Five new species of Bradina Lederer (Lepidoptera, Crambidae) from China, with remarks on the morphology of the genus

Abstract

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.

Full text

Five new species of Bradina Lederer
(Lepidoptera, Crambidae) from China,
with remarks on the morphology of the genus
Jia-Ming Guo1, Xi-Cui Du1
1College of Plant Protection, Southwest University, Chongqing, China
Corresponding author: Xi-Cui Du (duxicui@hotmail.com; duxicui@swu.edu.cn)
Academic editor: R. Mally|Received 30 December 2022|Accepted 30 March 2023|Published 19 April 2023
https://zoobank.org/A3AD0A1D-9178-4B91-BC2C-0C1A1AC65312
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
Abstract
Bradina is a species-rich genus that diers 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 rst time. e images of the habitus and genitalia of
these eight species are provided, with a key to their identication.
Keywords
Genitalia, identication key, morphology, Pyraloidea, Spilomelinae, taxonomy
Introduction
Bradina Lederer, 1863 is the most species-rich genus in the Spilomelinae tribe Steniini
Guenée, 1854, redened by Mally et al. (2019). So far, there are 89 species recorded
in this genus worldwide (Nuss et al. 2003–2023) and they are mainly distributed in
the Oriental and Australian regions. Numerous endemic species are present in the
Australian and Pacic islands.
ZooKeys 1158: 49–67 (2023)
doi: 10.3897/zookeys.1158.99411
https://zookeys.pensoft.net
Copyright Jia-Ming Guo & Xi-Cui Du. This is an open access article distributed under the terms of the Creative Commons Attribution License
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Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
50
In Spilomelinae, Bradina can be dierentiated from most other genera by the fore-
wings with Rs1 anastomosed with Rs2+s3 at the base, which is common in Acentropinae.
erefore, this genus was placed in Hydrocampinae (= Acentropinae Stephens, 1835) for
a long time (Hampson 1896; Hampson 1897; Rothschild 1915; Schaus 1924; Caradja
1925). Inoue (1955) transferred the genus to Pyraustinae (s. l.). Systematics research on
Bradina is inadequate globally besides some early studies (Hampson 1896; Hampson
1897; Yamanaka 1984; Du 2008). Seizmair (2021) recorded Bradina from the Arabian
Peninsula and divided the genus into seven groups according to wing pattern.
Species identication of Bradina is dicult because of their very similar appear-
ance, so the genitalia characteristics are necessary in the identication of most species.
e large spinose crescent-shaped signum of the female genitalia is a diagnostic char-
acteristic of the genus, but shows little dierence among species. In male genitalia, the
valvae and uncus are very diverse in morphology, which is very valuable for interspe-
cic identication. Before this study, 13 species were recorded in China (Lu and Guan
1953; Wang and Speidel 2000; Du 2008). In the present study, eight Bradina species
with externally similar adults and male genitalia morphology are recorded, including
ve new species and two newly recorded species from China.
Materials and methods
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 (NKU). e corresponding author
examined many specimens of Bradina deposited in Natural History Museum, London,
United Kingdom (NHMUK), including some types.
e 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) at-
tached to a stereomicroscope (Leica M205 A).
e preparation of genitalia mainly follows Li and Zheng (1996). Morphological
terminology mainly refers to Maes (1995) as well as Mally and Nuss (2011).
Taxonomic account
Bradina Lederer, 1863
Bradina Lederer, 1863: 424. Type species: Bradina impressalis Lederer, 1863; subse-
quent designation by Hampson 1896.
Erilita Lederer, 1863: 426. Type species: Erilita modestalis Lederer, 1863, by monotypy.
Pleonectusa Lederer, 1863: 426. Type species: Botys admixtalis Walker, 1859; subse-
quent designation by Moore 1884.

