Citation

urn:lsid:arphahub.com:pub:F7FCE910-8E78-573F-9C77-7788555F8AAD

urn:lsid:zoobank.org:pub:66878968-5FA3-48E4-A0CA-743FBBB0315D

PhytoKeys

1314-2011 1314-2003

Pensoft Publishers

10.3897/phytokeys.266.170977

170977

Research Article

Angiospermae Fabaceae

Taxonomy

Cenozoic

Asia

Astragalus

zibaishanensis

sp. nov. (

Fabaceae

), a new species from Shaanxi, China

Liu

Pei-Liang

https://orcid.org/0000-0001-6566-7824 1 Xun

Lu-Lu

https://orcid.org/0000-0002-6861-5883 2 Su

Chun

2 Lu

Yuan

https://orcid.org/0000-0002-0329-0803 2 Li

Bin

https://orcid.org/0000-0001-9254-8003 2 Yue

Ming

yueming@nwu.edu.cn https://orcid.org/0000-0002-8618-1126 1 2

Key Laboratory of Resource Biology and Biotechnology in Western China, Ministry of Education, Northwest University, Xi’an, Shaanxi, China

Northwest University

Xi'an

China 2

Shaanxi Engineering Research Centre for Conservation and Utilization of Botanical Resources, Xi’an Botanical Garden of Shaanxi Province (Institute of Botany of Shaanxi Province), Xi’an, Shaanxi, China

Shaanxi Engineering Research Centre for Conservation and Utilization of Botanical Resources, Xi’an Botanical Garden of Shaanxi Province (Institute of Botany of Shaanxi Province)

Xi'an

China

Corresponding author: Ming Yue (yueming@nwu.edu.cn)

Academic editor: Stephen Boatwright

2025

19 11 2025

266 267 280 C9F4F344-0835-5E2E-9225-726AE6BF5D1D 17668914 03 09 2025 03 11 2025

Pei-Liang Liu, Lu-Lu Xun, Chun Su, Yuan Lu, Bin Li, Ming Yue

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.

Abstract

Astragalus zibaishanensissp. nov. (Fabaceae) is described and illustrated from Shaanxi, China. This new species is similar to Astragalus monadelphus, A. xitaibaicus and A. neomonodelphus by having yellow petals, pseudomonadelphous androecium, and stipitate legumes, but can be distinguished by its stem 0.9–1.5 mm in diameter; leaflets 5–7, hairy adaxially and abaxially; calyx teeth triangular, ca. 0.5 mm long; standard 17–20 mm long; wings 18–21 mm long; keels 17–20 mm long; pistil glabrous; legume glabrous, with a stipe 10–11 mm long. Astragalus zibaishanensis is unique in Astragalus by having pseudomonadelphous androecium. Phylogenetic analyses confirm that the new species is a member of A. sect. Cenantrum. The nuclear ITS tree shows that A. zibaishanensis forms a clade with A. henryi, A. umbellatus, A. chilienshanensis, A. moellendorffii, A. przewalskii, A. monadelphus, A. xitaibaicus and A. neomonodelphus.

Key words:

Fabaceae

new species phylogeny taxonomy

Basic Research Program of the Shaanxi Academy of Sciences (grant number 2023K-14)

Citation

Liu P-L, Xun L-L, Su C, Lu Y, Li B, Yue M (2025) Astragalus zibaishanensis sp. nov. (Fabaceae), a new species from Shaanxi, China. PhytoKeys 266: 267–280. https://doi.org/10.3897/phytokeys.266.170977

Introduction

Astragalus L. (Fabaceae) is the largest plant genus, and more than 3200 species have been recognized worldwide (Moonlight et al. 2024). The genus is distributed in various habitats in temperate northern hemisphere, with less species in the southern hemisphere. The genus exhibits outstanding species richness in arid and semi-arid areas in Asia and North America with great ecological importance (Folk et al. 2024). Molecular phylogenetic analyses have revealed intergeneric and intrageneric relationships of Astragalus (Wojciechowski et al. 1999; Kazempour Osaloo et al. 2003; Wojciechowski 2005; Duan et al. 2024; Folk et al. 2024), and 11 intrageneric clades have been recognized (Azani et al. 2017, 2019; Su et al. 2021a; Buono et al. 2025). Within Astragalus, the clade Phaca is an early-divergent group which includes perennial herbaceous species with basifixed hairs, large yellow petals, mostly bilocular or semi-bilocular legumes (Azani et al. 2017, 2019; Su et al. 2021a; Folk et al. 2024). Within the Phaca clade, Astragalus sect. Cenantrum Bunge is generally characterized by well-developed stems, free stipules, imparipinnate leaves, peduncled racemes, campanulate calyx and stipitate unilocular papery legumes (Podlech and Zarre 2013). In this section, there are 41 species in Eurasia (Podlech and Zarre 2013), and one species, A. americanus (Hook.) M. E. Jones, is endemic to North America (Isely 1998).

