Research Article |
Corresponding author: Xi-Cui Du ( duxicui@hotmail.com ) Academic editor: Richard Mally
© 2023 Jia-Ming Guo, Xi-Cui Du.
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:
Guo J-M, Du X-C (2023) Five new species of Bradina Lederer (Lepidoptera, Crambidae) from China, with remarks on the morphology of the genus. ZooKeys 1158: 49-67. https://doi.org/10.3897/zookeys.1158.99411
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Bradina is a species-rich genus that differs from most other Spilomelinae genera because of its distinctive wing venation. Most species of this genus are very similar in appearance. In this study, we have studied morphological characteristics of the genus and eight closely related species from China. Among them, B. falciculata Guo & Du, sp. nov., B. fusoidea Guo & Du, sp. nov., B. spirella Guo & Du, sp. nov., B. ternifolia Guo & Du, sp. nov. and B. torsiva Guo & Du, sp. nov. are described as new to science. Bradina megesalis (Walker, 1859), B. translinealis Hampson, 1896 and B. subpurpurescens (Warren, 1896) are redescribed based on their holotypes and additional material, and the latter two are newly recorded from China and their genitalia are described for the first time. The images of the habitus and genitalia of these eight species are provided, with a key to their identification.
Genitalia, identification key, morphology, Pyraloidea, Spilomelinae, taxonomy
Bradina Lederer, 1863 is the most species-rich genus in the Spilomelinae tribe Steniini Guenée, 1854, redefined by
In Spilomelinae, Bradina can be differentiated from most other genera by the forewings with Rs1 anastomosed with Rs2+s3 at the base, which is common in Acentropinae. Therefore, this genus was placed in Hydrocampinae (= Acentropinae Stephens, 1835) for a long time (
Species identification of Bradina is difficult because of their very similar appearance, so the genitalia characteristics are necessary in the identification of most species. The large spinose crescent-shaped signum of the female genitalia is a diagnostic characteristic of the genus, but shows little difference among species. In male genitalia, the valvae and uncus are very diverse in morphology, which is very valuable for interspecific identification. Before this study, 13 species were recorded in China (
Specimens examined, including the types of new species, are deposited in the College of Plant Protection, Southwest University, Chongqing, China (SWU) except for two holotypes and 38 paratypes which are deposited in the Insect Collection of the College of Life Science, Nankai University, Tianjin, China (
The photographs of the adults were taken with a digital camera (Canon EOS 5D), and those of the genitalia were obtained with a digital camera (Leica DFC 450) attached to a stereomicroscope (Leica M205 A).
The preparation of genitalia mainly follows
Bradina
Lederer, 1863: 424. Type species: Bradina impressalis Lederer, 1863; subsequent designation by
Erilita Lederer, 1863: 426. Type species: Erilita modestalis Lederer, 1863, by monotypy.
Pleonectusa
Lederer, 1863: 426. Type species: Botys admixtalis Walker, 1859; subsequent designation by
Trematarcha
Meyrick, 1886: 233. Type species: Marasmia erilitalis Felder, Felder & Rogenhofer, 1875; subsequent designation by
Head
(Fig.
Male genitalia. Uncus diverse, apex with setae dorsally. Valva narrow or broad, some with well-developed setal cluster. Saccus developed. Phallus long and cylindrical.
Female genitalia. Papillae anales densely setose. Apophyses anteriores c. twice length of apophyses posteriores. Ostium bursae well developed. Corpus bursae rounded or oval, inside usually densely studded with tiny spines; signum crescent and densely spinose, spines on concave side developed.
The generic characteristics were summarized by
The bodies of Bradina species are usually brown, yellowish brown, or pale brown, except for a few species with white bodies, and have nearly identical wing markings. The male genitalia, on the other hand, are very diverse among species of this genus. We found that the male genitalia can be divided into three types according to the morphological characteristics of the valvae. The first type, represented by B. admixtalis, have long and narrow valvae; the second type, represented by B. melanoperas, have short and broad valvae; the third type, represented by B. megesalis, have broad valvae in which the costa is arched near the base or middle. Species in the present study have the third valva type, accompanied by the following common characteristics: body brown of various shades. Uncus broad, distal part bilobed, with dense short setae dorsally. Valva broad; costa arched near base or middle and accompanied by a cluster of long setae, usually followed by a depression. Saccus nearly trapezoidal, slightly concaved terminally. Juxta nearly rounded, split posteriorly. Phallus distinctly inflated at anterior end. Apophyses anteriores expanded at c. one third from base.