Five new species of Bradina, with remarks on the morphology of the genus 51
Trematarcha Meyrick, 1886: 233. Type species: Marasmia erilitalis Felder, Felder &
Rogenhofer, 1875; subsequent designation by Klima 1937.
Diagnosis. Head (Fig. 1A). Frons rounded. Antenna annulated, male with short cilia
ventrally. Labial palpus obliquely upturned, second segment with broad scales ven-
trally, third joint minute and forward, apex blunt. Maxillary palpus liform. orax.
Forewing long and narrow usually; length of cell c. half of wing; discocellulars in-
curved; R from cell at c. four fths above; Rs1 anastomosed with Rs2+s3 at base and
with a long stalk c. two fths of Rs3; Rs2 and Rs3 stalked c. three fths of Rs3; basal
half of Rs4 straight and clearly separated from Rs1+s2+s3; M2, M3 and CuA1 uniformly
from posterior angle of cell at base (except for B. diagonalis Hampson, 1896); CuA2
from cell at three quarters below. Hindwing with length of cell c. one third of wing;
discocellulars incurved; Sc+R and Rs stalked c. one third of Rs; Rs and M1 from ante-
rior angle of cell; M2, M3 and CuA1 uniformly from posterior angle of cell; CuA2 from
cell at four fths below (Fig. 1B). Legs long and slender; middle tibia with outer distal
spur c. half-length of inner spur. Abdomen. Male abdomen long and slender (except
for B. melanoperas Hampson, 1896).
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 round-
ed or oval, inside usually densely studded with tiny spines; signum crescent and dense-
ly spinose, spines on concave side developed.
Remarks. e generic characteristics were summarized by Khan (2000), but the
description about wing venation was incomplete. In addition, the lengths of the maxil-
lary palpus and labial palpus were identical in Khan’s description, while we found that
both have their ends at the same height but they are of dierent lengths. erefore, the
generic characteristics are revised in the present study.
e 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.
e 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. e rst type, represented by B. admix-
talis, 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 character-
istics: 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 trapezoi-
dal, slightly concaved terminally. Juxta nearly rounded, split posteriorly. Phallus
distinctly inated at anterior end. Apophyses anteriores expanded at c. one third
from base.

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
52
Key to Bradina species in the present study based on genitalia
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 pos-
teriorly. 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 me-
dially ................................................................................ B. falciculata sp. nov.
Bradina megesalis (Walker, 1859)
Figs 1A, 2A, 3A–E
Botys megesalis Walker, 1859: 663. Type locality: North China. Type depository:
NHMUK.
Bradina megesalis: Hampson, 1897: 200.
Material examined. Holotype, ♂ North China, from Mr Fortune’s collection (Walker,
1859), genitalia slide no. 8735 (NHMUK).
Additional material. 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 ♀♀, Da-
dong 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

Five new species of Bradina, with remarks on the morphology of the genus 53
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.
Redescription. Adult (Figs 1A, 2A). Wingspan 31.0–39.0 mm, forewing length
15.0–19.0 mm. Body and wings pale brown. Frons brown, with lateral sides white
above. Vertex yellowish white. Antenna brownish yellow, with black ring dorsally; ven-
tral cilia c. one quarter length of diameter of male agellomeres. Labial palpus with
basal two thirds white, black-brown distally. Maxillary palpus yellowish brown, white
at base. Patagium pale brown, yellowish. orax white ventrally. Forewing pale brown,
darker along basal two thirds costa, stigmata and lines brown; discoidal stigma crescent;
postmedial line at c. two thirds of wing, straight and nearly parallel to terminal margin.
Hindwing with postmedial line and discoidal stigma pale brown, inconspicuous usu-
ally; discoidal stigma crescent. Cilia pale brown, yellowish white on inner margin of
hindwing. Legs pale yellow, coxae and femora with white gloss. Middle tibia brown;
Figure 1. Bradina megesalis, male A head B wing venation, wing slide no. GJM21001. Scale bars:
0.5 mm (A); 1.0 mm (B).

