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Research Article
Sinocrassula obliquifolia (Crassulaceae), a new species from China
expand article infoRong-Juan Li, Jing Zhao, Shao-Li Fang§, Chuan-Jie Huang, Miao Luo, Zhao-Rong He, Xin-Mao Zhou, Jia-Guan Wang
‡ Yunnan University, Kunming, China
§ Kunming Coal Design and Research Institute, Kunming, China
Open Access

Abstract

Based on a comprehensive morphological and molecular data analysis, we have confirmed and described a new species within the genus Sinocrassula, which is distributed in Sichuan Province, China. Morphologically, the new species resembles Sinocrassula diversifolia and S. indica, but it is distinctly different from them in its asymmetrical leaves, leaf apex with glands, triangular petals that are white at the base and adorned with dense purple-red stripes and spots on the surfaces upward, and rectangular nectar scales. A phylogenetic analysis utilizing four plastid markers and one nuclear marker supports the conclusion that the new species is sister to S. ganluoensis.

Key words:

Hengduan mountains, phylogeny, rosette, species diversity, taxonomy

Introduction

Crassulaceae are a morphologically diverse and systematically complex group of angiosperms, comprising 35 genera and approximately 1,400 species (Berger 1930; Eggli 2003; Thiede and Eggli 2007; Messerschmid et al. 2020). Known for their heat resistance and rapid growth, members of the Crassulaceae family are highly adaptable and easy to cultivate. This adaptability makes them excellent for soil improvement and essential components in vertical gardens and green roofs (Zhu 2015). Sinocrassula A. Berger is a small genus within the family Crassulaceae, comprising approximately ten species distributed across various regions, including Bhutan, China, India, Nepal, Pakistan, Sikkim, and Vietnam (Wang et al. 2012, 2022; Fu and Ohba 2013; Averyanov et al. 2014). Despite the distinct characteristics of Sinocrassula, defining monophyletic groups within the genus has proven to be extremely challenging due to the frequent intergradation of morphological traits among taxa, as well as the extensive diversity in morphology and growth habits (Fu and Ohba 2013). Some species of Sinocrassula (e.g., S. indica) have been used as traditional medicine, or enjoyed as a vegetable and infused in herbal teas (Zhao et al. 2004; Xie and Yoshikawa 2012). In recent years, an increasing number of species of Sinocrassula have been discovered in Asia (e.g., S. jiaozishanensis Chao Chen, J. Guan Wang & Z.R. He, S. vietnamensis Aver. & V.V. Byalt) (Averyanov et al. 2014; Wang et al. 2022). The discovery of new species may provide more medicinal resources and increase the species diversity of Sinocrassula.

In July 2023, we conducted two field surveys in Sichuan Province, where we identified two populations of Sinocrassula in Shimian and Hanyuan counties. After carefully culturing the plants and performing a morphological study, we discovered that these populations are distinctly different from all other species within the genus. Additionally, we conducted a phylogenetic analysis, which indicated that these populations form a well-supported clade that is sister to S. ganluoensis (Li et al. 2024). We propose those populations as a new species of Sinocrassula.

Materials and methods

Morphological studies

The living plants of the new species were cultivated in the greenhouse at Yunnan University. Plant morphologies were photographed using a Nikon SMZ1270 stereo microscope (Nikon). Morphological comparisons among the new species and its related species were from field observations, herbarium investigations, and the literature (e.g., Fu and Ohba 2013; Averyanov et al. 2014; Wang et al. 2022). Voucher specimens of the new species have been deposited in the Herbarium of Yunnan University (YUKU; herbarium acronyms follow Index Herbarium by Thiers 2024).

Taxonomic sampling

In order to clarify the phylogenetic position of the new species, two samples of the new species and its putative closely related taxa of Sinocrassula from China were included in the phylogenetic analysis. In total, 22 accessions representing 12 species of Sinocrassula were used for the phylogenetic analysis. Based on previous phylogenetic studies (Messerschmid et al. 2020; Wang et al. 2022), Kungia aliciae (Raym.–Hamet) K. T. Fu was selected as the outgroup.

DNA extraction, amplification, and sequencing

Total genomic DNA was extracted from silica-dried material using the TIANGEN plant genomic DNA extraction kit (TIANGEN Biotech., Beijing, China) following the manufacturers’ protocols. Four chloroplast DNA markers (psbA-trnH, trnL-F, rbcL, matK)] and one nuclear marker (ITS) were amplified and sequenced using previous primers and protocols (Wang et al. 2022).