1 | Sacculus with a cluster of long setae near middle | 2 |
– | Sacculus without long setal cluster | 5 |
2 | Phallus with one fusiform cornutus composed of short and blunt spines, and one subcircular sclerotized cornutus | B. fusoidea sp. nov. |
– | Phallus with various cornuti, but not as above | 3 |
3 | Phallus with three long leaf-like cornuti tapered apically | B. ternifolia sp. nov. |
– | Cornuti not as above | 4 |
4 | Posterior phallus with two developed spear-like cornuti | B. megesalis |
– | Phallus with one short and spiral band-like cornutus tapered at posterior end, along with two lamellar cornuti in posterior half | B. spirella sp. nov. |
5 | Valva nearly rectangular, broad distally | B. subpurpurescens |
– | Valva nearly elliptical or narrowed distally | 6 |
6 | Phallus with one broad and spiral band-like cornutus | B. torsiva sp. nov. |
– | Phallus with cornutus not as above | 7 |
7 | Phallus with one fusiform cornutus medially, and with two lamellar cornuti posteriorly. Ductus bursae slender, membranous | B. translinealis |
– | Phallus with one small fusiform cornutus anteriorly, and with two sickle-shaped cornuti posteriorly. Ductus bursae relatively thick, slightly sclerotized medially | B. falciculata sp. nov. |
Botys megesalis
Walker, 1859: 663. Type locality: North China. Type depository:
Bradina megesalis: Hampson, 1897: 200.
Holotype
, ♂ North China, from Mr Fortune’s collection (Walker, 1859), genitalia slide no. 8735 (
China, Chongqing Municipality, 2 ♂♂, 6 ♀♀, Bashan Town, Chengkou County, alt. 900 m, 10 July 2017, Ji-Ping Wan leg.; 2 ♂♂, 4 ♀♀, Jinfo Mountain, alt. 918 m, 27 August 2019, You Zeng leg.; 1 ♂, Simian Mountain, alt. 1280 m, 12 July 2012, Gui-Qing He leg.; Guangdong Prov., 2 ♂♂, 2 ♀♀, Dadong Mountain, Lian County, 5–8 July 2008, Feng-Xia He leg., genitalia slide no.: GJM21078 ♂; 1 ♂, 2 ♀♀, Qianjin Conservation Station, Shimentai Nature Reserve, Qingyuan City, alt. 523 m, 26 May 2021, Xing-Hai Zuo leg., genitalia slide no.: GJM21073 ♂; Guangxi Zhuang Autonomous Region, 1 ♂, Yinshan Park, Dayao Mountain, alt. 1564 m, 8 July 2013, Xiao-Hua Chen leg.; Guizhou Prov., 2 ♂♂, 4 ♀♀, Leigong Mountain, Leishan County, alt. 1198 m, 14–15 July 2013, Xiao-Hua Chen leg., genitalia slide no.: GJM21076 ♂; Hubei Prov., 18 ♂♂, 33 ♀♀, Dabie Mountain, alt. 590 m, 24–25 June 2014, Li-Jun Xu leg.; Hainan Prov., 1 ♂, 1 ♀, Bawangling National Forest Park, 8–10 June 2010, Li Kang leg.; Hunan Prov., 5 ♀♀, Wuyunjie Nature Reserve, alt. 178 m, 19 June 2019, Ying Yang leg.; Sichuan Prov., 2 ♂♂, 1 ♀, Longcanggou Forest Park, Xingjing County, alt. 1388 m, 17 June 2021, Shuai Yu leg., genitalia slide no.: GJM21074 ♂, GJM21080 ♀; Shaanxi Prov., 1 ♂, Zuoshui County, Shangluo City, alt. 810 m, 29 June 2021, Jin-Hang Han leg.; 1 ♂, 5 ♀♀, Hanyin County, Ankang City, alt. 410 m, 26 June 2021, Jin-Hang Han leg.; Yunnan Prov., 9 ♂♂, 6 ♀♀, Baihualing Village, Baoshan City, alt. 1487 m, 20–23 June 2020, Ying Yang & Hong Zhao leg.; 2 ♂♂, 1 ♀, Cuanlong Village, Mangba Town, Tengchong City, Baoshan City, alt. 1329 m, 8 August 2015, Jing-Xia Zhao & Hao Wei leg.; Zhejiang Prov., 6 ♂♂, 7 ♀♀, Jiulongshan Forest Park, 4–6 August 2011, Xiao-Bing Fu leg.; 6 ♂♂, 34 ♀♀, Tianmu Mountain, alt. 800 m, 29 July 2011, Xi-Cui Du & Xiao-Bing Fu leg.