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
54
hind tibia with outer middle spurs c. two thirds length of inner spurs. Abdomen pale
brown dorsally, with each segment pale terminally; yellowish white ventrally.
Male genitalia (Fig. 3A, B). Valva distally gradually narrowed and bearing dense
long setae; costa sharply arched near middle and accompanied by a cluster of long
curved setae; sacculus gradually narrowed to apex, with a cluster of long setae near
middle. Posterior phallus with two developed spear-like cornuti.
Female genitalia (Fig. 3C–E). Antrum broad. Ductus bursae posteriorly sharply
inated into a thorny irregular protrusion, adjoined posteriorly by crescent collicu-
lum laterally, anterior half gradually widened to corpus bursae. Corpus bursae nearly
rounded, with dense tiny spines inside, transverse signum crescent and densely spinose.
Distribution. China (Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou,
Hubei, Hainan, Hunan, Sichuan, Shaanxi, Yunnan, Zhejiang), Japan (Shibuya 1929).
Remarks. 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.
Bradina fusoidea Guo & Du, sp. nov.
https://zoobank.org/CEACA386-3CD0-4FFF-85F5-1140F4B86FEF
Figs 2B, 3F–K
Type material. Holotype, ♂ China: Sichuan Prov., Qingcheng Mountain, Dujiang-
yan city, alt. 860 m, 30°92'N, 103°50'E, 22 July 2021, Shuai Yu, Xiao-Ju Zhu, Di
Zhang leg. (NKU), genitalia slide no. GJM21117. Paratypes. China: Sichuan Prov.,
6 ♂♂, 1 ♀, other same data as holotype (NKU), genitalia slide no.: GJM21112 ♂,
GJM21118 ♀; 1 ♂, Emei Mountain, Leshan City, alt. 847 m, 22 July 2021, Shuai
Yu, Xiao-Ju Zhu, Di Zhang leg. (NKU), genitalia slide no.: GJM21113; Guangdong
Prov. , 2 ♂♂, 1 ♀, Heishiding Nature Reserve, Fengkai County, Zhaoqing City, 14–16
June 2009, Feng-Xia He leg., genitalia slide no.: GJM21114 ♂, GJM21180 ♀.
Diagnosis. is species is similar to B. megesalis. e dierence 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 inated but not form-
ing 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.
Description. Adult (Fig. 2B). Wingspan 31.0–33.0 mm, forewing length 15.0–
16.0 mm. Frons white, except brown on frontal base and middle near vertex. Vertex
white. Antenna yellow, with pale brown ring dorsally; ventral cilia c. half-length of
agellomere diameter of male. Labial palpus with basal two thirds white, black-brown
distally. Maxillary palpus black-brown. Patagium yellowish white. Tegula pale brown.
orax pale brown dorsally, white ventrally. Wings pale brown, gradually darkened
to terminal; stigmata and lines dark brown. Forewing black-brown along basal half of

Five new species of Bradina, with remarks on the morphology of the genus 55
costa; orbicular stigma very small; discoidal stigma crescent; postmedial line at c. two
thirds of wing, straight and unparallel to terminal margin. Hindwing with postmedial
line straight, towards to tornus, only middle part obvious. Cilia pale brown, with a
Figure 2. Habitus of Bradina species, male A B. megesalis B B. fusoidea sp. nov., holotype C B. spirella
sp. nov., holotype D B. torsiva sp. nov., holotype E B. subpurpurescens F B. falciculata sp. nov., holotype
G B. translinealis H B. ternifolia sp. nov., holotype. Scale bars: 0.5 cm.

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
56
white line at base, except yellowish white on inner margin of the hindwing. Legs pale
yellow. Front and middle tibiae brown; hind tibia with outer middle spurs c. two thirds
length of inner spurs. Abdomen with basal half pale brown and distal half dark brown
dorsally, each segment pale terminally; yellowish white ventrally.
Male genitalia (Fig. 3F–H). Valva distally gradually narrowed and bearing dense
long setae; costa arched near middle and accompanied by a cluster of long curved setae;
sacculus gradually narrowed to apex, with a cluster of long setae near middle. Phallus
with one fusiform cornutus medially, composed of short and blunt spines, c. one third
length of phallus, and one subcircular sclerotized cornutus posteriorly.
Female genitalia (Fig. 3I–K). Antrum broad. Posterior ductus bursae inated and
thorny, adjoined posteriorly by colliculum, posterior half inated, weakly sclerotized,
narrowed medially, then gradually widened to corpus bursae. Corpus bursae nearly
rounded, with dense tiny spines inside, transverse signum crescent and densely spinose.
Etymology. e specic name is derived from the Latin fusoideus (meaning ‘fusi-
form’), in reference to a fusiform cornutus.
Distribution. China (Guangdong, Sichuan).
Bradina spirella Guo & Du, sp. nov.
https://zoobank.org/2AF3F855-1103-42AA-94DD-D6634879BB67
Figs 2C, 3L–P
Type material. 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 ♀.
Diagnosis. is species is similar to B. megesalis. e dierence 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 spe-
cies, 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 inated into a thick nger-like protru-
sion. In B. megesalis, the phallus has two spear-like cornuti and is without spines on the
vesica; the posterior ductus bursae is inated into an irregular protrusion.
Description. Adult (Fig. 2C). Wingspan 29.0–32.0 mm, forewing length 14.0–
15.5 mm. Frons brown, with lateral sides yellowish white above. Vertex pale yellow.
Antenna brownish yellow, with black-brown ring dorsally, basal segments of agellum

Five new species of Bradina, with remarks on the morphology of the genus 57
Figure 3. 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, para-
type, 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).

