Research Article
Research Article
Phylogeny and classification of Endromidae (Lepidoptera: Bombycoidea) based on mitochondrial genomes
expand article infoMin Deng§, Andreas Zwick|, Qi Chen, Cheng-Qing Liao, Wei Wang§, Xing Wang§, Guo-Hua Huang
‡ Hunan Agricultural University, Changsha, China
§ Qiongtai Normal University, Haikou, China
| CSIRO, Australian National Insect Collection, Canberra, Australia
Open Access


The small, relict-like moth family Endromidae is well-established within the superfamily Bombycoidea, but relationships within the family have remained vague for the last decade, primarily due to very limited taxon sampling. This resulted in the explicit removal of all internal suprageneric classification by Zwick et al. (2011) when they synonymized Mirinidae and the bombycid subfamilies Oberthueriinae and Prismostictinae with Endromidae. Nucleotide and amino acid data sets of the 13 mitochondrial, protein-coding genes from representatives of 13 of the 16 accepted endromid genera were used to estimate phylogenetic relationships based on maximum likelihood and Bayesian inference methods. The results strongly support Endromidae as a monophyletic group and enable the establishment and diagnosis of four subfamilies (Endrominae, Mirininae stat. rev., Oberthueriinae stat. rev. and Prismostictinae stat. rev.). Within subfamily Oberthueriinae, we establish three tribes: Oberthueriini stat. rev., Andracini tribe nov. and Mustiliini tribe nov. We provide morphological diagnoses and a genus-level checklist for the three tribes. Promustilia yajiangensis Wang, X. & Zolotuhin, 2015 is transferred to Mustilizans as M. yajiangensis comb. nov. to establish reciprocal monophyly of the two genera, and Andraca gongshanensis is transferred to Pseudandraca as P. gongshanensis comb. nov. We also synonymize Andraca (Chrypathemola) syn. nov. with Andraca (Andraca), as the latter is deeply nested within the former.


Endromidae, mitochondrial genome, phylogenetic analysis, revision

1. Introduction

The moth family Endromidae Boisduval, 1828 is relatively species poor (72 species) and occurs primarily in Asia, with just a single species extending into Europe. This species, the very distinct and widespread Palearctic species Endromis versicolora (Linnaeus, 1758), is placed in the monobasic genus Endromis Ochsenheimer, 1810 and, for over a century, its own family Endromidae. It was regarded as an isolated taxon within the “Bombyces”, until Seitz (1911) tentatively included the then monobasic genus Mirina Staudinger, 1892. This action was followed by Kuznetsov & Stekolnikov (1985), who studied the muscles of Bombycoidea. Shortly after, Kozlov (1985) erected the family Mirinidae Kozlov, 1985 for Mirina. The concept of two closely related yet distinct families was then generally accepted due to the clear differences in adult morphology, larval appearance and host plant usage (e.g., Kozlov 1985; Minet 1986, 1994; Lemaire and Minet 1998; Zolotuhin and Witt 2000).

The use of DNA sequence data has greatly contributed to the phylogenetic hypotheses and consequential changes in the classification of Bombycoidea. Based on five protein-coding nuclear genes, phylogenetic analyses of the ‘bombycoid complex’ grouped Endromidae, Mirinidae and the bombycid subfamilies Oberthueriinae Kuznetsov & Stekolnikov, 1985 and Prismostictinae Forbes, 1955 into a single clade (Regier et al. 2008). Zwick et al. (2011) robustly corroborated the above result with increased taxon and gene sampling (50 bombycoid taxa, up to 20 nuclear, protein-coding genes), and genetic variation within this clade was found to be less than in other bombycoid families. However, the internal branches between endromid genera were very short and taxon sampling was minimal with only four species, resulting in weak and sometimes conflicting statistical support for relationships within the clade. Consequently, Zwick et al. (2011) synonymized Mirinidae, Oberthueriinae and Prismostictinae with Endromidae, intentionally removing all suprageneric structure within Endromidae in recognition of the uncertain relationships and poor taxon sampling. Although the previous, independent systematic family status was preferred by many workers (e.g., Wang et al. 2011a; Zolotuhin and Than 2011; Zolotuhin 2012; Zolotuhin and Wang 2013), other authors agreed with the revised concept of Endromidae sensu Zwick et al. (2011) and followed or further supported it in their respective studies (e.g., Wang et al. 2011b; Regier et al. 2013; Hamilton et al. 2019; Wang et al. 2019). Similarly, the previous and clearly polyphyletic (Regier et al. 2008; Zwick, 2008; Zwick et al. 2011) concept of Bombycidae Latreille, [1802] sensu Minet (1994), which united Oberthueriinae, Prismostictinae, Bombycinae, Phiditiidae Minet, 1994 and Apatelodidae Neumoegen & Dyar, 1894, was retained by some authors as “Bombycidae sensu lato” (e.g., Zolotuhin 2007; Zolotuhin and Witt 2009; Zolotuhin and Tran 2011; Zolotuhin 2012; Zolotuhin and Wang 2013; Wang et al. 2015; Wu and Chang 2016).

