Research Article
Research Article
A new species of the genus Ivela Swinhoe (Lepidoptera, Erebidae, Lymantriinae) from Guangdong, China
expand article infoLin-Zhe Xie, Kun-Yuan Li§, Liu-Sheng Chen§, Hou-Shuai Wang
‡ South China Agricultural University, Guangzhou, China
§ Guangdong Academy of Forestry, Guangzhou, China
Open Access


Ivela yini sp. nov., is described from Guangdong, China based on morphological characters and molecular data. Adults, including genitalia and wing venation, and pupa are illustrated and compared to those of similar species. A key to Chinese Ivela species is provided. Assignment of the new species to Ivela Swinhoe is based primarily on a molecular phylogenetic analysis and is corroborated by morphology. Life histories of I. yini and Dendrophleps semihyalina Hampson are discussed.


adults, Dendrophleps, Leucomini, molecular data, phylogenetic analyses, pupae


The tribe Leucomini of Lymantriinae (Erebidae), proposed by Hollоway (1999), is mainly distributed in the Oriental tropics and contains approximately 60 species in four genera (Wang et al. 2015). Adults of this tribe can be recognized by their appearance, often pale white, with translucent areas in male wings, and asymmetric male genitalia (Holloway 1999). Prior to this study, Ivela Swinhoe contained three species: I. auripes (Butler), I. ochropoda (Eversmann), and I. eleuterioides (Semper). Of these, I. auripes and I. ochropoda occur in China.

We report the discovery of a previously unrecognized species of tussock moth that resembles Dendrophleps semihyalina Hampson. Based on multiple morphological characters of adults and pupae and molecular data of four gene regions, we describe it as a species of Ivela.

Materials and methods

Collecting and morphology

All examined specimens were collected in light traps. They are deposited in the Insect Collection of Department of Entomology, South China Agricultural University (SCAU), Guangzhou, China. Adults and genitalia were treated following Wang et al. (2010, 2014). Terminology follows Holloway (1999) and Chao (2003).

Molecular taxa sampling

We sampled six species, including the type species of all genera of Leucomini, with two species of Lymantria Hübner as outgroups. Most sequences of Leucomini and those of the outgroup taxa were downloaded from NCBI. The detailed sampling data for molecular analyses are provided in Table 1.

Molecular data analyses

DNA was extracted from two or three legs of adult specimens using the TIANGEN DNA extraction kit following the manufacturer’s instructions. One mitochondrial gene, DNA barcode region of cytochrome c oxidase subunit I (COI), and three nuclear genes, Elongation factor-1 alpha (EF-1α), ribosomal protein S5 (RpS5), and wingless (WNT), were amplificated and sequenced following Folmer et al. (1994) and Wahlberg and Wheat (2008). Concatenation and sequence alignment was performed using MEGA X (Kumar et al. 2018).

A neighbor-joining (NJ) analysis of DNA barcode was performed with MEGA X under the Kimura 2-Parameter (K2P) model (Kimura 1980), and bootstrap values were calculated with 1,000 replicates. A maximum-likelihood (ML) analysis was performed using IQ-TREE (Nguyen et al. 2015) with 1,000 bootstrap replicates, and the best-fitting model was automatically selected by ModelFinder (Kalyaanamoorthy et al. 2017) implemented in IQ-TREE. A Bayesian-inference (BI) analysis was performed using MrBayes 3.2.6 (Ronquist et al. 2012) under the GTR + F + G4 model, with two parallel runs for 2,000,000 generations. The first 25% of trees were discarded as burn-in, and the remaining trees were used to calculate posterior probabilities (PP).

Table 1.

Sampling data used for molecular analyses in this study.

Specimen voucher no. Taxa Locality GenBank accession no.
LE114 Ivela yini sp. nov. Guangdong, China OM242956 #
LE074 Ivela yini sp. nov. Guangdong, China OM242952 #
LE118 Ivela yini sp. nov. Guangdong, China OM242955 #
H340 Ivela yini sp. nov. China KP081829.1 KP082270.1 KP082761.1
LE124 Ivela auripes* Guangdong, China OM242951 #
H49 Ivela auripes* China KP081830.1 KP082302.1 KP082762.1
H181 Perina nuda* Guangdong, China KP081831.1 KP082248.1 KP082623.1 KP082763
LE014 Dendrophleps semihyalina* Guangdong, China OM250083 # OM328195 # OM328197 # OM328196 #
LE115 Dendrophleps semihyalina* Guangdong, China OM242954 #
LE116 Dendrophleps semihyalina* Guangdong, China OM242953 #
GD385 Dendrophleps semihyalina* Guangdong, China OM242949 #
H377 Leucoma sp. China KP081825.1 KP082289.1 KP082620.1 KP082757.1
H351 Leucoma salicis* China KP081826.1 KP082276.1 KP082621.1 KP082758.1
H127 Lymantria dissoluta China KP081854.1 KP082225 KP082643.1 KP082781
H58 Lymantria similis China KP081855.1 KP082304.1 KP082644.1 KP082782.1