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
58
black-brown dorsally; ventral cilia c. one third length of agellomere diameter of male.
Labial palpus with basal two thirds white, black-brown distally. Maxillary palpus
black-brown. Patagium and tegula dark brown. orax dark brown dorsally, white
ventrally. Wings dark brown, stigmata and lines black-brown. Forewing black-brown
along costa, slightly paler distally; orbicular stigma very small; discoidal stigma cres-
cent; postmedial line at c. two thirds of wing, inconspicuously waved and unparallel
to terminal margin. Hindwing pale at base; postmedial line straight, only middle part
obvious. Cilia greyish brown. Legs pale yellow. Front and middle tibiae dark brown;
hind tibia with outer middle spurs c. same length as inner spurs. Abdomen brown
dorsally, with each segment pale terminally; yellowish white ventrally.
Male genitalia (Fig. 3L, M). Valva with the distal part gradually narrowed and bear-
ing dense long setae; costa arched near middle and accompanied by a cluster of long curved
setae; sacculus gradually narrowed to apex, with a cluster of long setae near middle. Juxta
narrowed distally. Phallus with one short and spiral band-like cornutus tapered at posterior
end, along with two lamellar cornuti in posterior half; some tiny spines on vesica medially.
Female genitalia (Fig. 3N–P). Antrum broad. Ductus bursae slender, posteriorly
inated into a thick nger-like protrusion, adjoined posteriorly by colliculum. Corpus
bursae nearly oval, without spines inside, transverse signum crescent and densely spinose.
Etymology. e specic name is derived from the Latin spirellus (meaning ‘small
spiral-like’), in reference to a short and spiral band-like cornutus.
Distribution. China (Hunan, Jiangxi).
Remark. 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.
Bradina torsiva Guo & Du, sp. nov.
https://zoobank.org/3DF230C3-6BA8-4D7F-AB54-61C46C1CE689
Figs 2D, 4A–E
Type material. 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., genita-
lia 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 Na-
ture Reserve, alt. 980 m, 19 May 2021, Xing-Hai Zuo leg. (NKU), genitalia slide no.:
GJM21099 ♂; 5 ♂♂, 2 ♀♀, Yangmeikeng Village, Shimentai National Nature Reserve,
alt. 870 m, 27 May 2021, Xing-Hai Zuo leg. (NKU), genitalia slide no.: GJM21104 ♀.
Diagnosis. is species is similar to B. megesalis. e dierence in appearance is that
wings of this species are darker in color, and hindwings are slightly broader; the postme-

Five new species of Bradina, with remarks on the morphology of the genus 59
dial 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 duc-
tus bursae with slightly sclerotized elongate ination, but not forming irregular protru-
Figures 4. 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 ).

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
60
sion. In B. megesalis, the sacculus has long setal cluster near middle, and the phallus has
two spear-like cornuti; the posterior ductus bursae is inated into an irregular protrusion.
Description. Adult (Fig. 2D). Wingspan 36.0–37.0 mm, forewing length 17.5–
18.0 mm. Frons brown, with lateral sides yellowish white above. Vertex yellowish white.
Antenna brownish yellow, with black-brown ring dorsally, ventral cilia c. half-length of
agellomere diameter of male. Labial palpus with basal two thirds white, black-brown
distally. Maxillary palpus black-brown or brown. Patagium yellowish white. Tegula
dark brown. orax dark brown dorsally, white ventrally. Wings dark brown, stigmata
and lines black-brown. Forewing black-brown along costa, slightly paler distally; or-
bicular stigma very small; discoidal stigma crescent; postmedial line at c. two thirds
of wing, unparallel to terminal margin. Hindwing slightly pale at base; postmedial
line inconspicuously waved, only middle part obvious. Cilia pale yellow, with a black-
brown line at base, except black-brown on inner margin. Legs yellow. Front and mid-
dle tibiae dark brown; hind tibia with outer middle spurs c. three fths length of inner
spurs. Abdomen brown dorsally, each segment pale terminally; paler ventrally.
Male genitalia (Fig. 4A, B). Valva with the distal part gradually narrowing and
with dense long setae; costa arched near middle and accompanied by a cluster of long
curved setae; sacculus narrowed distally, without long setal cluster. Juxta narrowed dis-
tally. Phallus slightly inated, with one broad and spiral band-like cornutus posteriorly.
Female genitalia (Fig. 4C–E). Ductus bursae broad, with posterior third inated,
sclerotized, and thorny, adjoined posteriorly by colliculum, gradually widened to cor-
pus bursae. Corpus bursae nearly oval, truncated terminally, with dense tiny spines
inside, transverse signum crescent and densely spinose.
Etymology. e specic name is derived from the Latin torsivus (meaning 'spiral'),
in reference to a spiral band-like cornutus.
Distribution. China (Guangdong, Hunan).
Remark. 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.
Bradina subpurpurescens (Warren, 1896)
Figs 2E, 4F, G
Pleonectusa subpurpurescens Warren, 1896: 147. Type locality: India. Type depository:
NHMUK.
Bradina subpurpurescens: Hampson, 1896: 227.
Material examined. Holotype, ♂ india: Khasis, X. 1894, Nat. Coll (NHMUK).
Additional material. 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.

