Research Article
Research Article
A new species of Hiptage (Malpighiaceae) from northwest Yunnan (China) based on molecular and morphological data
expand article infoTong-Tong Zhang, Shu-Yun Yang, Ke Tan§, Ming-Xun Ren
‡ Hainan University, Haikou, China
§ Guangxi Institute of Botany, Guilin, China
Open Access


Hiptage stenopterum K.Tan & M.X.Ren, a new species of Hiptage collected from a deep valley close to the Nujiang Gorge, northwest of Yunnan Province, China, is described and illustrated based on molecular and morphological data. The new species was found isolated in an entrenched valley of the Laowo River, a tributary of the Nujiang River, at the northern edge of the distribution range of the genus. H. stenopterum shares some morphological similarities with the narrowly endemic H. incurvatum and H. lushuiensis. However, H. stenopterum is easily distinguished by its oblanceolate lateral wing of winged mericarp, 10 to 12 calyx glands, and branchlets densely rusty tomentose. The new species status is also supported by molecular phylogenetic analyses based on nuclear ribosome internal transcribed spacer (nrITS), which showed distinct systematic boundaries from the most morphologically similar species, H. incurvatum and their morphological relatives, H. lushuiensis.

Key words

flora of Yunnan, Hengduan Mountains, Hiptage, Malpighiaceae, taxonomy


Hiptage Gaertn. (Gaertner 1791) is a large genus of Malpighiaceae, currently comprising ca. 40 species of woody shrubs or lianas growing at shrub forests or valleys on limestone hills or riverbanks of tropical Asia, such as Indo-China Peninsular, Malay Archipelago and Southern China (Sirirugsa 1991; Hô 1992; Srivastava 1992; Chen and Funston 2008; Ren 2015; Yang et al. 2018; Tan et al. 2019; Dong et al. 2020; Wei et al. 2022). Hiptage is derived from the Greek hiptamai, which means “to fly” and refers to its unique three-winged fruit. Such three-winged fruit means it is easily dispersed over long distances, explaining its widespread distribution in tropical Asia (Sirirugsa 1991; Srivastava 1992; Hô 1992; Chen and Funston 2008). Nonetheless, most species of this genus are narrow and endangered endemics, especially in the ravine and mountain peaks, such as H. multiflora F.N.Wei, H. lushuiensis S.P.Dong, K.Tan & M.X.Ren, and H. incurvatum K. Tan & M.X. Ren (Wei 2018; Tan et al. 2019; Dong et al. 2020). In recent years, five new species of Hiptage have been described. One species was found in Southern India (Sujana and Vadhyar 2020), and four were collected in Southwestern China (Yang et al. 2018; Tan et al. 2019; Dong et al. 2020).

Hiptage is currently classified within the Tetrapteroid clade, which is among the ten primary lineages identified for Malpighiaceae (Davis and Anderson 2010). Within that phylogenetic framework, Hiptage included six species and was resolved as closely related to Flabellariopsis R.Wilczek, a genus endemic to Africa. Hiptage and Flabellariopsis comprise a clade with weak support, sister to the Neotropical genus Carolus W.R.Anderson (Davis and Anderson 2010). Despite relying on a single marker (ITS) and yielding mainly weakly supported clades, recent molecular phylogenies of Hiptage indicate that H. stellulifera Arènes stands as the basal group within the genus (Tan et al. 2019; Almeida and van den Berg 2022).

During our fieldwork in the Hengduan Mountains, northwest of Yunnan Province, China, we collected specimens from a population of a distinct morphotype of Hiptage growing on a hillside in the valley of Laowo River, close to the Nujiang Gorge. After detailed analyses of morphological characters and molecular data of individuals from this population, we found out that this population is most morphologically similar to H. incurvatum and H. lushuiensis. But it differs in the number of calyx glands and the morphology of mericarp wings, both of which are critical morphological traits for species taxonomy in Hiptage (Chen and Funston 2008; Ren 2015). Therefore, we describe it as a new species, depicted and illustrated here.

Materials and methods


Measurements and observations of morphological characters of the new species were based on fresh and herbarium specimens. Each character was measured across five individuals. Photographs of fresh plants and floral characteristics were taken in the field. The new species was compared with all type specimens of Hiptage in morphology from protologues and specimens in the herbarium (at the herbaria IBK, KUN, and IBSC). Images of type specimens of all Hiptage species were consulted from JSTOR Global Plants (, the Chinese Virtual Herbarium (, and taxonomic literature (e.g., Yang et al. 2018; Tan et al. 2019; Dong et al. 2020). The morphological terminology follows Niedenzu (1924), Jacobs (1955), Anderson et al. (2006), Chen and Funston (2008), Ren (2015), and Almeida and Morais (2022). The conservation status assessments of the new species H. stenopterum were based on the International Union for Conservation of Nature guidelines (IUCN 2022).

