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. 2023 Mar 10;221:117–129. doi: 10.3897/phytokeys.221.97495

A new species of Sedum (Crassulaceae) from Mount Danxia in Guangdong, China

Yan-Shuang Huang 1, Kai-Kai Meng 1, Yuan-Yuan Sun 1, Zai-Xiong Chen 2, Qiang Fan 1,✉
PMCID: PMC10209613  PMID: 37250354

Abstract

Sedumjinglanii, a new species of Crassulaceae from Mount Danxia in Guangdong, China, is described and illustrated. Phylogenetic analysis based on the internal transcribed spacer (ITS) region of nrDNA suggests that the new species belongs to S.sect.Sedum sensu Fu and Ohba (2001) in the “Flora of China”, and is sister to a clade comprising S.alfredi and S.emarginatum with high support values (SH-aLRT = 84, UFBS = 95) but is distantly related to S.baileyi. The new species is morphologically similar to S.alfredi but it can be distinguished from the latter in its opposite leaves (vs. alternate leaves), its usually wider leaves (0.4–1.2 cm vs. 0.2–0.6 cm), its usually shorter petals (3.4–4.5 mm vs. 4–6 mm), its shorter nectar scales (0.4–0.5 mm vs. 0.5–1 mm), its shorter carpels (1.5–2.6 mm vs. 4–5 mm), and its shorter styles (0.6–0.9 mm vs. 1–2 mm). The new species can be easily distinguished from S.emarginatum which both have opposite leaves by its short, erect or ascending rhizome (vs. long and prostrate rhizome in the latter), shorter petals (3.4–4.5 mm vs. 6–8 mm) and shorter carpels (1.5–2.6 mm vs. 4–5 mm). It can also be easily distinguished from S.baileyi by its short, erect or ascending rhizome (vs. long and prostrate rhizome) and its shorter style (0.6–0.9 mm vs. 1–1.5 mm).

Keywords: Danxia landscape, morphology, nrITS, Sedum sect. Sedum

Introduction

According to Fu and Ohba (2001) in the “Flora of China”, Sedum Linnaeus is the most species-rich genus of the family Crassulaceae, comprising about 470 species. However, as presently circumscribed, the genus is highly polyphyletic, and a monophyletic circumscribed genus Sedum s.l. would comprise approximately 755 species by inclusion of all 14 genera currently recognized in tribe Sedeae into it (Messerschmid et al. 2020). The genus is distributed in temperate and subtropical environments, and the diversity center is in the Mediterranean Sea, Central America, the Himalayas and East Asia (Stephenson 1994; Thiede and Eggli 2007). In China, 121 Sedum species are documented, amongst which 91 species are endemic (Fu and Ohba 2001).

During the past 20 years, about seventeen Sedum species have been newly described from China, including S.hoi X.F.Jin & B.Y.Ding (Wang et al. 2005), S.plumbizincicola X.H.Guo & S.B.Zhou (Wu et al. 2012), S.fanjingshanense C.D.Yang & X.Yu Wang (Yang et al. 2012), S.kuntsunianum X.F.Jin, S.H.Jin & B.Y.Ding (Jin et al. 2013), S.tarokoense H.W.Lin & J.C.Wang (Lu et al. 2013), S.spiralifolium D.Q.Wang, D.M.Xie & L.Q.Huang (Xie et al. 2014), S.peltatum M.L.Chen & X.H.Cao (Chen et al. 2017), S.kwanwuense H.W.Lin, J.C.Wang & C.T.Lu and S.taiwanalpinum H.W.Lin, J.C.Wang & C.T.Lu (Lu et al. 2019), S.ichangense Y.B.Wang (Wang and Xiong 2019), S.lipingense R.B.Zhang, D.Tan & R.X.Wei (Zhang et al. 2019), S.nanlingense Yan Liu & C.Y.Zou (Zou et al. 2020), S.cirenianum S.S.Ying, S.shaoakouense S.S.Ying and S.shengkuangense S.S.Ying (Ying 2022a), and S.parviflorum S.S.Ying and S.tachingshuianum S.S.Ying (Ying 2022b).