Phylogenetic relationships

The genetic distances of the DNA barcode data (a 658 bp region of the COI gene) of Leucomini species in China are given in Appendix 1. The interspecific genetic distances within Ivela ranged from 10.6 to 12.2% (I. yini sp. nov. and I. auripes); the intraspecific genetic distances from 1.1% (I. yini) to 1.9% (D. semihyalina); and the intergeneric genetic distances within Leucomini ranged from 12.0% (I. yini and Perina nuda (Fabricius)) to 19.3% (Leucoma salicis (Linnaeus) and D. semihyalina). The concatenated dataset of four genes consists of 2,851 nucleotide positions (658 bp for COI, 400 bp for WNT, 600 bp for RPS5 and 1,193 bp for EF-1α). The NJ analysis of the DNA barcode data indicates that the new species and I. auripes (the type species of Ivela) form a clade in Leucomini (Fig. 1). This clade is strongly supported by both BI and ML analyses of the concatenated dataset (Fig. 2: BP = 1.00, PP = 87).

A key to Ivela from China

1 Forewings with R3 and R4 coincident I. auripes
Forewings with R3 and R4 separated at near apex 2
2 Palpi white I. yini
Palpi yellow I. ochropoda

Species accounts

Ivela yini Xie & Wang, sp. nov.

Figs 3–6, 9–11, 14, 15–16, 21–24


This new species is diagnosed by a combination of characters. Superficially, the thorax and abdomen of the adult are white without black markings (Figs 3–6, 9, 10), the palpi are white (Fig. 11), and the forelegs are orange with white rings on the tarsal segments (Figs 9–11). In the male, the asymmetrical valvae are wide and truncated, with a deeply concave cucullus (Fig. 15). The uncus of I. yini (Fig. 15) is more than twice as long as the uncus of I. auripes (Fig. 17) and I. ochropoda (Inoue 1956: fig. 25). The female corpus bursae of I. yini has a pair of caudal projections (Fig. 16). The pupa has white hairs on the prothorax, on segments A2 and A3, and near the posterior end (Figs 21–24).

Figure 1. 

NJ tree of the selected samples of Leucomini based on DNA barcode data. Numbers near nodes represent support values.

Figure 2. 

BI tree of the selected species of Leucomini inferred from the combined COI, EF-1α, RPS5, and WNT genes. Posterior probabilities from BI analysis and bootstrap values from ML analysis are indicated above the branches as PP/BP.

Dendrophleps semihyalina has black markings on the thorax and abdomen (Figs 7, 8, 12, 13), and its valvae are long, narrow, and without a deeply concave cucullus (Fig. 19). The corpora bursae of I. auripes and D. semihyalina lack caudal projections (Figs 18, 20).


Adult (Figs 3–6, 9–11, 1416).

Head (Fig. 11). Antennae bipectinate, fuscous; frons and vertex covered densely with white hairs; labial palpi white, short.

Thorax (Figs 3–6, 9, 10, 14). Dorsum and venter covered with white scales, tegula white. Forewing length: 39‒42 mm male, 48‒50 mm female. Forewings translucent with dense white scales at basal area in male, white in female; R1 and R2 almost parallel, R3, R4, and R5 stalked, M1 arising from upper angle of discal cell, M2 and M3 arising from the lower angle of discal cell respectively, Cu1 and Cu2 approximately parallel, fringe white. Hindwings white, with a transparent area near apex in male; Rs and M1 short stalked, M2 and M3 short stalked in male but arising separately from the lower angle of discal cell in female, fringe white. Forelegs densely covered with orange scales, tarsi with white rings; mid- and hindlegs white, tarsi yellow with white rings but inconspicuous in male.

Abdomen. Male genitalia (Fig. 15). Uncus hook-shaped apically; tegumen broad; valvae moderately symmetric, left valva smaller than right, broad, extremely short, cucullus concave medially, densely covered with setae on the dorsal and ventral parts of cucullus; saccus well developed; aedeagus tubular, distal gradually slightly curve toward distal area; vesica simple, without cornuti.

Figures 3–8. 

Adults 3‒6 Ivela yini sp. nov. (3 male, holotype 4 female, paratype 5 male, paratype 6 female, paratype) 7, 8 Dendrophleps semihyalina (7 male 8 female). Scale bars: 10 mm.