Phylogenetic analysis

To verify the taxonomic status of the new species within Hiptage, we conducted a phylogenetic analysis using the nuclear ribosomal internal transcribed spacer (ITS) region. We downloaded ITS sequence data for 42 samples from GenBank (Table 1) encompassing 16 species of Hiptage and, based on the study of Almeida and van den Berg (2022), four species from the Tetrapteroid clade serving as outgroups [i.e. Callaeum psilophyllum (A.Juss.) D.M.Johnson, Heteropterys brunnea R.Sebast. & Mamede, Niedenzuella multiglandulosa (A.Juss.) W.R.Anderson and N. stannea (Griseb.) W.R.Anderson] (Table 1). Dried leaf material of the proposed new species was collected from the type locality in a valley of the Laowo River, close to Lushui City (Yunnan, China). Five samples from the new species and three samples from H. lushuiensis were newly sequenced in this study to provide molecular evidence (Table 1). Total genomic DNA was isolated from dried leaf samples using a CTAB protocol adapted from Doyle and Doyle (1987). PCR amplification, in 25 μl reactions, comprised 1 μl of sample DNA, 12.5 μl of 2× Taq PCR Master Mix (Aidlab Biotechnologies Co. Ltd), 1μl of each primer (10 μmol/ml), with double distilled water making up the remainder. Amplification of the ITS region utilized primers ITS17SE and ITS26SE as per Sun et al. (1994). The amplification regime involved a 5 min initial denaturation at 94 °C, then 35 cycles of 40s at 94 °C, 20s at 69 °C, 1 min at 72 °C, and concluded with a 10 min extension at 72 °C. The resulting PCR products were bidirectionally sequenced on an ABI3730XL platform.

Table 1.

Taxa, GenBank accession numbers, voucher numbers of Hiptage used in this study.

