Re-Examination: No True Tabidia Snellen, 1880 (Lepidoptera: Crambidae) in China, with Descriptions of Two New Genera and Three New Species
Jia-Xin Wang, Jun Wu, Wan-Lu Liu, Yun-Li Xiao
TL;DR
This study finds no true Tabidia moths in China and describes two new genera and three new species based on physical and genetic evidence.
Contribution
The paper introduces two new genera and three new species, correcting the taxonomy of moths previously classified under Tabidia in China.
Findings
No true Tabidia species are found in China based on morphological and molecular data.
Two new genera, Scintilla and Melanoleuca, are established and placed in the tribe Agroterini.
Three new cryptic species within Melanoleuca are described and identified.
Abstract
Based on morphological and molecular evidence, this study revises the crambid genus Tabidia Snellen, 1880, confirming that no true species of this genus are distributed in China. Two new genera, Scintilla gen. nov. and Melanoleuca gen. nov., are established and assigned to the tribe Agroterini Acloque, 1897. Additionally, three new species are described within Melanoleuca gen. nov. This study clarifies the taxonomic boundaries of this group in China and provides a foundation for related systematic research. Many species within the genus Tabidia Snellen, 1880 exhibit significant differences in wing pattern and genital morphology, which are inconsistent with the definition of Tabidia, indicating that the genus is not monophyletic. To address this, the present study revises the taxonomy of the Chinese species previously placed in Tabidia based on wing morphological characteristics,…
Genes, proteins, chemicals, diseases, species, mutations and cell lines named across the full text — each resolved to its canonical identifier and authoritative record.
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Figure 15| Tribe | Species | Location | NCBI and BOLD Accession No. |
|---|---|---|---|
| Outgroup | Italy, South Tyrol | TLMF Lep 17359 | |
| Outgroup | Australia, Queensland | ANIC Gen No. 001009 | |
| Unplaced | United States, Arizona | ||
| Wurthiini | China, Shanxi | Pyr000807 | |
| Agroterini | Australia, Queensland | ||
| Unplaced | Papua New Guinea, Madang | USNM ENT 00204919 | |
| Unplaced | Papua New Guinea, Madang | USNM ENT 00704561 | |
| Agroterini | China, Hubei, Wuxue | ||
| Agroterini | China, Anhui, Qianshan | ||
| Agroterini | China, Hubei, Yingshan | ||
| Agroterini | China, Jiangxi, Guanshan | ||
| Agroterini | China, Hubei, Yingshan | ||
| Agroterini | China, Hubei, Wuxue | ||
| Agroterini | China, Hubei, Wuxue | ||
| Trichaeini | China, Hubei | Pyr000776 | |
| Unplaced | China, Hubei | Pyr000683 | |
| Agroterini | China, Hainan | ||
| Agroterini | China, Hubei, Yingshan | ||
| Agroterini | China, Anhui, Lujiang | ||
| Agroterini | China, Anhui, Huoshan | ||
| Agroterini | China, Yunnan | ||
| Nomophilini | China, Tianjin | Pyr001404 | |
| Wurthiini | China, Hubei | Pyr000232 | |
| Nomophilini | China, Guizhou | Pyr001026 | |
| Unplaced | Australia, Queensland | 10ANIC-10498 | |
| Herpetogrammatini | Costa Rica, Guanacaste | ||
| Herpetogrammatini | Australia, Queensland | ||
| Unplaced | Indonesia, Kalimantan Timur | RMNH.INS.19697 | |
| Herpetogrammatini | Malaysia, Bangi | ||
| Herpetogrammatini | China, Hubei | Pyr000626 |
| Family | Subfamily | Species | GenBank Acc. No. | References |
|---|---|---|---|---|
| Pyralidae | Pyralinae | [ | ||
| [ | ||||
| [ | ||||
| Epipaschiinae | [ | |||
| [ | ||||
| Phycitinae | [ | |||
| [ | ||||
| [ | ||||
| Crambidae | Glaphyriinae | [ | ||
| [ | ||||
| Odontiinae | [ | |||
| [ | ||||
| [ | ||||
| Scopariinae | [ | |||
| Crambinae | [ | |||
| [ | ||||
| Unpublished | ||||
| [ | ||||
| Schoenobiinae | Unpublished | |||
| Acentropinae | [ | |||
| Unpublished | ||||
| [ | ||||
| [ | ||||
| Pyraustinae | [ | |||
| Unpublished | ||||
| [ | ||||
| [ | ||||
| [ | ||||
| [ | ||||
| [ | ||||
| Spilomelinae | [ | |||
| [ | ||||
| Unpublished | ||||
| This study. | ||||
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| [ |
| Generic Characters | |||
|---|---|---|---|
|
| Pale yellowish | White | White |
|
| Rounded | Rounded | Rounded |
|
| Upturned, curved; third segment long, tapering | Upturned, curved; third segment long, tapering | Sickle-shaped, curved, long, slender, acutely pointed |
|
| Minute, filiform | Minute, filiform | Small |
|
| Without a group of large scales | Without a group of large scales | With a dorsal group of recumbent large scales from base of median nervure |
|
| Smooth, rounded; no distinct anal angle | Smooth, rounded; no distinct anal angle | Slightly wavy; distinct anal angle |
|
| Dark brown or black speckles or fine stripes | Brown or black blotches | Black banded or zigzag markings |
|
| Origin near 3/4 distal of median cell | Origin near 2/3 distal of median cell | Origin near base of median cell |
| Genus | Species | Type Locality |
|---|---|---|
| Russia, Amur [Amurland], Khabarovsk [Chabarofka] [ | ||
| China, Hubei [ | ||
| China, Hubei; Henan; Yunnan | ||
| China, Anhui, Qianshan | ||
| China, Hubei; Jiangxi | ||
| India, Meghalaya, Khasi Hills [Khasias] [ | ||
| China, Hubei, Wuxue [ | ||
| Indonesia, Sulawesi [Celebes], Balangnipa; Makassar; Maros [ | ||
| Indonesia, Sulawesi [Celebes], Maros; Bantaeng [Bonthain] [ | ||
| Indonesia, Sula [ | ||
| Indonesia, Pulo Laut [ | ||
| India, Meghalaya, Khasi Hills [Khasias] [ | ||
| New Guinea, Dinner Island [Dinner-eiland] [ | ||
| Indonesia, Maluku, Ambon Island [Amboina]. Papua New Guinea, Humboldt Bay; D’Entrecasteaux Islands, Fergusson Island. Australia, Queensland [ | ||
| Malaysia, Penang [ | ||
| Sri Lanka [Ceylon], Ratnapura [ |
- —National Science & Technology Fundamental Resources Investigation Program of China
- —National Natural Science Foundation of China
- —Science and Technology Research Project for Young Talents of Hubei Provincial Department of Education
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Taxonomy
TopicsLepidoptera: Biology and Taxonomy · Genomics and Phylogenetic Studies · Insect Pheromone Research and Control
1. Introduction
The genus Tabidia Snellen, 1880 belongs to the subfamily Spilomelinae (Lepidoptera: Crambidae). It was established by Snellen (1880) [1] with Tabidia insanalis as the type species; however, its taxonomic placement at the tribal level remains unstudied [2]. To date, 13 species are known worldwide, primarily distributed in the Oriental and Australasian regions [3,4,5,6,7,8,9,10,11,12,13,14,15,16]. Apart from the original descriptions, taxonomic research on this genus has been limited globally, with most studies focusing on Chinese fauna. Du & Li (2014) revised the Chinese species of Tabidia, describing the new species T. obvia Du & Li, 2014 and redescribing T. strigiferalis Hampson, 1900 and T. candidalis (Warren, 1896) [15]. Recently, Wang et al. (2024) described T. longatuba Wang, Du & Xiao, 2024 [16]. Consequently, four species of Tabidia are currently recorded in China.