Molecular data unambiguously demonstrate the polyphyletic nature of Sedum with its species placed in four major crown clades of the crassulacean tree, for example, Acre, Aeonium, Leucosedum, and Sempervivum. There is no agreement between specialists regarding the infrageneric structure of Sedum (reviewed in Nikulin et al. 2016). According to Fu and Ohba (2001), Chinese Sedum are divided into three sections, including S.sect.Sedum, S.sect.Oreades (Fröderström) K.T. Fu, and S.sect.Filipes (Fröderström) S.H. Fu. S.sect.Sedum is distinguishable from the two latter sections by the adaxially gibbous carpels and follicles, while S.sect.Oreades differs from S.sect.Filipes in the spurred leaf base and yellow petals (vs. spurless leaf base and white or reddish-purple petals) (Fu and Ohba 2001). During our investigations in Danxiashan National Park, Guangdong Province, China, an unknown Sedum species with opposite leaves was collected. After several years of field observations, comprehensive literature studies and molecular analysis, we confirmed that it was a new species and it is described and illustrated here.

Materials and method

Field investigations and observations were conducted during the flowering and fruiting periods of the putative new species. We obtained morphological data of this putative species by measurements based on 6–8 living samples. Mean values of these statistical data were calculated and then were compared with six other related species (Table 2). The specimens were collected in Danxiashan National Park, Renhua County, Guangdong Province, China. Voucher specimens were deposited in the herbarium of Sun Yat-sen University (SYS).

Table 2.

Morphological comparisons between S.jinglanii, S.alfredi, S.baileyi, S.emarginatum, S.kuntsunianum, S.makinoi, and S.satumense.

Characters S.jinglanii S.alfredi † S.baileyi ‡ S.emarginatum § S.kuntsunianum | S.makinoi S.satumense #
Leaf blade Spatulate or obovate Linear-cuneate, spatulate or obovate Obovate-spatulate Spatulate-obovate to broadly obovate Widely obovate or suborbiculate, spatulate Obovate or obovate-spatulate Narrowly obovate or spatulate
Leaf size (cm) 0.8–2.9 × 0.4–1.2 1.2–3.0 × 0.2–0.6 1–2.5 × 0.6–0.8 1–2.5 × 0.5–1.2 1.4–2.0 × 0.9–1.5 1–2 × 0.6–0.8 1.0–2.2 × 0.6–0.9
Phyllotaxy Opposite Alternate Opposite Opposite Opposite, or rarely alternate at base Opposite Opposite
Rhizome Short, erect or ascending Short, erect or ascending Long, prostrate Long, prostrate Absent Short, erect or ascending Short, erect or ascending
Sepal length (mm) 2–3.1 2–5 1.5–2.5 2–5 5–9 2–3 6–7
Petal length (mm) 3.4–4.5 4–6 4–5 6–8 7–8 4–5 7–8
Stamen length (antepetalous) (mm) 2.2–2.6 2.5–3.5 2–3 3–4 ca. 5 2.5–3.2 –
Stamen length (antesepalous) (mm) 3.2–3.3 3.8–4.5 3–4 4–5 ca. 6 2.8–4.5 –
Nectar scale length (mm) 0.4–0.5 0.5–1 0.4–0.6 0.6–0.8 ca. 0.5 0.5–0.7 ca. 0.5
Carpel length (mm) 1.5–2.6 4–5 2–3 4–5 ca. 5 4–5 –
Style length (mm) 0.6–0.9 1–2 1–1.5 1.5–2 ca. 1 1–2 1.0–1.5
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†,‡,§, Based on Fu and Ohba (2001) and own measurements at IBSC and SYS; |Jin et al. (2013); #Ohba (2003).