Species Locality GenBank Accession Voucher Number
Hiptage benghalensis (L.) Kurz Yangjie, Yunnan, China MH718400 M. X. Ren & L. Tang 128 (HUTB)
Menglian County, Yunnan, China MH718422 S. P. Dong 131 (HUTB)
Daxin County, Guangxi, China MH718414 K. Tan & S. P. Dong 95 (HUTB)
Lekang County, Guizhou, China MH718415 K. Tan, S. P. Dong, & M. X. Ren 82 (HUTB)
Singapore MH718399 T. W. Yam 3334 (HUTB)
Phatthaya, Thailand MH718408 K. Tan, S. P. Dong, & M. X. Ren 3344 (HUTB)
Chiangmai, Thailand MH718410 K. Tan, S. P. Dong, & M. X. Ren 3336 (HUTB)
H. multiflora F.N.Wei Nonggang National Nature Reserve, Guangxi, China MH718424 K. Tan & S. P. Dong 52 (HUTB)
H. stenopterum K. Tan & M.X. Ren Lushui City, Yunnan, China OQ968812 tank 190402001 (HUTB)
OR417356 23tk041401(IBK)
OR417357 23tk041402(IBK)
OR417358 23tk041403(IBK)
H. incurvatum K. Tan & M.X. Ren Pingpo Town, Yunnan, China MK967956 K. Tan, H. L. Zheng, & M. X. Ren 201903309 (HUTB)
MK967957 K. Tan, H. L. Zheng, & M. X. Ren 201903310 (HUTB)
MK967958 K. Tan, H. L. Zheng, & M. X. Ren 201903305 (HUTB)
MK967959 K. Tan, H. L. Zheng, & M. X. Ren 201903306 (HUTB)
H. monopteryx Sirirugsa Phatthaya, Thailand MH718407 K. Tan, S. P. Dong, & M. X. Ren 3337 (HUTB)
H. marginata Arènes Hue, Vietnam MH718413 K. Tan & Q. Yang 3363 (HUTB)
H. detergens Craib KuiBuri, Thailand MH718404 K. Tan, S. P. Dong, & M. X. Ren 3328 (HUTB)
Sam Roi Yot, Thailand MH718405 K. Tan, S. P. Dong, & M. X. Ren 3326 (HUTB)
H. lucida Pierre Phatthaya, Thailand MH718406 K. Tan, S. P. Dong, & M. X. Ren 38 (HUTB)
Xishuangbanna, Yunnan, China MH718418 Z. N. Qian & S. P. Dong120 (HUTB)
subglabra Arènes Nui Chua National Park, Phan Rang, Vietnam MH718427 K. Tan & S. J. Ling 3364 (HUTB)
H. bullata Craib Lampang, Thailand MH718412 K. Tan, S. P. Dong, & M. X. Ren 3320 (HUTB)
H. minor Dunn Lekang County, Guizhou, China MH718398 K. Tan, S. P. Dong, & M. X. Ren 79 (HUTB)
Wenshan City, Yunnan, China MH718423 K. Tan, S. P. Dong, & M. X. Ren 94 (HUTB)
Lushui City, Yunnan, China MH718401 K. Tan, S. P. Dong, & M. X. Ren 88 (HUTB)
H. ferruginea Y.H.Tan & Bin Yang Xishuangbanna, Yunnan, China MH718402 S. P. Dong 116 (HUTB)
Xishuangbanna, Yunnan, China MH718403 S. P. Dong 117 (HUTB)
H. pauciflora Y.H.Tan & Bin Yang Menglian County, Yunnan, China MH718420 S. P. Dong 73 (HUTB)
H. umbellulifera Arènes Cana, Phan Rang, Vietnam MH718426 K. Tan & S. J. Ling 3386 (HUTB)
Phan Rang, Vietnam MH718430 K. Tan & S. J. Ling 3399 (HUTB)
Nui Chua National Park, Phan Rang, Vietnam MH718428 K. Tan & S. J. Ling 3385 (HUTB)
H. luzonica Merr. Luzon Island, Philippines MH718425 K. Tan, W. Q. Xiang & M. X. Ren 20191181436 (HUTB)
Palawan Island, Philippines MH718432 K. Tan, W. Q. Xiang & M. X. Ren 3305 (HUTB)
Cebu Island, Philippines MH718431 K. Tan & W. Q. Xiang 3301(HUTB)
H. candicans Hook. Chiangmai, Thailand MH718409 K. Tan, S. P. Dong, & M. X. Ren 3328 (HUTB)
Chomthong, Thailand MH718411 K. Tan, S. P. Dong, & M. X. Ren 3330 (HUTB)
H. stellulifera Arènes NhaTrang, Vietnam MH718429 K. Tan & S. J. Ling 3376 (HUTB)
H. lushuiensis S.P.Dong, K.Tan & M.X.Ren Lushui City, Yunnan, China OR471605 S. P. Dong 176 (HUTB)
OR471606 S. P. Dong 177 (HUTB)
OR471607 S. P. Dong 178 (HUTB)
Heteropterys brunnea R.Sebast. & Mamede OK284366 RFAlmeida 579 (HUEFS)
Callaeum psilophyllum (A.Juss.) D.M.Johnson OK268022 RFAlmeida 734 (HUEFS)
Niedenzuella multiglandulosa (A.Juss.) W.R.Anderson OK271417 RFAlmeida 639 (HUEFS)
Niedenzuella stannea (Griseb.) W.R.Anderson OK271412 Pott 1816 (HUEFS)

The sequencing results of ITS fragments were evaluated with PhyDE (Müller et al. 2010) for base confirmation and contiguous sequence editing. All sequences were manually aligned in MEGA v.7 (Kumar et al. 2016). Some erroneous sequencing results were excluded from the alignment range. The Bayesian inference (BI) and Maximum likelihood (ML) analyses were used in Phylosuite (Zhang et al. 2020) to construct the phylogenetic tree of Hiptage. The SYM+G4 model and the K2P+G4 model as the best-fit substitution model for ML and BI analysis, respectively, using ModelFinder (Kalyaanamoorthy et al. 2017) with Corrected Akaike Information Criterion (AICc). Bayesian Inference started with a random tree as a simulated tree, used a Markov chain Monte Carlo (MCMC) to run simulations for 10 million generations and sampled every 1000 generations, and was performed using MrBayes v3.2.5 (Ronquist et al. 2012). The first 2500 trees (25% of the total trees) were discarded as burn-in samples. The maximum likelihood (ML) analyses were accomplished with IQ-TREE v.2.0.6 (Nguyen et al. 2015) with 1000 bootstrap replicates. The final constructed phylogenetic tree was visualized in FigTree v.1.4.3 (


