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Korean J. Pl. Taxon > Volume 56(1); 2026 > Article
KIM, JANG, KANG, LEE, CHUNG, and SON: A new species, Inula minipetala (Asteraceae), from Korea, formerly treated as an invalid variety

Abstract

The name Inula salicina var. minipetala Y. N. Lee was originally published in 1996 in Flora of Korea with a brief Latin diagnosis, but without indication of the herbarium where the type specimen is conserved. This omission constitutes non-compliance with Article 40.7 of the International Code of Nomenclature for algae, fungi, and plants (ICN), thereby rendering the name not validly published (nomen invalidum). Although recent field surveys have not confirmed its continued presence at the original locality in Jeonui-myeon (Sejong City), where it was first reported, this morphologically distinct taxon has repeatedly been reported under the vernacular name ‘Jeon-ui-geum-bul-cho’ from several localities throughout central Korea. It is consistently distinguished by its conspicuously short ligulate ray florets (lamina 3–4 mm), linear-lanceolate to narrowly lanceolate entire leaves with appressed pilose hairs and glandular dots on the abaxial surface, a pappus shorter than the corolla of the disc florets, and a compact, subglobose involucre composed of four series of broadly lanceolate phyllaries. In particular, the outermost phyllaries are notably shorter than the inner ones, and the cypselae are glabrous or only sparsely pubescent at the apex. These features collectively constitute a diagnostic combination of characters that clearly distinguishes this species from other Korean Inula. Although formerly associated with Inula salicina, phylogenetic analyses based on nuclear ribosomal internal transcribed spacer sequences indicate that the lineage forms a well-supported and independent clade more closely allied to the I. japonica complex. As Lee’s original material is no longer traceable, we designate a newly collected specimen as the holotype and hereby propose this entity as a new species endemic to Korea: Inula minipetala Y. N. Lee ex H. B. Kim & D. C. Son, sp. nov.

INTRODUCTION

The genus Inula L. (Asteraceae, Inuleae) comprises over 100 species distributed primarily across Eurasia and North Africa, many of which inhabit temperate grasslands and montane wetlands (Shekhar et al., 2013; Kim, 2020). It is distinguished from other genera within the Inuleae by its herbaceous phyllaries, radiate capitula, and florets bearing a capillary pappus (Koyama, 1995; Chen et al., 2011). In Korea, native representatives of Inula include Inula salicina var. asiatica Kitam., I. japonica Thunb., I. japonica var. ramosa (Kom.) C. Y. Li, and I. linariifolia Turcz., with each adapted to specific ecological niches such as grassy areas, riparian meadows, roadside banks, forest margins, and mountain slopes (Korea National Arboretum, 2020). Owing to substantial morphological variation within and among taxa, several specific and infraspecific names have historically been described, although most have subsequently been reduced to synonymy (POWO, 2025).
Among these, a morphological variant characterized by conspicuously shorter ligulate florets was first described by Y. N. Lee in 1996 under the name Inula salicina var. minipetala, published in Flora of Korea, one of his most influential floristic works. However, he did not specify the herbarium in which the type specimen was conserved in the protologue—a requirement stipulated by Article 40.7 of the International Code of Nomenclature for algae, fungi, and plants (ICN) (Turland et al., 2025). As a result, the name was not validly published (nomen invalidum), and has since been excluded from national checklists of the vascular flora of Korea, floristic databases, field guides, and regional floristic surveys (Kim et al., 2018; Kim, 2020; Korea National Arboretum, 2020; Sung and Kang, 2020; Jang and Lee, 2022; Son et al., 2025).
Despite its nomenclatural invalidity, this taxon—commonly referred to by the vernacular name ‘Jeon-ui-geum-bul-cho’—has been consistently reported not only from its originally recognized locality in Jeonui-myeon, Sejong City but also from various regions of central Korea through public online records and citizen science platforms (e.g., Naturing). It is morphologically distinguished from other native Korean Inula taxa by a unique combination of features, including notably short ligulate ray florets, linear-lanceolate to narrowly lanceolate leaves with appressed pilose hairs and glandular dots on the abaxial surface, and a compact, subglobose involucre composed of four series of broadly lanceolate phyllaries that are erect and closely appressed (the outermost series are markedly shorter than the ones in the interior). The cypselae are glabrous or only sparsely pubescent at the apex.
Although Lee (1996) did not designate a herbarium for the type of Inula salicina var. minipetala, his type materials were reportedly donated by his family to the National Institute of Biological Resources (KB) (Son et al., 2016). Despite extensive efforts to trace the type specimen—including searches at KB and other major Korean herbaria, such as the Ewha Womans University (EWH) and Korea National Arboretum (KH)—no original material referable to this name has been located.
Accordingly, we designate a holotype based on the newly collected material and formally describe the taxon as a new species under the name Inula minipetala.