Five new species of Bradina, with remarks on the morphology of the genus 61
Redescription. Adult (Fig. 2E). Wingspan 26.0–33.0 mm, forewing length 14.5–
16.0 mm. Frons brown, with lateral sides white above. Vertex brownish yellow. An-
tenna brownish yellow, with pale brown ring dorsally; length of ventral cilia c. one
quarter of the agellomere diameter of male. Labial palpus with basal two thirds white,
black-brown distally. Maxillary palpus black-brown, pale brown at base. Patagium and
tegula brown. orax pale brown distally, white ventrally. Wings dark brown, stigmata
and lines black-brown. Forewing black-brown along basal costa; orbicular stigma very
small; discoidal stigma crescent; postmedial line at c. three quarters of wing, nearly
parallel to terminal margin, slightly excurved near costa. Hindwing with postmedial
line slightly beyond basal half of wing, only middle part obvious. Cilia pale brown on
forewing, yellowish white on hindwing; a brown line at base. Legs pale yellow. Front
and middle tibiae yellowish brown; hind tibia with outer middle spurs c. half-length
of inner spurs. Abdomen pale yellowish brown dorsally, each segment pale terminally.
Male genitalia (Fig. 4F, G). Valva nearly rectangular, broad distally; costa sharply
arched near base and accompanied by a cluster of long curved setae, then slightly
concave towards apex; sacculus gradually narrowed distally, without long setal cluster.
Phallus narrow medially, signicantly inated anteriorly and posteriorly, posterior half
sclerotized; cornutus absent.
Distribution. China (Yunnan), India.
Remarks. is species is recorded in China for the rst time, and its male genitalia
are also described for the rst 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. e female of this species is unknown.
Bradina falciculata Guo & Du, sp. nov.
https://zoobank.org/9F1E4111-1E29-42C2-AD1D-300EAC54E761
Figs 2F, 4H–M
Type material. 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. (NKU), genitalia slide no. GJM21084. Paratypes. China: Tibet Autonomous
Region, 5 ♂♂, 3 ♀♀, other same data as holotype, genitalia slide no.: GJM21082
♂; 5 ♂♂, 8 ♀♀, Bomi-Medog Highway, Medog County, alt. 880 m, 14 August
2003, Xin-Pu Wang & Huai-Jun Xue leg. (NKU), genitalia slide no.: GJM21083 ♂,
GJM21085 ♀, GJM21086 ♀.
Diagnosis. is species is similar to B. subpurpurescens. e dierence in appear-
ance 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 lat-
ter. It also can be distinguished by an elliptical valva and the phallus inconspicuously
inated distally. In B. subpurpurescens, the valva is nearly rectangular and the phallus is
signicantly inated distally.
Description. Adult (Fig. 2F). Wingspan 25.0–27.0 mm, forewing length 12.0–
13.0 mm. Frons brown, with lateral sides yellowish white above. Vertex yellowish white. An-