Two representative individuals from different populations were selected for further molecular experiments, one from Bazhai of Mount Danxia (Y. S. Huang 21040301) and another one from Yanyan of Mount Danxia (Q. Fan et al. DNPC 2873). Fresh leaves of the two individuals were collected and stored with silica gel in zip-lock plastic bags until use. Total DNA was extracted using the modified CTAB method (Doyle and Doyle 1987). The region of the partial internal transcribed spacer 1, 5.8S ribosomal RNA gene and partial internal transcribed spacer 2 was amplified using previously reported primers ITS1 and ITS4 (White et al. 1990). PCR amplifications were performed following Huang et al. (2021).

In order to explore the phylogenetic position of the putative new species in Sedum, ITS sequences of 56 accessions representing 46 Sedum taxa and three outgroup species (Aeoniumlancerottense, Aeoniumviscatum, and Greenoviaaizoon) were downloaded from the Genbank public database at the National Center for Biotechnology Information (NCBI) (Table 1). The sequences were aligned using ClustalW 1.8 (Thompson et al. 1994) and then adjusted manually. Besides, to improve the credibility, we also aligned the sequences using MAFFT v.7.402 (Katoh and Standley 2013), and the alignments generated from the two methods were consistent. The best-fit nucleotide substitution model was determined by ModelFinder (Kalyaanamoorthy et al. 2017). Based on the maximum likelihood (ML) method, the phylogenetic tree was constructed using IQ-Tree v. 2.0.3 (Nguyen et al. 2015) by executing 5,000 replicates of SH approximate likelihood ratio test (SH-aLRT) and ultrafast bootstrap (UFBS) (Hoang et al. 2018). Finally, the tree file was visualized by the online tool of Interactive Tree Of Life (iTOL) v5 (Letunic and Bork 2021).

Table 1.

Taxa, voucher information, and GenBank accession numbers of the sequences used in this study.

Taxon Voucher Accession numbers References
SedumSect.Oreades
S.oreades Rao 090803-03 KF113733 Zhang et al. 2014
S.trullipetalum Miyamoto et al. 9420132 AB088630 Mayuzumi and Ohba 2004
S.bergeri Ni et al. AY352897 Ni et al. unpublished
S.erici-magnusii Ito 2077 LC229235 Ito et al. 2017a
SedumSect.Sedum
S.jinglanii Huang 21040301 OP288035 This study
Fan et al. DNPC 2873 OQ162326 This study
S.actinocarpum Ito 1749 LC229265 Ito et al. 2017a
S.alfredi Kokubugata 17190 AB930259 Ito et al. 2014a
Kokubugata 17191 AB930260 Ito et al. 2014a
Kokubugata 17192 AB930261 Ito et al. 2014a
WUK415208 FJ919953 Wang and Shu unpublished
S.baileyi LBG0064555 FJ919935 Wang and Shu unpublished
S.bulbiferum Ito 416 LC229234 Ito et al. 2017a
130514hs41 KM111166 Xie et al. 2014
130524qz09 KM111165 Xie et al. 2014
S.emarginatum 130512hs27 KM111145 Xie et al. 2014
S.erythrospermum Tsutsumi 1504 AB906473 Ito et al. 2014b
S.formosanum Ito 1260 LC229279 Ito et al. 2017a
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.senanense Ito 2200 LC229238 Ito et al. 2017a
S.oryzifolium Ito 2285 LC229239 Ito et al. 2017a
S.japonicumvar.pumilum Ito 2287 LC229240 Ito et al. 2017a
S.japonicumssp.uniflorum Ito 447 LC229241 Ito et al. 2017a
S.boninense Ito 2371 LC229242 Ito et al. 2017a
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.lungtsuanense Ito 3563 LC260131 Ito et al. 2017b
S.makinoi Kokubugata 16730 AB906476 Ito et al. 2014b
S.morrisonense Ito 2765 LC229290 Ito et al. 2017a
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.yabeanum Ito 396 AB906490 Ito et al. 2014b
S.polytrichoidesvar.setouchiense Ito 2298 LC229253 Ito et al. 2017a
S.polytrichoides CMQ1057 LC229251 Ito et al. 2017a
S.rupifragum Ito 2070 LC229254 Ito et al. 2017a
S.sarmentosum Ito 978 LC229255 Ito et al. 2017a
S.satumense Ito 2295 LC229256 Ito et al. 2017a
S.subtile Shimizu 1999 AB088622 Mayuzumi and Ohba 2004
Ito 2259 LC229257 Ito et al. 2017a
S.subtile Ito 624 AB930277 Ito et al. 2014a
S.taiwanianum Ito 2770 LC229297 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
S.lipingense * ZRB1479 MN150061 Zhang et al. 2019
S.mexicanum * Ito 647 LC229247 Ito et al. 2017a
S.truncatistigmum * Ito 3254 LC229306 Ito et al. 2017a
S.zentaro-tashiroi * Ohba 1998 AB088619 Mayuzumi and Ohba 2004
Outgroups
Aeoniumlancerottense Mort 1518 AY082143 Mort et al. 2002
Aeoniumviscatum Mort 1432 AY082154 Mort et al. 2002
Greenoviaaizoon Mort 1425 AY082112 Mort et al. 2002
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*Not recorded in Fu and Ohba (2001).