With further studies on the phylogeny of Bombycoidea (Hamilton et al. 2019; Wang et al. 2019), the composition of Endromidae has gradually been clarified and stabilized. Hamilton et al. (2019) included nine genera of Endromidae, but even with up to 571 nuclear loci from anchored hybrid enrichment, relationships and statistical support within Endromidae varied depending on the type of data and method of analysis. The currently recognized endromid taxa are detailed in a global checklist of Bombycoidea (Kitching et al. 2018) and subsequent publications (Chandra et al. 2019; Andraca yauichui Wu & Chang, 2016 and Mustilizans zolotuhini Chandra, 2019). At present, a mere 72 species are recognized as valid in 16 genera, and three of these genera are monobasic (Endromis; Prismostictoides Zolotuhin & Than Thieu, 2011; Falcogona Zolotuhin, 2007). However, no objective phylogeny-based subfamily or tribal classification exists for the genera of Endromidae, despite the very different appearances of the moths and caterpillars, which has led to their long taxonomic separation and hinders the recognition of natural relationships. These deep morphological differences, combined with very limited species diversity in most genera and restricted distributions, indicate a relictual nature of this family. Indeed, the Endromidae are the Asian representatives of a more inclusive relictual lineage, the “CAPOPEM” group of Regier et al. (2008), which includes additionally the monobasic Carthaeidae Common, 1966 in the SW corner of Australia, the small Austral-New Guinean family Anthelidae Turner, 1904 and the even smaller Neotropical family Phiditiidae Minet, 1994.

Several endromid genera (e.g., Dalailama Staudinger, 1896, Sesquiluna Forbes, 1955 and Falcogona) are rarely collected and underrepresented in collections, making it difficult to obtain comprehensive taxon sampling for molecular phylogenetic studies. DNA sequencing of old collection specimens helps to improve taxon sampling of rarely collected species (Zimmermann et al. 2008; Burrell et al. 2015), and mitochondrial (mt) genomes are particularly easy to obtain from strongly degraded DNA samples due to their high copy number (Duan et al. 2018). Furthermore, mt-genomes are characterized by a simple genetic structure, small size, strictly orthologous genes, reduced recombination and fast evolutionary rates (Zhang and Hewitt 1997; Boore 1999; Cameron 2014), which makes them attractive for some phylogenetic questions. Some studies found that mt-genomes were inadequate for resolving subfamily-level relationships, but given good taxon sampling, could resolve lower-level phylogenetic relationships (Nie et al. 2020; Ghanavi et al. 2022). In other studies, mt-genomes have proven a useful data source for taxonomic and phylogenetic studies of arthropods (e.g., Xiao et al. 2012; Cameron 2014; Wang et al. 2018; Chen et al. 2020; Bian et al. 2020; Jin et al. 2020a, b). In this study, we utilize mt-genomes of 13 of the 16 genera (80% of lineage diversity) to investigate the phylogeny of the family Endromidae and to revise its internal classification.

2. Material and methods

2.1. Taxon sampling and DNA extraction

The mt-genomes of 26 taxa of Endromidae, belonging to 13 genera in four subfamilies (Table S1), were newly sequenced for this study. Following collection, the three right legs or thorax of each specimen were preserved in absolute ethanol and then stored at –20°C. The remainders of all specimens are deposited as vouchers in the Insect Museum of Hunan Agricultural University, Changsha City, Hunan Province, China.