About 430 species of Astragalus have been documented from China (Chai and Yan 2010; Podlech 2010; Xu and Podlech 2010; Liu et al. 2011; Zhao 2012; Li and Yu 2014; Wu 2015; Ri and Zhao 2018; Liu et al. 2020; Hu et al. 2021; Su et al. 2021b; Zhang et al. 2021; Xu et al. 2023; Ke et al. 2024; Yang et al. 2024; Wu 2025; Yang et al. 2025). A total of 28 species of Astragalus sect. Cenantrum are known from China (Podlech 2010; Xu and Podlech 2010; Podlech and Zarre 2013). In the fieldwork in Mt. Zibaishan, Shaanxi, China, we found a species belonging to A. sect. Cenantrum, which was revealed as a new species by detailed morphological comparison and phylogenetic analyses.

Material and methods Taxon sampling

Three accessions of the new species were sampled from the field. Because the new species is morphologically similar to A. monadelphus Bunge ex Maxim., A. xitaibaicus (K. T. Fu) Podlech & L. R. Xu and A. neomonodelphus H. T. Tsai & T. T. Yu, these three species were also sampled for phylogenetic analyses. We also sampled another 15 species in A. sect. Cenantrum to test the phylogenetic position of the new species in this section. Previous phylogenetic analysis showed that species of A. sect. Skythropos N. D. Simpson and A. sect. Ebracteolati N. D. Simpson were nested in A. sect. Cenantrum (Su et al. 2021a), therefore A. skythropos Bunge and A. craibianus N. D. Simpson were also sampled. Representative species of other clades of Astragalus (Su et al. 2021a) were included in phylogeny. Oxytropis bicolor Bunge was selected as the outgroup. Sample information was shown in Appendix 1.

DNA extraction, PCR and sequencing

Genomic DNA was extracted from silica-gel dried leaves by using the Qiagen DNeasy® Plant Mini Kit (Hilden, Germany). The nuclear ribosomal internal transcribed spacer (ITS), and the plastid psbA-trnH and trnL-F sequences were amplified by Polymerase chain reactions (PCR). Primers were ITS5 and ITS4 (White et al. 1990) for ITS, psbAF and trnHR (Sang et al. 1997) for psbA-trnH, c and f (Taberlet et al. 1991) for trnL-F sequences. PCR conditions followed Liu et al. (2017). Amplicons were sequenced in both directions by using the PCR primers. Sequences were assembled in Geneious Prime® 2025.0.2 (https://www.geneious.com). Newly generated sequences have been deposited in the GenBase (Bu et al. 2024) in National Genomics Data Center (CNCB-NGDC Members and Partners 2025), Beijing Institute of Genomics, Chinese Academy of Sciences/China National Center for Bioinformation. The GenBase accession numbers were listed in Appendix 1, and the data were publicly accessible at https://ngdc.cncb.ac.cn/genbase.

Phylogenetic analysis

Phylogenetic analyses were conducted based on the newly generated sequences together with the data from Su et al. (2021a) and several sequences from GenBank (Appendix 1). Multiple sequence alignments were performed in Geneious Prime®. The best-fit nucleotide substitution model for each alignment was determined by using jModelTest v.2.1.7 with the Bayesian Information Criterion (BIC) (Darriba et al. 2012). The models for the ITS, psbA-trnH and trnL-F sequences were K80+G, F81+I+G and F81+G, respectively. The nuclear and the plastid data were analyzed separately. The plastid psbA-trnH and trnL-F sequences were concatenated and partitioned in phylogenetic reconstructions by using respective models. MrBayes v.3.2.5 (Ronquist and Huelsenbeck 2003; Ronquist et al. 2012) was used to conduct Bayesian inference (BI). BI was run for 10,000,000 generations, and trees were sampled every 1,000 generations. The first 2,500 trees were discarded, and the remaining 7,500 trees were used to build a 50% majority-rule consensus tree and posterior probabilities (PP). The maximum parsimony (MP) and maximum likelihood (ML) analyses were conducted by using PAUP* 4.0a169 (Swofford 2002) and RAxML v.8.2 (Stamatakis 2014), respectively. The MP and ML bootstrap analyses were performed with 1,000 replicates. Bootstrap support percentages (BSMP, BSML) from the MP and ML analyses were labeled on the corresponding branches of the BI trees.

Results Nuclear data

In the nuclear tree (Fig. 1A), three accessions of the new species clustered together (PP = 0.94, BSMP = 63%, BSML = 76%). The new species and A. henryi Oliv., A. umbellatus Bunge, A. chilienshanensis Y. C. Ho, A. moellendorffii Bunge, A. przewalskii Bunge, A. monadelphus, A. xitaibaicus and A. neomonodelphus formed a well-supported clade (PP = 1, BSMP = 85%, BSML = 92%). Within this clade, A. monadelphus, A. xitaibaicus and A. neomonodelphus clustered together (PP = 0.82, BSMP = 66, BSML = 65). Astragalus skythropos (A. sect. Skythropos) and A. craibianus (A. sect. Ebracteolati) nested with species of A. sect. Cenantrum, and they formed a well-supported clade (PP = 1, BSMP = 95%, BSML = 98%). The Phaca clade was well-supported (PP = 0.99, BSMP = 77%, BSML = 89%).