Results and discussion

The alignment length of the ITS sequences was 624 bp, amongst which 340 were parsimony-informative. Within the new species, only one variable site was detected, but 40 variable sites were detected between the new species and S.alfredi and 40 variable sites were detected between the new species and S.emarginatum, indicating that pronounced genetic differentiation existed between the new species and S.alfredi as well as S.emarginatum. The best-fit nucleotide substitution model was estimated as SYM+I+G4 according to the Bayesian Information Criterion (BIC).

As the ML phylogenetic tree shows (Fig. 1), seven subclades were resolved with moderate to high support values. Accessions of the putative new species, S.alfredi, S.emarginatum, and S.lungtsuanense together formed subclade 7 with high support values (SH-aLRT = 92, UFBS = 98), all belonging to S.sect.Sedum sensu Fu and Ohba (2001).

Figure 1.

Figure 1.

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Maximum Likelihood tree based on ITS sequences for Eastern Asian species of Sedumsect.Sedum, four species of S.sect.Oreades, and three outgroups. Numbers near the branches are SH approximate likelihood ratio test (SH-aLRT) and ultrafast bootstrap (UFBS) support values. The new species is highlighted in bold.

Morphologically, the putative new species is similar to S.alfredi from which it can be easily distinguished by its opposite leaves (vs. alternate leaves in the latter). Furthermore, the leaves of the putative new species are usually wider than those of S.alfredi (0.4–1.2 mm vs. 0.2–0.6 mm), the petals are usually shorter (3.4–4.5 mm vs. 4–6 mm), the nectar scales shorter (0.4–0.5 mm vs. 0.5–1 mm), the carpels shorter (1.5–2.6 mm vs. 4–5 mm) and the styles shorter (0.6–0.9 mm vs. 1–2 mm) (Table 2). Phylogenetically, the putative new species is closely related to S.emarginatum. Although the leaves of both species are opposite, it can be easily distinguished from the latter by its short, erect or ascending rhizome (vs. long and prostrate rhizome), shorter petals (3.4–4.5 mm vs. 6–8 mm) and shorter carpels (1.5–2.6 mm vs. 4–5 mm). The putative new species was distantly related to S.baileyi in the phylogenetic tree although both are morphologically similar (Table 2). Also, it can be easily distinguished from the latter by its short, erect or ascending rhizome (vs. long and prostrate rhizome) and its shorter style (0.6–0.9 mm vs. 1–1.5 mm).