Figures 9–13. 

Field images of adults 9‒11 Ivela yini sp. nov. male (9 dorsal view 10 lateral view 11 ventral view of head) 12, 13 Dendrophleps semihyalina (12 male, dorsal view 13 female, dorsal view).

Female genitalia (Fig. 16). Anterior apophysis almost as long as posterior apophysis; anal papillae larger; ostium larger; ductus bursae short, sclerotized; corpus bursae with a pair of terminal projections.

Pupa (Figs 21–24). Head white; prothorax white, with long, white hairs; mesothorax and metathorax chestnut-colored on dorsal surface, with dark brown hairs; wings white, with two black lines dorsally; forelegs and midlegs yellow, hindlegs white, with dark yellow dot terminally. Abdomen pale green laterally and ventrally, with chestnut-colored dots and spots dorsally; segments A1–A6 with a pair of black setae; segments A2 and A3 and terminal of abdomen with white hairs.

A single pupa of I. yini was discovered on Idesia polycarpa Maxim. (Salicaceae) (Fig. 24). This suggests that this is the foodplant of this species.


Forest zone 1000‒1315 m elevation.

Materials examined

Holotype : ♂, Nanling National Nature Reserve, Ruyuan County, Guangdong, 25.VI.2008, leg. Min Wang. Paratypes: 1♂, same data as holotype, altitude 1315 m, 12.VII.2010, leg. Min Wang. 1♀, same data as holotype, 3.VII.2011, leg. Min Wang. 1♂, same data as holotype, altitude 1000 m, 10.VII.2019, leg. Ran Yin & Xiao-juan Xing. 1♀, same data as holotype, 11‒14.VI.2019, leg. Hou-shuai Wang.


China (Guangdong).


The species is named after Ran Yin, who discovered the pupa of the new species. The name is in the genitive case.


The female genitalia of I. auripes (Fig. 18) and D. semihyalina (Fig. 20) have to our knowledge not been illustrated previously. They are illustrated here for comparative purposes.

The early stages of D. semihyalina are also newly reported as below (Figs 25–30):

Host plant of D. semihyalina (Fig. 25): Indocalamus tessellatus (Munro) Keng f. (Poaceae).

Last instar larva of D. semihyalina (Figs 26, 27): body white laterally and ventrally; dorsally black, with scattered white dots. A1 and A2 with reddish orange tufts dorsally. Verrucae pale yellow laterally, black dorsally, with long white or black hairs.

Figure 14. 

Wing venation of Ivela yini sp. nov. (male, paratype).

Figures 15–20. 

Genitalia 15, 16 Ivela yini sp. nov. (15 male, holotype 16 female, paratype) 17, 18 I. auripes (17 male 18 female) 19, 20 Dendrophleps semihyalina (19 male 20 female).

Figures 21–24. 

Pupa of Ivela yini sp. nov. 21 dorsal view 22 lateral view 23 ventral view 24 pupa on Idesia polycarpa Maxim.

Figures 25–30. 

Immature stages and host plant of Dendrophleps semihyalina 25 host plant: Indocalamus tessellatus (Munro) Keng f. 26, 27 last instar larva on the host plant 28‒30 pupa (28 dorsal view 29 lateral view 30 ventral view).

Pupa of D. semihyalina (Figs 28–30): body white laterally and ventrally, green-brown dorsally. Thorax and abdomen with irregular black spots on lateral and ventral surfaces. A1–A3 with a pair of green patches on dorsal surface. Wings with some mixed orange and black veins.


Ivela yini is superficially similar to several tussock moths with which it is sympatric. We illustrated it with Dendrophleps semihyalina and Ivela auripes for comparative purposes. These species can be distinguished reliably by the combinations of superficial characters outlined above. The identification can be confirmed by dissection of the male and female genitalia if required.

The genetic distance values of DNA barcode data between Lepidoptera species are ordinarily greater than 3% (Hebert et al. 2003). Our analysis of Chinese Leucomini indicates that the DNA barcode of I. yini is 10.6% from the closest species I. auripes (Appendix 1). The NJ tree also strongly supports the validity of the new species (Fig. 1). Our phylogenetic analyses show that I. yini and I. auripes are a monophyletic clade (PP = 1.00, BP = 87) and strongly suggest that I. yini belongs in Ivela (Fig. 2). This arrangement is supported by morphology. All Ivela, including I. yini, share wide valvae, and their hindwings lack a row of oblique veinlets (accessory veins) between A2 and the dorsal margin. These veinlets are considered diagnostic for Dendrophleps (Holloway 1999; Mackey 2019).