Phylogenetic analyses

Sequences from GenBank and the newly generated data (Table 1) were aligned using MAFFT v7.450 (Katoh and Standley 2013), followed by manual refinement in BioEdit (Hall 1999). Single alignments were concatenated to a matrix using the ‘Concatenate Sequence’ plugin in PhyloSuite v1.2.2 (Zhang et al. 2020). ModelFinder (Kalyaanamoorthy et al. 2017) was used to select the best-fitting likelihood model for Maximum Likelihood (ML) and Bayesian Inference (BI) using the bias-corrected Akaike information criterion (AICc). Maximum likelihood bootstrapping was performed with 5,000 rapid bootstrap (BS) analyses followed by a search for the best-scoring tree in a single run through IQ-tree v2.1.2 (Nguyen et al. 2015). Bayesian inference was conducted for the combined dataset using MrBayes v3.1.2 (Huelsenbeck and Ronquist 2001) with two runs of four Markov chain Monte Carlo (MCMC) chains, each beginning with a random tree and sampling every 1000 generations for 20,000,000 generations. Finally, the concatenated trees were generated and visualized with their Maximum-Likelihood Bootstrap Support values (ML-BS) and Bayesian Inference Posterior Probability (BI-PP) in Figtree v1.4.3 (Rambaut 2017).

Table 1.

Species information and corresponding GenBank accession numbers of Sinocrassula and close relative genus used in this study.

Species Voucher Location trnL-trnF psbA-trnH rbcL ITS matK Reference
Kungia aliciae Mayuzumi CH00061 (TI) Sichuan, China AB480632 AB480591 Mayuzumi and Ohba 2009
Sinocrassula ambigua Chen et al. YUS12973 (YUKU) Deqin, Yunnan, China PQ629032 PQ629054 PQ629039 PQ611189 PQ629047 Xu et al. 2025
S. ambigua Chen et al. YUS12672 (YUKU) Deqin, Yunnan, China PQ629030 PQ629055 PQ629038 PQ611188 PQ629046 Xu et al. 2025
S. ambigua Chen et al. YUS6698 (YUKU) Deqin, Yunnan, China PQ629035 PQ629059 PQ629040 PQ611190 PQ629048 Xu et al. 2025
S. densirosulata Chang XC19075 (SZ) China MW206800 MW206800 MW206800 MW206800 Unknown
S. diversifolia Chen et al. YUS9407 (YUKU) Gongshan, Yunnan, China PQ629070 PQ629074 PQ629066 PQ623396 PQ629062 This study
S. diversifolia Chen et al. YUS9477 (YUKU) Gongshan, Yunnan, China PQ629071 PQ629075 PQ629067 PQ623397 PQ629063 This study
S. ganluoensis Zhao et al. YUS6699 (YUKU) Ganluo, Sichuan, China PQ505691 PQ505693 PQ505695 PQ496498 PQ505697 Li et al. 2024
S. ganluoensis Zhao et al. YUS13920 (YUKU) Kangding, Sichuan, China PQ505692 PQ505694 PQ505696 PQ496499 PQ505698 Li et al. 2024
S. holotricha Zhao et al. YUS13475 (YUKU) Jiulong, Sichuan, China PQ629034 PQ629056 PQ629042 PQ611192 PQ629050 Xu et al. 2025
S. holotricha Zhao et al. YUS12867 (YUKU) Yuexi, Sichuan, China PQ629031 PQ629057 PQ629043 PQ611193 PQ629051 Xu et al. 2025
S. indica zjq20160061 (SANU) Xizang, China MN794334 MN794334 MN794334 MN794334 Zhao et al. 2020
S. indica var. obtusifolia Wang et al. YUS13936 (YUKU) Deqin, Yunnan, China PQ505699 PQ505701 PQ505703 PQ496500 PQ505705 Li et al. 2024
S. indica var. obtusifolia Wang et al. YUS13959 (YUKU) Deqin, Yunnan, China PQ505700 PQ505702 PQ505704 PQ496501 PQ505706 Li et al. 2024
S. jiaozishanensis Chen et al. JZS001 (YUKU) Luquan, Yunnan, China MZ343264 MZ343262 MZ343263 MZ343261 Wang et al. 2022
S. jiaozishanensis Chen et al. JZS002 (YUKU) Luquan, Yunnan, China MZ343269 MZ343267 MZ343268 MZ343266 Wang et al. 2022
S. jiaozishanensis Chen et al. YUS5900 (YUKU) Luquan, Yunnan, China PQ629036 PQ629058 PQ629044 PQ611194 PQ629052 Xu et al. 2025
S. obliquifolia Zhao et al. YUS9064 (YUKU) Hanyuan, Sichuan, China PQ629072 PQ629076 PQ629068 PQ623398 PQ629064 This study
S. obliquifolia Huang et al. YUS9366 (YUKU) Shimian, Sichuan, China PQ629073 PQ629077 PQ629069 PQ623399 PQ629065 This study
S. yunnanensis Chen s.n. (HIB) Yunnan, China KC988288 KC988295 Chen et al. 2014
S. yunnanensis Mayuzumi C00115 (TI) Yunnan, China AB480669 AB088582 Mayuzumi and Ohba 2004
S. yunnanensis Chen et al. YUS6697 (YUKU) Heqing, Yunnan, China PQ629037 PQ629060 PQ629045 PQ611195 PQ629053 Xu et al. 2025
S. yunnanensis Chen et al. YUS13776 (YUKU) Heqing, Yunnan, China PQ629033 PQ629061 PQ629041 PQ611191 PQ629049 Xu et al. 2025