Adult
(Figs
Male genitalia
(Fig.
Genitalia of Bradina species A–E B. megesalis A, B male, slide no. GJM21074 C–E female, slide no. GJM21080 F–K B. fusoidea sp. nov. F–H male holotype, slide no. GJM21117 I–K female, paratype, slide no. GJM21118 L–P B. spirella sp. nov. L, M male, holotype, slide no. GJM21161 N–P female, paratype, slide no. GJM21164 B, G, H, M partial enlargement of phallus D, J, O partial enlargement of ductus bursae E, K, P signum. Scale bars: 0.2 mm (E, G, K, P); 0.5 mm (A, B, D, F, H, J, L, M, O); 1.0 mm (C, I, N).
Female genitalia
(Fig.
China (Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Hainan, Hunan, Sichuan, Shaanxi, Yunnan, Zhejiang), Japan (
We found that the coremata of some male individuals of this species were protruded out of the body, forming a cluster of white hairs at the end of the abdomen.
Holotype
, ♂ China: Sichuan Prov., Qingcheng Mountain, Dujiangyan city, alt. 860 m, 30°92'N, 103°50'E, 22 July 2021, Shuai Yu, Xiao-Ju Zhu, Di Zhang leg. (
This species is similar to B. megesalis. The difference in appearance is that the distance between discoidal stigma and postmedial line on forewing of this species is longer than that of B. megesalis; the postmedial line and terminal margin of forewing is obviously unparallel in this species, while is nearly parallel in the latter. It also can be distinguished by the setal cluster near middle of sacculus being shorter than that of the latter, phallus with one fusiform cornutus and one subcircular cornutus (two spear-like cornuti in B. megesalis), posterior ductus bursae inflated but not forming irregular protrusion as in B. megesalis, plus subposterior section of ductus bursae widened along half of ductus length, a feature that is absent in B. megesalis.
Adult
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
The specific name is derived from the Latin fusoideus (meaning ‘fusiform’), in reference to a fusiform cornutus.
China (Guangdong, Sichuan).
Holotype , ♂ China: Hunan Prov., Xianchijie, Wuyunjie National Nature Reserve, alt. 720 m, 28°90'N, 111°48'E, 24 June 2019, Ying Yang leg., genitalia slide no. GJM21161. Paratypes. China: Hunan Prov., 9 ♂♂, 4 ♀♀, Bamian Mountain Nature Reserve, Guidong County, alt. 973 m, 16 June 2015, Kai Chen leg., genitalia slide no.: GJM21160 ♂; 5 ♂♂, 2 ♀♀, Zhushan Village, Wuyunjie National Nature Reserve, alt. 100 m, 15 June 2019, Ying Yang leg.; 34 ♂♂, 14 ♀♀, Jindongjie, Wuyunjie National Nature Reserve, alt. 178 m, 16–19 June 2019, Ying Yang leg.; 66 ♂♂, 22 ♂♂, 18–24 June 2019, other same data as holotype, genitalia slide no.: GJM21048 ♂, GJM21049 ♂, GJM21162 ♂, GJM21163 ♂, GJM21164 ♀; Jiangxi Prov., 4 ♂♂, 2 ♀♀, Jinggang Mountain, 30 June 2011, Jin-Wei Li leg., genitalia slide no.: GJM21159 ♂, GJM21183 ♀.
This species is similar to B. megesalis. The difference in appearance is that wings of this species are darker in color and hindwings are slightly broader; the postmedial line and terminal margin of forewing are obviously unparallel in this species, while nearly parallel in the latter. It also can be distinguished by phallus with one short, spiral band-like cornutus and two lamellar cornuti, and some tiny spines on the vesica medially; the posterior ductus bursae is inflated into a thick finger-like protrusion. In B. megesalis, the phallus has two spear-like cornuti and is without spines on the vesica; the posterior ductus bursae is inflated into an irregular protrusion.
