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Research Article
The new genus Purpurata (Lepidoptera, Crambidae, Spilomelinae), with descriptions of two new species from China
expand article infoXin-Lei Xue§, Xiao-Qiang Lu|§, Xi-Cui Du§
‡ Yibin Academy of Southwest University, Yibin, China
§ Southwest University, Chongqing, China
| Guangyuan Center for Disease Control and Prevention, Guangyuan, China
Open Access

Abstract

The male genitalia characters of four species, Botys iopasalis Walker, 1859, Pleuroptya obfuscalis Yamanaka, 1998, Botys plagiatalis Walker, 1859 and Patania shompen Singh et Ahmad, 2022, placed in the genus Patania Moore, 1888 before the present study, do not conform to the diagnosis of Patania. A new genus, Purpurata gen. nov., is established for these four species, and two new species, Purpurata directa sp. nov. and Purpurata lurida sp. nov. are described based on their external morphology and genitalia characters. Purpurata directa sp. nov. is designated as the type species of the new genus. Five species of the new genus were clearly separated from Patania species in the Maximum likelihood phylogenetic tree constructed based on COI sequence data. Compared to Patania, the new genus Purpurata exhibits distinctive characters in male genitalia: the uncus is short, broad, and arc-shaped posteriorly; the gnathos is present and setose, or reduced; and the fibula is very small and setose. In addition, Patania clava (Xu & Du), syn. nov. is synonymized with Purpurata iopasalis comb. nov. An identification key to species of the new genus is presented based on morphological characters of habitus and genitalia. Images of the habitus and genitalia are provided.

Key words

DNA barcode, identification key, morphology, moths, taxonomy

Introduction

The genus Patania Moore, 1888 was assigned to the Spilomelinae tribe Agroterini Acloque, 1897 by Mally et al. (2019). Many species of Patania were previously recorded in Pleuroptya Meyrick, 1890 (Munroe 1983, 1995; Shaffer et al. 1996; Leraut 2005; Du 2009; Kirpichnikova 2009; Heppner 2012; Sasaki and Yamanaka 2013), which was synonymized with Patania by Kirti and Gill (2007). During our study of the genus Patania, we found that four species, P. iopasalis (Walker, 1859), P. obfuscalis (Yamanaka, 1998), P. plagiatalis (Walker, 1859) and P. shompen Singh et Ahmad, 2022, exhibited some distinctive characters in the genitalia, such as on the uncus, gnathos, fibula, and apophyses anteriores, which did not correspond to the diagnostic characters of Patania. Additionally, some genitalia characters of these four species, such as the fibula, cornutus, and corpus bursae, did not correspond to the characters of another similar genus, i.e., Nagiella Munroe, 1976. Therefore, after systematically comparing the morphological characters of these four species with Patania and Nagiella species, and based on the phylogenetic analysis of DNA barcode data, we establish a new genus, Purpurata gen. nov., for these four known species and describe two new species.

Materials and methods

Taxon sampling

Specimens were collected by light trap at night and killed by ammonium hydroxide or ethyl acetate. The examined pinned specimens, including all type specimens of the new species, are deposited in the Insect Collection, Southwest University, Chongqing, China (SWU), except for six specimens of two known species, which are deposited in the Insect Collection of Nankai University (NKU). The corresponding author examined comparative specimens of Botys iopasalis and some Patania species, which were deposited in the Natural History Museum, London, United Kingdom (NHMUK). We extracted and obtained 11 sequences from five species of the new genus Purpurata, seven sequences from three Patania species, and five sequences from two Nagiella species (Table 1). All these sequences have been uploaded to NCBI. In addition, four sequences of two Agrotera species, one sequence of one Patania species, two sequences of one Pycnarmon species and six sequences of two Nosophora species were downloaded from NCBI and BOLD. Among them, Agrotera is the type genus of Agroterini (Mally et al. 2019), Patania and Nagiella are similar genera to Purpurata, and Pycnarman pantherata (Butler, 1878) and Nosophora are closely related to Botys iopasalis (Matsui et al. 2022), one of the focal taxa in the present study. We included the type species of Patania, Botys concatenalis Walker, 1866, in the analysis to deduce the monophyly of the new genus Purpurata. Therefore, the five genera Patania, Nagiella, Agrotera, Pycnarmon, and Nosophora are used as outgroups in this study. The information on specimens used for mitochondrial COI gene sequencing and phylogenetic analysis is provided in Table 1.

Table 1.

Sample information for species of Purpurata gen. nov. and the outgroups.

Species Sequence ID Location (China, except for last four) NCBI and BOLD accession no.
Purpurata iopasalis comb. nov. XD1401067 Wuzhishan, Hainan PQ463661
LXQ180076 Diaoluoshan, Hainan PQ463660
XD1401014 Diaoluoshan, Hainan PQ463662
Purpurata directa sp. nov. XD1401032 Wuzhishan, Hainan PQ463664
XD1401110 Diaoluoshan, Hainan PQ463663
Purpurata plagiatalis comb. nov. XD1500022 Mulun, Guangxi PP865067
Purpurata obfuscalis comb. nov. LXQ180070 Haugaoxi, Sichuan PP865068
LXQ180071 Haugaoxi, Sichuan PP865069
XD1405336 Huagaoxi, Sichuan KU143853
Purpurata lurida sp. nov. XD1401079 Diaoluoshan, Hainan PP865070
LXQ180262 Changyang, Hubei PP865071
Patania balteata XD1405399 Haugaoxi, Sichuan KU143838
XD1405300 Haugaoxi, Sichuan KU143837
XD1405441 Haugaoxi, Sichuan KU143839
Patania chlorophanta XD1404265 Jinzhongshan, Guangxi KU058652
XD1404239 Jinzhongshan, Guangxi KU058653
XD1401035 Wuzhishan, Hainan KU058654
Patania concatenalis XD1401058 Diaoluoshan, Hainan KU143840
Pyr000116 Lingshui, Hainan CNPYA116-10
Pycnarmon pantherata Pyr000126 Yuanqu, Shanxi CNPYA126-10
Pyr000127 Yuanqu, Shanxi CNPYA127-10
Nagiella hortulatoides LXQ180100 Huanglianshan, Yunnan PQ463665
LXQ180099 Huanglianshan, Yunnan PQ463666
LXQ180217 Huanglianshan, Yunnan PQ463667
Nagiella quadrimaculalis XD1405327 Haugaoxi, Sichuan KU143854
XD1402131 Bawangling, Hainan PP865072
Nosophora dispilalis Pyr001429 Nanning, Guangxi CNPYD1429-10
Pyr001430 Nanning, Guangxi CNPYD1430-10
Pyr001431 Nanning, Guangxi CNPYD1431-10
Nosophora semitritalis Pyr000949 Yuanqu, Shanxi CNPYD949-10
Pyr000950 Yuanqu, Shanxi CNPYD950-10
Pyr000951 Yichang Wufeng, Hubei CNPYD951-10
Agrotera nemoralis ODOPE217-11 Bavaria, Germany KX045648
PHLAC175-10 South Tyrol, Italy JF859792
Agrotera basinotata ANICN229-10 Queensland, Australia HQ952613
ANICN230-10 Queensland, Australia HQ952614