10.3897/phytokeys.266.170977.figure1

17669032

855AA868-77EB-5183-A9A6-E4C1504815C8

Figure 1.

Bayesian 50% majority-rule consensus trees based on the nuclear ITS sequence A. and the combined plastid psbA-trnH and trnL-F sequences B. The Bayesian posterior probabilities (PP) are below the branches, the maximum parsimony and the maximum likelihood bootstrap support percentages (BSMP, BSML) are above the branches. A dash (–) indicates a branch that is not found in the maximum parsimony tree.

https://binary.pensoft.net/fig/1469679 Plastid data

In the plastid tree (Fig. 1B), three accessions of the new species clustered together (PP = 1, BSMP = 73%, BSML = 89%). Astragalus monadelphus and A. xitaibaicus formed a clade (PP = 0.92, BSMP = 85%, BSML = 90%). The Phaca clade was also supported (PP = 0.93, BSMP = 53%, BSML = 54%), but the relationships among species of the Phaca clade were largely unresolved in a polytomy.

Taxonomy

Taxon classification

Plantae

Fabales

Fabaceae

74EA866E-7C02-5E96-8E6E-6C707FDBB0BA

Astragalus

zibaishanensis

urn:lsid:ipni.org:names:77372165-1

P.L. Liu & L.L. Xun, sp. nov. (A. sect. Cenantrum Bunge)

Figs 2 , 3

Type.

China • Shaanxi Province, Fengxian County, north slope of Mt. Zibaishan, 33°41'49.6"N, 106°45'21.3"E, on stony slope in grasses, 2311 m above sea level (a. s. l.), 13 June 2023, P. L. Liu 1458 (Holotype, WUK!; Isotypes, WUK!, WNU!).

10.3897/phytokeys.266.170977.figure2

17669034

46A51176-45B2-550A-B4E1-1F854E019BF3

Figure 2.

Illustration of Astragalus zibaishanensis. a. Root with basal stems; b. Upper part of a flowering plant; c. Upper part of a fruiting plant; d. Leaflet, adaxial view; e. Leaflet, abaxial view; f. Split calyx, inner side; g. Standard; h. Wing; i. Keels; j. Androecium; k. Pistil; l. Seed. Drawn by Yi-Fan Li.

https://binary.pensoft.net/fig/1469680

Diagnosis.

This new species is similar to A. monadelphus, A. xitaibaicus and A. neomonodelphus by having yellow petals, pseudomonadelphous androecium, and stipitate legumes, but can be distinguished by its stem 0.9–1.5 mm in diameter; leaflets 5–7, hairy adaxially and abaxially; calyx teeth triangular, ca. 0.5 mm long; standard 17–20 mm long; wings 18–21 mm long; keels 17–20 mm long; pistil glabrous; legume glabrous, with a stipe 10–11 mm long (Table 1).

Table 1.

Morphological comparison of Astragalus zibaishanensis, A. monadelphus, A. xitaibaicus and A. neomonodelphus.

	A. zibaishanensis	A. monadelphus	A. xitaibaicus	A. neomonodelphus
Stem diameter	0.9–1.5 mm	2–4 mm	2–4 mm	2–3 mm
Leaflets	5–7; hairy adaxially and abaxially	9–17; glabrous adaxially, hairy abaxially	11–15; glabrous adaxially, glabrous or hairy abaxially	9–15; glabrous adaxially and abaxially
Calyx teeth	triangular, ca. 0.5 mm long	subulate, 2.0–4.0 mm long	subulate, 2.5–3.0 mm long	two adaxial teeth triangular, 1.0–1.5 mm long, three abaxial teeth narrowly triangular, 1.5–3.0 mm long
Standard	17–20 mm long	10–13 mm long	ca. 11 mm long	16–17 mm long
Wings	18–21 mm long	10–12 mm long	ca. 11 mm long	15–16 mm long
Keels	17–20 mm long	10–11 mm long	ca. 10 mm long	15–16 mm long
Pistil	glabrous	hairy	hairy	hairy
Legume	glabrous; stipe 10–11 mm long	hairy; stipe 5–6 mm long	hairy; stipe 5–6 mm long	hairy; stipe 7–8 mm long

10.3897/phytokeys.266.170977.figure3

17669036

05E6A853-250F-552C-B6B1-2BC316A09DC0

Figure 3.

Photos of Astragalus zibaishanensis from the field. A. Habitat; B. Upper part of a flowering plant; C. Raceme; D. A dissected flower (s, standard; w, wings; k, keels; a, androecium); E. Young legumes; F. Mature legumes; G. Seeds. Photoed by Lu-Lu Xun.

https://binary.pensoft.net/fig/1469681

Description.