Additionally, four representatives of Sedumsect.Oreades sensu Fu and Ohba (2001) (S.oreades, S.trullipetalum, S.bergeri, and S.erici-magnusii) were also included in our analysis. However, these four species were nested within species belonging to S.sect.Sedum sensu Fu and Ohba (2001), thus showing that S.sect.Sedum might not be monophyletic. This result is consistent with previous studies (Nikulin et al. 2016; Zhang et al. 2019; Messerschmid et al. 2020).

Through numerous scientific investigations, more than a dozen new species were found on Mount Danxia in Guangdong in recent years, and most are endemic to it such as Lespedezadanxiaensis Q.Fan, W.Y.Zhao & K.W.Jiang (Zhao et al. 2021), Aspleniumdanxiaense K.W.Xu (Xu et al. 2022), Pileadanxiaensis L.F.Fu, A.K.Monro & Y.G.Wei (Fu et al. 2022), Wikstroemiafragrans W.B.Liao, Q.Fan & J.R.Chen (Chen et al. 2022), and Commelinadanxiaensis Q.Fan, Long Y.Wang & W.Guo (Wang et al. 2023). As a World Heritage site and tourist attraction, Danxia landform possesses special and complicated habitat differences at a small scale, which might contribute to the plant endemism at Mount Danxia (Peng et al. 2018).

Taxonomic treatment

. Sedum jinglanii

Yan S.Huang & Q.Fan sp. nov.

35584965-9C8A-516F-823A-890BDDE1C6C8

urn:lsid:ipni.org:names:77315511-1

Type.

China. Guangdong Province, Renhua County, Mount Danxia, Bazhai, in the cliff of steep slopes, 25°00'N, 113°39'E, 520 m a.s.l., 3 April 2021, Y. S. Huang 21040301 (holotype: SYS; isotype: SYS) (Figs 2, 3).

Figure 2.

Figure 2.

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Sedumjinglanii sp. nov. A habit B flower with sepals, petals, stamens and carpels C petals and stamens D sepal E leaves. Illustration by Yuan-Yuan Sun based on living field-collected material (Y. S. Huang 21040301).

Figure 3.

Figure 3.

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Sedumjinglanii sp. nov. A habit B flower, front view, showing brownish-red anthers C carpels and a nectar scale (red arrow) D abaxial leaf surface E young unripe fruits F young sterile individuals A, B, D photographed by Qiang Fan in the field (Pingtouzhai, 6 April 2022, Q. Fan et al., DNPC 1953) C photographed by Min Lin in the lab (7 April 2022, Q. Fan et al., DNPC 1953) E, F photographed by Yan-Shuang Huang (Bazhai, 3 April 2021, Y. S. Huang 21040301).

Diagnosis.

This new species is similar to S.alfredi, but differs from the latter in its opposite leaves (vs. alternate leaves), its usually wider leaves (0.8–2.9 × 0.4–1.2 cm vs. 1.2–3.0 × 0.2–0.6 cm), usually shorter petals (3.4–4.5 mm vs. 4–6 mm), shorter nectar scales (0.4–0.5 mm vs. ca. 0.5–1 mm), and shorter carpels (1.5–2.6 mm vs. 4–5 mm). Although the leaves of this new species and of S.emarginatum are opposite, it can be easily distinguished from the latter by its short, erect or ascending rhizomes (vs. long and prostrate rhizomes), shorter petals (3.4–4.5 mm vs. 6–8 mm) and shorter carpels (1.5–2.6 mm vs. 4–5 mm).

Description.