The aligned matrix of ITS sequences consisted of 690 bp, of which 466 sites were identical, 132 (19.1%) were parsimony informative, and 92 parsimony-uninformative variable characters. The phylogenetic analysis showed that Hiptage is a monophyletic group (PP/BS=1/100), with H. stellulifera (PP/BS=1/100) being the first lineage to diverge, consistent with previous studies (Tan et al. 2019; Almeida and van den Berg 2022). The five samples of the proposed new species, H. stenopterum, formed a clade with strong support (PP/BS = 0.79/94) sister to a well-supported subclade alongside H. incurvatum (PP/BS=0.90/91) (Fig. 3). The results of molecular phylogenetics align with morphological observations, with the new species closely resembling H. incurvatum due to the presence of multiple glands. However, H. stenopterum is distinguished by its oblanceolate wings, a more significant number of calyx glands, and branchlets that are densely covered with rusty tomentose. Other features, like pedicels, differentiate the two species, as detailed in Table 2. The new species is also similar to H. lushuiensis due to the elliptic leaf blade and lanceolate bracteoles. However, there are significant discrepancies between the morphological and molecular phylogenetic findings (Fig. 2, Table 2). Especially in phylogenetics, H. lushuiensis was resolved as sister to H. minor (PP/BS = 1/99) (Fig. 2).

Figure 1. 

Hiptage stenopterum A habit B flowering branch C flower in side view D flowers in frontal view E–G calyx glands (E calyx glands with secretions attracting ants) H young branch (showing densely rusty tomentose) I petals J winged mericarp K young leaf. Photos A, B by M.X. Ren, E by T.T. Zhang, and C, D, F–K by K. Tan.

Figure 2. 

Molecular phylogenetic tree of Hiptage based on ITS sequences. Bayesian posterior probability (PP) and ML bootstrap values (BS) are shown above branches as PP/BS (only shown if BS > 50%).

Figure 3. 

Distribution map showcasing the new species, Hiptage stenopterum, alongside the other seven species of the genus found in adjacent regions

Table 2.

Morphological comparison of key characteristics in H. stenopterum, H. incurvatum, H. lushuiensis, and the most widespread H. benghalensis.

Character stenopterum H. incurvatum H. lushuiensis H. benghalensis
Leaf blade elliptic to oblong, 4.0–12.0 × 2.0–5.0 cm, 2 glands elliptic, 6–12 × 5–4.5 cm, 10–16 glands elliptic, 9–16 × 3.5–5.0 cm, 2 or more glands at the base oblong, elliptic-oblong, or ovate-lanceolate, 9–18 × 3–7 cm, 2 glands
Branchlet densely rusty tomentose white to grey sericeous white to grey sericeous densely yellowish brown or silver-gray sericeous
Bracteole lanceolate, ca. 0.5 cm long lanceolate, 0.3–0.5 cm long lanceolate, ca. 1.5 mm long acute, 0.2–0.4 cm long
Pedicel yellow-brown sericeous, 1.5–2.0 cm long white sericeous, 1.5–2.5 cm long white sericeous, ca. 1.5 cm long sericeous, 0.8–2.5 cm long
Calyx gland 10(-12), two large, ca. 3.0 ×1.0 mm, basally coalescent two smaller adnate to the apex of the two big glands separately, 0.4–0.5 mm diam.; remaining glands 1.8–2.0 × 0.8–1.0 mm, oblong; not decurrent to the pedicel 4-(6), two large, ca. 3.0 × 1 mm, basally connate, remaining glands 0.4–0.5 mm diam., round; not decurrent to the pedicel 2, ca. 1 mm diam., free, sometimes with additional smaller glands on the other sepals, not decurrent along the pedicel 1, very large, oblong, lanceolate to oblanceolate; more or less 1/2 adnate onto the pedicel
Sepal elliptic to oblong, ca. 0.5 cm long elliptic to oblong, ca. 0.5 cm long elliptic, 0.4–0.5 cm long broadly elliptic or ovate, 0.5–0.6 cm long
Petal suborbicular, white to light white-pink, 1.0–1.5 × 0.8–1.2 cm suborbicular, white to light white-pink, ca. 1 × 0.8 cm suborbicular, pink, ca. 1 cm long ovate-oblong to suborbicular, white, ca. 1 cm long
Claw 1 mm long, pubescent 1 mm long, pubescent 1 mm long, pubescent 1–2 mm long, glabrous
Posterior lateral wing oblanceolate, ca. 4.2 × 0.8 cm ovate, ca. 3.6 ×1.3 cm ovate, ca. 3.5 × 1.3 cm oblong, elliptic or obovate-lanceolate, 3–5 × 1–1.6 cm
Anterior lateral wing lanceolate to linear, straight, ca. 2.5 × 0.4 cm lanceolate, arcuate back to the middle, ca. 2.3 × 0.7 cm lanceolate, ca. 2.0 × 0.9 cm lanceolate-oblong, 1.5–3 cm long