Significant differences in external appearance and genital morphology were observed among the species of this genus, indicating that it might not be monophyletic. This suspicion is supported by traditional morphological characteristics: Hampson (1896) mentioned in the generic diagnosis that males possess a tuft of large, decumbent scales on the dorsum at the base of the median vein on the forewing [5]. However, Du & Li (2014) found that this character is present only in some species, such as T. insanalis Snellen, 1880, T. aculealis (Walker 1866), and T. craterodes Meyrick, 1894, but is absent in T. candidalis (Warren, 1896), T. fuscifusalis Hampson, 1917, T. strigiferalis Hampson, 1900 and the newly described T. obvia Du & Li, 2014 [15].
To address these taxonomic uncertainties and test the hypothesis of non-monophyly, the present study conducted a comprehensive re-examination of the Chinese species previously assigned to Tabidia. We integrated evidence from adult morphology, genitalia structure, mitochondrial COI barcoding, and complete mitogenome phylogenetics. The primary aims of this work are to: evaluate the monophyly of Tabidia with respect to Chinese fauna; clarify the phylogenetic relationships and tribal placement of these taxa; and establish a robust taxonomic framework for this group in China. The findings are expected to provide a foundation for a future global revision of Tabidia and related genera.
2. Materials and Methods
2.1. Taxon Sampling
Specimens examined in this study were collected mainly with light traps in the Chinese provinces of Hubei, Anhui, Henan, and Yunnan. Legs were removed from freshly collected individuals for molecular analysis. We generated COI DNA barcode data for the Chinese species formerly assigned to Tabidia, in addition to sequencing the complete mitochondrial genomes of the type species of the two new genera: Scintilla strigiferalis (Hampson, 1900) comb. nov. (PQ821889) and Melanoleuca luteamacula sp. nov. (PQ631180). All newly produced sequences have been deposited in GenBank, with the mitogenomes accessible under the accession numbers provided above, and COI barcode sequences under accession numbers OR371985–OR371989, OR596415, OR596416, PQ801142, PQ801143, and PQ817972 (see Table 1 for further details). To complement the dataset, additional sequences of related genera were obtained from BOLD (http://www.boldsystems.org, accessed on 22 September 2025) and GenBank (https://www.ncbi.nlm.nih.gov/genbank/, accessed on 22 September 2025) for phylogenetic reconstruction. Detailed collection information, BOLD sample IDs, and GenBank accession numbers are listed in Table 1 and Table 2. The holotype and paratype specimens of the new species described in this work are deposited in the Biological Specimen Museum, Huanggang Normal University, Huanggang, China (HGNU).
2.2. Morphological Methods
Genitalia were dissected and prepared following the protocol established by Li and Zheng (1996) [17]. Morphological terminology conforms to the standards outlined by Maes (1995) [18] and Kristensen (2003) [19]. Adult and genitalia images were captured using a Canon EOS 70D camera (Canon Inc., Tokyo, Japan) equipped with an EF 180mm f/3.5L USM macro lens and a Nexcope NE930 microscope (Yongxin Optics Co., Ltd., Ningbo, China), respectively. Image refinement was performed using Adobe Photoshop^®^ 2020 (Adobe Inc., San Jose, CA, USA).
2.3. DNA Barcoding Acquisition
Genomic DNA was extracted from the legs of dried adult specimens using the TIANamp Genomic DNA Kit (DP304, Tiangen Biotech Co., Ltd., Beijing, China). A 658 bp fragment of the cytochrome c oxidase subunit I (COI) gene was amplified with the primers LepF1 and LepR1 [20]. PCR products were sent to Tianyi Huayu Gene Co., Ltd., (Wuhan, China) for bidirectional Sanger sequencing using the same primers. The PCR program included an initial denaturation at 95 °C for 3 min; 35 cycles of 95 °C for 30 s, 55 °C for 30 s, and 72 °C for 1 min, with a final extension at 72 °C for 5 min.
2.4. Mitogenome Extraction, Assembly, and Annotation
Total genomic DNA was isolated from specimen legs using a DNA extraction kit (CWBIO, Jiangsu Cowin Biotech Co., Ltd., Taizhou, China). Sequencing libraries with an average insert size of 350 bp were prepared using the Nextera XT DNA Library Preparation Kit (Illumina, San Diego, CA, USA) and sequenced on the Illumina NovaSeq 6000 platform (Illumina, San Diego, CA, USA). Raw reads were processed with the NGS QC Toolkit (National Institute of Plant Genome Research, New Delhi, India) [21] to obtain high quality clean data. Assembly was performed de novo using SPAdes v3.14.1 (Center for Algorithmic Biotechnology, St. Petersburg State University, St. Petersburg, Russia) [22]. The complete mitochondrial genome was annotated with MITOS (http://mitos.bioinf.uni-leipzig.de, developed by the research team at Leipzig University, Leipzig, Germany, among others). Quality metrics including sequencing depth, GC content, Q20, and Q30 were assessed. Experimental procedures were conducted by Tianyi Huayu Gene Co., Ltd., (Wuhan, China) (https://www.tyhygene.com/).