Perennial herbs, 20–30 cm tall; main root 6–10 mm in diameter. Stems cespitose, ascending, unbranched, slightly zigzag, slender, 0.9–1.5 mm in diameter; internodes sparsely pubescent or glabrous. Leaves imparipinnate, alternate, 7–11 cm long; stipules free, often reflexed, lanceolate or ovate, 6–9 × 1–3 mm, brown or green, glabrous; rachises sparsely pubescent or glabrous; leaflets 5–7, opposite; petiolules 1.0–1.5 mm long, pubescent; leaflet blades ovate or oblong, 15–30 × 7–15 mm, with white appressed simple hairs adaxially and abaxially, base wide cuneate or rounded, apex retuse or obtuse. Peduncles axillary, 3–8 cm long, sparsely pubescent or glabrous; racemes with 9–15 flowers; pedicels 4–5 mm long, sparsely pubescent; bracts lanceolate, 7–9 × 1.5–2.5 mm, glabrous. Calyx 6–7 mm long, inner side of calyx teeth with brown hairs, other parts of calyx glabrous; calyx teeth 5, triangular, equal, ca. 0.5 mm long. Petals light yellow; standard obovate, 17–20 × 7–8 mm, apex retuse or rounded, base attenuate; wings 18–21 × 1.8–2.0 mm, auricle narrowly triangular, ca. 1 mm long, claw 11–14 mm long; keels 17–20 × 2.5–3.0 mm, auricle triangular, ca. 0.8 mm long; androecium pseudomonadelphous, 18–21 mm long, the vexillary stamen (the single stamen which is close to the standard) is attached to other nine fused filaments at middle, and free from the fused filaments at base and apex; pistil 17–21 mm long, glabrous, stipe 7–10 mm long, style 4–5 mm long. Legume glabrous, with a stipe 10–11 mm long, inflated, unilocular, outline lanceolate, 18–22 mm long (excluding stipe), 5–6 mm wide. Seeds black-brown, 3.5–4.0 × 2.0–2.5 mm.

Phenology.

Flowering in May and June; fruiting in June and July; seeds mature in July.

Distribution and habitat.

Astragalus zibaishanensis is distributed on the north slope of Mt. Zibaishan in Fengxian County, Shaanxi, China (Fig. 4). It grows on stony slope in forest, 2300–2400 m a. s. l.

10.3897/phytokeys.266.170977.figure4

17669038

A855E8C3-9435-5B86-809C-D4EE9A012F87

Figure 4.

Distributions of Astragalus zibaishanensis (red dot), A. monadelphus (green dots), A. xitaibaicus (black dots) and A. neomonodelphus (blue dot). Maps from map.tianditu.gov.cn

https://binary.pensoft.net/fig/1469682

Etymology.

The epithet zibaishanensis derives from the type locality of the new species, Mt. Zibaishan (紫柏山). The Chinese vernacular name for this new species is 紫柏山黄耆 (zǐ bǎi shān huáng qí).

Other specimens checked (Paratypes).

China • Shaanxi Province, Fengxian County, north slope of Mt. Zibaishan, 33°41'47.0"N, 106°45'18.7"E, 2380 m a. s. l., 24 May 2022, P. L. Liu 1454 (WUK!, WNU!); • same locality, 33°41'43.0"N, 106°45'10.3"E, 2391 m a. s. l., 24 May 2022, P. L. Liu 1455 (WUK!); • same locality, 33°41'47.0"N, 106°45'18.3"E, 2385 m a. s. l., 16 July 2025, P. L. Liu 1757 (WUK!).

Citations of comparative species.

Astragalus monadelphus Bunge ex Maxim., Bull. Acad. Imp. Sci. Saint-Pétersbourg 24: 32. 1878. Lectotype (Podlech and Sytin 1996): West China, Prov. Kansu, vii 1872, Prezewalski (LE, sheet marked as lectotypus; drawing and fragments at K!); isolectotypes: LE, P, PE!.

Astragalus xitaibaicus (K. T. Fu) Podlech & L. R. Xu, Novon 14 (2): 222. 2004. Holotype: Shaanxi, Xitaibai Shan, Dingpeng Shan, alt. 3300 m, in grassland on mountain ridge, 9 August 1957, Kun-Tsun Fu 10321 (WUK!); isotype: WUK!.

Astragalus neomonodelphus H. T. Tsai & T. T. Yu, Bull. Fan Mem. Inst. Biol. Bot. 9: 263. 1940. Holotype: Yunnan, Chungtien, Sianrentung, alt. 3400 m, grassy slope, herb, fl. yellow, 12 July 1937, T. T. Yü 12120 (KUN); isotypes: PE!, HUH!.

Discussion

Astragalus zibaishanensis is unique in Astragalus by having pseudomonadelphous androecium. The vexillary stamen is attached to other nine fused filaments in the middle, and free from the fused filaments at base and apex (Fig. 2j). In contrast, a monadelphous androecium is characterized by the completely joined filaments forming a tube of ten stamens (Rodríguez-Riaño et al. 1999). In Fabaceae, monadelphous androecium is found in tribe Genisteae, the genus Ononis L., and Galega officinalis L., while pseudomonadelphous androecium is found in the genera Coronilla L., Lathyrus L., Medicago L., Vicia L., Pisum L., Trifolium L., Robinia L. and others (Rodríguez-Riaño et al. 1999; Sinjushin et al. 2022). In Astragalus, most species have diadelphous androecium, and only A. monadelphus, A. xitaibaicus, A. neomonodelphus and A. changmuicus C. C. Ni & P. C. Li had been recorded to have monadelphous androecium (Li and Ni 1979; Xu and Podlech 2010; Podlech and Zarre 2013). However, careful observation showed that A. monadelphus has pseudomonadelphous androecium (Liu et al. 2010), which is similar to that of A. zibaishanensis. Considering the close phylogenetic relationship of A. monadelphus, A. xitaibaicus and A. neomonodelphus (Fig. 1A), further observations of the androecium type of A. xitaibaicus and A. neomonodelphus are needed.