MATERIALS AND METHODS

Morphological data were obtained from specimens collected in the field in 2025. The specimens were pressed and dried or preserved in 70% ethanol and deposited in the KH herbarium. Measurements and descriptions of reproductive characters were based on fresh and ethanol-preserved materials, whereas vegetative characters were examined using fresh materials and dried herbarium specimens. Floral morphology, as well as surface characters of leaves and cypselae, were observed under a stereomicroscope (SZ51, Olympus Corp., Tokyo, Japan). For comparison, morphological data of Inula species distributed in the Republic of Korea (I. japonica, I. linariifolia, and I. salicina var. asiatica) were obtained from fresh materials and herbarium specimens (Appendix 1). Phylogenetic analyses were performed using total DNA extracted from young leaves of plants collected in the field and from herbarium specimens (Appendix 1), using the DNeasy Plant Mini Kit (Qiagen, Hilden, Germany). The nuclear ribosomal internal transcribed spacer region was amplified by PCR and sequenced, and sequence alignment was performed in Geneious Prime v.2025.2.2 (Biomatters Ltd., Auckland, New Zealand). The analysis model was tested using PhyloSuite v.2.0.2 (Zhang et al., 2020; Xiang et al., 2023), and the best-fit model for maximum-likelihood (ML) analysis was selected with ModelFinder v.3.0.1 (Kalyaanamoorthy et al., 2017). ML analysis was conducted in IQ-TREE v.2.2.0 (Nguyen et al., 2015) under the TNe model with 5,000 ultrafast bootstrap replicates (Minh et al., 2013). The resulting phylogenetic tree was visualized using FigTree v.1.4.4 (http://tree.bio.ed.ac.uk/software/figtree).

