Research Article |
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Corresponding author: Zhao-Rong He ( zhrhe@ynu.edu.cn ) Corresponding author: Xin-Mao Zhou ( xinmao.zhou@ynu.edu.cn ) Corresponding author: Jia-Guan Wang ( wangjiaguan@ynu.edu.cn ) Academic editor: Takuro Ito
© 2025 Rong-Juan Li, Jing Zhao, Shao-Li Fang, Chuan-Jie Huang, Miao Luo, Zhao-Rong He, Xin-Mao Zhou, Jia-Guan Wang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Li R-J, Zhao J, Fang S-L, Huang C-J, Luo M, He Z-R, Zhou X-M, Wang J-G (2025) Sinocrassula obliquifolia (Crassulaceae), a new species from China. PhytoKeys 255: 103-112. https://doi.org/10.3897/phytokeys.255.142079
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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.
Hengduan mountains, phylogeny, rosette, species diversity, taxonomy
Crassulaceae are a morphologically diverse and systematically complex group of angiosperms, comprising 35 genera and approximately 1,400 species (
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 (
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.,
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 (
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 (
Sequences from GenBank and the newly generated data (Table
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 | – |
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| Sinocrassula ambigua | Chen et al. YUS12973 ( |
Deqin, Yunnan, China | PQ629032 | PQ629054 | PQ629039 | PQ611189 | PQ629047 |
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| S. ambigua | Chen et al. YUS12672 ( |
Deqin, Yunnan, China | PQ629030 | PQ629055 | PQ629038 | PQ611188 | PQ629046 |
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| S. ambigua | Chen et al. YUS6698 ( |
Deqin, Yunnan, China | PQ629035 | PQ629059 | PQ629040 | PQ611190 | PQ629048 |
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| S. densirosulata | Chang XC19075 (SZ) | China | MW206800 | MW206800 | MW206800 | – | MW206800 | Unknown |
| S. diversifolia | Chen et al. YUS9407 ( |
Gongshan, Yunnan, China | PQ629070 | PQ629074 | PQ629066 | PQ623396 | PQ629062 | This study |
| S. diversifolia | Chen et al. YUS9477 ( |
Gongshan, Yunnan, China | PQ629071 | PQ629075 | PQ629067 | PQ623397 | PQ629063 | This study |
| S. ganluoensis | Zhao et al. YUS6699 ( |
Ganluo, Sichuan, China | PQ505691 | PQ505693 | PQ505695 | PQ496498 | PQ505697 |
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| S. ganluoensis | Zhao et al. YUS13920 ( |
Kangding, Sichuan, China | PQ505692 | PQ505694 | PQ505696 | PQ496499 | PQ505698 |
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| S. holotricha | Zhao et al. YUS13475 ( |
Jiulong, Sichuan, China | PQ629034 | PQ629056 | PQ629042 | PQ611192 | PQ629050 |
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| S. holotricha | Zhao et al. YUS12867 ( |
Yuexi, Sichuan, China | PQ629031 | PQ629057 | PQ629043 | PQ611193 | PQ629051 |
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| S. indica | zjq20160061 (SANU) | Xizang, China | MN794334 | MN794334 | MN794334 | – | MN794334 |
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| S. indica var. obtusifolia | Wang et al. YUS13936 ( |
Deqin, Yunnan, China | PQ505699 | PQ505701 | PQ505703 | PQ496500 | PQ505705 |
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| S. indica var. obtusifolia | Wang et al. YUS13959 ( |
Deqin, Yunnan, China | PQ505700 | PQ505702 | PQ505704 | PQ496501 | PQ505706 |
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| S. jiaozishanensis | Chen et al. JZS001 ( |
Luquan, Yunnan, China | MZ343264 | MZ343262 | MZ343263 | – | MZ343261 |
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| S. jiaozishanensis | Chen et al. JZS002 ( |
Luquan, Yunnan, China | MZ343269 | MZ343267 | MZ343268 | – | MZ343266 |
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| S. jiaozishanensis | Chen et al. YUS5900 ( |
Luquan, Yunnan, China | PQ629036 | PQ629058 | PQ629044 | PQ611194 | PQ629052 |
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| S. obliquifolia | Zhao et al. YUS9064 ( |
Hanyuan, Sichuan, China | PQ629072 | PQ629076 | PQ629068 | PQ623398 | PQ629064 | This study |
| S. obliquifolia | Huang et al. YUS9366 ( |
Shimian, Sichuan, China | PQ629073 | PQ629077 | PQ629069 | PQ623399 | PQ629065 | This study |
| S. yunnanensis | Chen s.n. (HIB) | Yunnan, China | – | – | – | KC988288 | KC988295 |
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| S. yunnanensis | Mayuzumi C00115 (TI) | Yunnan, China | AB480669 | – | – | AB088582 | – | Mayuzumi and Ohba 2004 |
| S. yunnanensis | Chen et al. YUS6697 ( |
Heqing, Yunnan, China | PQ629037 | PQ629060 | PQ629045 | PQ611195 | PQ629053 |
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| S. yunnanensis | Chen et al. YUS13776 ( |
Heqing, Yunnan, China | PQ629033 | PQ629061 | PQ629041 | PQ611191 | PQ629049 |
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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.
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 |
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 (
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).
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.
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.
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 (
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)’.
We thank Yi Liu for help in the field. Three anonymous reviewers and an editor are thanked for their helpful suggestions.
The authors have declared that no competing interests exist.
No ethical statement was reported.
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).
Data curation: ZRH. Formal analysis: SLF. Resources: CJH, ML. Writing – original draft: RJL, JZ. Writing – review and editing: XMZ, JGW.
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
All of the data that support the findings of this study are available in the main text.