In morphology, A. zibaishanensis can be distinguished from A. monadelphus, A. xitaibaicus (Fig. 5) and A. neomonodelphus by the characters listed in Table 1. Besides, these four species are allopatric in geographic distribution. Astragalus monadelphus is widely distributed in Gansu, Qinghai and Sichuan Provinces; A. neomonodelphus is found in northwestern Yunnan Province; A. xitaibaicus is found in Taibai and Yangxian Counties in Shaanxi Province; A. zibaishanensis is only known from Mt. Zibaishan in Shaanxi Province at present (Fig. 4).

10.3897/phytokeys.266.170977.figure5

17669041

C9E74AEA-3E6D-56A6-9313-8251CFD1AAEF

Figure 5.

Photos of Astragalus monadelphus (A–C) and A. xitaibaicus (D–F) from the field. A, D. Upper part of a flowering plant; B, E. Raceme; C, F. Legumes. A–C photoed by Lang-Ran Xu; D photoed by Lu-Lu Xun; E photoed by Yuan Lu; F photoed by Jian-Quan Tang.

https://binary.pensoft.net/fig/1469683

In the monograph of Old World Astragalus, Podlech and Zarre (2013) offered a practical key to 41 species in A. sect. Cenantrum (pages 171–174). We could insert the 42^nd species, A. zibaishanensis, to the key, and modify the key as follows (begins with the original numbering).

Modified key to the species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Astragalus">Astragalus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="infraspecific-rank">sect.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="section" reg="Cenantrum">Cenantrum</tp:taxon-name-part></tp:taxon-name> in Old World

4	Leaflets 3–5; inflorescence paniculate	A. henryi
–	Leaflets at least 5; inflorescence racemose	5
5	Leaflets (11–) 13–25; bracts 2–6 mm	6 (original contents, omitted)
–	Leaflets 5–13; bracts 5–20 mm	10
10	Leaflets hairy adaxially and abaxially; androecium pseudomonadelphous	A. zibaishanensis
–	Leaflets glabrous adaxially, hairy abaxially; androecium diadelphous	species in the original key numbering 10 to 13 (omitted)

Acknowledgements

We thank Mr. Jian-Quan Tang for his help with fieldwork. We are grateful to the curators of WNU, WUK and XBGH for their kind help in checking the specimens. We thank Mr. Yi-Fan Li for preparing the illustration.

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 research was funded by the Basic Research Program of the Shaanxi Academy of Sciences (grant number 2023K-14).

Author contributions

Conceptualization: PLL, MY. Data curation: PLL, LLX, CS. Funding acquisition: BL, MY. Investigation: PLL, LLX, YL. Methodology: PLL, CS. Project administration: MY. Resources: LLX, BL, MY. Supervision: MY. Visualization: PLL, LLX. Writing - original draft: PLL, LLX. Writing - review and editing: CS, YL, BL, MY.

Author ORCIDs

Pei-Liang Liu https://orcid.org/0000-0001-6566-7824

Lu-Lu Xun https://orcid.org/0000-0002-6861-5883

Yuan Lu https://orcid.org/0000-0002-0329-0803

Bin Li https://orcid.org/0000-0001-9254-8003

Ming Yue https://orcid.org/0000-0002-8618-1126

Data availability

The newly generated DNA sequences have been deposited in GenBase and the accession numbers can be found in Appendix 1.

References

Azani

Bruneau

Wojciechowski

Zarre

(2017) Molecular phylogenetics of annual

Astragalus

(

Fabaceae

) and its systematic implications.Botanical Journal of the Linnean Society184(3): 347–365. https://doi.org/10.1093/botlinnean/box032

Azani

Bruneau

Wojciechowski

Zarre

(2019) Miocene climate change as a driving force for multiple origins of annual species in

Astragalus

(

Fabaceae

Papilionoideae

).Molecular Phylogenetics and Evolution137: 210–221. https://doi.org/10.1016/j.ympev.2019.05.008

Zheng

Zhao

Bai

Jia

Chen

Hao

Xiao

Zhang

Zhao

Tang

Bao

(2024) GenBase: A Nucleotide Sequence Database. Genomics, Proteomics & Bioinformatics 22(3): qzae047. https://doi.org/10.1093/gpbjnl/qzae047

Buono

Kadereit

Liston

Zarre

Morales‐Briones

(2025) Building a robust backbone for

Astragalus

using a clade‐specific target enrichment bait set. American Journal of Botany 112(8): e70084. https://doi.org/10.1002/ajb2.70084

