Research Article |
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Corresponding author: Hui-Lin Han ( hanhuilin@aliyun.com ) Academic editor: Christian Schmidt
© 2025 Ming-Xu Han, Hui-Lin Han.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Han M-X, Han H-L (2025) A taxonomic review of the genus Eversmannia Staudinger, 1871 (Lepidoptera, Uraniidae) from China, with descriptions of three new species and three new combinations. ZooKeys 1251: 275-291. https://doi.org/10.3897/zookeys.1251.145353
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Seven species of the genus Eversmannia Staudinger, 1871 are recognized from China. Among them, three species are described as new species: E. atromarginata sp. nov., E. spiralis sp. nov., and E. zhangorum sp. nov. New combinations are proposed for three species: E. bicaudata (Moore, [1868]) comb. nov., E. himala (Butler, 1880), comb. nov., and E. fuscifrons (Warren, 1896), comb. nov. Eversmannia bicaudata and E. fuscifrons are reported for the first time from China. Adults and genitalia are illustrated, and a key for identifying Chinese Eversmannia species is provided.
Epipleminae, key, swallowtail moth, taxonomy
The genus Eversmannia was established as a monotypic genus from Kazan, Russia by
In this study, we describe three new species, redescribe two newly recorded species from China, and transfer three species from Epiplema Herrich-Schäffer, 1855 to Eversmannia based on external and genital characters. This increases the number of species within Eversmannia to seven. Updated generic and species diagnoses, illustrations, a key to Chinese Eversmannia species, and distributional data are also provided.
Study specimens were collected in China, using 220V/450W mercury lamp and DC black (UV) light. Standard methods for dissection and preparation of the genitalia slides were used (
HGNU Huanggang Normal University, Huanggang, China
NEFU Northeast Forestry University, Harbin, China
SHNU Shanghai Normal University, Shanghai, China
Eversmannia Staudinger, 1871, in Staudinger & Wocke, Catalog der Lepidopteren des Europaeischen Faunengebiets: 159. Type species: Idaea exornata Eversmann, 1837. Type locality: Kazan, Russia.
In external appearance, this genus is similar to Europlema Holloway, 1998 but can be distinguished by the following characters: (1) male genitalia: uncus hook- or spoon-shaped; valva bar-shaped, editum-costa complex bifurcate; in Europlema uncus triangular or lanceolate and covered with hair, valva triangular, transtilla long and curved; (2) female genitalia: papillae anales conical or hoof-shaped; ductus bursae long and tubular, membranous; in Europlema, papillae anales flat and broad; ductus bursae very short, sclerotized.
Adult. Head white; labial palpus up-curved; in some species male with plumose antenna, female antenna filiform. Thorax and tegula white. Forewing ground color white, with brown or black patches; subbasal line and postmedial line brown or black. Hindwing white, with brown or yellow patches; outer margin with short extension at Rs and M3 veins. Abdomen covered with white scales.
Male genitalia. Uncus hook- or spoon-shaped, sclerotized. Gnathos trident- or spike-shaped. Tegumen triangular. Valva bar-shaped, covered with dense hair; costa long and strap-like, extending to cucullus. Editum-costa complex bifurcate, one branch short, finger-shaped, another long and bar-shaped. Juxta membranous, horseshoe-shaped. Vinculum V- or U-shaped. Saccus broad and V or U-shaped. Aedeagus cylindrical shaped, slightly curved or strongly curved; vesica with several spine-like cornuti.
Female genitalia. Papillae anales conical or hoof-shaped, covered with short setae. Apophysis posterioris longer than apophysis anterioris. Ostium bursae flat, sclerotized. Ductus bursae membranous, slightly curved or spiral. Corpus bursae oval or elongate and drop-shaped, in some species with separate tubular and globular parts; usually with variously shaped signum, sometimes absent.