TAXONOMIC TREATMENT

Inula minipetala Y. N. Lee ex H. B. Kim & D. C. Son, sp. nov. (Fig. 1).TYPE: KOREA. Chungcheongnam-do, Geum-san-gun, Chubu-myeon, Seongdang-ri 231-3, elev. 189 m, 36.233721°N, 127.524885°E; 22 Aug 2025, H.B. Kim et al. 250822-1 [holotype KH (Fig. 2); isotypes: KH (2 sheets), KB (1 sheet)].
Korean name: Jeon-ui-geum-bul-cho (전의금불초).
Herbs, perennial, arising from short rhizomes. Stems erect, 40–70 cm tall, sparsely appressed pilose, branched in the upper part. Leaves alternate; basal and lower cauline leaves withering at anthesis; middle cauline leaves sessile; leaf blades linear-lanceolate to narrowly lanceolate, 4.3–8.5 cm long, 0.5–1.2 cm wide, apex acute, base attenuate, margin entire, adaxial surface glabrous, abaxial surface with appressed pilose and glandular dots; upper cauline leaves progressively reduced upward, linear-lanceolate. Synflorescences loosely corymbiform. Capitula solitary at stem apices, 1.2–1.7 cm in diam. Involucre subglobose, 5–8 mm in diameter; phyllaries arranged in 4 series; outer phyllaries broadly lanceolate, 2.3–3.6 × 0.8–1.5 mm, apex acute, margin with pilose and glandular trichomes, adaxially sparsely pilose, without glandular dots, abaxially glabrous; inner phyllaries broadly lanceolate to lanceolate, 3.4–5.6 × 0.5–1.2 mm, apex acute, margin with pilose and glandular trichomes, both surfaces glabrous. Ray florets numerous; lamina linear, 3–4 mm long, yellow; disc florets yellow, corolla 3.2–3.7 mm long. Cypselae cylindrical, 0.7– 1 mm long, 10-ribbed, glabrous or sparsely pubescent apically; pappus composed of numerous barbellate bristles, connate at base and persistent; bristles white, 2.4–3 mm long.
Distribution: This species is regarded as endemic to Korea, with confirmed occurrences in Geumsan-gun, Chungcheongnam-do, central Korea.
Etymology: The specific epithet minipetala derives from Latin mini- (‘small’) and petalum (‘petal’). It was originally proposed by Lee (1996) as Inula salicina var. minipetala; however, the name was not validly published because no holotype was designated, in violation of Art. 40.7 of the ICN. In the present treatment, the epithet is formally adopted and validated as a species (sp. nov.), in recognition of Lee’s original taxonomic concept, which emphasizes the conspicuously short ligulate ray florets characteristic of this species. Accordingly, the authorship follows ICN Art. 46.10 as ‘Y. N. Lee ex H. B. Kim & D. C. Son’, acknowledging Lee’s initial proposal of the name. Retaining the epithet preserves taxonomic continuity and clearly reflects the species’ diagnostic floral morphology, notably the exceptionally short ligulate ray florets relative to its congeners.
This vernacular name reflects the type locality of the species, Jeonui-myeon in Sejong City, where the taxon was first discovered by Lee (1996). Although recent field surveys and herbarium investigations have not confirmed its current presence in Jeonui-myeon, the species’ historical occurrence in the area is supported by a photographic record published in Flora of Korea (Lee, 1996; pl. 2492). The continued informal use of this vernacular name among Korean botanists and floristic workers underscores the cultural and scientific significance of the original collection site in the species’ naming history.
Note: Inula minipetala is readily distinguishable from other native Korean Inula by a diagnostic combination of traits: conspicuously short ligulate ray florets (lamina 3–4 mm), linear-lanceolate to narrowly lanceolate leaves with entire margins, bearing appressed pilose hairs and glandular dots restricted to the abaxial surface, a pappus shorter than the corolla of the disc florets, and a compact, subglobose involucre comprising four series of broadly lanceolate phyllaries, with the outermost distinctly shorter than the inner series. The cypselae are glabrous or only sparsely pubescent apically (Fig. 3). The persistence of these morphological differences across both vegetative and reproductive traits reinforces the taxonomic distinctiveness of I. minipetala and supports its recognition as a distinct species within the Korean Inula complex (see Table 1).
Furthermore, our nuclear ribosomal internal transcribed spacer phylogenetic analyses reveal that the lineage corresponding to this taxon does not cluster with Inula salicina var. asiatica but instead forms a well-supported, distinct clade (maximum likelihood bootstrap = 100%) that is more closely related to the I. japonica complex (I. japonica and I. linariifolia) (Fig. 4). This clear phylogenetic pattern conflicts with the historical taxonomic treatment of this taxon as a variety of I. salicina. Although bootstrap values within the subclades of the I. japonica complex were generally low, I. japonica, I. linariifolia, and I. minipetala were each resolved as distinct clades (Fig. 4). In addition, this taxon exhibits clear morphological differentiation in reproductive characters, including the morphology and micromorphological characters of the cypselae, which have been regarded as key diagnostic traits in Asteraceae and Inula taxonomy (Pandey et al., 1983; Abid and Qaiser, 2002, 2007; Abid and Zehra, 2007; Shekhar et al., 2011), as well as the length of ray florets (lamina) and the shape of phyllaries, thereby supporting its distinctiveness at the rank of species (Fig. 3, Table 1).
Meanwhile, the Inula complex has undergone substantial taxonomic reevaluation, with several species—most notably Inula britannica—transferred to the genus Pentanema based on combined nuclear and plastid phylogenetic evidence (Gutiérrez-Larruscain et al., 2018). These reclassifications have been partially adopted by global databases such as POWO, which now assign many Eurasian Inula species to Pentanema. Notably, Korean taxa—such as Inula salicina var. asiatica, I. japonica, I. japonica var. ramosa, and I. linariifolia—have previously been considered infraspecific variants of I. britannica (Koyama, 1995; Chen et al., 2011; Kim et al., 2018; Kim, 2020), suggesting potential future taxonomic realignment. However, the phylogenetic sampling in Gutiérrez-Larruscain et al. (2018) remains geographically limited, as northeastern Asian taxa—particularly those from Korea—were not included in their analyses. Recent studies have further argued for retaining certain lineages within a broader circumscription of Inula, citing unresolved paraphyly between Inula and Pentanema and insufficient molecular representation of East Asian species (Yuan et al., 2024).
In light of these uncertainties and the taxon’s stable morphological features and repeated occurrence across multiple regions in Korea, we conservatively assign Inula minipetala to Inula rather than transfer it to Pentanema, thereby facilitating further comprehensive molecular analyses.
Additional specimen examined (paratypes): KOREA. Chungcheongnam-do: Geumsan-gun, Chubu-myeon, Seongdangri, 231-3, 18 Sep 2025, H.B. Kim et al. KHB1663379, KHB1663380, KHB1663381, KHB1663382 (KH).