Chai

Yan

(2010)

Astragalus

beitashanensis, a New Species of

Leguminosae

from Xinjiang, China.Novon20: 21–22. https://doi.org/10.3417/2008017

CNCB-NGDC Members and Partners

(2025) Database Resources of the National Genomics Data Center, China National Center for Bioinformation in 2025. Nucleic Acids Research 53(D1): D30–D44. https://doi.org/10.1093/nar/gkae978

Darriba

Taboada

Doallo

Posada

(2012) jModelTest 2: More models, new heuristics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109

Duan

Wen

Y-W

Jiang

Zhang

Charboneau

JLM

Q-F

Wang

F-G

T-Y

S-J

Jiang

K-W

Fang

Y-H

Chen

H-F

(2024) New insights into the phylogenetic relationships of tribe Astragaleae (

Fabaceae

subfamily

Papilionoideae

) and

Astragalus

—The largest genus of angiosperm.Biological Diversity1(3–4): 136–146. https://doi.org/10.1002/bod2.12021

Folk

Charboneau

JLM

Belitz

Singh

Kates

Soltis

Guralnick

Siniscalchi

(2024) Anatomy of a mega‐radiation: Biogeography and niche evolution in

Astragalus

. American Journal of Botany 111(3): e16299. https://doi.org/10.1002/ajb2.16299

F-B

Xin

Guo

Zhao

L-Q

(2021) Seven newly recorded species from Xizang and Xinjiang in China.Shengwu Duoyangxing29(9): 1265–1270. https://doi.org/10.17520/biods.2021181

Isely

(1998) Native and Naturalized

Leguminosae

(

Fabaceae

) of the United States: (exclusive of Alaska and Hawaii). Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah, 213.

Kazempour Osaloo

Maassoumi

Murakami

(2003) Molecular systematics of the genus

Astragalus

L. (

Fabaceae

): Phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacers and chloroplast gene ndhF sequences.Plant Systematics and Evolution24: 1–32. https://doi.org/10.1007/s00606-003-0014-1

Z-W

Gan

Q-L

(2024)

Astragalus

duheyuanensis

(

Fabaceae

), a new species from central China.Annales Botanici Fennici61(1): 1–5. https://doi.org/10.5735/085.061.0101

P-C

C-C

(1979) New taxa of the genus

Astragalus

L. from Tibet.Zhiwu Fenlei Xuebao17(2): 104–112.

J-X

X-L

(2014)

Astragalus

wulingensis

(

Leguminosae

), a new species from Hunan China.Phytotaxa159(4): 279–286. https://doi.org/10.11646/phytotaxa.159.4.4

Liu

X-L

Liu

P-L

Chang

Z-Y

L-R

(2010) Pseudomonadelphus Characters of

Astragalus

monadelphus

Maxim. and its taxonomical Significances.Xibei Zhiwu Xuebao30(9): 1834–1836.

Liu

X-L

Yue

X-N

Chang

Z-Y

L-R

(2011) A New Species of

Astragalus

(

Leguminosae

) from Northwestern Xinjiang, China.Novon21(2): 216–218. https://doi.org/10.3417/2009083

Liu

P-L

Wen

Duan

Arslan

Ertuğrul

Chang

Z-Y

(2017)

Hedysarum

L. (

Fabaceae

: Hedysareae) is not monophyletic – Evidence from phylogenetic analyses based on five nuclear and five plastid sequences. PLoS One 12(1): e0170596. https://doi.org/10.1371/journal.pone.0170596

Liu

Z-Y

Shi

W-G

(2020)

Astragalus

shaerqinensis

(

Fabaceae

), A New Species from Inner Mongolia, China.Annales Botanici Fennici57(1–3): 181–183. https://doi.org/10.5735/085.057.0125

Moonlight

Baldaszti

Cardoso

Elliott

Särkinen

Knapp

(2024) Twenty years of big plant genera. Proceedings. Biological Sciences 291: 20240702. https://doi.org/10.1098/rspb.2024.0702

Podlech

(2010) Four new

Astragalus

species (

Leguminosae

) from China and Bhutan. Pakistan Journal of Botany 42 (SI): 27–34.

Podlech

Sytin

(1996) Typification of Russian and some other species of

Astragalus

I.Sendtnera3: 149–176.

Podlech

Zarre

(2013) A Taxonomic revision of the genus

Astragalus

L. (

Leguminosae

) in the Old World. Naturhistorisches Museum Wien, Vienna, 170–212.