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
62
tenna yellow, with pale brown ring dorsally; ventral cilia c. half-length of agellomere diam-
eter of male. Labial palpus with basal two thirds white, dark brown distally. Maxillary palpus
dark brown. Patagium and tegula pale brown. orax pale brown dorsally, white ventrally.
Wings yellowish brown, stigmata and lines brown. Forewing with discoidal stigma crescent;
postmedial line at basal two thirds of wing, slightly excurved and nearly parallel to terminal
margin. Hindwing with postmedial line straight, usually not obvious. Cilia pale brown,
with a darker line at base, except greyish white on inner margin of the hindwing. Legs pale
yellow. Front and middle tibiae black-brown on distal half; hind tibia with outer middle
spurs c. half-length of inner spurs. Abdomen pale brown dorsally, yellowish white ventrally.
Male genitalia (Fig. 4H–J). Valva elliptical; costa arched near middle and accom-
panied by a cluster of long curved setae; sacculus gradually narrowed to apex, without
long setal cluster. Juxta narrowed distally. Phallus with one small fusiform cornutus
anteriorly, composed of short and blunt spines, c. one sixth length of phallus, and with
two sickle-shaped cornuti posteriorly.
Female genitalia (Fig. 4K–M). Antrum broad. Ductus bursae with one oval scle-
rotized piece close to crescent-shaped colliculum, then inated and thorny, slightly
sclerotized medially. Corpus bursae nearly oval, with dense tiny spines inside, trans-
verse signum crescent and densely spinose.
Etymology. e specic name is derived from the Latin falciculatus (meaning
'falcate'), in reference to two sickle cornuti of posterior phallus.
Distribution. China (Tibet).
Bradina translinealis Hampson, 1896
Figs 2G, 5A–F
Bradina translinealis Hampson, 1896: 228. Type locality: N. W. Himalayas. Type de-
pository: NHMUK.
Material examined. Holotype, ♂. Moore Coll. 94–106, genitalia slide no. 8734.
(NHMUK).
Additional material. 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 ♀.
Redescription. Adult (Fig. 2G). Wingspan 25.0–30.0 mm, forewing length
12.0–13.0 mm. Frons brown, with lateral sides yellowish white above. Vertex yellow-
ish white. Antenna brownish yellow, with brown ring dorsally; ventral cilia c. quarter
length of agellomere diameter of male. Labial palpus with basal two thirds white,
black-brown distally. Maxillary palpus black-brown, yellowish white at base. Patagium
and tegula pale brown. orax pale brown dorsally, white ventrally. Forewing yellowish
brown, dark brown along basal half of costa; stigmata and lines black-brown; discoidal
stigma reniform; postmedial line at c. two thirds of wing, slightly excurved near costa.
Hindwing pale yellowish brown dorsally, yellowish white to pale brown ventrally; paler

Five new species of Bradina, with remarks on the morphology of the genus 63
at base; postmedial line pale brown, inconspicuous, extending beyond basal half of
wing. Cilia pale brown on forewing, yellowish white on hindwing, a brown line at
base. Legs yellowish white. Front and middle tibiae brown; hind tibia with outer mid-
dle spurs c. half-length of inner spurs. Abdomen pale yellowish brown dorsally, yellow-
ish white ventrally.
Male genitalia (Fig. 5A–C). Valva nearly elliptical, bearing dense long setae distal-
ly; costa arched near middle and accompanied by a cluster of long setae; sacculus grad-
ually narrowed distally, without long setal cluster. Phallus with one fusiform cornutus
medially, composed of short and blunt spines, c. one fth length of phallus; posterior
half slightly inated, and with two weakly sclerotized lamellar cornuti posteriorly.
Figure 5. 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 ).

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
64
Female genitalia (Fig. 5D–F). Antrum broad and sclerotized, strongly sclerotized
laterally, anteriorly adjoined by colliculum. Posterior ductus bursae slender, inated,
widened close to corpus bursae. Corpus bursae nearly oval, truncated terminally; with
dense tiny spines inside, transverse signum crescent and densely spinose.
Distribution. China (Yunnan), N. W. Himalayas.
Remarks. is species is rst recorded in China and its genitalia are described
for the rst time. It can be distinguished by the valva being nearly elliptical, the phal-
lus with one fusiform cornutus medially and two weakly sclerotized lamellar cor-
nuti posteriorly.
Bradina ternifolia Guo & Du, sp. nov.
https://zoobank.org/F49AC3BF-5605-4538-A1AF-145EA7B517FF
Figs 2H, 5G–K
Type material. 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 holo-
type, genitalia slide no.: GJM21174 ♂, GJM21175 ♂, GJM21176 ♀, GJM21177 ♀,
GJM21178 ♀, GJM21181 ♀; 2 ♀♀, 5 August 2007, other same data as holotype.
Diagnosis. is species is similar to B. translinealis. It can be distinguished by
postmedial line of forewing straight; valva elongated and narrowed comparatively, sac-
culus 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.
Description. Adult (Fig. 2H). Wingspan 31.0–33.0 mm, forewing length 15.0–
16.0 mm. Frons brown, except black-brown above. Vertex yellowish white mixed
with brownish yellow. Antenna pale yellow, with pale brown ring dorsally; ventral
cilia c. half-length of agellomeres diameter of male. Labial palpus with basal half
yellowish white, brown or black-brown distally. Maxillary palpus brown or black-
brown. Patagium and tegula dark brown. orax dark brown dorsally, white ven-
trally. Wings brown, stigmata and lines black-brown. Forewing black-brown along
costa; discoidal stigma reniform; postmedial line at c. two thirds of wing, straight
and nearly parallel to terminal margin. Hindwing with discoidal stigma crescent;
postmedial line beyond basal half of wing, usually inconspicuous. Cilia pale brown
on forewing, greyish white on hindwing, a darker line at base. Legs pale yellow. Front
and middle tibiae brown; hind tibia with outer middle spurs c. three fths length of
inner spurs. Abdomen dark brown dorsally, each segment pale terminally; slightly
pale ventrally.
Male genitalia (Fig. 5G, H). Valva gradually narrowed and bearing dense long
setae distally; costa arched near base and accompanied by a cluster of short and curved
setae; sacculus gradually narrowed to apex, with a cluster of long setae near middle.
Posterior phallus inated slightly, with three long leaf-like cornuti tapered apically.