Key to the species of Inula in Korea

  • 1. Middle cauline leaves abaxially glabrous, without glandular dots ··································· I. salicina var. asiatica

  • 1. Middle cauline leaves abaxially appressed pilose with glandular dots.

    • 2. Involucre composed of 5 series of phyllaries ···································································· I. japonica

    • 2. Involucre composed of 4 series of phyllaries.

      • 3. Heads ca. 1.8–2.5 cm across; ray florets lamina 7–10 mm long; disc florets corolla as long as pappus; outer phyllaries adaxially pilose with glandular dots; cypselae pubescent ······················ I. linariifolia

      • 3. Heads ca. 1.2–1.7 cm across; ray florets lamina 3–4 mm long; disc florets corolla longer than pappus; outer phyllaries adaxially sparsely pilose, without glandular dots; cypselae glabrous or only sparsely pubescent at apex ······················· I. minipetala

NOTES

ACKNOWLEDGMENTS
This study was supported by the project “Enhancement of the checklist of Korean Peninsula based on the names (KNA1-1-30-25-1),” which was funded by the Korea National Arboretum.
CONFLICTS OF INTEREST
The authors declare that there are no conflicts of interest.

Fig. 1
Inula minipetala. A. Habit. B. Root. C. Stem. D. Cauline leaves. E, F. Adaxial surface of cauline leaves. G, H. Abaxial surface of cauline leaves. I. Inflorescence. J. Flower. K. Involucre. L, M. Phyllary (L, adaxial surface; M, abaxial surface). N. Ray florets. O. Disc florets. P, Q. Cypselae. Photographs by Hye Been Kim and Kang-Hyup Lee.
kjpt-56-1-56f1.jpg
Fig. 2
Holotype specimen of Inula minipetala (KH barcode KHB1663375) collected from Chubu-myeon, Geumsan-gun, Chungcheongnam-do.
kjpt-56-1-56f2.jpg
Fig. 3
Comparative photographs of the flower, cauline leaves, disc florets, involucre, phyllary (left: outermost phyllary, right: innermost phyllary), cypselae of Inula minipetala (A–F), I. japonica (G–L), I. linariifolia (M–R), and I. salicina var. asiatica (S–X). Photographs by Hye Been Kim and Kang-Hyup Lee.
kjpt-56-1-56f3.jpg
Fig. 4
Phylogenetic tree based on internal transcribed spacer (ITS) regions of Korean Inula. The numbers above the branches are bootstrap values (BS > 50%) by the maximum likelihood method. Newly collected and generated sequences in this study are shown with an asterisk, and the new species is marked with a red box. The voucher information of all samples used in the analysis is indicated after the scientific names.
kjpt-56-1-56f4.jpg
Table 1
Comparison of the major morphological characteristics of Inula minipetala and related Korean Inula taxa.
Characters I. minipetala I. japonica I. linariifolia I. salicina var. asiatica
Cauline leaves Shape Linear-lanceolate to narrowly lanceolate Lanceolate to oblong Linear to linear-lanceolate Lanceolate or oblong
Length (cm) 4.3–8.5 5.0–12.6 4.0–9.0 4.5–7.6
Width (cm) 0.5–1.2 1.4–3.0 0.5–1.0 1.0–1.7
Surface Adaxially glabrous
Abaxially appressed pilose with glandular dots
Adaxially sparsely appressed pilose or subglabrous
Abaxially appressed pilose with glandular dots
Adaxially glabrous
Abaxially appressed pilose with glandular dots
Glabrous on both surfaces
Capitula Diameter (cm) 1.2–1.7 3–4 1.8–2.5 2–4
Involucre Number of series 4 series 5 series 4 series 4 series
Outer phyllaries Shape Broadly lanceolate Linear-lanceolate Lanceolate Spatulate-oblong
Length (mm) 2.3–3.6 8–10 3.6–5 5.5–6
Surface Adaxially sparsely pilose without glandular dots
Abaxially glabrous
Adaxially pilose with glandular dots
Abaxially glabrous
Adaxially pilose with glandular dots, Abaxially glabrous
Glabrous on both surfaces
Inner phyllaries Shape Broadly lanceolate to lanceolate Linear-lanceolate Lanceolate Linear-lanceolate
Length (mm) 3.4–5.6 8–10 3.8–5.1 5–9.5
Surface Glabrous on both surfaces Glabrous on both surfaces Glabrous on both surfaces Adaxially densely pubescent
Abaxially glabrous
Ray florets (lamina) Length (mm) 3–4 8–15 7–10 10–15
Disc florets (corolla) Length (mm) 3.2–3.7 3–5 3.6–4.3 5–9
Cypselae Surface Glabrous or only sparsely pubescent at apex Pubescent Pubescent Glabrous
Pappus Length (mm) 2.4–3 4–5 3.6–4.5 7–8