Total genomic DNA was extracted from the legs or thoracic tissue of each specimen using a TaKaRa MiniBEST Universal Genomic DNA Extraction Kit Ver.5.0 (Shiga Prefecture, Kusatsu City, Japan). Purified DNA was preserved at –20°C prior to sequencing.

2.2. Genome sequencing and assembly

Illumina TruSeq libraries with 350 bp insert size were prepared for each species, and these Whole Genome Shotgun libraries were sequenced by Novogene (Beijing, China) on the Illumina Hiseq platform with 150 bp paired-end reads. For each library, 6 Gbp of clean data were obtained after removing reads containing adaptor contamination, poly-Ns (>15 bp Ns), or >75 bp bases with quality scores ≤ 20. Cleaned reads were assembled into contigs and scaffolds using IDBA (v1.1.3; Peng et al. 2010, 2011, 2012), with kmer values ranging between 60 and 160 bp. The mt-genomes were annotated with ORF finder ( and compared against annotated sequences in NCBI using BLAST ( Then, the 22 tRNA and two rRNA sequences were annotated with MITOS web server ( (Bernt et al. 2013). The two rRNA subunits and all protein-coding genes (PCGs) were annotated by alignment with homologous genes from the same genus or subfamily using Geneious R8 (Kearse et al. 2012). MEGA 10.1.5 was used to calculate the AT and GC content and P-distances (Kumar et al. 2018). The annotated sequences of the 22 newly sequenced species are deposited in GenBank, and accession numbers (OQ472264OQ472285) and the other 13 sequences, which are incomplete, can be accessed from Zenodo ( The collection data are detailed for all new endromid samples in Table S1.

2.3. Phylogenetic analyses

Twelve publicly available mt-genomes were obtained from NCBI GenBank (, including nine ingroup species and three lasiocampid species used as outgroups. All mt-genome sequences were imported and standardized in Geneious R8. All PCGs were exported from Geneious R8. The 13 PCGs were aligned with the TranslatorX server ( (Castresana 2000), with the “ALL”-parameter. Concatenation of single gene alignments was performed in Geneious R8, resulting in 2 datasets: 1) 13 protein-coding genes (13PCGs); 2) 13 PCGs as amino acids (13PCGs-AA). Partitionfinder 2.1.1 was used to search the optimal partitioning scheme and models for each data set. IQ-Tree (v1.5.5; Nguyen et al. 2016) was used to estimate a maximum likelihood (ML) tree with 1,000 non-parametric bootstrap replicates to estimate branch support. Nodes with a bootstrap percentage (BP) of at least 70% were considered well supported in the ML analyses (Hillis et al. 1993). A Bayesian (BI) tree was calculated with MrBayes (v3.2.6) on XSEDE ( (Nguyen et al. 2014; Ronquist et al. 2003), with the Markov Chain Monte Carlo analysis run for 10,000,000 generations, sampled every 1,000th generation and with a burn-in of 25%. Bayesian posterior probabilities (PP) > 0.95 were interpreted as strongly supported (Erixon et al. 2003). The phylogenetic trees were drawn using the software FigTree (v1.4.3; Rambaut 2016).

3. Results

3.1. Characteristics of endromid mtgenomes

Our study provides the first mt-genomes for six endromid genera, i.e., Endromis, Mirina, Pseudandraca Miyata, 1970, Smerkata Zolotuhin, 2007, Dalailama and Promustilia Zolotuhin, 2007. Most mt-genomes, except Andraca lawa_21, Mustilizans dierli and Prismosticta tiretta_32, used in this study comprised a total of 37 genes (13 PCGs, 22 tRNAs and 2 rRNAs), and the total length of all sequences ranges from an incomplete 6,350 bp (Andraca lawa_21) to 15,880 bp (Andraca olivacea−GD) (Figure S1). As is usual for the mt-genomes of Lepidoptera (Arnoldi et al. 2007; Yang et al. 2013; Amaral et al. 2016; Yuan et al. 2019), those of Endromidae have a significant bias towards adenine and thymine, ranging from 77.7% to 81.1% (average 79.8%; Table S2). For almost all samples, the AT-skew is greater than 0 and GC-skew is less than 0, except Mirina confucius Zolotuhin & Witt, 2000, which has a negative AT-skew (Table S2). Moreover, the uncorrected pairwise distance P (proportion of nucleotide sites at which two compared sequences are different) shows that the COX1 gene, widely adopted as the DNA barcode marker, possesses the smallest amount of interspecies genetic variation, while ATP8 possesses the highest (Figure S2). The G + C content for each PCG reveals that ATP8 possesses the lowest G + C content, and ATP6 the highest (Figure S3).