Five new species of Bradina, with remarks on the morphology of the genus 65
Female genitalia (Fig. 5I–K). Antrum membranous. Ductus bursae slender, adjoined
posteriorly by crescent colliculum laterally, then inated and bent. Corpus bursae nearly
oval, with dense tiny spines inside, transverse signum crescent and densely spinose.
Etymology. e specic name is derived from the Latin ternifolius (meaning
'trifoliate'), in reference to three long leaf-like cornuti.
Distribution. China (Yunnan).
Remark. 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.
Discussion
e 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. e cornuti of the phallus of these species are diverse and therefore useful for in-
terspecic identication. Seizmair (2021) divided Bradina into seven groups according
to wing pattern characteristics, such as forewing and hindwing of the same color or of
dierent colors, and whether the postmedial line is absent or present, and in the latter
case whether it is straight or curved. In our opinion, however, it is dicult to divide the
genus based only on wing pattern and color, and it would be more convincing to divide
it by combining the appearance and morphology of the genitalia. In addition, most spe-
cies of the B. diagonalis group in Seizmair’s study are similar to the species in the present
study both in appearance and genitalia. However, we found that B. diagonalis, as repre-
sentative species of the B. diagonalis group, is dierent from most other species of this
group in wing venation and genitalia. erefore, further study on more species globally
and in more detail is needed to clarify this confusion. In addition, the transverse cres-
cent signum of Bradina is a distinctive feature. Like Bradina, the Steniini genera Diath-
rausta and Perisyntrocha also have an arched, transverse signum in the female genitalia’s
corpus bursae (Mally et al. 2019). is could indicate an evolutionary relatedness of
these three genera, which is expected to be addressed in the future research.
Acknowledgements
We sincerely thank the other members of our laboratory as well as to Jian-Yue Qiu and
Hao Xu for their eorts 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 Mu-
seum, 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 review-
er for their academic suggestion and detailed revision of our manuscript. e research was
funded by the National Natural Sciences Foundation of China (No. 31772500) and the
Natural Sciences Foundation Project of Chongqing (No. CSTB2022NSCQ-MSX1164).