Results and discussion

A total of 23 accessions representing 12 species from the genus Sinocrassula, along with one outgroup (Kungia aliciae), were included in the phylogenetic analysis. The concatenated dataset was 6,433 bp in length and GTR + F + I + G4 was selected as the best evolutionary model of nucleotide substitutions. The inferred phylogenetic trees from the ML and BI analyses revealed identical topologies. Two samples of the new species formed a highly supported clade (ML-BS = 99; BI-PP = 1.00, Fig. 1), and were found to be sister to Sinocrassula ganluoensis (ML-BS = 100, BI-PP = 1.00, Fig. 1). Then, they together are sister to S. diversifolia (ML-BS = 74, BI-PP = 0.94, Fig. 1). Although the new species is closely related to S. ganluoensis and S. diversifolia, it can be easily distinguished from these two species by its asymmetrical leaves and triangular petals, which are white at the base with dense purple-red stripes and spots on the both surfaces upward. Morphologically, the new species is most similar to S. indica (Table 2). However, the phylogenetic analysis showed that they have a distant relationship (Fig. 1).

Table 2.

Morphological comparison of Sinocrassula diversifolia, S. ganluoensis, S. indica, and S. obliquifolia.

Character S. diversifolia S. ganluoensis S. indica S. obliquifolia
Life cycle Perennial Biennial Biennial
Plant surface Glabrous, many parts purple-spotted Glabrous Glabrous Glabrous, many parts purple-spotted
Basal leaves Rosette lax, many parts brown-spotted, broadly obovate Rosette compact, orbicular- lanceolate Rosette, spatulate-oblong Rosette, asymmetrical leaves ovoid to lanceolate
Stem leaves Alternate, dimorphic Alternate, Linear, lanceolate Alternate, broadly oblanceolate, subobovate, ovate-orbicula Alternate alternate, lanceolate, many parts with purple spots
Sepals Triangular-lanceolate Ovate-lanceolate Broadly triangular Broadly triangular
Bracts Resembling distal stem leaves but smaller Lanceolate Resembling distal stem leaves but smaller Lanceolate
Inflorescences Corymbiform Corymbiform Paniculate, often corymbiform Corymbiform
Length of flowering stems 5–11.5 cm 5–60 cm or shorter 10–18 cm
Petals Lanceolate, yellow, spotted with purple Broad triangular, yellowish, deeply purplish red upward Lanceolate to ovate, red, reddish, yellow or greenish yellow Triangular, base white, spotted with purple upward
Nectar scales Broadly quadrate Quadrate Quadrate, apex emarginate Rectangler
Nectar scales size 0.3–0.5 × 0.5–0.7 mm 0.5 × 0.9 mm 0.3–0.5 × 0.2–0.3 mm
Length of styles 0.5–1.0 mm 0.6–1 mm Less than 1.0 mm 1.0–1.5 mm
Figure 1. 

The maximum likelihood phylogeny of Sinocrassula and its allies based on four chloroplast markers (psbA-trnH, trnL-F, rbcL, and matK) and one nuclear marker (ITS). Values associated with branches are Maximum Likelihood Bootstrap Support (ML-BS) and Bayesian Inference Posterior Probability (BI-PP). Bold indicated it received full ML-BS and BI-PP support.

The new species exhibits some typical characteristics of Sinocrassula including compact rosette, stem leaves alternate and lanceolate, bracts resembling distal stem leaves but smaller, flowers 5-merous, broadly triangular sepals, and ovoid carpels (Fu and Ohba 2013). A comparison of morphological characters among the new species and its morphologically similar species is shown in Table 2. The new species can easily distinguished from other species by its unique combination of asymmetrical leaves, rectangler nectar scales, and triangular petals which are white at the base and gradually transition to purple toward the tips (Fig. 2).

Figure 2. 

Sinocrassula obliquifolia A, B habit C, D flowers E stamen F, G petals H carpel I sepal J nectar scale K basal leaves L stem leaves M gland of basal leaf N leaves that abscise and bud during the flowering period, left: basal leaf, right: stem leaf.

Taxonomic treatment

Sinocrassula obliquifolia Jing Zhao, J.Guan Wang & X.M.Zhou, sp. nov.

Figs 2, 3

Type.