The preparation of genitalia slides primarily followed Li and Zheng (1996). The habitus images were taken with a digital camera (Canon EOS 5D), and the genitalia images were taken with a digital camera (Leica DFC 450) attached to a stereomicroscope (Leica M205 A).

DNA extraction, PCR amplification, sequencing

A total of 15 species, including five species of Purpurata gen. nov. and ten outgroup species, were included for PCR analysis and DNA sequencing (Table 1). Total DNA was extracted from dried legs using the TIANGEN DNA extraction kit according to the manufacturer’s instructions. The 658-base pair (bp) barcode region of COI was PCR amplified using primers LepF1 and LepR1 (Hajibabaei et al. 2006). After verifying PCR products by running them on a 1% agarose gel, sequencing was conducted by Shanghai Sangon Biotechnology Co., Ltd (Shanghai, China) using the same primers as those used in PCR.

Data analysis

All COI sequences were aligned manually using Align-Muscle in MEGA 7.0, and were translated into amino acid sequences for visual correction. Intraspecific and interspecific genetic distances were calculated based on the Kimura 2-parameter (K2P) distance model (Kimura 1980). Phylogenetic analysis was performed based on a phylogenetic tree constructed using the Maximum likelihood (ML) method with 1,000 bootstrap replications (Saitou and Nei 1987), in which three Patania species, two Nagiella species, one Pycnarmon species, two Nosophora species and two Agrotera species were chosen as the outgroups.

Results

DNA sequence analysis

A total of 11 COI sequences from five species of Purpurata gen. nov. were obtained. In total, 36 COI sequences from Purpurata species and the outgroup species were analyzed. The dataset contained no obvious pseudogenes, indicating the correct target gene sequence was amplified and sequenced.

The ML phylogenetic tree shows five monophyletic branches for Purpurata gen. nov., corresponding to the five morphological species P. iopasalis comb. nov., P. lurida sp. nov., P. obfuscalis comb. nov., P. directa sp. nov., and P. plagiatalis comb. nov. (Fig. 1). These branches form a well-supported (98% Bootstrap support) monophylum that is sister to Nosophora. Patania, with three sampled species, including the type species P. concatenalis, is polyphyletic, with Nagiella, Pycnarmon, and Agrotera nested within. The intraspecific genetic distances within Purpurata range from 0.00% to 0.01%, and interspecific genetic distances range from 3.58% to 9.49% (Table 2). The interspecific genetic distances between the ingroup species and outgroup species range from 9.15% to 13.54% (Table 2).

Figure 1. 

Phylogenetic hypothesis of relationships among five species of Purpurata gen. nov. and ten species of five outgroup genera inferred from a Maximum likelihood (ML) analysis of the DNA barcode data. Numbers near the branches are bootstrap support values based on 1000 replicates.

Table 2.

Kimura 2-parameter genetic distances (as percent) calculated within (in bold) and between Purpurata species and the outgroups.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
1. Purpurata iopasalis comb. nov. 0.00
2. Purpurata obfuscalis comb. nov. 8.72 0.01
3. Purpurata directa sp. nov. 6.93 9.49 0.00
4. Purpurata plagiatalis comb. nov. 6.84 8.36 5.45 0.00
5. Purpurata lurida sp. nov. 3.58 6.52 8.28 7.85 0.00
6. Patania concatenalis 9.91 12.76 10.90 10.80 11.87 0.00
7. Patania balteata 10.59 13.09 10.25 9.38 12.36 10.60 0.00
8. Patania chlorophanta 9.15 12.50 11.36 11.06 12.07 12.07 8.53 0.00
9. Nosophora dispilalis 10.95 11.72 11.40 10.94 11.31 10.25 12.01 10.53 0.00
10. Nosophora semitritalis 10.77 11.14 10.34 10.07 10.25 11.51 12.36 11.41 7.20 0.00
11. Pycnarmon pantherata 10.42 12.23 11.21 10.95 11.30 10.77 8.87 9.39 10.07 11.84 0.00
12. Nagiella hortulatoides 11.30 11.43 12.47 10.43 12.85 10.90 9.21 10.95 10.89 11.84 9.40 0.00
13. Nagiella quadrimaculalis 11.39 12.29 11.58 10.68 12.64 10.69 8.95 10.98 10.33 11.58 8.96 4.47 0.00
14. Agrotera nemoralis 11.92 13.54 12.29 11.21 13.18 10.86 9.81 11.98 11.56 11.03 11.21 11.03 11.65 0.00
15. Agrotera basinotata 10.24 12.92 10.43 10.25 10.94 9.74 7.18 11.15 11.93 10.96 9.22 10.43 9.46 8.37 0.00

Taxonomic account

Purpurata gen. nov.

Type species

Purpurata directa sp. nov., here designated.