2.5. Phylogenetic Analysis
Based on COI sequences, a phylogenetic tree was constructed, including 51 species across 51 genera from 7 tribes (excluding unplaced taxa) within Spilomelinae, using Ostrinia nubilalis and O. furnacalis as outgroups (Table 1). The COI gene tree was reconstructed using Bayesian Inference (BI) in PhyloSuite v1.2.3 (developed by the research team led by Dong Zhang at Lanzhou University, in collaboration with Fujian Agriculture and Forestry University and the Institute of Hydrobiology, Chinese Academy of Sciences, China) [23,24]. The analysis was performed with MrBayes v3.2.7 (developed by Ronquist et al.) [25] under the GTR+I+G+F model for 2 million generations, with the first 25% of samples discarded as burn-in. Numbers in Figure 1 at nodes indicate Bayesian posterior probabilities. Habitus images of some Tabidia species were obtained from the BOLD database with permission. Pairwise genetic distances based on the COI sequences were calculated using the Kimura 2-parameter (K2P) model in MEGA v11.0.13 [26].
For mitogenome based phylogeny, 13 protein coding genes (ND2, COX1, COX2, ATP8, ATP6, COX3, ND3, ND5, ND4, ND4L, ND6, CYTB, ND1) and two rRNA genes (rrnS, rrnL) were used. Eight pyraloid species served as outgroups. Additional mitogenomes of crambid taxa, particularly from Spilomelinae, were retrieved from GenBank (Table 2). Maximum likelihood phylogenies were reconstructed using IQ-TREE v2.2.0 [27] under the Edge-linked partition model. Branch support was evaluated with 5000 ultrafast bootstrap replicates [28], along with the approximate Bayes test [29] and the Shimodaira–Hasegawa-like approximate likelihood-ratio test [30]. All analyses were performed using PhyloSuite v1.2.3 [23,24]. Phylogenetic visualization was performed using Chiplot (https://www.chiplot.online/) [31].
3. Results
3.1. Phylogenetic Analysis Based on COI
A phylogenetic tree was constructed using available COI sequences (Table 1, Figure 1 and Figure 2). Tabidia insanalis (the type species of Tabidia) clustered with Agrotera barcealis (Agroterini). Tabidia inconsequens clustered with Patania concatenalis (Agroterini). The species now assigned to Scintilla gen. nov. (S. strigiferalis and S. obvia) formed a monophyletic clade. The species assigned to Melanoleuca gen. nov. (excluding M. candidalis (Warren, 1896) comb. nov. for which data was unavailable) also formed a monophyletic clade. The Scintilla gen. nov. and Melanoleuca gen. nov. clades were located on branches distinct from those containing T. insanalis and T. inconsequens.
Phylogenetic relationships among Scintilla gen. nov., Melanoleuca gen. nov., Tabidia, and related genera, inferred from COI sequences using Bayesian Inference (BI).
Kimura 2-parameter genetic distances (as percent) calculated between Scintilla gen. nov., Melanoleuca gen. nov., Tabidia, and related genera.
3.2. Mitogenome Characteristics
The complete mitochondrial genomes of the type species Scintilla strigiferalis (Hampson, 1900) comb. nov. and the type species Melanoleuca luteamacula sp. nov. were sequenced and assembled (Figure 3, Supporting Information, Tables S1 and S2). The mitogenome of S. strigiferalis was a circular molecule of 15,351 bp, with a GC content of 19.3% and a gene repertoire of 13 PCGs, 22 tRNAs, and 2 rRNAs. The mitogenome of M. luteamacula was 15,443 bp in length, with a GC content of 19.5%, and contained the same set of 37 genes. The gene order in both mitogenomes was identical to the inferred ancestral arrangement for Lepidoptera. The A+T-rich control region was located between rrnS and trnM, measuring 360 bp (A+T content: 95.8%) in S. strigiferalis and 334 bp (A+T content: 94.9%) in M. luteamacula.
3.3. Phylogenetic Analysis Based on Mitogenomes
A higher-level phylogenetic tree was reconstructed using mitochondrial genome data (Table 2, Tables S1 and S2, Figure 3 and Figure 4). Within Crambidae, the eight subfamilies were divided into two major clades. Within Spilomelinae, the nine tribes formed a topology as follows: Lineodini + Agroterini + (((Hymeniini + Herpetogrammatini) + (Trichaeini + Nomophilini)) + (Spilomelini + Unplaced) + Margaroniini).
The type species of the two new genera, Scintilla strigiferalis (Hampson, 1900) comb. nov. and Melanoleuca luteamacula sp. nov., were both placed within the tribe Agroterini. Their relationship to other Agroterini was recovered as: (S. strigiferalis + (Nagiella inferior + Haritalodes derogata)) + (M. luteamacula + (Patania balteata + Notarcha quaternalis)). Tylostega tylostegalis formed a separate branch outside the core Agroterini clade in our analysis.
Geographically, the revised genus Tabidia exhibits a tropical Asia-Australasia pattern with no confirmed records in China, while Scintilla gen. nov. and Melanoleuca gen. nov. are primarily distributed within East Asia (Figure 5). Notably, the three new Melanoleuca species described herein are endemic to specific mountainous regions of Central, Southwestern, and Eastern China (Figure 6).
Maximum likelihood phylogeny based on complete mitochondrial genomes, illustrating the positions of Scintilla gen. nov. and Melanoleuca gen. nov. within Crambidae.
Distribution of Scintilla gen. nov., Melanoleuca gen. nov., and Tabidia. Yellow, red, and blue symbols represent two species of Scintilla; five species of Melanoleuca, and nine species of Tabidia, respectively.
Type locality map of three new species in the genus Melanoleuca gen. nov.