The zygomorphic, mirror-image, and floral structure of Hiptage represents a distinct evolutionary trait of biological significance that promotes adaptability (Ren et al. 2013; Qian et al. 2016). Currently, the presence, shape, and number of calyx glands, as well as their attachment to the pedicel, are critical diagnostic features for species identification in Hiptage (Sirirugsa 1991; Srivastava 1992; Chen and Funston 2008; Yang et al. 2018; Tan et al. 2019; Almeida and van den Berg 2022). However, Almeida and van den Berg (2022) contend that the presence and number of calyx glands in Hiptage are highly homoplasic and unsuitable for circumscribing infrageneric ranks. Calyx glands in Hiptage might be an evolutionary adaptation to attract myrmecophilous interactions for herbivore defense (Ren et al. 2013; Qian et al. 2016). Therefore, the absence and number of calyx glands may be related to local herbivorous insects, which are a significant factor in promoting speciation (McCall and Irwin 2006; Ramos and Schiestl 2019). Based on long-term field observations, we believe that although there is a slight probability of variation in the number of calyx glands, the overall characteristic remains relatively stable. In summary, we believe that the absence and number of calyx glands still hold significance in the infrageneric classification of Hiptage. However, it should be properly carried out in future phylogenetic studies to establish a new monophyletic infrageneric classification system.

Morphologically, H. stenopterum shares specific characters with H. incurvatum and H. lushuiensis in having elliptic leaf blades, lanceolate bracteoles, white to light pink flowers, and suborbicular petals. Based on the phylogeny tree, H. incurvatum is closely related to the new species, while H. lushuiensis clusters with H. minor, which has non-reflexed petals. Not only do the petals differ in posture, but there are also significant morphological differences between H. lushuiensis and H. minor such as the number of calyx and leaf blade glands, and leaf size (Chen and Funston 2008; Dong et al. 2020). Their clustering together might be due to the limited genetic information from just the ITS region and more comprehensive molecular data will be needed in the future to explain this curious relationship.

Malpighiaceae is characterized by an unparalleled diversity of winged fruits in angiosperms (Anderson et al. 2006; Manchester and O’Leary 2010; Tan et al. 2018). Winged mericarps are an adaption for wind dispersal of fruits (Tan et al. 2018), suggesting that such morphological adaptations have significantly facilitated long-distance dispersal and speciation (Davis et al. 2001, 2002; Tan et al. 2018, 2019). Notably, the wings of H. stenopterum distinctly deviate from its phylogenetically proximate taxa, H. incurvatum. Moreover, these two taxa are partitioned by the Hengduan Mountains, implying that differing aerodynamic conditions, perhaps governed by localized wind patterns, might have driven morphological divergence. Furthermore, the type localities of H. stenopterum and H. incurvatum, encircled by high mountains and deep gorges, constitute a distinct isolated habitat, which restricts gene flow and facilitates speciation.


Hiptage stenopterum K.Tan & M.X.Ren, sp. nov.

Fig. 1


Hiptage stenopterum is most similar to H. incurvatum K.Tan & M.X.Ren by branchlets densely rusty tomentose (vs white to grey sericeous), leaf blades with 2 glands near the base (vs 10–16 glands), 10(-12) calyx glands [vs 4 (-6)], the posterior lateral wing oblanceolate (vs ovate), anterior lateral wings straight, lanceolate to linear (vs arcuate back to the middle, lanceolate).


China. Yunnan Province: Lushui City, Laowo River, 25°50′08″N, 98°54′28″E, 1071 m alt., 2 Apr. 2019, K. Tan 190402001 (Holotype: HUTB!; Isotype: HUTB!, IBK00450922!).


