Four New Species of the Genus Eoneureclipsis (Trichoptera: Psychomyiidae) from China Inferred from Morphology and DNA Barcodes †
1
Lab of Insect Taxonomy & Aquatic Insects, College of Plant Protection, Nanjing Agricultural University, Nanjing 210095, China
2
Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment of China, Nanjing 210042, China
*
Authors to whom correspondence should be addressed.
†
The LSID (Life Science Identifier) for this publication is: LSIDurn:lsid:zoobank.org:pub:0BB1597F-5405-404D-A990-A6E00B54E556.
Insects 2023, 14(2), 158; https://doi.org/10.3390/insects14020158
Submission received: 3 January 2023
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Revised: 25 January 2023
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Accepted: 2 February 2023
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Published: 4 February 2023
(This article belongs to the Special Issue A Themed Issue in Honor of Prof. Dr. Xin-Hua Wang on the Occasion of His 70th Birthday: Diversity, Ecology and Evolution of Aquatic Insects)
Abstract
:Simple Summary
The family Psychomyiidae Walker, 1852 currently includes 8 genera and more than 600 extant species, and is represented by three subfamilies, Psychomyiinae, Tinodinae, and Eoneureclipsinae. Subfamily Eoneureclipsinae was established by Mey in 2013, with the genus Eoneureclipsis Kimmins, 1955, based on morphological features, which were significant enough to separate it from the other genera of family Psychomyiidae. The new species in this study provide additional distribution data to the genus.
Abstract
Four new species of the genus Eoneureclipsis Kimmins, 1955 from China are described, illustrated, and diagnosed based on male genitalia: Eoneureclipsis jianfenglingensis sp. nov. from Hainan, E. foraminulatus sp. nov. from Guangxi, E. spinosus sp. nov. from Guangxi and Guangdong, and E. gei sp. nov. from Fujian. A dichotomous key to Chinese adult males of Eoneureclipsis is provided. A distribution map for all Eoneureclipsis species is also presented. The DNA barcodes (partial mtCOI sequences) of E. jianfenglingensis sp. nov., E. gei sp. nov., and E. hainanensis Mey, 2013 have been generated and compared with all existing sequences of Eoneureclipsis species.
1. Introduction
The genus Eoneureclipsis Kimmins, 1955 [1] is a small genus in the family Psychomyiidae Walker, 1852 [2,3], containing 19 [4,5,6] extant species with two species distributed in the Eastern Palearctic Region, Japan [7] and 17 in the Oriental Region, of which, four species are distributed in Thailand [5,8,9,10,11], three in Vietnam [12,13,14], three in China [6,15,16], two species each in India [17] and Japan [7], and one species each in Laos [18], Myanmar [17], and Malaysia [1]. Their distributions are shown in Figure 1.
Recognition of the Eoneureclipsis larvae occurred much later than that for adults. Larvae suspected to be of the genus Eoneureclipsis were first recorded in 2005 by Thamsenanupap et al. [19]; they found that the larvae of two species from northern Thailand had a larger body size than the larvae of other genera among the psychomyiids and treated them as Eoneureclipsis querquobad Malicky & Chantaramongkol, 1989 and Eoneureclipsis alekto Malicky & Chantaramongkol, 1997 according to their distributions. Torii and Nakamura [20] in 2016 associated a putative Eoneureclipsis larva with its adult by using mtCOI sequence data and described it in detail.
Up to now, three Eoneureclipsis species have been reported in China. The first one was recognized by Mey [15] from Hainan; in his research, he established a new subfamily Eoneureclipsinae to accommodate all Eoneureclipsis species. The second and third species were described as from Taiwan [16] and Guangdong [6] by Malicky (Table 1).
In this study, we describe four new species from Oriental China based on male adults, bringing the world’s Eoneureclipsis fauna to 23 (Table 1).
2. Materials and Methods
2.1. Sample Collection
A total of 8 adult specimens of the new species were collected in 2004, 2019, and 2022 with light traps (Table 2). After collection, they were preserved in 95% ethanol in the field. A photograph of a mobile light trap and photographs of the ecosystems (Figure 2) were taken with an Olympus TOUGH TG-5 digital camera.
2.2. Morphological Study
The methods used for the preparation of specimens followed Xu et al. [21]. Male abdomens used for illustrations were cleared with 10% NaOH solution and heated to 80 °C for 20 min to remove all the non-chitinous tissues. Then the cleaned genitalia were rinsed in distilled water and mounted on a depression slide with lactic acid for examination. Genitalic structures of males were traced with the pencil using a Nikon Eclipse 80i microscope equipped with a camera lucida. Pencil drawings were scanned with an Epson Perfection (V30 SE) scanner, then placed as templates in Adobe Photoshop (Version: CC 2018 19.0) and inked digitally with a Wacom CTL-671 tablet to produce the final illustrations. Then each abdomen was stored in a microvial together with the remainder of the specimen in 95% ethanol.