Diagnosis

Purpurata is readily distinguished from its phylogenetic sister genus Nosophora (type species: Botys dispilalis Hampson, 1896) by the yellow wings with their purple-brown lines and patches, while in Nosophora, the wings are dark shiny brown and feature a large, comma-shaped spot in the forewing. Furthermore, the vertex of the head is hollowed out in males of N. dispilalis (see Lederer 1863: pl. 4 fig. 24 (misidentified as N. chironalis); Hampson 1896: 288), whereas it is rounded in Purpurata species (Fig. 2). In Purpurata, compared to the similar genera Patania and Nagiella, the uncus is arc-shaped on the posterior margin, the valva is shorter and broader than those of the latter two, and the fibula is shorter and smaller in male genitalia; the apophyses anteriores is not broadened or very slightly broadened near the base in the female genitalia. Comparatively, in Patania and Nagiella, the uncus is generally trapezoidal (except that N. bispina has a rather short and broad uncus, with rounded posterior margin), and the fibula are generally well developed and sclerotized; the apophyses anteriores broaden rhomboidally near the base. In addition, in Purpurata, the gnathos is present and setose apically, generally undeveloped (except that P. iopasalis and P. shompen have well-developed gnathos). Patania species usually lack a gnathos, and a few species have a gnathos but no setae at the apex. Nagiella species generally have a gnathos, but no setae at the apex. In Purpurata, the phallus has a protruding sclerotized structure at the posterior end, with a thick needle-like or spine-like cornutus and a brush-like cornutus. Comparatively, the phallus of Patania and Nagiella lack a protruding sclerotized structure posteriorly. Patania species have, or lack, a cornutus (if present, diverse morphologically), and Nagiella species generally lack a cornutus (except for N. bispina with a curved hook cornutus). The compared characters here are present in the type species of Patania and Nagiella, Botys concatenalis, and Nagia desmialis respectively, which were fully investigated in our study.

Description

Habitus. Body and wings yellow, with purple-brown wing markings. Frons rounded. Labial palpus upturned, exposed 3rd joint short and blunt (Fig. 2). Antenna filiform, male with ventral cilia. Forewing with length of cell ~ 1/2 of wing; discocellulars incurved; R from cell at ~ 5/6 above; Rs2 anastomosed with Rs3 ~ 3/5 of Rs3 beyond cell; Rs1 close to Rs2+s3; Rs4 slightly curved towards Rs2+s3 at base; M2, M3 and CuA1 from posterior angle of cell and uniformly spaced at the base; CuA2 from cell at ~ 2/3 below. Hindwing with length of cell ~ 1/3 of wing; discocellulars incurved; Sc+R anastomosed with Rs ~ 1/5 beyond the cell; M2, M3 and CuA1 separately from posterior angle cell; CuA2 from cell at 2/3 below (Figs 3, 4). Abdomen yellow dorsally; 1st and 2nd tergites with black spots laterally and 7th tergites black posteriorly in male. Tympanal organs with praecinctorium strongly bifid (Fig. 5).

Figures 2–5. 

Purpurata directa sp. nov. 2 head ♂ 3 wing venation ♂ 4 wing venation ♀ 5 tympanal organ. Scale bars: 0.5 mm (2, 5); 1.0 mm (3, 4).

Male genitalia. Uncus short and broad, with arc-shaped posterior margin, except for P. iopasalis with narrowed apex of uncus. Gnathos present and bearing setae apically, or vestigial to a transverse band. Valva ligulate, relatively broad and short generally. Fibula small and membranous, with setae. Sacculus undeveloped. Phallus cylindrical, with a protruding sclerotized structure posteriorly, with a thick needle-like or spine-like cornutus and a brush-like cornutus composed of a spine cluster, and a granular sclerotized area near the middle.

Female genitalia. Apophyses anteriores longer than apophyses posteriores, occasionally slightly broadened near base. Antrum sclerotized. Ductus bursae usually long and slender. Corpus bursae round or oval, usually with a round signum.

Distribution

This genus is mainly distributed in the Palaearctic and Oriental realms. And P. plagiatalis is also found in the Neotropical and Australasian realms, P. iopasalis is also found in the Australasian realms.

Etymology

The genus name Purpurata is derived from the Latin word purpuratus meaning purple, indicating the distinctive purple-brown coloration of wing markings. The gender is feminine.

Remarks

This new genus comprises four known species from the genus Patania of Spilomelinae, and two new species described in the present study. It corresponds to the main diagnostic characters of Crambidae: forewing with vein RS4 free and with oval sclerotization costad at base of vein 1A+2A; bullae tympani open cephalad; tympanum and conjunctivum lying at a blunt angle; praecinctorium present; and male genitalia without uncus arms. Further, it corresponds to the main diagnostic characters of Spilomelinae: fornix tympani projecting ventrad from tympanic frame; praecinctorium bilobed; retinacular hook absent; and females with two frenular bristles. Moreover, based on the Agroterini characters stated in Mally et al. (2019), labial palps upturned and 3rd labial palpomere directed dorsally; uncus with a broad base, head chaetae simple and unsplit; the ratio between saccus length and sacculus breadth > 1, we place this genus in the tribe Agroterini.

Key to Purpurata species based on morphology of habitus and genitalia

1 Postmedial line of forewing punctiform between M2 and CuA2 (Figs 7, 11); phallus posteriorly with a nail head-like, rectangular, or rounded protruding sclerite (Figs 14c, 19c) 2
Postmedial line of forewing dentate or linear between M2 and CuA2 (Figs 6, 8–9, 10); phallus posteriorly with a finger-like or oval protruding sclerite (Figs 12c, 15c, 17c) 4
2 Gnathos vestigial to a narrow band (Fig. 19a); sacculus with a lamellar projection near distal end (Fig. 19); phallus posteriorly with a rounded protruding sclerite, and a short thorn-like cornutus near end, besides needle-like cornutus and brush-like cornutus (Fig. 19c) P. plagiatalis comb. nov.
Gnathos thick finger-like (Fig. 14a); sacculus without lamellar projection; phallus posteriorly with a nail head-like or rectangular sclerite, and without short thorn-like cornutus near distal end, besides needle-like cornutus and brush-like cornutus (Fig. 14c) 3
3 Postmedial line of hindwing punctiform between M2 and CuA2 (Fig. 7); phallus posteriorly with a nail head-like sclerite (Fig. 14c) P. iopasalis comb. nov.
Postmedial line of hindwing linear between M2 and CuA2; phallus posteriorly with a rectangular sclerite P. shompen comb. nov.
4 Postmedial line of forewing linear between M2 and CuA2 (Figs 8, 9); phallus posteriorly with an oval sclerite (Fig. 15c); gnathos transverse lamina (Fig. 15a) P. directa sp. nov.
Postmedial line of forewing dentate between M2 and CuA2 (Figs 6, 10); phallus posteriorly with a finger-like sclerite (Figs 12c, 17c); gnathos vestigial to a narrow band (Figs 12a, 17a) 5
5 Uncus short and broad (Fig. 17a), triangular; valva with distal 1/3 narrowed very gradually and rounded apically (Fig. 17) P. lurida sp. nov.
Uncus semicircular (Fig. 12a); valva with distal 1/3 narrowed gradually and pointed apically (Fig. 12) P. obfuscalis comb. nov.