3.2. Phylogenetic relationships among Endromidae

Based on the 13PCGs dataset, the two phylogenetic trees estimated with BI (Figure S4) and ML analyses (Figure S5) are almost identical. Only the relationships between the three clades of Oberthuriinae and the relationships between species of Primosticta differ. And the Bayesian tree of the 13-PCGs data set is used to label the values of other trees. Statistical support (PP and BP) is strong for 34 of the 40 nodes, with weak support restricted to backbone nodes and within a clade of Oberthueriinae stat. rev. Within the limits of taxon sampling, the results strongly support the monophyly of the family Endromidae, as well as a division into four subfamilies, with Prismostictinae stat. rev. sister to all other taxa (PP = 1, BP = 100%). The phylogenetic relationship between Endrominae and Mirininae stat. rev., which are nested between the two other subfamilies, is strongly supported in the BI tree (PP = 0.992). Within Prismostictinae stat. rev., the genera Prismosticta Butler, 1880 and Prismostictoides are sister to each other (PP = 1, BP = 99.5%). A distinct monophyletic group, subfamily Oberthueriinae stat. rev. comprises three major, well-supported clades (PP = 1, BP = 100%). Clade 1 includes the genera Pseudandraca Miyata, 1970 and Andraca Walker, 1865. In addition, Andraca gongshanensis and Pseudandraca flavamaculata are shown as sisters with strong support (PP = 1, BP = 99.8%). The clades (Andraca apodecta + Andraca melli) and (Andraca trilochoides + (Andraca draco + Andraca lawa) are grouped together (PP = 1, BP = 100%). Clade 2 includes four genera, with Comparmustilia Wang, X. & Zolotuhin, 2015 sister to the other genera. Clade 3 is divided into two major groups comprising the monophyletic genus Oberthueria Kirby, 1892 (PP = 1, BP = 100%) and the genus Mustilizans Yang, 1995, which is a paraphyletic relative to the species Promustilia yajiangensis Wang, X. & Zolotuhin, 2015. The relationship of these three clades is shown as (Clade 1 + (Clade 2 + Clade 3)) (PP = 0.771, BP = 49.8). Although both methods of analysis of the different datasets resulted in largely congruent topologies, there are still obvious differences compared to the analysis results of the 13PCGs-AA dataset (BI in Figure S6, ML in Figure S7, and combined in Figure 1), which show the relationships of the three clades within Oberthueriinae as (Clade 3 + (Clade 1 + Clade 2)) in the BI tree (Figure S6) with good support (PP = 0.917).

4. Discussion

All the newly sequenced Endromidae mitochondrial genomes have the same gene order as in the other known Lepidoptera (Cao et al. 2012; Timmermans et al. 2014; Zou et al. 2017). The particularly low divergences among the COX1 sequences indicate the genetic stability of the gene, which is used for species identification in many studies (e.g., Rodrigues et al. 2017; Liao et al. 2021).

Mitochondrial genomes are widely used for studying population genetics, comparative and evolutionary genomics, the reconstruction of phylogenetic relationships, and evolutionary biology (e.g., Boore 1999; Vilhelmsen 2019; Jin et al. 2020b; Quicke et al. 2020). Our analyses of mt-genomes resulted in ML and BI trees that overall differ very little from each other (only in the unstable position of three clades of Oberthueriinae). Furthermore, the study of Hamilton et al. (2019), which has by far the greatest nuclear gene sampling for nine of the 16 endromid genera, recovered exactly the same taxonomic groupings as our mt-genome analyses, albeit with greatly reduced taxon sampling (10 vs 35 endromid species). This congruence lends credibility to our results and the placement of the four additional genera Prismostictoides, Smerkata, Dalailama and Promustilia.