Several hardwoods were reported as foodplants for I. auripes: Corylopsis multiflora Hance (Hamamelidaceae), Cornus controversa Hemsley (Cornaceae), C. brachypoda C.A. Mey (Cornaceae), Styrax japonicus Sieb. et Zucc (Styracaceae), and S. obassis Siebold et Zucc (Styracaceae) (Inoue 1956; Chao 2003). While our discovery of a pupa of I. yini on Idesia polycarpa is less than absolute proof that it is the foodplant of this moth, it does suggest that I. yini feeds on a broadleaved tree. In contrast, D. semihyalina was discovered to be a grass feeder. These foodplant differences support placement of these similar-appearing moths into different genera.


We are grateful to three reviewers for their constructive feedback on our manuscript. We are thankful Prof. Min Wang from South China Agricultural University for collecting the specimens. We are also thankful to Mr Ran Yin and Ms Xiao-juan Xing for discovering the pupa of the new species. This work was supported by a grant from National Natural Science Foundation of China to HSW (no. 31601879) and Natural Science Foundation of Guangdong (no. 2022A1515012050).


  • Chao CL (2003) Lepidoptera Lymantriidae. Fauna Sinica, Insecta 30. Science Press, Beijing, 484 pp.
  • Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3(5): 294–299.
  • Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings. Biological Sciences 270(1512): 313–321.
  • Holloway JD (1999) The moths of Borneo [Part 5]: Family Lymantriidae. Malayan Nature Journal 53: 1–188.
  • Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589.
  • Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16(2): 111–120.
  • Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35(6): 1547–1549.
  • Mackey P (2019) A note on Dendrophleps lobipennis (Swinhoe, 1892) (Lepidoptera, Erebidae, Lymantriinae) with a description of the female. Suara Serangga Papua 12(1): 1–7. [SUGAPA digital]
  • Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542.
  • Wahlberg N, Wheat CW (2008) Genomic outposts serve the phylogenomic pioneers: Designing novel nuclear markers for genomic DNA extractions of Lepidoptera. Systematic Biology 57(2): 231–242.
  • Wang HS, Xiong W, Wang M (2010) Two new species of the genus Longipenis (Lepidoptera: Lecithoceridae) from China. The Florida Entomologist 93(3): 352–356.
  • Wang HS, Fan XL, Owada M, Wang M, Nylin S (2014) Phylogeny, systematics and biogeography of the genus Panolis (Lepidoptera: Noctuidae) based on morphological and molecular evidence. PLoS ONE 9(3): e90598.
  • Wang HS, Wahlberg N, Holloway JD, Bergsten J, Fan XL, Janzen DH, Hallwachs W, Wen LJ, Wang M, Nylin S (2015) Molecular phylogeny of Lymantriinae (Lepidoptera, Noctuoidea, Erebidae) inferred from eight gene regions. Cladistics 31(6): 579–592.

Appendix 1

Table A1.

Kimura 2-parameter genetic distances based on COI barcodes among 13 samples of Leucomini and two outgroups.

Species code Species name 1 2 3 4 5 6 7 8 9 10 11 12 13 14
LE114 Ivela yini sp. nov.
LE074 Ivela yini sp. nov. 0.006
LE118 Ivela yini sp. nov. 0.000 0.006
H340 Ivela yini sp. nov. 0.011 0.011 0.011
LE124 Ivela auripes 0.122 0.119 0.122 0.120
H49 Ivela auripes 0.110 0.106 0.110 0.108 0.020
H181 Perina nuda 0.126 0.122 0.126 0.120 0.157 0.144
LE116 Dendrophleps semihyalina 0.154 0.156 0.154 0.154 0.183 0.165 0.170
LE115 Dendrophleps semihyalina 0.152 0.154 0.152 0.152 0.181 0.163 0.168 0.002
GD385 Dendrophleps semihyalina 0.144 0.146 0.144 0.144 0.167 0.150 0.161 0.019 0.017
LE014 Dendrophleps semihyalina 0.142 0.144 0.142 0.142 0.165 0.148 0.159 0.015 0.014 0.003
H377 Leucoma sp. 0.169 0.173 0.169 0.171 0.174 0.155 0.172 0.187 0.187 0.182 0.181
H351 Leucoma salicis 0.159 0.161 0.159 0.161 0.163 0.149 0.182 0.193 0.191 0.188 0.187 0.127
H127 Lymantria dissoluta 0.138 0.135 0.138 0.142 0.153 0.136 0.131 0.157 0.155 0.149 0.147 0.159 0.167
H58 Lymantria similis 0.133 0.129 0.133 0.133 0.139 0.121 0.137 0.154 0.154 0.146 0.145 0.144 0.163 0.071
login to comment