Purpurata obfuscalis (Yamanaka, 1998), comb. nov.

Figs 6, 12, 12a–c, 13

Pleuroptya obfuscalis Yamanaka, 1998: 106. Type locality: Nepal.

Patania obfuscalis: Nuss et al. 2003–2024. Global Information System on Pyraloidea.

Material examined

ChinaChongqing Municipality, 2 ♂♂, 1 ♀, Jinyun Mountain, alt. 550 m, 29 July 2010, Xi-Cui Du & Chao-Wei Bi leg., genitalia slide no.: XLJ14221 ♀ • 1 ♂, Simian Mountain, alt. 1120 m, 17 July 2010, Xi-Cui Du & Li-Fang Song leg. • 2 ♂♂, JinYin Mountain, Qianjiang District, alt. 1100 m, 25–26 July 2012, Jun Zhang & Li-Jun Xu leg. • 1 ♂, Small South Sea, Qianjiang District, alt. 370 m, 21 July 2012, Jun Zhang & Li-Jun Xu leg. • Sichuan Prov., 4 ♂♂, Huagaoxi Nature Reserve, Guandou Village, alt. 763 m, 30 August 2014, Dan-Xu & Xue-Li Wei leg., genitalia slide no.: XDJ15048 ♂ • Gansu Prov., 2 ♂♂, Bifeng Valley, Wen County, alt. 860 m, 2005. 9–10 July 2012, Hai-Li Yu leg. (NKU) • Yunnan Prov., 2 ♂♂, Pianma Village, Lushui County, Nujiang Prefecture, alt. 1889 m, 18 August 2015, Xue-Li Wei leg., genitalia slide no.: LXQ18279 • 2 ♂♂, Cangyuan County, Lincang City, alt. 1242 m, 25 July 2015, Xue-Li Wei leg. • 1 ♂, Jinuo Township, Xishuangbanna, alt. 1100 m, 15 May 2018, Xi-Cui Du leg. • 2 ♂♂, Xishuangbanna Tropical Botanical Garden, alt. 659 m, 28 May 2015, Man-Fei Tao leg. • 2 ♂♂, Daxichang Village, Malipo County, alt. 1465 m, 5 June 2015, Man-Fei Tao leg. • 2 ♂♂, Huanglian Mountain, Honghe Prefecture, alt. 900 m, 23 May 2018, Xi-Cui Du & Xiao Qiang-Lu leg. • Xizang Autonomous Region, 1 ♂, Zhangmu Town, alt. 2300 m, 5 August 2017, Jian-Yue Qiu & Hao Xu leg., genitalia slide no.: XXL23275 ♂ • Guangxi Zhuang Autonomous Region, 1 ♂, Nonggang National Nature Reserve, Longzhou County, alt. 188 m, 3 August 2011, GuiQing-He leg. • Guizhou Prov., 2 ♂♂, 3 ♀♀, Baishao, Kuankuoshui, alt. 800 m, 10–11, 13 August 2010, Xi-Cui Du leg. • 1 ♂, WengAng village, LiBo County, alt. 1345 m, 20 July 2015, Ji-Ping Wan leg. • Hainan Prov., 4 ♀♀, Wuzhi Mountain, alt.795 m, 18 May 2014, Li-Jun Xu & Dan Xu leg. • 1♂, Diaoluo Mountain, alt. 500 m, 23 May 2014, Li-Jun Xu & Dan Xu leg. • Shaanxi Prov., 1 ♂, Ningshan County, Xunyangba Town, alt. 1400 m, 3 August 2014, Hai-Li Yu leg. • Hubei Prov., 1♂, Pingbaying National Forest Park, Xianfeng County, alt. 280 m, 21 July 1999, Hou-Hun Li leg. (NKU) • 5 ♂♂, 2 ♀♀, Hejiaping Town, Changyang County, alt. 800 m, 18 June 2018, Xi-Cui Du & Xiao Qiang-Lu leg., genitalia slide no.: LXQ18140 ♀ • Anhui Prov., 1 ♂, Tang Kou Town, Huangshan City, 4 August 2004, Jia-Sheng Xu & Jia-Liang Zhang leg. (NKU) • Zhejiang Prov., 1 ♂, Longtang Mountain, alt. 500 m, 22 May 2012, Xiao-Bing Fu leg. • 2 ♂♂, Tianmu Mountain, alt. 400 m, 24–25 July 2011, Xi-Cui Du & Xiao Bing-Fu leg. • Hong Kong Special Administrative Region, 1 ♂, Kadoorie Farm, alt. 21 m, 13 April 2007, Hou-Hun Li leg. (NKU).

Diagnosis

This species is distinguished by wings with postmedial line dentate and excurved between M2 and CuA2 (Fig. 6); gnathos vestigial to a narrow band (Fig. 12a); valva with distal 1/3 narrowed gradually and pointed apically, costa arched medially and bearing a cluster of long setae (Fig. 12); fibula a triangular lamina, with setae medially (Fig. 12b); phallus posteriorly with a finger-like protruding sclerite, a thick, spine-like cornutus and a brush-like cornutus (Fig. 12c).

Redescription

Habitus (Fig. 6). Forewing length 8.0–15.0 mm, wingspan 25.0–33.0 mm. Frons and vertex yellowish brown. Labial palpus with 1st segment yellowish white ventrally, the remainder yellowish brown. Maxillary palpus brown. Antenna yellowish brown, with ventral cilia ~ 1/3 in length of diameter of flagellomere in male. Patagium and tegula yellow, with brown patches. Thorax yellow dorsally, white ventrally. Legs yellowish white, distal end of front tibia black. Wings yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line slightly wavy, accompanied by a large elliptical pale patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally; postmedial line obliquely inward from costa, dentate and excurved between M2 and CuA2, then incurved to discoidal stigma below and sinuous to inner margin; an irregular large patch between anterior postmedial line and terminal margin, another irregular large patch below discoidal stigma and extended to tornus; a line of small spots along terminal margin. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA2, not apparent afterwards; an irregular large patch near apex beyond anterior postmedial line; a misty band below discoidal stigma, accompanied by another irregular misty wide band extended to tornus. Abdomen yellow dorsally; 1st and 2nd tergites with pale black spots laterally; 7th tergites black posteriorly in male; terminally black in some female individuals.