Adult
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
The specific name is derived from the Latin spirellus (meaning ‘small spiral-like’), in reference to a short and spiral band-like cornutus.
China (Hunan, Jiangxi).
Coremata of some male individuals of this species were protruded out of the body, forming a cluster of white hairs at the end of the abdomen.
Holotype
, ♂ China: Hunan Prov., Chenzhou Nature Reserve, alt. 1233 m, 25°78'N, 113°01'E, 3 June 2019, Xiao-Qiang Lu & Ying Yang leg., genitalia slide no. GJM21102. Paratypes. China: Hunan Prov., 4 ♂♂, other same data as holotype, genitalia slide no.: GJM21103 ♂; 1♂, Zhushan Village, Taoyuan County, Changde City, alt. 100 m, 15 June 2019, Xiao-Qiang Lu & Ying Yang leg., genitalia slide no.: GJM21101; 2 ♂♂, Maozhu River, Shimen County, Changde City, alt. 350 m, 6 June 2017, Jian-Yue Qiu & Hao Xu leg.; Guangdong Prov., 2 ♂♂, 1 ♀, Dadong Mountain, Lianzhou City, alt. 650 m, 21 June 2004, Dan-Dan Zhang leg., genitalia slide no.: GJM21100 ♂; 1 ♀, Qingyuan City, alt. 270 m, 7 June 2019, Xiao-Qiang Lu & Ying Yang leg.; 1 ♂, 2 ♀♀, Babaoshan Conservation Station, Nanling National Nature Reserve, alt. 980 m, 19 May 2021, Xing-Hai Zuo leg. (
This species is similar to B. megesalis. The difference in appearance is that wings of this species are darker in color, and hindwings are slightly broader; the postmedial line and terminal margin of forewing is obviously unparallel in this species, while it is nearly parallel in the latter. It also can be distinguished by the sacculus without long setal cluster, the phallus with one broad, spiral, band-like cornutus; the posterior third of ductus bursae with slightly sclerotized elongate inflation, but not forming irregular protrusion. In B. megesalis, the sacculus has long setal cluster near middle, and the phallus has two spear-like cornuti; the posterior ductus bursae is inflated into an irregular protrusion.
Adult
(Fig.
Male genitalia
(Fig.
Genitalia of Bradina species A–E B. torsiva sp. nov. A, B male, holotype, slide no. GJM21102 C–E female, paratype, slide no. GJM21104 F, G B. subpurpurescens male, slide no. GJM21018 H–M B. falciculata sp. nov. H–J male, holotype, slide no. GJM21084 K–M female, paratype, slide no. GJM21085 B, G, I, J partial enlargement of phallus D, L partial enlargement of ductus bursae E, M signum. Scale bars: 0.2 mm (D, E, J, M); 0.5 mm (A, B, F–I, L); 1.0 mm (C, K).
Female genitalia
(Fig.
The specific name is derived from the Latin torsivus (meaning 'spiral'), in reference to a spiral band-like cornutus.
China (Guangdong, Hunan).
Coremata of some male individuals of this species were protruded out of the body, forming a cluster of white hairs at the end of the abdomen.
Pleonectusa subpurpurescens
Warren, 1896: 147. Type locality: India. Type depository:
Bradina subpurpurescens: Hampson, 1896: 227.
Holotype
, ♂ India: Khasis, X. 1894, Nat. Coll (
China: Yunnan Prov., 1 ♂, Meizihu Park, Simao District, Pu’er City, alt. 1400 m, 11 May 2018, Xi-Cui Du & Xiao-Qiang Lu leg., genitalia slide no.: GJM21019; 1 ♂, Yunpan Mountain, Pu’er City, alt. 1400 m, 9 July 2013, Zhen-Guo Zhang leg., genitalia slide no.: GJM21018; 1 ♂, Taiyanghe National Forest Park, Puer City, alt. 1659 m, 29 June 2021, Yao Shen & Ci Tang leg., genitalia slide no.: GJM21020.
Adult
(Fig.
Male genitalia
(Fig.
China (Yunnan), India.
This species is recorded in China for the first time, and its male genitalia are also described for the first time. It can be distinguished from the other species (except B. falciculata) in the present study by the forewing relatively narrower; cilia pale brown on forewing, yellowish white on hindwing. The female of this species is unknown.