Endromid subfamilies

Our results are likewise in agreement with morphologically recognized group. Our analyses recovered both the family Endromidae and its four major lineages as monophyletic and strongly supported. These four lineages correspond to the morphologically recognized (sub)families that were synonymized with Endromidae (Zwick et al. 2011). Consequently, on this basis, we here re-instated these lineages as valid subfamilies: Endrominae, Mirininae stat. rev., Oberthueriinae stat. rev. and Prismostictinae stat. rev. All the different data sets recovered the same topology as Hamilton et al. (2019), i.e., Prismostictinae + (Endrominae + (Mirininae + Oberthueriinae)).


Endrominae, which is just consisting of one species, has typical characteristics different from other endromid moths as follows: forewing with three triangular white spots, thorax and abdomen bright-colored (Figure 1. Endromis), uncus and valva tongue-shaped, and gnathos absent. Its larval host is also different from other endromid species, mainly feeding Betula sp., Corylus sp. Cytisus sp., Quercus spp. (Waring and Townsend 2003; Pérez et al. 2009).

Figure 1. 

Phylogeny of Endromidae inferred from different data sets (13PCGs-AA, 13PCGs) using Bayesian inference and maximum likelihood analyses. Numbers above branches are posterior probabilities (BI PP), beneath which are bootstrap percentages (ML BP) for 1000 replicates; nodes with maximum support values are marked with a black dot instead. Dashed arrows (two in total) identify alternative topologies (relative to the topology shown) that receive at least 70% bootstrap support by one or more of the approaches. The asterisks indicate newly sequenced mitochondrial genomes.

Mirininae stat. rev

Mirininae stat. rev. is consisting of only one genus, Mirina, which was been controversial. Some scholars considered that it should be a separate family (Minet 1994; Zolotuhin and Witt 2000; Huang and Wang 2003; Regier 2008; Zolotuhin et al. 2011), but others recognized that it should belong to the family Endromidae (Zwick et al. 2008; Zwick et al. 2011). Although Mirina and Endromis are different from larval morphology, host and adult appearance (Zolotuhin and Witt 2000), subsequent molecular phylogenetic studies had continued to find good support for Endromidae sensu Zwick et al. (2011). And then they were listed in the family Endromidae in the global checklist of the Bombycoidea (Kitching et al. 2018). In this paper, their relationships had been supported based on anchored hybrid enrichment (Hamilton et al. 2019) and showed as Endrominae nest to Mirininae.

Prismostictinae stat. rev

The subfamily Prismostictinae stat. rev., which is sister to a clade comprising all the other subfamilies, consists of only two morphologically similar genera, Prismosticta and Prismostictoides. The monobasic genus Prismostictoides, with the type species Prismosticta unihyala Chu & Wang, 1993, was separated from Prismosticta based on a broad postmedial line and broad, dark yellow submarginal band on the forewing upperside, uncus with a long uncuslike projiection arising from the base of uncus, and valva asymmetrical. Otherwise, the two genera are rather similar and share a characteristic transparent spot near the apex of the forewing (Zolotuhin and Tran 2011; Wang et al. 2015), which might be used to diagnose the subfamily. Our molecular data place Prismostictoides as sister to the five sampled species of Prismosticta.

Oberthueriinae stat. rev

Oberthueriinae stat. rev., the largest subfamily of Endromidae, is divided into three major clades that are strongly supported as monophyletic groups. Based on our results, we treat these clades as three tribes: Andracini tribe nov. (Clade 1), Mustiliini tribe nov. (Clade 2) and Oberthueriini Kuznetzov & Stekolnikov, 1985 stat. rev. (Clade 3). The sister relationship between Andraca and Pseudandraca in Andracini tribe nov. is consistent with findings of previous studies (Wang et al. 2012, 2015; Hamilton et al. 2019). The genus Pseudandraca was distinguished from Andraca, the type genus of Andracini, based on forewing pattern and male genital structures (Zolotuhin and Witt 2009; Wang et al. 2012). However, our phylogenetic analysis results indicate that Andraca gongshanensis should be transferred to Pseudandraca as Pseudandraca gongshanensis comb. nov. The two subgenera of Andraca, Andraca (Andraca) Walker, 1865 and Andraca (Chrypathemola) Zolotuhin, 2012, were established based on the structure of the uncus in male genitalia. However, none of our analyses support the distinction of these two subgenera. The two representatives of Andraca (Andraca), A. (A.) lawa and A. (A.) draco, are deeply nested within Andraca (Chrypathemola) and therefore we here synonymize Andraca (Chrypathemola) syn. nov. with Andraca (Andraca).