Male genitalia (Fig. 12). Uncus semicircular. Gnathos vestigial to a narrow band (Fig. 12a). Valva with distal 1/3 narrowed gradually, and pointed apically; costa arched medially and bearing a cluster of long setae. Fibula a triangular lamina, with setae medially (Fig. 12b). Saccus oval. Juxta a broad plate, invaginated anteriorly. Phallus posteriorly with a finger-like protruding sclerite, with a thick spine-like cornutus and a brush-like cornutus composed of a spine cluster (Fig. 12c).

Female genitalia (Fig. 13). Apophyses anteriores ~ 2 × length of apophyses posteriores. Ostium bursae relatively large, antrum broad. Ductus bursae ~ 2 × length of corpus bursae. Corpus bursae nearly rounded, with a round signum.

Distribution

China (Chongqing, Sichuan, Gansu, Xizang, Yunnan, Guangxi, Guizhou, Hainan, Shanaxi, Hubei, Anhui, Zhejiang, Xianggang), Nepal (Yamanaka 1998).

Purpurata iopasalis (Walker, 1859), comb. nov.

Figs 7, 14, 14a–c

Botys iopasalis Walker, 1859: 652. Type locality: India (Hindustan). Type depository: NHMUK.

Boty boteralis Walker, 1859: 716. Type locality: Malaysia (Sarawak).

Sylepta [sic] marcidalis Swinhoe, 1906: 382.

Pleuroptya iopasalis: Inoue 1982: 1: 343, 2: 234, 454.

Patania clava Xu & Du, 2016: 130, figs 1–4, 9–10. Syn. nov. Type locality: China (Hainan).

Material examined

Holotype of Patania clava • 1 ♂, China, Hainan Prov., Diaoluo Mountain, alt. 900 m, 23 May 2014, Li-Jun Xu & Dan Xu leg., genitalia slide number XD15056. Paratype of P. clava • 1 ♀, same data as holotype, genitalia slide number XD15050.

Figures 6–11. 

Habitus of Purpurata species 6 P. obfuscalis7 P. iopasalis8 P. directa sp. nov. holotype ♂ 9 P. directa sp. nov. paratype ♀ 10 P. lurida sp. nov. holotype ♂ 11 P. plagiatalis ♂. Scale bars: 0.5 cm.

Additional material

ChinaHainan Prov., 1 ♂, Wuzhi Mountain, alt. 795 m, 20 May 2014, Li-Jun Xu & Dan Xu leg., genitalia slide no.: XXL23277.

Diagnosis

This species is similar to P. obfuscalis in appearance, but can be distinguished by the larger size (forewing length 13.0–19.0 mm, wingspan 33.0–35.0 mm), antenna with ventral cilia approximately as long as flagellomere diameter in male, postmedial line of fore and hind wings punctiform between M2 and CuA2 (Fig. 7); uncus nearly triangular (Fig. 14), gnathos thick, finger-like (Fig. 14a), costa of valva without a cluster of setae medially, fibula densely covered with setae (Fig. 14b), phallus posteriorly with a nail head-like protruding sclerite (Fig. 14c) in male genitalia (Fig. 14); ductus bursae ~ 4 × length of corpus bursae, signum absent in female genitalia. In P. obfuscalis, the forewing length is 8.0–15.0 mm and wingspan is 25.0–33.0 mm, the ventral cilia of antenna is ~ 1/3 length of flagellomere diameter in male, the postmedial lines of the fore and hind wings are dentate between M2 and CuA2 (Fig. 6). In male genitalia, the uncus of P. obfuscalis is semicircular (Fig. 12), the gnathos is reduced to a narrow band (Fig. 12a), the costa of valva bears a cluster of long setae medially (Fig. 12), the fibula bears setae medially (Fig. 12b), the phallus posteriorly has a finger-like protruding sclerite (Fig. 12c); in female genitalia, the ductus bursae is ~ 2 × as long as the corpus bursae and a rounded signum is present (Fig. 13).

Distribution

China (Yunnan, Hainan, Fujian, Guangdong, Taiwan) (Lu and Guan 1953; Xu and Du 2016), Japan, Indonesia, India, Myanmar, Sri Lanka, Malaysia, Thailand, Timor Leste, Philippines, Caroline Islands, Papua New Guinea, Pakistan, Australia (Hampson 1896; Shibuya 1928, 1929; Klima 1939; Inoue 1982; Wang and Speidel 2000).

Remarks

The uncus and fibula of this species are narrower than in other species of this genus. We found that the external morphology and male genitalia characters of Patania clava were the same as those of Purpurata iopasalis. Their distinctive characters were that the postmedial lines of the fore and hind wings were punctiform between M2 and CuA2, the gnathos was thick finger-like and setose apically (Walker 1859; Inoue 1982, Yamanaka 1998; Xu and Du 2016). Therefore, Patania clava is synonymized with Botys iopasalis. For a detailed description of this species see Walker (1859) and Xu and Du (2016). Moreover, Xu and Du (2016) described that the labial palpus of Patania clava was pale yellow with a brown patch near the base, while we observed that this species had a subapical brown spot beside this basal spot, and the subapical brown spot was obscured in some individuals.

Purpurata directa sp. nov.

Figs 2, 8, 9, 15, 15a–c, 16

Patania iopasalis: Xu and Du 2016: 132, figs 5–8. Type locality: India (misidentification).

Diagnosis

This species is very similar to P. iopasalis in appearance, but can be distinguished by ventral cilia ~ 1/3 length of flagellomere diameter in male, antemedial line of forewing relatively straight and slightly inclined outward, postmedial line of fore and hind wings smooth linear between M2 and CuA2 (Figs 8, 9); uncus semicircular, gnathos a transverse lamina, and valva broader than the latter in male genitalia (Fig. 15); a signum present in female genitalia (Fig. 16). In P. iopasalis, the ventral cilia are approximately equal in length of flagellomere diameter in male, the antemedial line is slightly wavy, the postmedial line of fore and hind wings punctiform between M2 and CuA2 (Fig. 7); the uncus is nearly triangular, and the gnathos is finger-like in the male genitalia (Fig. 14a); and the signum is absent in the female genitalia.

