which 91 are endemic. These species are classified into three sections: sect. Oreades Fröd., sect. Sedum, and sect. Filipes Fröd., with the majority primarily distributed in southwestern China. Among them, Sect. Sedum represents the most species-rich group (Zou et al. 2025). It is characterized by convex ventral surfaces of carpels and follicles.
From 2005 to 2024, 18 new species of the genus Sedum have been described from China (Dai et al. 2025). Between 2024 and October 2025, an additional five species—S. orientalichinense Q. Fan & P. Li (Dai et al. 2025), S. guangxiense Yan Liu & C.Y.Zou (Zou et al. 2025), S. simingshanense Y.L.Xu (She et al. 2025a), S. yongkangense Y.L.Xu et Z.H.Chen (She et al. 2025b), and S. shunhuangense L.Wu & Z.L.Feng (Feng et al. 2025) —were published. As of 2025, the genus Sedum comprises 144 species in China, including 114 endemic species. During field surveys in Guangdong Province, southern China, this species was discovered growing on rocky slopes in Qingyuan City. The species morphologically resembles S. alfredii Hance and S. emarginatum Migo but is distinguished by its smaller sepals, creeping sterile stems, and conspicuous pseudopetioles. Phylogenetic analysis based on nrITS sequencing supports its status as an independent lineage. Morphological comparisons and molecular evidence confirm that S. qingyuanense represents a previously undescribed species within Sedum sect. Sedum. This paper provides a comprehensive morphological description, taxonomic discussion, and ecological illustrations of the new species.
Methods
Morphological study
To obtain fresh material for morphological analysis, detailed field investigations were conducted during the flowering period of the new species and its close relatives: S. emarginatum (Hangzhou), S. alfredii (Hangzhou and Guangzhou), S. jinglanii (ShaoGuan). Collected 4 specimens were thoroughly examined, dissected, and photographed. Living specimens of this new species were collected from TaiHe Grotto, Qingyuan City, Guangdong Province, China.
Digital specimens of related species were examined via the Chinese Virtual Herbarium (https://www.cvh.ac.cn/); and the Global Biodiversity Information Facility (https://www.gbif.org/) to obtain comparative morphological data. The holotype was collected from TaiHe Grotto, Qingyuan City, and is deposited in the Herbarium of Sun Yat-sen University (SYS!).
Molecular study
Fresh leaves from two individuals of S. qingyuanense were collected at TaiHe Grotto, Qingyuan City, and dried in silica gel. Total DNA was extracted using a modified CTAB method (Doyle and Doyle 1987). The partial internal transcribed spacer 1 (ITS1), the 5.8S ribosomal RNA gene, and the partial internal transcribed spacer 2 (ITS2) region were amplified using the primers ITS1 and ITS4 (White et al. 1990), following PCR protocols from Huang et al. (2021).
We downloaded 74 ITS sequences from GenBank, representing 53 Sedum species (including subspecies and variants) and 3 outgroups (Table 1). Greenovia aizoon, Aeonium viscatum and A. lancerottense were selected as outgroups (Huang et al. 2023). Ultimately, a total of 79 sequences were used to construct the phylogenetic tree, including the sequence of S. qingyuanense and our newly sequenced S. alfredii and S. emarginatum sequences.
Table 1.
Taxa, voucher information, GenBank accession numbers and references for ITS sequences of Sedum (S.) species and three outgroups used for phylogenetic analyses in this study.