Zhao

L-Q

(2018)

Astragalus

nuoergongensis

(

Fabaceae

), a new species from Inner Mongolia and Ningxia, China.Annales Botanici Fennici55(4–6): 405–407. https://doi.org/10.5735/085.055.0421

Rodríguez-Riaño

Ortega-Olivencia

Devesa

(1999) Types of Androecium in the

Fabaceae

of SW Europe.Annals of Botany83(2): 109–116. https://doi.org/10.1006/anbo.1998.0808

Ronquist

Huelsenbeck

(2003) MrBayes 3: Bayesian phylogenetic inference under mixed models.Bioinformatics (Oxford, England)19(12): 1572–1574. https://doi.org/10.1093/bioinformatics/btg180

Ronquist

Teslenko

van der Mark

Ayres

Darling

Höhna

Larget

Liu

Suchard

Huelsenbeck

(2012) MrBayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice Across a Large Model Space.Systematic Biology61(3): 539–542. https://doi.org/10.1093/sysbio/sys029

Sang

Crawford

Stuessy

(1997) Chloroplast DNA phylogeny, reticulate evolution, and biogeography of

Paeonia

(

Paeoniaceae

).American Journal of Botany84(8): 1120–1136. https://doi.org/10.2307/2446155

Sinjushin

Ploshinskaya

Maassoumi

Mahmoodi

Bagheri

(2022) Variations in Structure among Androecia and Floral Nectaries in the Inverted Repeat-Lacking Clade (

Leguminosae

Papilionoideae

).Plants11(5): 649. https://doi.org/10.3390/plants11050649

Stamatakis

(2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies.Bioinformatics (Oxford, England)30(9): 1312–1313. https://doi.org/10.1093/bioinformatics/btu033

Duan

Liu

P-L

Liu

Chang

Z-Y

Wen

(2021a) Chloroplast phylogenomics and character evolution of eastern Asian

Astragalus

(

Leguminosae

): Tackling the phylogenetic structure of the largest genus of flowering plants in Asia. Molecular Phylogenetics and Evolution 156: 107025. https://doi.org/10.1016/j.ympev.2020.107025

Rather

Wang

Chang

Z-Y

(2021b)

Astragalus

verus

(

Fabaceae

)—A New Record for the Flora of China.Shokubutsu Kenkyu Zasshi96(2): 119–121. https://doi.org/10.51033/jjapbot.96_2_11086

Swofford

(2002) PAUP* Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland.

Taberlet

Gielly

Pautou

Bouvet

(1991) Universal primers for amplification of three non-coding regions of chloroplast DNA.Plant Molecular Biology17: 1105–1109. https://doi.org/10.1007/BF00037152

White

Bruns

Lee

Taylor

(1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis

Gelfand

Sninsky

White

(Eds) PCR protocols: A guide to methods and applications.Academic Press, San Diego, 315–322.

Wojciechowski

(2005)

Astragalus

(

Fabaceae

): A molecular phylogenetic perspective. Brittonia 57(4): 382–396. https://doi.org/10.1663/0007-196X(2005)057[0382:AFAMPP]2.0.CO;2

Wojciechowski

Sanderson

J-M

(1999) Evidence on the Monophyly of

Astragalus

(

Fabaceae

) and its Major Subgroups Based on Nuclear Ribosomal DNA ITS and Chloroplast DNA trnL Intron Data.Systematic Botany24(3): 409–437. https://doi.org/10.2307/2419698

Y-H

(2015) Flora Kunlunica, Vol. 2.Chongqing Press, Chongqing, 771 pp.

Y-H

(2025) Qinghai Plant Species Areal-Types System.Qinghai People’s Publishing House, Xining, 580 pp.

L-R

Podlech

(2010)

Astragalus

L. In: Wu

Raven

Hong

(Eds) Flora of China, Vol.10. Science Press, Beijing, Missouri Botanical Garden Press, St. Louis, 328–453.

S-Z

Gan

Q-L

X-W

(2023)

Astragalus

bashanensis

(

Leguminosae

), a new species from Central China.PhytoKeys219: 49–55. https://doi.org/10.3897/phytokeys.219.96916

Yang

Z-Z

Liu

Q-R

Liu

Xiang

J-S

X-L

(2024)

Astragalus

liuaiminii

, a new species of

Astragalus

(

Fabaceae

) from Xinjiang, China.PhytoKeys243: 209–214. https://doi.org/10.3897/phytokeys.243.119707

Yang

Wang

Wei

Liang

(2025)

Astragalus

helanshanensis

, a new plant species from the arid region of the Helan Mountains in northwestern China.Phytotaxa690(1): 99–106. https://doi.org/10.11646/phytotaxa.690.1.8

Zhang

Ren

S-S

Yan

(2021)

Astragalus

taledensis

, a new species of

Leguminosae

from Xinjiang, China.Phytotaxa524(3): 199–204. https://doi.org/10.11646/phytotaxa.524.3.4

Zhao

(2012) Classification and its Florristic Ecological Geographic Distributions of Vascular Plants in Inner Mongolia. Inner Mongolia University Press, Hohhot, 856 pages.

Appendix 1

Taxon name, geographical locality, voucher, herbarium code, and GenBase (https://ngdc.cncb.ac.cn/genbase) accession number for the newly generated sequences (indicated by an asterisk *). For each sample, accession numbers are given for the ITS, psbA-trnH, trnL-F sequences. A dash (–) indicates a missing sequence. Sequences acquired from the GenBank (https://www.ncbi.nlm.nih.gov/) are listed with taxon names and accession numbers.