Type material

Holotype : ♂ pinned, with genitalia in a separate slide. ChinaHunan Prov., Huping Mountain, Shimen County, alt. 350 m, 6 June 2017, Jian-Yue Qiu & Hao Xu leg., genitalia slide number: LXQ18315. Paratypes: pinned, some with genitalia in the separate slides, respectively. China • 6 ♂♂, same data as holotype • Hubei Prov., 1 ♂, Hejiaping Town, Changyang County, alt. 800 m, 18 May 2018, Xian-Qiang Lu & Xi-Cui Du leg. • Chongqing Municipality, 2 ♀♀, Daheba, Jinfo Mountain, alt. 600 m, 14, 17 July 2017, Shi-Man Bu leg • Sichuan Prov., 2 ♂♂, Emei Mountain, alt. 863 m, 17 July 2011, Jian-Bo Cao leg. • Guizhou Prov., 3 ♂♂, Baishao, Kuankuoshui, alt. 800 m, 10 August 2010, Xi-Cui Du leg., genitalia slide number: HGQ13018, HGQ13019 • 3 ♂♂, 1 ♀, Wengang village, Libo County, alt. 1345 m, 20 July 2015, Ji-Ping Wan leg. • Yunnan Prov., 1 ♂, Baihualing Village, Baoshan City, alt. 1520 m, 13 July 2007, Dan-Dan Zhang leg. • 1 ♂, Daxichang Village, Malipo County, alt. 1465 m, 7 August 2007, Man-Fei Tao leg. • Hainan Prov., 1 ♂, Bawangling National Forest Park, 11 June 2010, Li Kang leg., genitalia slide number: XLJ14105 • 3 ♂♂, 1 ♀, Wuzhi Mountain, alt. 795 m, 18, 19, 21 May 2014, Li-Jun Xu & Dan Xu leg., genitalia slide number: XD15024 ♂, XLJ14146 ♂, XLJ14147 ♀ • 1 ♂, Diaoluo Mountain, alt. 500 m, 25 May 2014, Li-Jun Xu & Dan Xu leg., genitalia slide number: XD15049 • Guangxi Zhuang Autonomous Region, 3 ♂♂, Nonggang National Nature Reserve, Longzhou County, alt. 188 m, 25, 27, 30 July 2011, Gui-Qing He leg., genitalia slide number: XLJ14083 (being identified as Patania iopasalis by Xu and Du (2016)), XLJ13199 • 3 ♂♂, Nonggang National Nature Reserve, Longzhou County, alt. 188 m, 2 August 2011, Gui-Qing He leg., genitalia slide number: XLJ13164 • 5 ♂♂, Mulun Nature Reserve, alt. 288 m, 21 July 2015, Dan Xu leg. • Guangdong Prov., 5 ♂♂, Renhua County, Danxia Mountain, alt. 408 m, 31 May 2018, Feng-Xia He leg.

Description

Habitus (Figs 2, 8, 9). Forewing length 9.0–13.5 mm, wingspan 21.0–30.0 mm. Frons and vertex yellow. Labial palpus yellowish white, with distal part of 2nd segment brown. Maxillary palpus yellowish white basally, brown near distal end. Antenna yellowish brown, with brown spots on scape, ventral cilia ~ 1/3 in length of diameter of flagellomere in male. Patagium and tegula yellow, with brown patches. Thorax yellowish brown dorsally, white ventrally. Legs pale yellow, front coxa and middle tibia with outer sides black at the base and distal end, front tibia with distal end black. Wings yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line relatively straight, slightly inclined outward, accompanied by a large elliptical pale patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally, connected to postmedial line posteriorly; postmedial line slightly obliquely inward from costa, straightly excurved between M2 and CuA2, then incurved to discoidal stigma below and sinuous to inner margin; an irregular large patch between anterior postmedial line and terminal margin, another irregular large patch below discoidal stigma and extended to tornus; a line of small spots along terminal margin. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA2, not apparent afterwards; an irregular large patch near apex beyond anterior postmedial line; a band below discoidal stigma slightly inclined towards tornus, accompanied by another irregular misty wide band extended to tornus. Cilia of fore and hind wings purple-brown, white basally. Abdomen yellow dorsally, white ventrally; 1st and 2nd tergites with black spots laterally; 7th and 8th tergites black posteriorly in male; terminally black in female.

Male genitalia (Fig. 15). Uncus semicircular. Gnathos a transverse lamina, with posterior end arc-shaped and setose (Fig. 15a). Valva broad tongue-shaped, with long setae along distal costa; sacculus a narrowed band; fibula a triangular lamina, setose basally (Fig. 15b). Saccus cylindrical, with rounded end. Juxta a narrowed plate. Phallus posteriorly with an oval protruding sclerite, with a thick, needle-like cornutus and a brush-like cornutus composed of a spine cluster (Fig. 15c).

Female genitalia (Fig. 16). Apophyses anteriores ~ 2 × length of apophyses posteriores. Ductus seminalis originating from antrum. Ductus bursae ~ 2 × length of corpus bursae. Corpus bursae nearly rounded, with a short transverse bar-like signum.

Distribution

China (Chongqing, Sichuan, Guizhou, Yunnan, Hainan, Hubei, Hunan, Guangdong, Guangxi).

Etymology

The species name directa is derived from the Latin word directus, an adjective, meaning straight, indicating the antemedial line of forewing relatively straight.

Purpurata lurida sp. nov.

Figs 10, 17, 17a–c, 18

Diagnosis

This species is similar to P. obfuscalis in appearance and genitalia, but can be distinguished by the body color paler than the latter (Fig. 10); uncus short and broad triangular (Fig. 17a); valva shorter and broader than the latter, with distal 1/3 narrowed very gradually and rounded apically (Fig. 17). In P. obfuscalis, the uncus is semicircular (Fig. 12a); the distal 1/3 of the valva is narrowed gradually and pointed apically (Fig. 12).