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APPENDICES

Appendix 1

Specimens examined for comparison and phylogenetic analyses.
Inula japonica: KOREA. Seoul: Eunpyeong-gu, Nokbeon-dong, 13 Jul 2025, H.B. Kim KHB1663385, KHB1663386 (KH); Mapo-gu, Sangam-dong, Pyeonghwa Park, 5 Aug 2015, S.H. Park KHB1559751 (KH). Gangwon-do: Jeongseon-gun, Jeongseon-eup, Mt. Gariwangsan, 7 Sep 2011, I.C. Hwang & Y.J. Yun KHB1548778 (KH); Pyeongchang-gun, Daegwallyeong-myeon, Byeongnae-ri, 23 Aug 2010, Y.C. Kim KHB1327372 (KH). Chungcheongbuk-do: Danyang-gun, Mt. Taehwasan, 25 Sep 2005, B.U. Oh KHB1144443 (KH). Gyeongsangbuk-do: Yeongyang-gun, Seokbo-myeon, Mt. Maengdongsan, 28 Sep 2006, G.Y. Chung KHB1230810 (KH).
Inula linariifolia: KOREA. Gyeonggi-do: Yeoncheon-gun, Yeoncheon-eup, Tonghyeon-ri, 14 Aug 2025, H. B. Kim et al. KHB1663383, KHB1663384 (KH). Gyeongsangbuk-do: Sangju-si, Hwaseo-myeon, Hasong-ri, Mt. Songnisan, 8 Aug 2006, G.Y. Chung KHB1130125, KHB1130126 (KH). Gyeongsangnam-do: Changnyeong-gun, Daehap-myeon, North of Uponeup (wet land), 21 Aug 2006, KHB1121306 (KH); Geoje-si, Sadeung-myeon, Changho-ri, Gajodo, 21 Jul 2010, G.Y. Chung KHB1361255 (KH). Jeollanam-do: Sinan-gun, Daeheuksando, 5 Jul 2007, B.U. Oh et al. KHB1617433 (KH).
Inula salicina var. asiatica: KOREA. Incheon: Ongjin-gun, Daecheong-myeon, Daecheong-ri, Daecheongdo, 23 Aug 2025, K.H. Lee KHB1663387, KHB1663388, KHB1663389 (KH). Gyeonggi-do: Hwaseong-si, Seoshin-myeon, Jebudo, 6 Jun 2009, E.S. Jeon KHB1268382 (KH). Chungcheongbuk-do: Jincheon-gun, Mt. Dutasan, 26 Jul 2005, K.O. Yoo KHB1143195 (KH). Chungcheongnam-do: Seosan-si, Jigok-myeon, Daeyo-ri, 28 May 2012, W.K. Paik KHB1434199 (KH). Gyeongsangbuk-do: Andong-si, Pungcheon-myeon, Gwangdeok-ri, 20 Jul 2011, S.T. Yoo & B.D. Kim KHB1344734 (KH); Gumi-si, Mt. Cheonghwasan, 13 Jul 2006, B.U. Oh et al. KHB1605621 (KH)
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