| Taxon | Voucher | Accession number | Reference |
|---|---|---|---|
| S. actinocarpum | Ito 1749 | LC229265 | Ito et al. 2017a |
| S. alfredii | IBK194564 | FJ919949 | Wang and Shu unpublished |
| IBK114924 | FJ919951 | Wang and Shu unpublished | |
| – | PX560094 | Present Study | |
| S. arisanense | Ito 1842 | LC229273 | Ito et al. 2017a |
| Ito 1836 | LC229272 | Ito et al. 2017a | |
| S. baileyi | LBG0064555 | FJ919935 | Wang and Shu unpublished |
| S. bergeri | – | AY352897 | Ni et al. unpublished |
| S. boninense | Ito 2371 | LC229242 | Ito et al. 2017a |
| S. bulbiferum | Ito 416 | LC229234 | Ito et al. 2017a |
| 130514hs41 | KM111166 | Xie et al. 2014 | |
| 130524qz09 | KM111165 | Xie et al. 2014 | |
| S. brachyrinchum var. brachyrinchum | Ito 1359 | LC229274 | Ito et al. 2017a |
| S. danjoense | Ito 3658 | LC260127 | Ito et al. 2017b |
| S. emarginatum | 130512hs27 | KM111145 | Xie et al. 2014 |
| 130529hz03 | KM111146 | Xie et al. 2014 | |
| 130503jz21 | KM111147 | Xie et al. 2014 | |
| 24041001 | PP981038 | Dai et al. 2025 | |
| 24052201 | PP981037 | Dai et al. 2025 | |
| AYJT-Nanjing-0308-08 | EU592006 | – | |
| – | PX560095 | Present Study | |
| – | PX560096 | Present Study | |
| S. erici-magnusii | Ito 2077 | LC229235 | Ito et al. 2017a |
| S. erythrospermum | Tsutsumi 1504 | AB906473 | Ito et al. 2014b |
| S. formosanum | Ito1115 | LC530813 | Ito et al. 2020 |
| Ito2296 | LC260124 | Ito et al. 2017b | |
| S. hakonense | Mayuzumi C00005 | AB088625 | Mayuzumi and Ohba 2004 |
| S. hangzhouense | Ito 2604 | LC260130 | Ito et al. 2017b |
| S. japonicum | Kokubugata 16749 | AB906475 | Ito et al. 2014b |
| S. japonicum var. oryzifolium | Ito 2285 | LC229239 | Ito et al. 2017a |
| S. japonicum var. pumilum | Ito 2287 | LC229240 | Ito et al. 2017a |
| S. japonicum var. senanense | Ito 2200 | LC229238 | Ito et al. 2017a |
| S. jinglanii | Y. S. Huang 21040301 | OP288035 | Huang et al. 2023 |
| DNPC 2873 | OQ162326 | Huang et al. 2023 | |
| S. jiulungshanense | Ito 76 | LC229243 | Ito et al. 2017a |
| S. kiangnanense | CMQ1030 | LC229244 | Ito et al. 2017a |
| S. lineare | Mayuzumi C00120 | AB088623 | Mayuzumi and Ohba 2004 |
| S. lipingense | ZRB1479 | MN150061 | Zhang et al. 2019 |
| S. lungtsuanense | Ito 3563 | LC260131 | Ito et al. 2017b |
| S. makinoi | Kokubugata 16730 | AB906476 | Ito et al. 2014b |
| S. mexicanum | Ito 647 | LC229247 | Ito et al. 2017a |
| S. microsepalum | Ito 2771 | LC229282 | Ito et al. 2017a |
| Ito 1965 | LC229281 | Ito et al. 2017a | |
| S. morrisonense | Ito 2765 | LC229290 | Ito et al. 2017a |
| Hornat S1 | LM993281 | Nikulin et al. 2016 | |
| S. multicaule | Miyamoto et al. TI9596136 | AB088631 | Mayuzumi and Ohba 2004 |
| S. nagasakianum | Ito 2064 | LC229249 | Ito et al. 2017a |
| S. nokoense | Kokubugata 10426 | AB906478 | Ito et al. 2014b |
| S. oligospermum | Ito 74 | LC229250 | Ito et al. 2017a |
| S. oreades | Rao 090803-03 | KF113733 | Zhang et al. 2014 |
| S. polytrichoides subsp. polytrichoides | CMQ1057 | LC229251 | Ito et al. 2017a |
| S. polytrichoides var. setouchiense | Ito 2298 | LC229253 | Ito et al. 2017a |
| S. qingyuanense | TH1001 | PX560092 | Present Study |
| TH1002 | PX560093 | Present Study | |
| S. rupifragum | Ito2070 | LC229254 | Ito et al. 2017a |
| S. sarmentosum | Ito 978 | LC229255 | Ito et al. 2017a |
| S. satumense | Ito 2295 | LC229256 | Ito et al. 2017a |
| S. sekiteiense | Ito 1456 | LC229295 | Ito et al. 2017a |
| S. simingshanense | - | PP464049 | She et al. 2025 |
| - | PP464048 | She et al. 2025 | |
| S. subtile | Ito 624 | AB930277 | Ito et al. 2014a |
| Shimizu 1999 | AB088622 | Mayuzumi and Ohba 2004 | |
| S. taiwanianum | Ito 2770 | LC229297 | Ito et al. 2017a |
| Ito 2770 | LC229296 | Ito et al. 2017a | |
| S. tetractinum | Ito 3623 | LC260135 | Ito et al. 2017b |