Jia-Ming Guo & Xi-Cui Du / ZooKeys 1158: 49–67 (2023)
66
References
Caradja AV (1925) Ueber Chinas Pyraliden, Tortriciden, Tineiden nebst kurze Betrachtun-
gen, zu denen das Studium dieser Fauna Veranlassung gibt (Eine biogeographische Skizze).
Memoriile Sectiunii Stiintice, Academia Romana seria 3 3(7): 257–383. [pls 1–2]
Du XC (2008) Taxonomic Study on Spilomelinae from China (Lepidoptera: Pyraloidea: Cram-
bidae). PhD esis, Nankai University, Tianjing, China.
Felder C, Felder R, Rogenhofer AF (1875) Heterocera. In: Felder C, Felder R, Rogenhofer AF,
Reise der österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859
unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer eil. Band
2. Abtheilung 2. Lepidoptera. Heft 5 (Schluss). Aus der Kaiserlich-Königliche Hof- und
Staatsdruckerei, Wien, 1–20. [pls 121–140]
Guenée MA (1854) Deltoïdes et Pyralites. In: de Boisduval JBAD, Guenée MA (Eds) Histoire
Naturelle des Insectes. Species Général des Lépidoptères, 8. Roret, Paris, 448 pp.
Hampson GF (1896) Moths. e Fauna of British India, including Ceylon and Burma, Lon-
don, Vol. 4, 594 pp.
Hampson GF (1897) On the classication of two subfamilies of moths of the family Pyralidae:
e Hydrocampinae and Scoparianae. e Transactions of the Entomological Society of
London 45(2): 127–240. https://doi.org/10.1111/j.1365-2311.1897.tb01679.x
Inoue H (1955) Pyralidae. Check List of the Lepidoptera of Japan, Vol. 2. Rikusuisha, Tokyo,
120–196.
Khan ZH (2000) Biosystematic Studies on Pyralid Pests of Rice Crop in India (Pyralidae:
Lepidoptera). PhD esis, Aligarh Muslim University, Aligarh, India. http://hdl.handle.
net/10603/52039
Klima A (1937) Pyralidae: Subfam.: Scopariinae et Nymphulinae. In: Bryk F (Ed.) Lepidop-
terorum Catalogus 84. Dr. W. Junk, ‘s-Gravenhage, 226 pp. https://doi.org/10.5962/bhl.
title.124187
Lederer J (1863) Beitrag zur Kenntniss der Pyralidinen. Wiener Entomologische Monatschrift
7(8, 10–12): 243–280, 331–504. [pls 2–18]
Li HH, Zheng ZM (1996) Methods and techniques of specimens of Microlepidoptera. Journal
of Shaanxi Normal University 24(3): 63–70. [natural science edn.]
Lu JR, Guan ZH (1953) List of Chinese pyralid insects, addendum of Hu “List of Chinese
Insects”, part 2, Nymphulinae, Scopariinae, Anerastiinae. Acta Entomologica Sinica 3(2):
203–244.
Maes KVN (1995) A comparative morphological study of the adult Crambidae (Lepidoptera,
Pyraloidea). Bulletin et Annales de la Société Royale Belge d’Entomologie 131: 383–434.
Mally R, Nuss M (2011) Molecular and morphological phylogeny of European Udea moths
(Insecta: Lepidoptera: Pyraloidea). Arthropod Systematics & Phylogeny 69(1): 55–71.
Mally R, Hayden JE, Neinhuis C, Jordal BH, Nuss M (2019) e phylogenetic systematics
of Spilomelinae and Pyraustinae (Lepidoptera: Pyraloidea: Crambidae) inferred from
DNA and morphology. Arthropod Systematics & Phylogeny 77(1): 141–204. https://doi.
org/10.26049/ASP77-1-2019-07

Five new species of Bradina, with remarks on the morphology of the genus 67
Meyrick E (1886) Descriptions of Lepidoptera from the South Pacic. e Trans-
actions of the Entomological Society of London 34(3): 189–296. https://doi.
org/10.1111/j.1365-2311.1886.tb01626.x
Moore F (1884) e Lepidoptera of Ceylon, Vol. 3. Reeve L and Co., London, 578 pp.
[pls 144–215]
Nuss M, Landry B, Mally R, Vegliante F, Tränkner A, Bauer F, Hayden J, Segerer A, Schouten
R, Li H, Tromova T, Solis MA Prins, De J, Speidel W (2003–2023) Global Information
System on Pyraloidea. http://www.pyraloidea.org [Accessed 2023-03-29]
Rothschild LW (1915) On the Lepidoptera from the islands of Ceram (Seram), Buru, Bali, and
Misol. Novitates Zoologicae 22: 209–227. https://doi.org/10.5962/bhl.part.1914
Schaus W (1924) New species of Pyralidae of the subfamily Nymphulinae from tropical America
(Lepid.). Proceedings of the Entomological Society of Washington 26(5): 93–130.
Seizmair M (2021) First records of the genus Bradina lederer, 1863 (Lepidoptera, Crambidae,
Spilomelinae) from the Arabian Peninsula with remarks on the taxonomy of the genus
and description of a new species. International Journal of Entomological Research 6(6):
307–312.
Shibuya J (1929) On the Japanese Hydrocampinae. Insecta Matsumurana 3(2–3): 120–136.
Stephens JF (1835) Illustrations of British Entomology; or, a synopsis of indigenous insects:
containing their generic and specic distinctions; with an account of their metamorphoses,
times of appearance, localities, food, and economy, as far as practicable. Baldwin and
Cradock, London, 3–240. [pls 28–34]
Walker F (1859) Pyralides. List of the Specimens of Lepidopterous Insects in the Collection of
the British Museum, London 18: 509–798.
Wang HY, Speidel W (2000) Pyraloidea (Pyralidae, Crambidae). Guide book to insects in
Taiwan (19). Shu Shin Books, Taipei, 295 pp.
Warren W (1896) New genera and species of Pyralidae, yrididae, and Epiplemidae. Annals &
Maga zine of Natu ra l Histor y 17(98): 131–150. https://doi.org/10.1080/00222939608680339
Yamanaka H (1984) Revisional study of some species of Bradina Lederer from Japan, China
and Taiwan (Lepidoptera: Pyralidae, Pyraustinae). Tinea 11(19): 161–176.