Woody shrubs; young branches densely rusty tomentose, hairs adpressed, older twigs glabrous, with white or greenish lenticels, rounded, coarse warts dotted. Leaves opposite; stipules absent; petiole ca. 0.5 cm long, round, tomentose, with yellowish brown hairs, eglandular; leaf blades 4.0–12.0 × 2.0–5.0 cm, coriaceous, elliptic to oblong; young leaves densely rusty tomentose on both surfaces; mature leaves green, glabrous, base obtuse or broadly cuneate, margin plane, apex acuminate, abaxially often with 2 marginal glands near the base; lateral veins in 5–8 pairs, both surfaces prominently. Thyrses, terminal or axillary; main axis 4.0–13.0 cm long, rusty tomentose; peduncle 1.0–2.5 cm, rusty sericeous; bracteoles ca. 0.5 cm long, lanceolate. Flowers white to slightly pink; pedicels 1.5–2.0 cm long, densely rusty tomentose; sepals 5, 4.5–5.5 × 1.5–2.5 mm, elliptic to oblong, apex obtuse, margin slightly revolute, abaxially densely rusty tomentose, adaxially glabrous. Calyx glands 10(–12), prominent, not decurrent to the pedicel, often 1 pair of glands at base abaxially; two large, 2.3–2.8 × 0.8–1.2 mm, connate at the base, two smaller glands, 0.4–0.5 diam., rounded, adnate to the apex of the two larger glands separately; remaining glands small and free, 0.5–0.6 × 0.8–1.0 mm, oblong, attached to the margins of other sepals, occasionally one or two glands lacking. Petals 5, 1.0–1.5 × 0.8–1.2 cm, white to light white to pink, basally yellow, extremely reflexed, suborbicular, margin ciliate, claws ca. 1 mm long, base subcordate to rounded, apex roundish, abaxially densely white tomentose, adaxially glabrous. Stamens 10, basally fused or free, glabrous, differing in size, pollen sacs rimose; one larger, filament 10–12 mm long, yellowish green, circinate, anther oblong, 1.8–2.0 × 0.7–1.0 mm; 9 smaller stamens, filament 4–6 mm long; anthers oblong, 1–1.3 × 0.7–0.9 mm. Ovary ca. 2 mm in diam., ovoid, white to rusty tomentose; style 1, yellowish green, 10–13 mm long, slightly curved upwards, deflected either to the left or right side, glabrous; stigma apical. Mericarps 3, wings pink with yellow-green base, rusty sericeous, posterior lateral wing 3.8–4.5 × 0.7–0.9 cm, oblanceolate, apex roundish or lobed slightly, base obtuse, anterior lateral wings 2.4–3.0 × 0.3–0.6 cm, lanceolate to linear; areole ca. 4–6 mm, approximately triangular. Seeds angular–globose, 3–5 mm, dark yellow or brown.


Flowering in March and fruiting from March to May.


Its specific epithet reflects the long and narrow mericarp wings of Hiptage stenopterum.

Vernacular name

Chinese: 狭翅风筝果(xiá chì fēng zhēng guǒ). The name’ xiá chì’ means its long and narrow wings, and ‘fēng zhēng gǔo’ is the Chinese name of Hiptage.

Habitat and distribution

H. stenopterum is only known from a valley of the Laowo River, a tributary of the Nujiang River, at an elevation ca. 1,000 m, near Lushui City, northwest of Yunnan Province, China (Fig. 2).

Conservation status

Since the only known population of H. stenopterum is in an entrenched valley of Nujiang River in the northwest Yunnan Province, we have not discovered the wild population outside of the abovementioned place, information known about the population status and natural distribution range of the new species is very limited. Currently, only about 20 individuals are found in the valley. Therefore, we suggest that the new species H. stenopterum should be considered Data Deficient (DD) according to current IUCN Red List Categories and Criteria (IUCN 2022).


We thank the two anonymous reviewers and extend special thanks to the subject editor, Rafael Felipe de Almeida, for his thoughtful and constructive criticisms.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.


This study was financially supported by the Hainan Provincial Natural Science Foundation of China (321QN185 & 422RC594), the Fund of Technology Innovation Alliance of Flower Industry (2020hhlm005), the Basic Research Fund of Guangxi Academy of Sciences (Grant No.CQZ-C-1901), and the Fund of Guangxi Key Laboratory of Plant Conservation and Restoration Ecology in Karst Terrain (No.22-035-26).

Author contributions

All authors have contributed equally.

Author ORCIDs

Tong-Tong Zhang

Shu-Yun Yang

Ke Tan

Ming-Xun Ren

Data availability

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


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