| S. tianmushanense | Ito 355 | LC229261 | Ito et al. 2017a |
| S. tosaense | Kokubugata 16726 | AB906483 | Ito et al. 2014b |
| S. triactina | 9596091 | AB088629 | Mayuzumi and Ohba 2004 |
| S. tricarpum | Ito 2269 | LC229259 | Ito et al. 2017a |
| - | PP989617 | - | |
| S. trullipetalum | Miyamoto et al. 9420132 | AB088630 | Mayuzumi and Ohba 2004 |
| S. truncatistigmum | Ito 3254 | LC229306 | Ito et al. 2017a |
| S. xunvense | - | PV101948 | Chai et al. 2024 |
| - | PV101947 | Chai et al. 2024 | |
| S. yabeanum | Ito 396 | AB906490 | Ito et al. 2014b |
| S. zentaro-tashiroi | Ohba 1998 | AB088619 | Mayuzumi and Ohba 2004 |
| Outgroups | |||
| Aeonium lancerottense | Mort 1518 | AY082143 | Mort et al. 2002 |
| Aeonium viscatum | Mort 1432 | AY082154 | Mort et al. 2002 |
| Greenovia aizoon | Mort 1425 | AY082112 | Mort et al. 2002 |
Sequence alignment was performed using MAFFT v7.505 (Katoh and Standley 2013) with the ‘--auto’ strategy. The initial alignment was manually inspected and refined in BioEdit v7.2.5 to correct misalignments in hypervariable regions. Maximum Likelihood (ML) phylogenetic inference was conducted using IQ-TREE 2.1.4 (Minh et al. 2020). The best-fit nucleotide substitution model was determined through the built-in ModelFinder utility, which selected the SYM+I+G4 model under the Bayesian Information Criterion (BIC). Branch support was assessed through standard non-parametric bootstrap analysis with 3,000 replicates. Bayesian inference (BI) analysis was conducted with MrBayes v.3.2.1 (nruns = 2, nchains = 4, mcmc = 30,000,000). The consensus tree was visualized and annotated in FigTree v1.4.4, with branches scaled by the expected number of substitutions per site and bootstrap support values (BS) displayed at the nodes.
Results and discussion
The final length of the aligned ITS sequences was 659 bp. Using these ITS data, we successfully constructed a Maximum Likelihood (ML) tree (Fig. 1). In the ML tree, Sedum qingyuanense was clearly distinguishable from its morphologically similar relatives, S. alfredii and S. emarginatum. It formed a distinct clade, positioned as a sister group to S. emarginatum, while S. alfredii was located on a separate branch of the tree. Notably, although S. qingyuanense appeared phylogenetically close to S. jinglanii in the ML tree, it can be unequivocally distinguished from this species morphologically. Moreover, their habitats are markedly distinct: S. jinglanii is endemic to the Danxia landscape of Danxiashan Mountain in Shaoguan City, whereas S. qingyuanense is restricted to the granite cliffs in TaiHe Grotto of Qingyuan City.
Figure 1.
Maximum Likelihood phylogenetic tree constructed based on ITS sequences of S. qingyuanense and closely related species with Bayesian posterior probabilities mapped onto the branches. Node support values are given as bootstrap support value (BS) and Bayesian posterior probabilities (PP). S. qingyuanense is highlighted in yellow and bolded. The newly sequenced S. alfredii and S. emarginatum from this study are marked in blue.
This new species closely resembles S. alfredii morphologically and may have previously been misidentified as such. However, it can be differentiated by its smaller sepals, creeping sterile stems, opposite leaves, emarginate leaf apices, conspicuously pseudopetiolate, and consistently spatulate leaf shape. Because of its emarginate leaves and opposite phyllotaxy, S. qingyuanense may also be confused with S. emarginatum. However, the two species are readily distinguishable by differences in sepal size, leaf shape and size, presence of creeping sterile stems, and conspicuous pseudopetiolate. Morphological analyses of the putative new species and its relatives reveal that, despite some shared characteristics, they can be clearly distinguished by several diagnostic traits.