3.4. Taxonomic Treatment of Scintilla Wang & Xiao, gen. nov.
Scintilla Wang & Xiao, gen. nov. [星火野螟属] https://zoobank.org/90DECB15-9FB8-466F-928E-020F4832BEC0
Type species. Scintilla strigiferalis (Hampson, 1900) comb. nov., redescription of the habitus and genitalia was provided by Du & Li (2014) [15].
Generic Characters. Adult (Figure 7A,C and Figure 8A): Body yellowish. Frons rounded. Labial palpus upturned and curved, third segment long, acuminate. Maxillary palpus minute, filiform. Male antenna ciliate ventrally. Wings pale yellow ground, scattered fine light brown striae. Forewing CuA_2_ origin near 3/4 distal discal cell, CuA_1_, M_3_, M_2_ separate from discal cell posterior angle; Rs_4_ straight, well separated from Rs_2_+Rs_3_; Rs_1_ approx to Rs_2_+Rs_3_. Hindwing CuA_1_, M_3_, M_2_ from discal cell posterior angle; M_1_ from discal cell anterior angle; Rs from distal 1/4 of Sc+R.
Male Genitalia (Figure 10A,B): Uncus and gnathos both reduced to a lip-like form; valvae well developed.
Female Genitalia (Figure 10C): Papillae anales narrow. Ductus bursae short and broad, wrinkled.
Distribution. This genus is mainly distributed in China, South Korea, and Russia.
Etymology. The genus name Scintilla is derived from the Latin word scintilla= spark, and refers to the spark like striped patterns on the wings.
Remarks. Phylogenetic trees constructed based on mitochondrial genomes revealed that Scintilla strigiferalis (Hampson, 1900) comb. nov. clusters within the clade of the tribe Agroterini Acloque, 1897. Furthermore, this species conforms to the diagnostic characters of Agroterini as described by Mally et al. (2019) [64], namely: labial palps upturned and third labial palpomere directed dorsally; uncus with a broad base; head chaetae simple and undivided; and the ratio of saccus length to sacculus breadth greater than one. Therefore, we place this genus in the tribe Agroterini.
Head and wing venation of the type species of Scintilla gen. nov. and Melanoleuca gen. nov. (A). Head of Scintilla strigiferalis (Hampson, 1900) comb. nov., male; (B). Head of Melanoleuca luteamacula sp. nov., male; (C). Wing venation of Scintilla strigiferalis (Hampson, 1900) comb. nov., male; (D). Wing venation of Melanoleuca luteamacula sp. nov., male.
Habitus of Scintilla species (A). Scintilla strigiferalis (Hampson, 1900) comb. nov., female; (B). Scintilla obvia (Du & Li, 2014) comb. nov., male. Scale bars: 1 cm.
Habitus of Melanoleuca species (A). Melanoleuca luteamacula sp. nov., holotype; male; (B). Melanoleuca qianshanensis sp. nov., holotype; male; (C). Melanoleuca yingshanensis sp. nov., holotype; male; (D). Melanoleuca candidalis (Warren, 1896) comb. nov., male (after Du & Li, 2014); (E). Melanoleuca longatuba (Wang, Du & Xiao, 2024) comb. nov., holotype, male; (F). Melanoleuca longatuba (Wang, Du & Xiao, 2024) comb. nov., paratype, female (after Wang et al., 2024). Scale bars: 0.5 cm.
Genitalia of Scintilla species (A). Scintilla strigiferalis (Hampson, 1900) comb. nov., male, genitalia slide no. Lep3599; (B). Scintilla obvia (Du & Li, 2014) comb. nov., genitalia slide no. WJX25love-16-1; (C). Scintilla strigiferalis (Hampson, 1900) comb. nov., female, genitalia slide no. Lep 3600. Scale bars: 0.5 mm (A,B); 1 mm (C).
Key to Scintilla gen. nov. species based on morphology of habitus and genitalia (After Du & Li, 2014, [15])
Subterminal line of hindwing interrupted and indistinct; posterior margin of valva with basal 2/3 arched, bearing a developed bundle of long hairs at middle; costa distally curved toward anterior margin........................Scintilla strigiferalis **comb. nov.**Subterminal line of hindwing uninterrupted and distinct; posterior margin of valva slightly concave, without bundle of long hairs; costa slightly concave.........................................................................................Scintilla obvia comb. nov.
3.4.1. Scintilla strigiferalis (Hampson, 1900) comb. nov. [细星火野螟]
Figure 7A,C, Figure 8A, Figure 10A,C and Figure 13ATabidia strigiferalis Hampson, 1900, Tr. Ent. Soc. Lond. 48: 386, pl. III, f. 30.Tabidia strigiferalis Hampson, 1900, Entomol. Fennica 25: 57–64, Figures 1c and 3a,b.
Note. Description of the habitus and genitalia was provided by Du & Li (2014) [15].
Material examined. 2 ♂♂, Potangyan Reservoir, Baihu Town, Lujiang County, Hefei City, Anhui Province, China, 31.2711° N, 117.3608° E, alt. 73.81 m, 2022-VIII-01, Jiaxin Wang, Peng Yu leg., specimen no. P42.1, genit. prep. Lep2131, Lep3599, molecular no. C4; 1 ♀, Baimajian, Dahuaiping Town, Huoshan County, Lu’an City, Anhui Province, China, 31.1377° N, 116.1877° E, alt. 794.41 m, 2023-VIII-09, Jiaxin Wang, Peng Yu leg., specimen no. H67, genit. prep. Lep3600, molecular no. H67.
Distribution: China (Heilongjiang, Jilin, Liaoning, Tianjin, Hebei, Shaanxi, Gansu, Henan, Anhui, Hubei, Chongqing, Sichuan, Guizhou, Zhejiang, Fujian, Guangdong, Hainan, Shandong, Shanxi, Inner Mongolia, Guangxi), Korea, Russia.
3.4.2. Scintilla obvia (Du & Li, 2014) comb. nov. [显星火野螟]
Figure 8B, Figure 10B and Figure 13BTabidia obvia Du & Li, 2014, Entomol. Fennica 25: 57–64, Figures 1d and 3c,d.
Note. Description of the habitus and genitalia was provided by Du & Li (2014) [15].