China • Sichuan: Hanyuan County, Yaan City, elev. ca. 837 m, 29.333262°N, 102.571721°E, on the granite crevices, 6 July 2023, Jing Zhao et al. YUS9064 (holotype YUKU!; isotype YUKU!).

Figure 3. 

Sinocrassula obliquifolia A habitat B, C habit.

Diagnosis.

Sinocrassula obliquifolia is similar to S. indica in having rosette, ovoid carpels, and broadly triangular sepals. However, this new species has shorter flowering stem (10–18 cm) (vs. 5–60 cm in S. indica), lanceolate stem leaves (vs. oblanceolate to ovate-orbicular stem leaves in S. indica), triangular petals (vs. lanceolate to ovate petals in S. indica), petals with a white base adorned with dense purple-red stripes and spots on the surfaces upward (vs. red, reddish, yellow, or greenish-yellow in S. indica) and rectangle nectar scales (vs. quadrate in S. indica). S. obliquifolia is also similar to S. diversifolia in having corymbiform inflorescences, petals, and stamens with purple spots. However, S. obliquifolia has a well-defined rosette (vs. less defined rosette in S. diversifolia), monomorphic stem leaves (vs. dimorphic stem leaves in S. diversifolia), triangular petals with dense purple-red stripes and spots on the surfaces upward (vs. lanceolate petals that are yellow with purple spots in S. diversifolia) and rectangle nectar scales (vs. broadly quadrate in S. diversifolia).

Description.

Perennial herbs, terrestrial or lithophytic, 5.0–20.0 cm tall, rosette 5.0–8.0 × 6.0–8.0 cm. Roots fibrous. Basal leaves rosette, spirally arranged, asymmetrically ovoid to lanceolate, apex with glands, 2.0–3.0 × 1.0–2.0 cm. Flowering stems terminal, 10.0–18.0 cm, glabrous with purple spots. Stem leaves alternate, lanceolate, many parts with purple spots. Bract resembling distal stem leaves but smaller, lanceolate, 3.0–3.5 × 0.5–1.0 cm. Inflorescences corymbiform, ca. 2.0–3.0 cm in diameter. Flowers small, ca. 3–6 mm in diameter. Sepals broadly triangular, purple with red spots, 1.5–2.0 × 0.5–1.2 mm. Petals triangular, base white, with dense purple-red stripes and spots on the surfaces upward, 2.0–4.0 × 1.0–2.0 mm. Stamens ca. 2–3 mm, white, anthers oblong-cordate, ca. 0.5 mm, pollen yellow. Nectar scales broadly rectangular, ca. 0.3–0.5 × 0.2–0.3 mm. Carpels 5, ovoid, clockwise rotation, 1.0–2.0 × 0.5–1 mm, styles 1.0–1.5 mm. Flowering June–October which coincides with the rainy season and the shedding of the leaves and the emergence of new buds.

Distribution and habitat.

Sinocrassula obliquifolia is currently known in central Sichuan Province, China. Two populations were found in granite crevices, as well as on dry stony or gravelly slopes at elevations ranging from 837 to 1140 m.

Additional specimens examined (paratypes).

China • Sichuan: Shimian County, Yaan City, elev. ca. 1140 m, 29.258594°N, 102.371583°E, on the granite crevices, 12 May 2023, Chuan-Jie Huang et al. YUS9366 (YUKU!).

Etymology.

The epithet obliquifolia refers to the asymmetrical leaves of the basal leaves, a unique characteristic for this species within Sinocrassula. Its Chinese name is suggested as ‘斜叶石莲(xie ye shi lian)’.

Acknowledgements

We thank Yi Liu for help in the field. Three anonymous reviewers and an editor are thanked for their helpful suggestions.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the National Science & Technology Fundamental Resources Investigation Program of China (Grant No. 2022FY100201) and the Postdoctoral Research Funds of Yunnan University (Grant No. W8163003).

Author contributions

Data curation: ZRH. Formal analysis: SLF. Resources: CJH, ML. Writing – original draft: RJL, JZ. Writing – review and editing: XMZ, JGW.

Author ORCIDs

Rong-Juan Li https://orcid.org/0009-0008-2667-398X

Jing Zhao https://orcid.org/0000-0001-7871-2209

Shao-Li Fang https://orcid.org/0000-0001-5942-0366

Chuan-Jie Huang https://orcid.org/0009-0000-3507-2766

Miao Luo https://orcid.org/0009-0006-9317-9151

Zhao-Rong He https://orcid.org/0000-0001-6492-3689

Xin-Mao Zhou https://orcid.org/0000-0003-3555-7784

Jia-Guan Wang https://orcid.org/0000-0003-0264-9180

Data availability

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

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Rong-Juan Li and Jing Zhao contributed equally to this work.
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