Based on our measurements and previous studies (Fu and Ohba 2001; Wu et al. 2012; Huang et al. 2023; Dai et al. 2025), we summarize and detail the morphological distinctions among these three species in Table 2.
Table 2.
Morphological comparisons of S. qingyuanense, S. alfredii, S. emarginatum, S. jinglanii.
| Character | Species | |||
|---|---|---|---|---|
| S. qingyuanense | S. alfredii | S. emarginatum | S. jinglanii | |
| Fertile stems | 4–15 cm | 10–20 cm | 10–27 cm | 10–15 cm |
| Phyllotaxy | opposite, sometimes alternate |
alternate | Opposite | opposite |
| Leaf apex | emarginate | obtuse sometimes emarginate | Emarginate | obtuse sometimes emarginate |
| Leaf blade | spatulate to broadly obovate | linear-cuneate, spatulate, or obovate | spatulate-obovate to broadly obovate | spatulate or obovate |
| Leaf size | 1.1–5.9 × 0.5–1.4 cm | 1.2–3 × 0.2–0.6 cm | 1–2 × 0.5–1 cm | 0.8–2.9 × 0.4–1.2 cm |
| Pseudopetiolate | conspicuously 8–24 mm | inconspicuously | Inconspicuously | inconspicuously |
| Sterile stem | creeping | ascending | absent | ascending |
| Sepal shape | linear-spatulate | linear-spatulate | lanceolate to narrowly oblong | linear-spatulate |
| Sepal size | 1.8–3.2 × 0.5–1.5 mm | 3–5 × 1–1.5 mm | 2–5 × 0.7–2 mm | 2–3.1 × 0.7–1.4 mm |
| Flowering | March–April | April–May | May–June | April–May |
Taxonomic treatment
. Sedum qingyuanense
X.W.Yi, Z.Y.Xie & Q.Fan sp. nov.
CAD17E2F-C962-523D-AF55-E9EA300D0A35
urn:lsid:ipni.org:names:77373284-1
Figure 2.
Sedum qingyuanense. A. Flower; B. Petals and epipetalous stamens; C. Episepalous stamens; D. Back view of flower (with sepals); E. Sepals; F. Frontal and lateral aspects of carpels; G. Unripe follicles; H. Opened follicles; I. Unripe seeds; J. Flower compared with S. alfredii (left: S. alfredii; right: S. qingyuanense); K. Sepals compared with S. alfredii (upper: S. alfredii; lower: S. qingyuanense). Photos: XiaoWei Yi.
Figure 3.
Sedum qingyuanense. A–B. Habitat; C. Leaves of a single S. qingyuanense; D. Immature cyme; E–G. Mature cyme. Photos: XiaoWei Yi & Ke Tang.
Figure 4.
Sedum qingyuanense. A–B. Flowers; C. Follicles; D. Petal with stamen; E. Seed; F. Sepals; G. Leave; H. Bract; I. Basal leaf; J. Fertile stems; K. Sterile stems. Drawing by RongEn Wu.
Figure 5.
Holotype of Sedum qingyuanense.
Chinese name.
清远景天 (Qīng yuăn JĬng tiān).
Type.
China • Guangdong Province, Qingyuan City, TaiHe Grotto; on rocky cliff; 23.7553°N, 112.9807°E; 238 m a.s.l.; 18 April 2025; X.W.Yi & K.Tang QY-1001 (holotype: SYS00237022).
Diagnosis.
The new species is distinguished from its congeners by the combination of creeping sterile stems, conspicuous pseudopetiolate, and small linear-spatulate sepals (1.8–3.2 × 0.5–1.5 mm). It differs from close species, S. jinglanii, S. alfredii and S. emarginatum in having creeping sterile stems (vs. ascending sterile stems), smaller sepals and more distinctly pseudopetiolate. In addition, we have also provided photographs of S. emarginatum and S. alfredii (Fig. 6).
Figure 6.
A–D.Sedum emarginatum from its type locality (LingYin Temple, HangZhou, ZheJiang, China); E, F.S. alfredii. Photos: YueLiang Xu.
Description.