Material examined. 1 ♂, Dahe Bridge, Longtan Gorge Rafting, Wujia Mountain, Yingshan County, Hubei Province, China, 31.0727° N, 115.8274° E, alt. 262.81 m, 2022-VIII-10, Jiaxin Wang, Peng Yu leg., specimen no. P87.6, genit. prep. Lep2132, molecular no. C6; 2 ♂♂, Huangshadi, Piaoba Village, Mugan Town, Daguan County, Zhaotong City, Yunnan Province, China, 2023-VII-22, Rentao Xu, Mingxu Han leg., specimen no. WJX25love-16-1, genit. prep. WJX25love-16-1.
Distribution. China (Gansu, Hubei, Chongqing, Sichuan, Guizhou, Zhejiang, Guangdong, Hong Kong).
3.5. Taxonomic Treatment of Melanoleuca Wang & Xiao, gen. nov.
Melanoleuca Wang & Xiao, gen. nov. [熊猫野螟属] https://zoobank.org/E39C0107-2FB6-4796-BABE-7F2A5E380312
Type species. Melanoleuca luteamacula sp. nov., here designated.
Generic Characters. Adult (Figure 7B,D and Figure 9): Body yellowish white or white. Forehead rounded. Labial palpus upturned and curved, third segment long, tapering. Maxillary palpus minute, filiform. Male antennae with cilia on ventral surface. Wings yellowish white or white, with dark maculae scattered on base. Forewing origin of CuA_2_ near 2/3 of distal middle chamber, CuA_1_, M_3_ and M_2_ each arising from hind angle of middle chamber; bases of M_2_ slightly separated from CuA_1_ and M_3_; Rs_4_ slightly curved, clearly separated from Rs_2_ + Rs_3_; Rs1 close to Rs_2_+3. Hindwing CuA_1_, M_3_ and M_2_ each arising from hind angle of middle chamber; M_1_ arising from anterior angle of middle chamber; Rs arising from 3/10 of distal Sc+R.
Male genitalia (Figure 11): Valva lingulate. Costa more sclerotized. No clasper. Base of saccus narrow. Phallus without cornuti.
Female genitalia (Figure 12): Papillae anales narrow. Ductus bursae slender and long. Corpus bursae without signum.
Distribution. This genus is mainly distributed in China and India.
Etymology. The generic name is derived from the Latin word melano-(black) and the Latin word -leuca (white), referring to the black markings on the wings of species in this genus, which resemble the black and white pelage of the giant panda, Ailuropoda melanoleuca.
Remarks. Phylogenetic trees constructed based on mitochondrial genomes revealed that Melanoleuca luteamacula sp. nov. falls within the clade of the tribe Agroterini Acloque, 1897. Moreover, it conforms to the characteristics of the tribe Agroterini as described by Mally et al. (2019) [64], namely the labial palps upturned and third labial palpomere directed dorsally; uncus with a broad base, head chaetae simple and unsplit; the ratio between saccus length and sacculus breadth more than one. Therefore, we assign this genus to the tribe Agroterini.
Key to Melanoleuca gen. nov. species based on morphology of habitus and genitalia
Body and wings yellowish white...............................Melanoleuca longatuba comb. nov.
-Body and wings white..........................................................................................................2
2.lower third of median area fulvous..............................Melanoleuca luteamacula sp. nov.
-lower third of median area black........................................................................................3
3.Hindwing with outer half interruptedly blackish..Melanoleuca qianshanensis sp. nov.
-Hindwing with outer third interruptedly blackish..........................................................4
4.Distal half of juxta sharply projecting bilaterally; distal half of saccussubrhombic........................................................Melanoleuca yingshanensis sp. nov.
-Saccus oval and juxta oval..........................................Melanoleuca candidalis comb. nov.
3.5.1. Melanoleuca luteamacula Wang & Xiao sp. nov. [黄斑熊猫野螟]
http://zoobank.org/9C321A83-2584-4B02-908A-6CBDDC6FDFD4 Figure 7B,D, Figure 9A, Figure 11A, Figure 12A,D,E and Figure 14A
Type material. HOLOTYPE: 1 ♂, Huangbai Mountain Scenic Area, Changzhuyuan Township, Shangcheng County, Henan Province, China, 31.4341° N, 115.3559° E, alt. 587.01 m, 2020-VIII-19, Yunli Xiao, Yue Fu leg., specimen no. WJ22.5, genit. prep. Lep2347. PARATYPES: 1 ♂, Qinggao Village, Henggang Mountain, Yuchuan Town, Wuxue City, Huanggang City, Hubei Province, China, 30.1726° N, 115.6647° E, alt. 193.61 m, 2023-VIII-10, Jiaxin Wang, Peng Yu leg., specimen no. J2, molecular no. J2; 1 ♀, Nibai Village, Wadi Township, Yuanjiang County, Yunnan Province, China, 2024-VIII-07, Wanlu Liu leg., specimen no. NB203-6-6, genit. prep. WJX25love-34-2; 1 ♂, Biluo River, Baijixun Township, Weixi Lisu Autonomous County, Diqing Tibetan Autonomous Prefecture, Yunnan Province, China, 2024-VII-31, Yuanyuan Jin leg., genit. prep. WJX25love-7-1; 1 ♂, Nuge Village, Baijixun Township, Weixi Lisu Autonomous County, Diqing Tibetan Autonomous Prefecture, Yunnan Province, China, 2024-VII-30, Yuanyuan Jin leg. Other material. 1 ♀, Nuozhadu Town, Lancang Lahu Autonomous County, Pu’er City, Yunnan Province, China, 2024-IX-09, Wanlu Liu leg., specimen no. NZ2.R2-76, genit. prep. WJX25love-31-2; 1 ♂, Nibai Village, Wadi Township, Yuanjiang County, Yunnan Province, China, 2024-VIII-07, Wanlu Liu leg., specimen no. NB203-5-1, genit. prep. WJX25love-27-1; 1 ♀, Nuozhadu Town, Lancang Lahu Autonomous County, Pu’er City, Yunnan Province, China, 2024-VIII-30, Wanlu Liu leg., specimen no. NZ2.R1-86, genit. prep. WJX25love-30-2.