Previously, Mustilia Walker, 1865 had been split into six separate genera (Zolotuhin 2007; Wang et al. 2005), i.e., Mustilia, Comparmustilia, Mustilizans, Promustilia, Smerkata and Falcogona, but not all of these belong in the tribe Mustiliini tribe nov. Within this tribe, the genera Mustilia and Comparmustilia were recovered as sister taxa in previous studies (Wang et al. 2019; Hamilton et al. 2019). However, the inclusion of the newly sequenced genera Smerkata and Dalailama provides a more comprehensive and different picture, with these two genera nested between Comparmustilia and Mustilia, and Dalailama being sister to Mustilia.

Our analyses place three genera in the tribe Oberthueriini, i.e., Oberthueria, Mustilizans and Promustilia, of which the latter two were previously included in Mustilia. The genus Oberthueria is monophyletic, with strong statistical support for the three species included in this study, and the six currently recognized species are morphologically very similar and show only moderate differences in COX1 barcode sequences (Zolotuhin and Wang 2013). In contrast, our results demonstrate clearly the paraphyly of the genus Mustilizans relative to the species Promustilia yajiangensis. Because our study lacks the type species of Promustilia, we do not, at this time, wish to synonymize Promustilia with Mustilizans. Instead, we transfer P. yajiangensis to Mustilizans as M. yajiangensis comb. nov. and retain Promustilia as a valid genus for the time being. The separation of Promustilia from Mustilizans is doubtful as the morphologies of P. andracoides (Zolotuhin, 2007) and M. yajiangensis comb. nov. are very similar, and the former also shares a similar bifid uncus and flat, apically blunt valva with Mustilizans (Zolotuhin 2007; Zolothuhin and Witt 2009; Wang et al. 2015). More detailed future studies should be undertaken to provide further molecular and morphological data to elucidate the relationships of these two genera.

Unfortunately, specimens of the remaining three endromid genera, Falcogona, Sesquiluna and Theophoba Fletcher & Nye, 1982 were unavailable to us and we lack mt-genomes for them. Based on similarities in adult morphology, such as eyes surrounded by setae, completely pectinate antennae and similar size, we tentatively include Sesquiluna and Theophoba in subfamily Prismostictinae stat. rev., as postulated by Wang et al. (2015). The genus Falcogona is difficult to place phylogenetically because of some significant morphological differences, especially the very long and strongly modified cuiller (Zolotuhin et al. 2007). In other characteristics, such as wing shape, the short and broad gnathos, and the tubular phallus (Zolotuhin et al. 2007; Wang et al. 2015), Falcogona is rather similar to Smerkata. Consequently, we tentatively include Falcogona in the tribe Mustiliini tribe nov. The phylogenetic relationships among these and related genera should also be investigated in future studies.

Taxonomy of Oberthueriinae

Oberthueriini Kuznetzov & Stekolnikov, 1985, stat. rev.

Type genus

Oberthueria Kirby, 1892


Members of this tribe share the following characters: 1) forewings long and narrow; 2) labial palpi of moderate length, about 2/3 of the vertical diameter of the compound eye; 3) distal half of antenna devoid of well-developed rami (Figure 2). In addition, the larvae also bear some distinct characters, such as a hairless body, moderately wider expansion of the thoracic tergites, and an extremely long anal horn (Figure 1: Oberthueria).

Figure 2. 

Cephalic characters of Oberthueriinae. A, B Andraca olivacea; C, D Mustilia undulosa; E, F Oberthueria jiatongae.)


Zolotuhin (2007) treated Promustilia as a subgenus of Mustilizans, before Wang et al. (2015) raised it to full genus level when they described Promustilia yajiangensis. Although P. yajiangensis is here transferred to Mustilizans to ensure monophyly of that genus, Promustilia is retained as a distinct genus as the type species, Mustilizans (Promustilia) andracoides, was not included in this study. If not even synonymous, Mustilizans and Promustilia are closely related, and Promustilia is here included in Oberthueriini stat. rev..