Perennial herb, entirely glabrous. Stems succulent; fertile stems suberect 5–20 cm long, apex erect, usually 3-branched above; sterile stems present, creeping, rooting at nodes and producing new rosettes. Leaves opposite, conspicuously pseudopetiolate; leaf blades spatulate to broadly obovate, margin entire, apex emarginate, base attenuate with a spur, 1.1–5.9 × 0.5–1.4 cm. Cymes 3–8 cm in diameter, usually 3-branched, multiflowered; bracts obovate with an obtuse apex. Flowers sessile, 7–10 mm long, pentamerous, actinomorphic. Sepals 5, linear-spatulate, 1.8–3.2 × 0.5–1.5 mm, base spurred. Petals 5, yellow, lanceolate to lanceolate-oblong, 4–6 × 1.1–1.7 mm, apex acuminate, base connate ca. 0.3 mm. Stamens 10 antepetalous stamens ca. 3 mm long, adnate to petals for ca. 0.3 mm; antesepalous stamens ca. 4.5 mm long. Carpels 5, lanceolate, erect, connate at base, 3–4 mm long. Follicles obliquely divergent, many-seeded; placentation marginal. Seeds ovoid, brown at maturity, 0.6–0.8 mm long.
Phenology.
Flowering from March–April; fruiting in May.
Etymology.
The specific epithet refers to the distribution of this species in Qingyuan City.
Distribution and habitat.
The new species is endemic to Qingyuan City, Guangdong Province, southern China, growing on rocky cliffs at 200–300 m a.s.l.
Conservation status.
The Extent of Occurrence (EOO) and Area of Occupancy (AOO) were calculated following the guidelines of the International Union for Conservation of Nature (IUCN 2024). All known occurrences of Sedum qingyuanense are currently restricted to a single locality—TaiHe Grotto, Qingxin District, Qingyuan City, Guangdong Province, China. The EOO was estimated using the minimum convex polygon method based on this single site, resulting in an EOO < 2 km2. The AOO was assessed using a standard 2 km × 2 km grid system, and as all individuals fall within one grid cell, the AOO is inferred to be ≤ 4 km2. Given that the entire population occurs in a heavily visited tourist area along roadsides and is subject to ongoing anthropogenic threats (e.g., trampling, habitat disturbance), and considering its extremely limited distribution, Sedum qingyuanense meets the criteria for Critically Endangered (CR) under B1ab(iii)+2ab(iii).
Supplementary Material
Acknowledgements
We are deeply grateful to the Qingyuan Forestry Bureau for their invaluable support throughout this research and extend our thanks to the Biological Museum of Sun Yat-sen University for their expert assistance in specimen preparation and conservation. Special acknowledgment goes to Yueliang Xu of the Zhejiang Museum of Natural History for providing high-quality samples and photographs of Sedum alfredii. We also extend our sincere thanks to Rong-En Wu for her skillful artwork in preparing the illustrations for this manuscript.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Use of AI
No use of AI was reported.
Funding
This work was supported by the 2024 Guangdong Province Ecological Quality Index (EQI) Monitoring Project(GPCGD241115FG155F)and the Foundation of Qingyuan Forestry Bureau (HT-99982024-0030).
Author contributions
Data curation: XWY. Funding acquisition: QF. Investigation: XGS, QGZ, XWY, WX, ZYX, KT. Project administration: QF. Resources: QF. Supervision: QF. Visualization: XWY, KT. Writing – original draft: XWY. Writing – review and editing: YSH.
Author ORCIDs
Xiao-Wei Yi https://orcid.org/0009-0004-8867-7418
Zhi-Yi Xie https://orcid.org/0009-0004-4443-4502
Xue-Gui Shen https://orcid.org/0000-0002-5619-7784
Wei Xu https://orcid.org/0000-0003-2907-0478
Qiu-Gen Zeng https://orcid.org/0009-0003-2250-4648
Ke Tang https://orcid.org/0009-0009-9108-5145
Qiang Fan https://orcid.org/0000-0003-4254-6936
Data availability
All of the data that support the findings of this study are available in the main text.
Supplementary materials
Sedum qingyuanense paratype
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Xiao-Wei Yi, Zhi-Yi Xie, Xue-Gui Shen, Wei Xu, Qiu-Gen Zeng, Ke Tang, Yan-Shuang Huang, Qiang Fan
Data type
jpg
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Sedum qingyuanense paratype
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Xiao-Wei Yi, Zhi-Yi Xie, Xue-Gui Shen, Wei Xu, Qiu-Gen Zeng, Ke Tang, Yan-Shuang Huang, Qiang Fan
Data type
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Data Availability Statement
All of the data that support the findings of this study are available in the main text.