Diagnosis. This species is similar to Melanoleuca candidalis (Warren, 1896) comb. nov., but can be distinguished by the presence of a squarish orange yellow patch between the posterior margin of vein CuA_1_ and the subterminal line outside the antemedial line on the forewing, as well as distinct morphological features of the juxta and saccus in the male genitalia. In this species, the juxta is subcylindrical with a width about half of its length, and the saccus has a short narrow base with the remaining portion elongated cylindrical. In contrast, Melanoleuca candidalis comb. nov. possesses a squarish blackish brown patch in the same forewing region, and both its juxta and saccus are ovate in shape.
Description. Adult (Figure 7B,D, Figure 9A and Figure 14A): Wingspan 15.2–15.8 mm. Body white. Frons white mixed with ochreous yellow. Vertex with a white tuft. Antenna ochreous yellow; ventral pili length nearly equal to antennal diameter. Labial palpus: first segment white, second and third segments yellowish white. Maxillary palpus small, ochreous yellow. Proboscis brown, its base scaled with yellowish white. Patagium and tegula yellowish white. Thorax and abdomen: ventrally white, dorsally white mixed with ochreous yellow. Wings white; cilia white. Forewing: base and costal one-third slightly ochreous yellow, faintly mixed with greyish brown; antemedial line blackish brown, slightly excavate, with a black spot on the outer edge of its upper half; postmedial line blackish brown, with a large reniform black spot on the costa, protruding between veins M_2_ and CuA_1_, greyish brown and diffuse on the lower median part; a rounded black ovate spot at the apex. Hindwing: base scattered with long greyish brown piliform scales; outer third irregularly black on a white ground color, forming a small notch medially; distal discoidal stigma black. Legs white.
Male Genitalia (Figure 11A): Anales tube nearly cylindrical, apex truncate. Uncus irregular, with a ridged longitudinal carina from the base, tapering to two-thirds. Valva linguiform, densely setose with fine setae, terminal setae stronger. Costa of valva sclerotized, narrowing from base to apex. Transtilla sclerotized, separated medially. Sacculus sclerotized, slender except for a bulge at the basal 1/5. Saccus: basal one-fourth transversely small and ovate; from 1/4 to 1/2 narrow and stalk like; distal half nearly rhomboid, with a sharp spiniform process medially on the anterior margin. Juxta: distal half with sharply protruding lateral edges; basal half slightly narrower, nearly semicircular. Phallus uniformly cylindrical, apex obliquely truncate, its length approximately 1.3 times the length of the valva.
Female Genitalia (Figure 12A,D,E): Apophysis anterioris about 1.5 times as long as apophysis posterioris, expanded and acutely projected in I-shaped at about basal 1/4. Ductus bursae slender and elongated, length approximately 4.5 times as long as apophysis anteriors. Corpus bursae oval; signum absent.
Etymology: The specific name is derived from the Latin word lutea (yellow) and the Latin word macula (patch), referring to the squarish orange yellow patch between the posterior margin of vein CuA_1_ and the subterminal line outside the antemedial line on the forewing.
Distribution: China (Hubei, Henan, Yunnan).
Living adults of Scintilla species (A) Scintilla strigiferalis (Hampson, 1900) comb. nov., photographed by Xiao Shan in Wuhan, China. (B) Scintilla obvia (Du & Li, 2014) comb. nov., photographed by Lawrence Hylton in Hong Kong, China.
Living adults of Melanoleuca species (A). Melanoleuca luteamacula sp. nov., photographed by Gao Fan in Yunnan, China. (B). Melanoleuca sp., photographed by Li Jianbo in Zhejiang, China.
3.5.2. Melanoleuca qianshanensis Wang & Xiao, sp. nov. [潜山熊猫野螟]
http://zoobank.org/D199A326-1517-4C92-A271-ACF0AF5772D4 Figure 9B and Figure11B
Type material. HOLOTYPE: 1 ♂, Zhengguo Group, Tianzhu Village, Shuihou Town, Qianshan City, Anqing City, Anhui Province, China, 30.7271° N, 116.4389° E, alt. 638.41 m, 2022-VIII-16, Jiaxin Wang, Peng Yu leg., specimen no. P56.6, genit. prep. Lep2346, molecular no. D31.
Diagnosis. This species is similar to Melanoleuca yingshanensis sp. nov. and Melanoleuca candidalis (Warren, 1896) comb. nov., but can be distinguished by distinct morphological differences in the juxta and saccus of the male genitalia. In this species, the juxta is ovate, and the distal three-fifths of the saccus is nearly triangular. In contrast, the juxta of Melanoleuca yingshanensis has sharply protruding lateral edges in its distal half, and the distal half of the saccus is nearly rhomboid. Both the juxta and saccus of Melanoleuca candidalis are ovate in shape.
Description. Adult (Figure 9B): Wingspan 16.8 mm. Body white. Antenna dorsally scaled yellowish brown, ventrally brown with pili subequal to antennal diameter. Labial palpus yellowish brown, second segment distally slightly blackish brown. Maxillary palpus small, yellowish brown. Proboscis yellowish brown, base scaled yellowish white. Patagium white mixed greyish brown. Tegula white with long piliform scales. Thorax white. Abdomen white slightly tinged greyish brown. Forewing white, base irregularly greyish brown; antemedial line blackish brown, slightly excavate, with black spot on outer edge of upper half; distal discoidal stigma black reniform; postmedial line greyish brown, with large black spot on costa and perpendicularly excavate between M_2_ and CuA_1_, greyish brown on lower median part; ovate black spot at apex; cilia white except at apex mixed greyish brown. Hindwing white, outer half irregularly pale black on white ground, forming small notch medially; distal discoidal stigma black; terminal cilia white mixed greyish brown. Legs white slightly mixed greyish brown.
Male Genitalia (Figure 11B): Anales tube semitubular, ends connected, apex truncate. Uncus distally inflated, apex slightly narrow. Valva linguiform, densely setose with fine setae, terminal setae stronger. Costa of valva sclerotized. Costa of valva widening from near base to basal one-fourth, narrowing from one-fourth to apex. Transtilla sclerotized, separated medially. Saccus basal two-fifths narrow, distal three-fifths nearly triangular, with small dentiform process medially on anterior margin. Juxta nearly cylindrical, length about twice width. Phallus cylindrical, basal half slightly narrow, distal half slightly broad, slightly longer than valva.