China, Russia, Japan, North Korea, Bhutan, Nepal, India, Bangladesh, Finland, Estonia, Latvia, Lithuania, Belarus.
| 1 | Ductus bursae spiral shaped |
E. spiralis sp. nov. (Fig. |
| – | Ductus bursae straight | 2 |
| 2 | Uncus spoon-shaped | 3 |
| - | Uncus hook-shaped | 4 |
| 3 | Ductus bursae long and slender |
E. bicaudata comb. nov. (Fig. |
| – | Ductus bursae short and wide |
E. himala comb. nov. (Fig. |
| 4 | Aedeagus bow-shaped, strongly curved | 5 |
| – | Aedeagus cylindrical, slightly curved | 6 |
| 5 | Gnathos spine-shaped |
E. fuscifrons comb. nov. (Fig. |
| – | Gnathos spear-shaped |
E. zhangorum sp. nov. (Fig. |
| 6 | Corpus bursae with three signa |
E. atromarginata sp. nov. (Fig. |
| – | Corpus bursae with one signum |
E. exornata (Fig. |
Epiplema bicaudata:
Holotype: China – Jiangxi Prov. • ♂; Huang’ao Country, Mt Jinggang, Xiaoxidong Forest farm; 1 Jul. 2011; WC. Xie leg.; genit. prep. hmx-152-1;
This new species is superficially similar to E. exornata (Eversmann, 1837) but can be distinguished from the latter by the following characters: (1) forewing: subterminal and terminal areas covered with dark-brown patches; in E. exornata, these areas with scattered black and brown spots or a short band; (2) male genitalia: valva slightly narrower; aedeagus slightly slenderer; in E. exornata, valva broader; aedeagus thicker; (3) female genitalia: corpus bursae with three signa: two small signa at base of corpus bursae and one large signum at middle of corpus bursae; E. exornata with only one large signum.
Adults of Eversmannia spp. (depositories of 1 in
Adult
(Figs
Male genitalia of Eversmannia spp. (depositories of all slides NEFU). 15 E. atromarginata sp. nov., paratype, slide hmx-232-1; 16 E. bicaudata comb. nov., slide hmx-141-1; 17 E. exornata, slide hmx-195-1; 18 E. fuscifrons comb. nov., slide hmx-216-1; 19 E. himala comb. nov., slide hmx-50-1; 20 E. spiralis sp. nov., paratype, slide hmx-250-1; 21 E. zhangorum sp. nov., holotype, slide hmx-249-1.
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Shaanxi, Jiangxi, Chongqing, Sichuan) (Fig.
The species name atromarginata is from the Latin words “atro” and “marginata”, meaning black and margin and referring to the terminal subterminal line regions of the forewing which are covered with black patches.
Acidalia bicaudata
Idaea bicaudata: Cotes et al. 1888 (1887–1889): 576.
Erosia bicaudata:
Dirades bicaudata:
Epiplema bicaudata:
Epiplema fuscifrons:
China – Jiangxi Prov. • 1 ♀; Jinggangshan City, Luofu Reservoir; 18 Sep. 2010; DD. Zhang & S. Zhao & B. Tong leg.; genit. prep. hmx-99-2;
Adult
(Figs
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Anhui, Fujian, Jiangxi, Yunnan, Xizang) (Fig.
This species was formerly placed in the genus Epiplema. However, it shares several typical characters with Eversmannia: (1) forewing ground color white; postmedial line double; crossvein extremely weak; (2) hindwing white; postmedial line double; M2 vestigial, forming crease; (3) editum-costa complex bifurcate; (4) corpus bursae oval.
Eversmannia bicaudata comb. nov. shows significant differences from the type species of the genus Epiplema, E. acutangularia Herrich-Schäffer, 1855: (1) wing ground color of white; in E. acutangularia, grayish brown; (2) outer margin of forewing smooth; in E. acutangularia, dentate; (3) in male genitalia, gnathos present as two blades, editum-costa complex bifurcate; in E. acutangularia, gnathos and editum-costa complex absent. Consequently, we formally transfer this species to the genus Eversmannia.
Female genitalia of Eversmannia spp. (depositories of all slides in NEFU). 22 E. atromarginata sp. nov., paratype, slide hmx-139-2; 23 E. bicaudata comb. nov., slide hmx-142-2; 24 E. exornata, slide hmx-197-2; 25 E. fuscifrons comb.nov., slide hmx-103-2; 26 E. himala comb. nov., slide hmx-109-2; 27 E. spiralis sp. nov., paratype, slide hmx-172-2; 28 E. zhangorum sp. nov., paratype, slide hmx-124-2.
Idaea exornata Eversmann, 1837: 65. Type locality: Kazan, Russia.