Type material

Holotype : ♂ pinned, with genitalia in a separate slide. ChinaGuizhou Prov., Kuankuoshui Nature Reserve, alt. 1500 m, 15 August 2010, Xi-Cui Du leg., genitalia slide number: XLJ14084. Paratypes: pinned, some with genitalia in separate slides. ChinaHubei Prov., 2 ♂♂, Hejiaping Town, Changyang County, alt. 800 m, 18 June 2018, Xiao-Qiang Lu & Xi-Cui Du leg. • 2 ♂♂, Xingdou Mountain, Maoba Town, Enshi, alt. 780 m, 30 July 2012, Jun Zhang & Xiao-Bing Fu leg. • Chongqing Municipality, 2 ♂♂, Jinyun Mountain, alt. 550 m, 22, July 2010, Li Kang & Xing-Fu Fu leg., genitalia slide number: XLJ14103 • 2 ♂♂, 1 ♀, Jinyun Mountain, 29 July 2010, Xi-Cui Du & Chao-Wei Bi leg., genitalia slide number: XLJ14104 ♀ • 1♂, Jinyun Mountain, 30 July 2012, Li-Jun Xu & Jian-Bo Cao leg. • 2 ♂♂, Jinyin Mountain, Qianjiang District, alt. 1100 m, 26 July 2012, Li-Jun Xu & Jun-Zhang leg. • Yunnan Prov., 1 ♂, Daxichang village, Malipo County, alt. 1465 m, 5 June 2015, Man-Fei Tao leg. • 2♀♀, Xishuangbanna Tropical Botanical Garden, alt. 659 m, 28 May 2015, Man-Fei Tao leg., genitalia slide number: LXQ18316 • Hainan Prov., 1 ♂, Diaoluo Mountain vocational village, alt. 500 m, 23 May 2014, Li-Jun Xu & Dan Xu leg. • Zhejiang Prov., 3 ♂♂, Tianmu Mountain, alt. 400 m, 24, 25, 30 July 2011, Xi-Cui Du & Xiao-Bing Fu leg.

Figures 12–16. 

Genitalia of Purpurata species 12, 13 P. obfuscalis 12 male, genitalia slide no. XXL23275 13 female, genitalia slide no. XLJ14221 14 P. iopasalis, male, genitalia slide no. XXL23277 15, 16 P. directa sp. nov. 15 male, holotype, genitalia slide no. LXQ18315 16 female, paratype, genitalia slide no. XLJ14147 12a, 14a, 15a gnathos 12b, 14b, 15b fibula 12c, 14c, 15c cornuti and posterior protrusion of phallus. Scale bars: 1.0 mm (12–16); 0.1 mm (12a–c, 14a–c, 15a–c).

Figures 17–20. 

Genitalia of Purpurata species 17, 18 P. lurida sp. nov. 17 male, holotype, genitalia slide no. XLJ14084 18 female, paratype, genitalia slide no. LXQ18316 19, 20 P. plagiatalis 19 male, genitalia slide no. HGQ13237 20 female, genitalia slide no. XLJ14204 17a, 19a gnathos 17b, 19b fibula 17c, 19c cornuti and posterior protrusion of phallus. Scale bars: 1.0 mm (17–20); 0.1 mm (17a–c, 19a–c).

Description

Habitus (Fig. 10). Forewing length 11.0–13.5 mm, wingspan 24.0–30.0 mm. Frons and vertex yellowish brown. Labial palpus with 1st segment yellowish white ventrally, the remainder yellowish brown. Maxillary palpus brown. Antenna yellowish brown, with ventral cilia ~ 1/3 in length of diameter of flagellomere in male. Patagium and tegula yellow. Thorax yellow dorsally, white ventrally. Legs yellowish white, distal end of front tibia black. Wings pale yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line slightly wavy, accompanied by a large elliptical pale patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally; postmedial line slightly obliquely inward from costa, dentate and excurved between M2 and CuA2, then incurved to discoidal stigma below and sinuous to inner margin; an irregular large patch near apex beyond anterior postmedial line, another misty patch near tornus below CuA2; a line of spots along marginal line. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA2, not apparent afterwards; an irregular large patch near apex beyond anterior postmedial line; a band below discoidal stigma, accompanied by another irregular misty wide band extended to tornus. Cilia of fore and hind wings yellowish white. Abdomen yellow dorsally, white ventrally; 1st and 2nd tergites with pale black spots laterally and 7th tergites black posteriorly in male.

Male genitalia (Fig. 17). Uncus short and broad triangular. Gnathos vestigial to a narrow band and bearing a few short setae (Fig. 17a). Valva broad tongue-shaped, with distal 1/3 narrowed very gradually, and rounded apically; costa arched medially and bearing a cluster of long setae; fibula a short lamina, setose basally (Fig. 17b). Saccus oval. Juxta a broad plate. Phallus posteriorly with a finger-like protruding sclerite, with a thick, needle-like cornutus and a brush-like cornutus composed of a spine cluster (Fig. 17c).

Female genitalia (Fig. 18). Apophyses anteriores ~ 2 × length of apophyses posteriores. Antrum broad, ductus seminalis originating from antrum. Ductus bursae ~ 2 × length of corpus bursae. Corpus bursae nearly oval, with a round signum.

Distribution

China (Chongqing, Guizhou, Yunnan, Hainan, Hubei, Zhejiang).

Etymology

The species name lurida is derived from the Latin word luridus, an adjective, meaning pale yellow, indicating the pale wing color.

Purpurata plagiatalis (Walker, 1859), comb. nov.

Figs 11, 19, 19a–c, 20

Botys plagiatalis Walker, 1859: 673. Type locality: Sri Lanka.

Pleuroptya plagiatalis: Inoue 1982: 1: 343, 2: 234.

Patania plagiatalis: Nuss et al. 2003–2024. Global Information System on Pyraloidea.

Material examined

Guangxi Zhuang Autonomous Region • 1 ♂, Nonggang National Nature Reserve, Longzhou County, alt. 188 m, 27 July 2011, Gui-Qing He leg., genitalia slide number: HGQ13237 • 1 ♂, Mulun National Nature Reserve, alt. 288 m, 22 July 2015, Dan Xu leg. • Hainan Prov., 1 ♀, Jianfengling Nature Reserve, alt. 770 m, 13 July 2014, Pei-Xin Cong, Lin-Jie Liu & Sha Hu leg. (NKU), genitalia slide number: XLJ14204.