Female. Unknown.
Etymology: The specific name is derived from the type locality, Qianshan City in Anhui Province, China.
Distribution: China (Anhui).
3.5.3. Melanoleuca yingshanensis Wang & Xiao, sp. nov. [英山熊猫野螟]
http://zoobank.org/21C8A12A-F570-4F86-83F9-5D5493651E46 Figure 9C, Figure 11C and Figure 12B,F,G
Type material. HOLOTYPE: 1 ♂, Longtan Gorge Power Station, Wujia Mountain, Yingshan County, Hubei Province, China, 31.0867° N, 115.8139° E, alt. 461.85 m, 2022-VIII-11, Jiaxin Wang, Peng Yu leg., specimen no. BP253, genit. prep. Lep3598, molecular no. WA1. PARATYPES: 1 ♀, Longtan Gorge Power Station, Wujia Mountain, Yingshan County, Hubei Province, China, 31.0867° N, 115.8139° E, alt. 461.85 m, 2022-VI-08, Jiaxin Wang, Peng Yu leg., specimen no. WJ22.4, genit. prep. Lep2345; 1♂, the second arc ditch down the path along Shigu Temple, Wujia Mountain, Yingshan County, Hubei Province, China, 31.1127° N, 115.7998° E, alt. 793.81 m, 2022-VIII-09, Jiaxin Wang, Peng Yu leg., specimen no. BP254; 1 ♂, Guanshan Nature Reserve, Jiangxi Province, China, 2017-VIII-26, Guangxin Wang, Wenjian Li leg., genit. prep. WJX25love-17-1.
Diagnosis. This species is similar to Melanoleuca qianshanensis sp. nov. and Melanoleuca candidalis (Warren, 1896) comb. nov., but can be distinguished by distinct morphological differences in the juxta and saccus of the male genitalia. In this species, the juxta has sharply protruding lateral edges in its distal half, and the distal half of the saccus is nearly rhomboid. In contrast, the juxta of Melanoleuca qianshanensis is ovate, and the distal 3/5 of the saccus is nearly triangular. Both the juxta and saccus of Melanoleuca candidalis are ovate in shape.
Description. Adult (Figure 9C): Wingspan 19.5–20.1 mm. Body white. Frons white mixed with ochreous yellow. Vertex with white tuft. Antenna ochreous yellow; ventral pili length subequal to antenna diameter. Labial palpus: first segment white, second and third segments yellowish white. Maxillary palpus small, ochreous yellow. Proboscis brown, scaled yellowish white basally. Patagium and tegula yellowish white. Thorax and abdomen: ventrally white, dorsally white mixed with ochreous yellow. Wings white; cilia white. Forewing: base and costal 1/3 slightly ochreous yellow, faintly mixed greyish brown; antemedial line blackish brown, slightly excavate; a black spot on outer edge of its upper half; postmedial line blackish brown, with large reniform black spot on costa, protruding between M_2_ and CuA_1_, greyish brown and diffuse on lower median part; a rounded black ovate spot at apex. Hindwing: base scattered with long greyish brown piliform scales; outer third irregularly black on white ground, forming a small notch medially; distal discoidal stigma black. Legs white.
Male Genitalia (Figure 11C): Anales tube nearly cylindrical, apex truncate. Uncus irregular, with a ridged longitudinal carina from base, tapering to 2/3. Valva linguiform, densely setose with fine setae, terminal setae stronger. Costa of valva sclerotized, narrowing from base to apex. Transtilla sclerotized, separated medially. Sacculus sclerotized, slender except for a bulge at basal 1/5. Saccus: basal 1/4 transversely small ovate; from 1/4 to 1/2 narrow stalk like; distal half nearly rhomboid, with a sharp spiniform process medially on anterior margin. Juxta: distal half with sharply protruding lateral edges; basal half slightly narrower, nearly semicircular. Phallus uniformly cylindrical, apex obliquely truncate, length approximately 1.3 times of valva.
Female Genitalia (Figure 12B,F,G): Apophysis anterioris about 1.5 times of apophysis posterioris, expanded and acutely projected in I-shaped at about basal 1/4. Ductus bursae slender and elongated, length approximately 7 times of apophysis anteriors. Corpus bursae small, elliptical; signum absent.
Etymology: The specific name is derived from the type locality, Yingshan County in Hubei Province, China.
Distribution: China (Hubei, Jiangxi).
3.5.4. Melanoleuca candidalis (Warren, 1896) comb. nov. [黑斑熊猫野螟]
Figure 9D and Figure 11DAripana? candidalis Warren, 1896, Ann. Mag. Nat. Hist. (ser. 6) 18: 169.Tabidia candidalis Hampson, 1898, Proc. Zool. Soc. Lond. 1898: 624.Tabidia candidalis (Warren, 1896), Entomol. Fennica 25: 57–64, Figures 1b and 2a,b.
Note. Description of the habitus and genitalia was provided by Du & Li (2014).
Distribution. China (Zhejiang, Guangdong), India (Du & Li, 2014).
3.5.5. Melanoleuca longatuba (Wang, Du & Xiao, 2024) comb. nov. [长管熊猫野螟]
Figure 9E,F, Figure 11E and Figure 12C,H,ITabidia longatuba Wang, Du & Xiao, 2024, Entomotaxonomia (2024) 46(1): 55–60, Figures 1–3.
Material examined. HOLOTYPE: 1 ♂, Henggang Mountain, Yuchuan Town, Wuxue City, Hubei Province, China, 30.1725° N, 115.6644° E, alt. 193 m, 2022-VIII-24, Jiaxin Wang, Peng Yu leg., specimen no. IOZ(E)224468, genit. prep. Lep3048, molecular no. WJXC3, GenBank accession no. OR371986. PARATYPES: 1 ♀, Wujia Mountain, Dabie Mountain National Nature Reserve, Yingshan County, Hubei Province, China, 31.0725° N, 115.8272° E, alt. 262 m, 2022-VI-06, Jiaxin Wang, Peng Yu leg., specimen no. IOZ(E)224469, genit. prep. Lep2200, molecular no. WJXC2, GenBank accession no. OR371985; 1 ♂, Sanjiao Mountain, Xishui County, Hubei Province, China, 30.4989° N, 115.5589° E, alt. 444 m, 2022-VIII-24, Jiaxin Wang, Peng Yu leg., specimen no. IOZ(E)224470, molecular no. WJXK2, GenBank accession no. OR596416.