Astragalus acaulis Baker: SAMN15406160;

Astragalus aksuensis Bunge: SAMN15406183;

Astragalus alopecurus Pallas: SAMN15406163 ;

Astragalus americanus (Hook.) M. E. Jones: SAMN15406142;

Astragalus annularis Forssk.: SAMN15406164 ;

Astragalus arnoldianus N. D. Simpson: China, Sichuan, Luding, P. L. Liu 1355 (WNU), C_AA122198*, C_AA121464*, C_AA121450*;

Astragalus chilienshanensis Y. C. Ho: China, Qinghai, Ledu, L. R. Xu & G. W Sheng 2009-003 (WUK), C_AA122199*, C_AA121465*, C_AA121451*;

Astragalus craibianus N. D. Simpson: SAMN15406173;

Astragalus discolor Bunge: SAMN15406200;

Astragalus ellipsoideus Ledeb.: SAMN15406192;

Astragalus floridulus Podlech: MZ198526, NC_083391;

Astragalus frigidus (L.) A. Gray: China, Shanxi, Wuzhai, P. L. Liu 1462-1 (WNU), C_AA122200*, C_AA121466*, C_AA121452*;

Astragalus glycyphyllus L.: SAMN15406130;

Astragalus graveolens Buch.-Ham. ex Benth.: SAMN15406162;

Astragalus henryi Oliv.: SAMN15406197;

Astragalus hoantchy Franch.: SAMN15406159;

Astragalus hoffmeisteri (Klotzsch) Ali: SAMN15406172;

Astragalus lepsensis Bunge: China, Xinjiang, Gongnaisi, L. R. Xu 1435 (WUK), –, C_AA121467*, C_AA121453*;

Astragalus lepsensis Bunge: AF359752;

Astragalus membranaceus Bunge: China, Shaanxi, Chang’an, B. Li LB24406 (XBGH), C_AA122201*, C_AA121468*, C_AA121454*;

Astragalus moellendorffii Bunge: China, Ningxia, Liupanshan, L. R. Xu 96-204 (WUK), C_AA122202*, C_AA121469*, C_AA121455*;

Astragalus monadelphus Bunge ex Maxim.: China, Qinghai, Ledu, L. R. Xu & G. W Sheng 2009-002 (WUK), –, C_AA121470*, –;

Astragalus monadelphus Bunge ex Maxim.: China, Gansu, Yuzhong, Z. Y. Chang et al. 2007021 (WUK), C_AA122203*, C_AA121471*, C_AA121456*;

Astragalus monadelphus Bunge ex Maxim., MH808432;

Astragalus muliensis Hand.-Mazz.: MZ198530, NC_083909;

Astragalus nakaianus Y. N. Lee: KC262198, NC_028171;

Astragalus neomonodelphus H. T. Tsai & T. T. Yu, SAMN15406153;

Astragalus oplites Benth. ex R. Parker: SAMN15406189;

Astragalus przewalskii Bunge: China, Qinghai, Xunhua, Z. Y. Chang et al. 2010103 (WUK), C_AA122205*, C_AA121473*, C_AA121458*;

Astragalus przewalskii Bunge: China, Gansu, Yuzhong, Z. Y. Chang et al. 2007020 (WUK), C_AA122204*, C_AA121472*, C_AA121457*;

Astragalus scaberrimus Bunge: SAMN15406155;

Astragalus shinanensis Ohwi: LC760273, LC760283, –;

Astragalus sinicus L.: SAMN15406198;

Astragalus skythropos Bunge: SAMN15406208;

Astragalus stipulatus D. Don: MZ198531, NC_083911;

Astragalus strictus Graham ex Benth.: SAMN15406213;

Astragalus tenuis Turcz.: SAMN15406203;

Astragalus tongolensis Ulbr.: China, Xizang, Jiacha, P. L. Liu 1531 (WNU), C_AA122206*, C_AA121474*, C_AA121459*;

Astragalus tumbatsicus C. Marquand & Airy Shaw: MZ198541, NC_083912;

Astragalus uliginosus L.: SAMN15406204;

Astragalus umbellatus Bunge: AF121683 & MG236337, –, AF126988;

Astragalus verus Oliv.: SAMN15406193;

Astragalus xitaibaicus (K. T. Fu) Podlech & L. R. Xu: China, Shaanxi, Yangxian, P. L. Liu 1651 (WNU), C_AA122207*, C_AA121475*, C_AA121460*;

Astragalus zibaishanensis P. L. Liu & L. L. Xun: China, Shaanxi, Fengxian, P. L. Liu 1454 (WNU), C_AA122208*, C_AA121476*, C_AA121461*;

Astragalus zibaishanensis P. L. Liu & L. L. Xun: China, Shaanxi, Fengxian, P. L. Liu 1455 (WNU), C_AA122209*, C_AA121477*, C_AA121462*;

Astragalus zibaishanensis P. L. Liu & L. L. Xun: China, Shaanxi, Fengxian, P. L. Liu 1458 (WNU), C_AA122210*, C_AA121478*, C_AA121463*;

Oxytropis bicolor Bunge: SAMN15406215.