Diagnosis

This species is distinguished by wings with postmedial line punctiform and excurved between M2 and CuA2 (Fig. 11); gnathos vestigial to a narrow band (Fig. 19a); sacculus narrowed medially, with a lamellar projection near distal end (Fig. 19); fibula ear-shaped, with long setae (Fig. 19b); phallus posteriorly with a rounded protruding sclerite, with a thick needle-like cornutus and a brush-like cornutus, and a short thorn-like cornutus near posterior end (Fig. 19c).

Redescription

Habitus (Fig. 11). Forewing length 8.0–9.5 mm, wingspan 18.0–22.0 mm. Frons and vertex yellowish brown. Labial palpus yellowish white, pale brown at distal end of 2nd segment. Maxillary palpus brown. Antenna yellowish brown, with ventral cilia ~ ½ in length of diameter of flagellomere in male. Patagium and tegula yellow, with pale yellowish brown patches centrally and basally respectively. Thorax yellow dorsally, white ventrally. Legs yellowish white, distal end of front tibia black. Wings yellow, with purple-brown lines and patches. Forewing with three small spots at base, another spot near basal dorsum; antemedial line slightly wavy, accompanied by a large elliptical patch inside; orbicular stigma a dark brown dot; discoidal stigma reniform, yellow centrally; postmedial line punctiform and excurved between M2 and CuA2; an irregular large patch near apex beyond anterior postmedial line, a nearly semicircular patch near tornus below CuA2; a line of small spots along terminal margin. Hindwing with discoidal stigma a short oblique stripe; postmedial line same as forewing before CuA2, not apparent afterwards; a large patch near apex beyond anterior postmedial line; a thin band below discoidal stigma, accompanied by an irregular misty wide band extended to tornus. Cilia of fore and hind wings yellowish white. Abdomen yellow dorsally, 1st and 2nd tergites with pale black spots laterally and 7th tergites black posteriorly in male.

Male genitalia (Fig. 19). Uncus short and broad. Gnathos vestigial to a narrow band and setose (Fig. 19a). Valva broad tongue-shaped with long setae along distal costa; fibula ear-shaped with long setae (Fig. 19b). Sacculus narrowed medially, with a lamellar projection near distal end. Saccus triangular. Juxta forcipate. Phallus posteriorly with a rounded protruding sclerite, with a thick needle-like cornutus and a brush-like cornutus composed of a spine cluster, and a short thorn-like cornutus near posterior end (Fig. 19c).

Female genitalia (Fig. 20). Apophyses anteriores ~ 2 × as long as apophyses posteriores. Antrum relatively long, with a long, sclerotized band. Ductus bursae ~ 4 × as long as corpus bursae. Corpus bursae nearly rounded, with a round signum.

Distribution

China (Xizang, Yunnan, Guangxi, Guangdong, Hainan, Fujian) (Lu and Guan 1953; Wang et al. 2003), Australia, Guatemala, Japan, India, Sri Lanka (Walker 1859; Inoue 1982; Klima 1939).

Host plants

Ipomoea spp. (Wang et al. 2003).

Purpurata shompen (Singh & Ahmad, 2022), comb. nov.

Patania shompen Singh & Ahmad in Singh et al. 2022: 14, figs 1–2, 5–7. Type locality: India (Great Nicobar Island).

Diagnosis

This species is similar to P. iopasalis, but can be distinguished by the postmedial line of forewing punctiform between M2 and CuA2, and the postmedial line of hindwing linear between M2 and CuA2 (Singh et al. 2022: fig. 1); phallus posteriorly with a rectangular protruding sclerite in male genitalia (Singh et al. 2022: fig. 7). In P. iopasalis, the postmedial line of the fore and hind wings are punctiform between M2 and CuA2 (Fig. 7); and phallus posteriorly with a nail head-like protruding sclerite in male genitalia (Fig. 14c).

Distribution

India (Great Nicobar Island) (Singh et al. 2022).

Remarks

This species is not found in China. The diagnosis is summarized based on the description and images of habitus and genitalia by Singh et al. (2022).

Discussion

According to the description and habitus images by Moore (1888), we found the external morphology of Syllepte leopardalis (Moore, 1888), distributed in India, very similar to the new genus Purpurata, especially in wing color and markings. But its antennae do not seem to be filiform according to the habitus illustration provided by Moore. The genitalia of S. leopardalis were not described or illustrated, so further study is needed to confirm its morphological characters and affiliation.

Acknowledgments

We are grateful to Li-Jun Xu for her preliminary work about the genus Patania, and the other graduate students and undergraduate in Laboratory of Lepidoptera Systematics (Southwest University, Chongqing, China) for their contributions to the specimen collection. We also give our thanks to Dr Jian-Yue Qiu and Dr Hao Xu (Mianyang Teachers’ College, Sichuan, China), to Dr Dan-Dan Zhang (Sun Yat-Sen University, China) for providing some specimens, to Prof. Hou-Hun Li (Nankai University, Tianjing, China) for lending us specimens, and to NHMUK for providing access to the corresponding author for examining some relevant specimens. We would also like to thank the language editor Dr Christopher Glasby for proofreading our manuscript. And special thanks are given to Dr Richard Mally (Czech University of Life Sciences, Czech Republic), Dr Bernard Landry (Muséum d’histoire naturelle, Switzerland), and another anonymous reviewer for reviewing our manuscript and their helpful suggestions. Cordial thanks are given to Dr Richard Mally again for professionally supplementing some important information to our revised manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study is sponsored by Natural Science Foundation of Chongqing, China (No. CSTB2022NSCQ-MSX1164), National Natural Science Foundation of China (No. 31772500), Shuangcheng cooperative agreement research grant of Yibin, China (No. XNDX2022020011) and Joint Fund of Henan Science and Technology R&D program (Advantageous Discipline De-velopment) (NO. 232301420119).

Author contributions

Conceptualization: XLX, XCD, XQL. Data curation: XLX, XCD. Funding acquisition: XCD. Investigation: XLX, XQL. Methodology: XLX, XCD, XQL. Project administration: XCD. Resources: XCD. Supervision: XCD. Writing - original draft: XQL, XLX. Writing - review and editing: XLX, XCD.

Author ORCIDs

Xin-Lei Xue https://orcid.org/0009-0000-5697-0872

Xiao-Qiang Lu https://orcid.org/0009-0009-3696-7778

Xi-Cui Du https://orcid.org/0000-0002-7796-7303

Data availability

All of the data that support the findings of this study are available in the main text.

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