Note. Description of the habitus and genitalia was provided by Wang et al. (2024) [16].
Distribution. China (Hubei).
3.6. Taxonomic Treatment of Tabidia Snellen, 1880
Tabidia Snellen, 1880 [条纹野螟属]
Type species. Tabidia insanalis Snellen, 1880
Distribution. This genus is range extends from South Asia to Australia, with no records in China, exhibiting a tropical Asia–Australasia distribution pattern.
Generic Characters. Adult: Body white. Frons rounded. Labial palpus sickle shaped, curved, long, slender, and acutely pointed. Maxillary palpus small. Male forewing with a dorsally located group of recumbent large scales from base of median nervure. Hindwing termen slightly wavy with a distinct anal angle. Wing pattern with black banded or zigzag markings. Forewing origin of CuA_2_ near base of median cell [1,5].
Remarks. Integrative evidence from morphology, DNA barcodes, and phylogenetic analyses of mitochondrial genomes indicates that the former concept of Tabidia requires division into two new genera. The three genera now recognized exhibit substantial diagnostic differences (Table 3). Furthermore, the species previously assigned to Tabidia do not form a monophyletic clade, justifying the taxonomic revision proposed here. This study clarifies that the true genus Tabidia, as redefined, is not distributed in China.
Following this revision, Tabidia comprises nine species (Table 4). However, this group remains non-monophyletic, as evidenced by distinct morphological disparities (Figure 15) and its polyphyly in phylogenetic trees (Figure 1) based on DNA barcodes. Due to limited specimen availability and molecular data, the taxonomic status of these nine species is provisionally maintained until more comprehensive morphological and molecular evidence becomes available.
Living adults of Tabidia species (A) Tabidia insanalis Snellen, 1880, photographed by Anthony Kurek at Babinda Boulders, Queensland, Australia. (B) Tabidia inconsequens (Warren, 1896), photographed by Gérard Chartier in Thma Bang, Kaôh Kong, Cambodia. (C) Tabidia aculealis (Walker, 1866), photographed by Mohammed Hayath in Dadadahalli, Karnataka, India. (D) Tabidia truncatalis Hampson, 1899, photographed by David White in Wonga, Queensland, Australia.
Table 4: A species list of the three genera: Scintilla gen. nov., Melanoleuca gen. nov., and Tabidia Snellen, 1880.
4. Discussion
4.1. Taxonomic Revision of Chinese Tabidia
Our integrated analysis, combining morphology, COI barcodes, and mitogenomic phylogeny, demonstrates that the Chinese species previously classified under Tabidia do not form a monophyletic group. This confirms the initial morphological suspicions regarding the heterogeneity of the genus. Consequently, we erect two new genera, Scintilla gen. nov. and Melanoleuca gen. nov., to accommodate these species. This revision is robustly supported by distinct morphological characters and stable molecular phylogenetic relationships. The conserved mitogenome structures of both new genera, showing the ancestral lepidopteran gene order, are consistent with patterns commonly observed in Spilomelinae.
4.2. Systematic Placement of the New Genera
The phylogenetic trees, particularly the robust mitogenomic analysis, consistently place the type species of both Scintilla and Melanoleuca within the tribe Agroterini. This finding aligns with previous studies based on molecular data [66,67] or mitochondrial genomic data [34,41,57,63,68], further confirming the tribal classification framework within Spilomelinae. However, they do not form a single clade but are recovered in two distinct subclades within Agroterini. This structure strongly supports their recognition as separate, valid genera within this tribe, rather than being merged into a single genus or placed elsewhere. The position of Tylostega tylostegalis outside the core Agroterini clade in our analysis highlights the potential non-monophyly of Agroterini as currently defined and warrants further investigation. Furthermore, the taxonomic status of the “Unplaced” group comprising Syllepte taiwanalis and Pycnarmon lactiferalis remains complex [64,69], reflecting the need for further exploration of higher-level phylogeny within Spilomelinae.
4.3. Delimitation and Composition of the New Genera
The monophyly of Scintilla gen. nov. (comprising S. strigiferalis comb. nov. and S. obvia comb. nov.) and Melanoleuca gen. nov. (comprising M. luteamacula sp. nov., M. qianshanensis sp. nov., M. yingshanensis sp. nov., M. candidalis comb. nov., and M. longatuba comb. nov.) is well-supported. Each genus possesses a unique combination of stable diagnostic characteristics in morphology and molecular sequences that distinguish them from each other and from other Agroterini genera. Species of Scintilla gen. nov. are mainly distributed in China, South Korea, and Russia, while those of Melanoleuca gen. nov., except for M. candidalis comb. nov., which is also found in India, are currently known only from China.
4.4. The Status of Tabidia Sensu Stricto and Remaining Species
After the transfer of the Chinese species, Tabidia sensu stricto comprises nine species. Our preliminary evidence suggests that this core group may also be non-monophyletic. For instance, the COI-based phylogeny and external morphology indicate that Tabidia defloralis likely belongs to the tribe Herpetogrammatini, clustering with Herpetogramma and Metoeca. This finding suggests that the genus Tabidia requires a comprehensive global revision. As noted by Yuan et al. (2019) [70], using a single gene locus (like COI) for reconstructing higher-level phylogeny in Spilomelinae may have limitations. Future studies should employ multi-gene data and detailed morphological comparisons (as used in Matsui et al. 2023 [71]; Tang & Du 2023 [72]) to validate these inferences. The taxonomic status of the remaining species, primarily known from original descriptions and distributed in the Australasian region, cannot be clarified without further morphological study and molecular data. Based on current knowledge, no definitive records of Tabidia sensu stricto exist in China. The revised Tabidia sensu stricto is mainly distributed in the Australasian region, including Papua New Guinea and Australia and coastal areas of Indonesia.
The reference list from the paper itself. Each links out to its DOI / PubMed record.
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