Oberthueria Kirby, 1892

Oberthueria Kirby, 1892, Syn. Cat. Lepid. Het., 1: 720. Type species: Euphranor caeca Oberthür, 1880, by monotypy

Oberthueria Staudinger, 1892, in Romanoff, Mém. Lépid.: 337 Type species: Euphranor caeca Oberthür, 1880, by monotypy (a junior homonym and junior objective synonym of Oberthueria Kirby, 1892)

Oberthüria : Staudinger, 1892, in Romanoff, Mémoires sur les lepidoptères (Mém. Lépid.) 6: 337. (incorrect original spelling)

Euphraor: Kirby, 1892, Syn. Cat. Lepid. Het. 1: 720 (incorrect subsequent spelling)

Euphranor Oberthür, 1880, Etudes d’Entomologie (Étud. ent.) 5: 40. Type species: Euphranor caeca Oberthür, 1880, by monotypy (a junior homonym of Euphranor Herrich-Schäffer, 1855 (Lepidoptera, Saturniidae))

Mustilizans Yang, 1995

Mustilizans Yang, 1995, Insects of Baishanzu Mountain, Eastern China: 355. Type species: Mustilizans drepaniformis Yang, 1995, by original designation

Promustilia Zolotuhin, 2007

Promustilia Zolotuhin, 2007, Neue ent. Nachr. 60: 199. Originally erected as a subgenus of Mustilizans Yang, 1993. Type species: Mustilizans (Promustilia) andracoides Zolotuhin, 2007, by original designation

Mustiliini tribe nov.

Type genus

Mustilia Walker, 1865


Mustiliini tribe nov. is similar to Oberthueriini stat. rev. in having narrow forewings and the distal half of the antenna with underdeveloped rami, but it can be easily distinguished by particularly short or completely reduced labial palpi. Larvae of this tribe also possess a hairless body, but the thoracic tergites are laterally strongly expanded, and the anal horn is relatively long and stout (Figure 1: Comparmustilia, Mustilia and Smerkata).


Based on the above morphological characteristics, we here establish the new tribe Mustiliini tribe nov., which is also supported by molecular data. Although we lack molecular data for Falcogona, it is included in this tribe because of its similarity in habitus and male genital structures to Smerkata.

Comparmustilia Wang, X. & Zolotuhin, 2015

Comparmustilia Wang, X. & Zolotuhin, 2015, Zootaxa, 3989: 79. Type species: Mustilia sphingiformis Moore, 1879, by present designation

Smerkata Zolotuhin, 2007

Smerkata Zolotuhin, 2007, Neue ent. Nachr. 60: 193. Originally proposed as a subgenus of Mustilia Walker, 1865. Type species: Mustilia phaeopera Hampson, 1910, by original designation

Dalailama Staudinger, 1896

Dalailama Staudinger, 1896, Dt. ent. Z. Iris 8 (2): 303. Type species: Dalailama bifurca Staudinger, 1896, by monotypy

Dailalama: Staudinger, 1901, Cat. Lepid. palaearct. Faunengeb. (1): 128. Incorrect subsequent spelling

Deilelamia: Pagenstecher, 1909, Geschichte eur. Schmett.: 433. Incorrect subsequent spelling

Mustilia Walker, 1865

Mustilia Walker, 1865, List Specimens lepid. Insects Colln Br. Mus. 32: 580. Type species: Mustilia falcipennis Walker, 1865, by monotypy

Falcogona Zolotuhin, 2007

Falcogona Zolotuhin, 2007, Neue ent. Nachr. 60: 199. Type species: Falcogona gryphea Zolotuhin, 2007, by original designation

Andracini tribe nov.

Type genus

Andraca Walker, 1865


Morphological synapomorphies supporting the monophyly of Andracini tribe nov. are the relatively broad forewings, the very long labial palpi (longer than the vertical diameter of the compound eye), and underdeveloped rami over the distal 1/3 of the antenna (Figure 2). The larvae of this tribe share non-expanded thoracic tergites, a very short or completely absent anal horn, and the body in most species is densely covered with short hairs (Figure 1: Andraca).