Fleshy herbs, perennial; stems glabrous, greenish, often with small reddish dots thus appearing more or less reddish, ascending; leaves opposite, usually deciduous, crowded distally on the stem, succulent; leaf blade spatulate or obovate, 8–29 mm long, 4–12 mm wide, base narrowly cuneate and spurred, apex obtuse and sometimes emarginate; inflorescence in dense terminal cymes, usually two to four branched; bracts leaflike, 1.7–2.4 mm long, 0.7–1.1 mm wide; flowers usually sessile, rarely with short pedicels to 0.8 mm long, unequally 5-merous; sepals green, linear-spatulate, 2–3.1 mm × 0.7–1.4 mm, base shortly spurred; petals yellow, lanceolate to lanceolate-oblong, 3.4–4.5 mm × 0.8–1.1 mm, base connate for 0.1–0.2 mm, apex mucronate; stamens 10, yellow, filiform, arranged in 2 whorls; antesepalous ones 3.2–3.3 mm, antepetalous ones 2.2–2.6 mm; anthers brownish red, long ellipsoid. Nectar scales yellow green, spatulate-quadrangular, 0.4–0.5 × 0.2–0.3 mm, apex obtusely truncate. Carpels yellow green, erect, ovoid-lanceolate, 1.5–2.6 mm long, 0.6–0.9 mm wide, adaxially gibbous, base shortly connate; styles 0.6–0.9 mm long. Follicles yellowish, obliquely divergent. Seeds numerous, brown, oblong, 0.5–0.6 mm, papillate.

Phenology.

Flowering from April to May. Fruiting from June to August.

Etymology.

Sedumjinglanii is named after Prof. Jing-Lan Feng (1898–1976), an academician of the Chinese Academy of Sciences and one of the founders of mineralogy in China. In 1928, he discovered and named the red beds and related strata in North Guangdong as “Danxia Formation” for the first time (Peng 2020).

Distribution and habitat.

Presently, this new species is only known from the type locality, Mount Danxia, Renhua County, Guangdong Province, China. It grows on the cliff of steep slopes at altitudes of 200–550 m a.s.l.

Conservation status.

Only five populations were found with no more than 1,000 mature individuals. Thus, the conservation status could be considered as Vulnerable (VU; D1), according to the IUCN Red List Criteria (IUCN Standards and Petitions Subcommittee 2022).

Additional specimens examined

(paratypes). China. Guangdong: Renhua County, Mount Danxia, Pingtouzhai, 25°00'N, 113°37'E, 536 m a.s.l., 6 April 2022, Q. Fan et al., DNPC 1953 (SYS); Renhua County, Mount Danxia, Yanyan, 25°02'N, 113°61'E, 263 m a.s.l., 27 December 2022, Q. Fan et al. DNPC 2873 (SYS).

Supplementary Material

XML Treatment for Sedum jinglanii

Acknowledgements

We appreciate Mrs. Jing-Min Dai for her assistance in preparing the manuscript. We thank Prof. Yan-Song Peng from Lushan Botanical Garden, Chinese Academy of Sciences for his help in collecting living material of Sedumbaileyi at the type locality. We thank Dr. Bin Chen, senior engineer at Shanghai Chenshan Botanical Garden, Chinese Academy of Sciences for his suggestions in nomenclature. We thank Mrs. Min Lin and Mrs. Li-Juan Liao for their help in the morphological analysis. We also kindly thank Zu-Hao Wang and Wen-Sheng Shu for unpublished sequence data, Yuan-Yuan Sun for the illustration, and the reviewers for their valuable comments. This study was supported by the Guangdong Provincial Special Research Grant for the Creation of National Parks (2021GJGY034) and the Foundation of Administrative Committee of Danxiashan National Park (K22-33000-060, 2016-0293).

Citation

Huang Y-S, Meng K-K, Sun Y-Y, Chen Z-X, Fan Q (2023) A new species of Sedum (Crassulaceae) from Mount Danxia in Guangdong, China. PhytoKeys 221: 117–129. https://doi.org/10.3897/phytokeys.221.97495

Funding Statement

This study was supported by the Guangdong Provincial Special Research Grant for the Creation of National Parks (2021GJGY034) and the Foundation of Administrative Committee of Danxiashan National Park (K22-33000-060, 2016-0293).

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Supplementary Materials

XML Treatment for Sedum jinglanii
phytokeys.221.97495-treatment1.xml (12.8KB, xml)

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