Epiplema exornataria: Herrich-Schäfer 1847: 29, fig. 418 (misspelled).
Eversmannia exornata:
Erosia rapha Butler, 1878: 403; Butler 1879: 42, pl. 51 fig. 5.
Epiplema exornata:
China – Heilongjiang Prov. • 1 ♂; Heihe City, Pingshan Forest farm; 5–7 Jul. 2024; HL. Han & J. Wu & XR. Yang & TT. Zhao leg.; genit. prep. hmx-196-1; NEFU • 2 ♂♂, 5 ♀♀; Huma County, G331, near 12 km Bridge; 8 Jul. 2024; J. Wu & XR. Yang leg.; hmx-187-1, hmx-189-2, hmx-193-2; NEFU • 1 ♀; Huma County, Jinshan Forest farm; 8–9 Jul. 2024; HL. Han & J. Wu & XR. Yang & TT. Zhao leg.; genit. prep. hmx-197-2; NEFU • 1 ♂; Daxing’anling Prefecture, Fendou Village; 12 Jul. 2024; J. Wu & XR. Yang & TT. Zhao leg.; genit. prep. hmx-195-1; NEFU.
China (Heilongjiang) (Fig.
Epiplema fuscifrons
Warren, 1896: 348. Type locality: Sikkim, India; type specimens: in coll.
Epiplema bicaudata:
China – Xizang Autonomous Region • 1 ♂, 4 ♀♀; Motuo County, Deergong Village; 26 May–4 Jun. 2021; HL. Han leg.; genit. prep. hmx-216-1, hmx-103-2, hmx-104-2, hmx-199-2; NEFU • 1 ♀; Motuo County, Renqingbeng Temple; 29 Aug. 2024; YT. Fu leg.;
Adult
(Figs
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Xizang) (Fig.
Eversmannia fuscifrons comb. nov. can be easily distinguished from E. acutangularia. (1) The ground color of wings is white; in E. acutangularia is grayish brown. (2) The outer margin of forewing is smooth; in E. acutangularia it is dentate. (3) In male genitalia, the is gnathos present as two spikes and the editum-costa complex is bifurcate; in E. acutangularia, both the gnathos and editum-costa complex are absent. Consequently, we formally transfer this species to the genus Eversmannia.
The species is reported for the first time from China. The male genitalia are described for the first time.
Erosia himala
Epiplema himala:
Epiplema himala evanescens
Epiplema evanescens:
China – Chongqing City • 1 ♂; Simianshan Town, Tudiyan Station; 4–5 Aug. 2023; RT. Xu & MX. Han leg.; genit. prep. hmx-49-1; NEFU • 1 ♀; Simianshan National Nature Reserve; 29 Apr.–1 May 2019; JJ. Fan, ZT. Wang leg.; genit. prep. hmx-109-2; NEFU. – Yunnan Prov. • 1 ♂; Dali City, Nanjian County; 30 May 2022; RT. Xu & JJ. Fan leg.; genit. prep. hmx-50-1 NEFU. – Sichuan Prov. • 1 ♂, 2 ♀♀; Mabian County, Yonghong Township; 23 Jul. 2004; YD. Ren leg.; genit. prep. hmx-159-1
Adult
(Figs
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Hubei, Shaanxi, Chongqing, Sichuan, Yunnan) (Fig.
The subspecies Epiplema himala evanescens was upgraded to a full species by
This species was formerly placed in the genus Epiplema. There are some discrepancies in the characteristic wing patterns of E. himala compared to typical Eversmannia species (the antemedial, medial, postmedial and subterminal lines of the forewing are straight), but this species still shares several typical characters with Eversmannia: (1) forewing crossvein extremely weak; (2) hindwing M2 vestigial, forming crease; (3) editum-costa complex bifurcate; (4) corpus bursae oval.
Eversmannia himala comb. nov. shows significant differences from E. acutangularia. (1) The ground color of wings is white; in E. acutangularia, it is grayish brown. (2) The outer margin of forewing is smooth; in E. acutangularia, it is dentate. (3) In the male genitalia, the editum-costa complex is bifurcate; in E. acutangularia, it is absent. Consequently, we formally transfer this species to the genus Eversmannia.
The male genitalia of this species are described for the first time.