Pseudandraca was established by Miyata (1970) with the type species Andraca gracilis Butler, 1885 based on a boot-shaped sacculus in the male genitalia. Wang et al. (2015) added Andraca flavamaculata Yang, 1995. Our molecular results also support these two genera as sister taxa. Andraca had been considered to have two subgenera (Zolotuhin 2012), but our analyses recover the subgenus Chrypathemola syn. nov. as a synonym of Andraca. The species Pseudandraca gongshanensis comb. nov. has a complex wing pattern, a relatively straight phallus and a foot-shaped apex of the valva, which are all characteristics shared with Pseudandraca (Figure 3).

Figure 3. 

Pseudandraca adult and male genitalia. A, B Pseudandraca flavamaculata; C, D P. gongshanensis comb. nov.

Andraca Walker, 1865

Andraca Walker, 1865, List Specimens lepid. Insects Colln Br. Mus. 32: 581. Type species: Andraca bipunctata Walker, 1865, by monotypy

Pseudoeupterote Shiraki, 1911, Catalogue Insectorum Noxiorum Formosarum: 48. Type species: Oreta theae Matsumura, 1909, by monotypy

Pseudandraca Miyata, 1970

Pseudandraca Miyata, 1970, Tinea. 8: 190. Type species: Andraca gracilis Butler, 1885, by original designation

5. Data availability statement

The complete sequences were uploaded to the NCBI (GenBank accession numbers: OQ472264OQ472285). The incomplete sequences provided in this article can be accessed from Zenodo, DOI:

6. Conflict of Interest

The authors have no conflicts of interests to declare.

7. Authors’ contributions

M.D. was responsible for drafting the manuscript, as well as the acquisition, analysis and interpretation of data. A.Z. provided part molecular sequences and contributed to the conception and design of the current study. Q.C. analyzed and interpreted the data. W.W. made suggestions and revised the manuscript. X.W. confirmed the insect species and revised the manuscript. G.-H.H. helped perform the analysis with constructive discussions and provided financial support. All authors read and approved the final manuscript.

8. Acknowledgements

The authors thank Prof. Min Wang, Dr. Hou-Shuai Wang (South China Agricultural University, China), Ms. Zhuang-Mei Chen, Meng-Yue Chen and Si-Jia Yi, Mr. Bin Chen and Lu Chen (Hunan Agricultural University) for their kind help in collecting the samples. This study was supported by the National Natural Science Foundation of China (31970450, 32111540167), and China Agriculture Research System (CARS-23-C08), and Guangdong Provincial Academy of Sciences Special Project of Science and Technology Development (2020GDASYL-20200102021).

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Supplementary materials

Supplementary material 1 

Table S1

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .docx

Explanation note: List of taxa (genera and species) examined for the study and sources of information.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (24.82 kb)
Supplementary material 2 

Table S2

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .docx

Explanation note: The characteristics of the mitochondrial genomes of Endromidae.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (31.34 kb)
Supplementary material 3 

Figure S1

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Characters of thirty-five newly sequenced endromid species’ mitochondrial genomes. Gene names are annotated using standard abbreviations; single letters are IUPAC amino acid abbreviations.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (1.27 MB)
Supplementary material 4 

Figure S2

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Boxplot showing P−distances between all 47 samples for each of the 13 genes analyzed. Outlier values are depicted as points outside of the box.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (134.15 kb)
Supplementary material 5 

Figure S3

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Boxplot showing the GC-ratios of all 47 samples for each of the 13 genes analyzed. Outlier values are depicted as points outside of the box.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (91.17 kb)
Supplementary material 6 

Figure S4

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Bayesian inference phylogram constructed with the 13PCGs data set of Endromidae.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (237.41 kb)
Supplementary material 7 

Figure S5

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Maximum likelihood phylogram constructed with the 13PCGs data set of Endromidae.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (247.78 kb)
Supplementary material 8 

Figure S6

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Bayesian inference phylogram constructed with the 13PCGs-AA data set of Endromidae.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (231.71 kb)
Supplementary material 9 

Figure S7

Deng M, Zwick A, Chen Q, Liao CQ, Wang W, Wang X, Huang GH (2023)

Data type: .pdf

Explanation note: Maximum likelihood phylogram constructed with the 13PCGs-AA data set of Endromidae.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (221.31 kb)
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