Holotype
, ♂ China: Tibet Autonomous Region, Medog County, alt. 1100 m, 29°32'N, 95°33'E, 14 August 2003, Xin-Pu Wang & Huai-Jun Xue leg. (
This species is similar to B. subpurpurescens. The difference in appearance is that wings are paler in color and the postmedial line of the forewing is placed at two thirds from the wing base in this new species, but at three quarters in the latter. It also can be distinguished by an elliptical valva and the phallus inconspicuously inflated distally. In B. subpurpurescens, the valva is nearly rectangular and the phallus is significantly inflated distally.
Adult
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
The specific name is derived from the Latin falciculatus (meaning 'falcate'), in reference to two sickle cornuti of posterior phallus.
China (Tibet).
Bradina translinealis
Hampson, 1896: 228. Type locality: N. W. Himalayas. Type depository:
Holotype
, ♂. Moore Coll. 94–106, genitalia slide no. 8734. (
China, Yunnan Prov., 5 ♂♂, 2 ♀♀, Baihualing Village, Baoshan City, alt. 1520 m, 11–13 August 2007, Dan-Dan Zhang leg., genitalia slide no.: GJM21165 ♂, GJM21166 ♂, GJM21167 ♀, GJM21182 ♀.
Adult
(Fig.
Male genitalia
(Fig.
Genitalia of Bradina species A–F B. translinealis A–C male, slide no. GJM21165 D–F female, slide no. GJM21167 G–K B. ternifolia sp. nov. G, H male, holotype, slide no. GJM21173 I–K female, paratype, slide no. GJM21178 B, C, H partial enlargement of phallus E, J partial enlargement of ductus bursae F, K signum. Scale bars: 0.2 mm (B, C, F, K); 0.5 mm (A, E, G, H, J); 1.0 mm (D, I).
Female genitalia
(Fig.
China (Yunnan), N. W. Himalayas.
This species is first recorded in China and its genitalia are described for the first time. It can be distinguished by the valva being nearly elliptical, the phallus with one fusiform cornutus medially and two weakly sclerotized lamellar cornuti posteriorly.
Holotype , ♂ China: Yunnan Prov., Dahaoping Village, Tengchong City, 25°02'N, 98°49'E, 6 August 2007, Dan-Dan Zhang leg., genitalia slide no. GJM21173. Paratypes. China: Yunnan Prov., 2 ♂♂, 7 ♀♀, other same data as holotype, genitalia slide no.: GJM21174 ♂, GJM21175 ♂, GJM21176 ♀, GJM21177 ♀, GJM21178 ♀, GJM21181 ♀; 2 ♀♀, 5 August 2007, other same data as holotype.
This species is similar to B. translinealis. It can be distinguished by postmedial line of forewing straight; valva elongated and narrowed comparatively, sacculus with long setal cluster near middle, posterior phallus with three long leaf-like cornuti; antrum membranous. In B. translinealis, postmedial line of forewing slightly excurved near costa; sacculus has no setal cluster, phallus has one fusiform cornutus medially and two lamellar cornuti posteriorly; antrum broader and sclerotized.
Adult
(Fig.
Male genitalia
(Fig.
Female genitalia
(Fig.
The specific name is derived from the Latin ternifolius (meaning 'trifoliate'), in reference to three long leaf-like cornuti.
China (Yunnan).
Coremata of some male individuals of this species were protruded out of the body, forming a cluster of white hairs at the end of the abdomen.
The species in the present study, represented by B. megesalis, have a broad valva, whose costa is arched near the base or middle and accompanied by a cluster of long curved setae. The cornuti of the phallus of these species are diverse and therefore useful for interspecific identification.
We sincerely thank the other members of our laboratory as well as to Jian-Yue Qiu and Hao Xu for their efforts in collecting specimens, to Dr Dan-Dan Zhang (Sun Yat-Sen University, China) for sending specimens, and to Prof. Hou-Hun Li (Nankai University, China) for lending us specimens. Cordial thanks are given to the Natural History Museum, London, United Kingdom) for providing access to the corresponding author for examining type specimens of Bradina. Our special thanks are given to Dr Richard Mally (Czech University of Life Sciences, Czech Republic) and to the other anonymous reviewer for their academic suggestion and detailed revision of our manuscript. The research was funded by the National Natural Sciences Foundation of China (No. 31772500) and the Natural Sciences Foundation Project of Chongqing (No. CSTB2022NSCQ-MSX1164).