Holotype: China – Chongqing City • ♂; Simianshan National Nature Reserve; 25 Apr. 2022; C. Zhang & XY. Zhang & D. Feng leg.; genit. prep. hmx-248-1; NEFU. Paratypes: China – Chongqing City • 1 ♂; same collection data as the holotype; genit. prep. hmx-250-1; NEFU • 2 ♀♀; Simianshan Town, Dawopu; 7 May 2019; JJ. Fan & ZT. Wang leg.; genit. prep. hmx-171-2, hmx-172-2; NEFU • 1 ♀; same collection data as for preceding; 4 May 2019; genit. prep. hmx-191-2; NEFU.
This new species can be easily distinguished from other species of the genus by the following characters: (1) forewing: with a big dark brown semicircle shaped patch at terminal area near apex; in other species only with small spots or patches near apex. (2) Male genitalia: the caecum of aedeagus strongly expanded; in other species the caecum is straight or slightly expanded. (3) Female genitalia: the ductus bursae is spiral shaped, signum absent; in other species the ductus bursae is slightly curved; signum absent.
Adult. (Fig.
Habitats and field images of Eversmannia spp. 29 E. atromarginata sp. nov.; Prov. Jiangxi, Mt Jinggang; 30 E. atromarginata sp. nov., E. spiralis sp. nov., E. zhangorum sp. nov.; Chongqing City, Mt Simianshan; 31 E. atromarginata sp. nov.; Prov. Shaanxi, Yangxian County (Photo by XR. Yang); 32 E. bicaudata comb. nov.; Prov. Yunnan, Kunming City (Photo by YK. You); 33 E. exornata; Prov. Heilongjiang, Huma County (Photo by J. Wu); 34 E. fuscifrons comb. nov.; Aut. Reg. Xizang, Motuo County (Photo by YT. Fu); 35 E. himala comb. nov.; Chongqing City, Mt Simianshan (Photo by C. Zhang); 36, 37 E. spiralis sp. nov.; Prov. Guizhou, Tongren City, Mt Fanjingshan; Prov. Zhejiang, Hangzhou City, Mt Tianmushan (Photo by Z. Peng); 38, 39 E. zhangorum sp. nov.; Prov. Hunan, Liuyang City, Mt Daweishan; Prov. Zhejiang, Ningbo City, Longguan Township (Photo by WX. Jiang and YJ. Hu).
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Zhejiang, Chongqing, Guizhou) (Fig.
The species name is from the Latin word “spiralis”, meaning spiraling, referring to the spiral corpus bursae.
Holotype: China – Chongqing City • ♂; Simianshan National Nature Reserve; 19 Aug. 2021; C. Zhang & XY. Zhang & L. Luo leg.; genit. prep. hmx-249-1: NEFU. Paratype: China – Guizhou Prov. • 1 ♀; Zunyi City, Shierbeihou Scenic Spot; 3–5 Aug. 2020; HL. Han & J. Wu leg.; genit. prep. hmx-124-2; NEFU.
This new species is similar to E. fuscifrons but can be distinguished from the latter by the following characters: (1) forewing: the postmedial line with a black spot at the middle; in E. fuscifrons, postmedial line without spot; (2) male genitalia: gnathos spear-shaped; in E. fuscifrons, gnathos spine-shaped; (3) female genitalia: signum reniform with one edge serrate; in E. fuscifrons, signum star-shaped.
Adult
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Zhejiang, Hunan, Chongqing, Guizhou) (Fig.
This species is dedicated to Mr Chao Zhang and Ms Xin-Yu Zhang, who collected the holotype.
We are grateful to Dr Dandan Zhang (
The authors have declared that no competing interests exist.
No ethical statement was reported.
No use of AI was reported.
The present study was supported by the National Natural Science Foundation of China (No. 31872261), the project of the Northeast Asia Biodiversity Research Center (2572022DS09) and the Fundamental Research Funds for the Central Universities (No. 2572021DJ08, 2572019CP11).
Conceptualization: HLH and MXH. Data curation: MXH. Investigation: MXH. Supervision: HLH. Writing-original draft: MXH. Writing-review and editing: HLH.
Ming-Xu Han https://orcid.org/0009-0002-9252-5981
Hui-Lin Han https://orcid.org/0000-0002-2045-6182
All of the data that support the findings of this study are available in the main text.