All specimens have been deposited in the Insect Collection, Nanjing Agricultural University, Nanjing, Jiangsu Province, China (NJAU).
2.3. Molecular Analysis
The left hind legs of three individuals (males) were taken from the body for DNA extractions. DNA extraction and PCR amplification followed the procedures of Zang [22]. The primers [23] (LCO1490/HCO2198) used to amplify the 658 bp fragment of the mitochondrial (mt) cytochrome c oxidase I unit (COI) are listed in Table 3.
Fragment sequencing and analysis followed the procedures of Ge [24]. Raw sequences were assembled and edited in Sequencher 4.5 (Gene Codes Corporation, Ann Arbor, MI, USA). A neighbor-joining (NJ) tree of 32 species within the family Psychomyiidae was constructed using MegaX v10.2.6 [25], with the following parameters: Kimura 2-parameter substitution model (K2P) [26], pairwise gap deletion, and others as defaults. The same software was used to calculate the K2P corrected p-distance of the 658 bp mtCOI fragment among all Eoneureclipsis species sequences available (Table S1). COI sequences of three Eoneureclipsis species were uploaded to GenBank. Accession numbers of the other analyzed Eoneureclipsis species are shown in Table S1.
3. Results
3.1. Taxonomy
Family Psychomyiidae Walker, 1852
Genus Eoneureclipsis Kimmins, 1955
3.1.1. Eoneureclipsis jianfenglingensis Peng, Zang & Sun, sp. nov. (Figure 3A–E and Figure 7A)
Description: Length of each forewing 6.9–7.0 mm (n = 2). Specimens (in alcohol) brown, with compound eyes black, antennae uniformly yellowish brown, palpi pale brown, legs and spurs yellowish brown, and wings dark brown. Both pairs of wings elliptical, each with rounded apex; each hind wing with costal margin, not sinuate and without projection as in other psychomyiid genera (Figure 7A). Venation typical for the genus. Forewings each with Forks I, II, III, IV, and V present, discoidal cell short, median and thyridial cells relatively elongate; irregular hyaline area across r-m, m, m-cu, and middle of M; two nygmas located at bases of R4 cell (Fork II) and thyridial cell (TC). Hind wings each with Forks I, II, III, and V present; one nygma present at base of R4 cell (Fork II).
Male genitalia: Tergite IX hairy, slender, nearly vertical, bent posterodorsad from segment’s mid-height in lateral view (Figure 3C); posterior margin produced into mesal triangular protrusion in dorsal view (Figure 3B). Sternite IX well developed, trapezoid in ventral view (Figure 3A); nearly rectangular with anterodorsal angle produced into projections in lateral view (Figure 3C). Segment X membranous, triangular, extending beyond apex of tergite IX in dorsal view (Figure 3B); irregular in lateral view (Figure 3C). Preanal appendages long, setose, straight, stick-shaped in dorsal view (Figure 3B); boomerang-shaped in lateral view (Figure 3C), each widest at middle, narrower at base and apex. Intermediate appendages originating laterally from anterolateral bases of sternite IX, inclining posteroventrad in lateral view (Figure 3C); straight, slender in dorsal view (Figure 3B), slightly shorter than preanal appendages, each with elongate process near base, straight in dorsal and lateral views (Figure 3B,C). Inferior appendages stout, setose, each with distal end of harpago extending beyond phallic apparatus in lateral view (Figure 3C); coxopodites closely appressed in ventral view (Figure 3A), each rectangular with anterodorsal angle produced internally into elongate apodeme (basal plate); and each with several stout spines at apicomesal corner (Figure 3C). Harpagones elongate, each with basal 1/3 fused with distal half of coxopodites in lateral view (Figure 3C); in ventral view (Figure 3A) with apex bent mesad, each with two spines apically. Phallic apparatus long, arched, with base slightly swollen, and distal 1/4 posteroventrad in lateral view (Figure 3C); basal half tubular, distal half swollen and suddenly narrowed, with apex having back hooks laterally in dorsal view (Figure 3B).
Diagnosis: The species is similar to Eoneureclipsis yaeyamaensis Torii & Nishimoto, 2011 from Japan in the intermediate appendages in which each has one short branch. However, the new species can be easily distinguished from the latter by the following: (1) the preanal appendages are elongate-triangular in lateral view, rather than clavate and curved at the base as in E. yaeyamaensis; (2) the intermediate appendages are branched near the base, rather than branched subapically as in E. yaeyamaensis; and (3) the coxopodites are rectangular in ventral view, rather than elliptical as in E. yaeyamaensis.
Holotype: Male, P.R. China, Hainan Province: Ledong Li Autonomous County, Jianfeng Town, Jianfengling National Forest Park, Jianfeng Parking Lot, 18.7107° N, 108.8758° E, alt. 706 m, 16 Apr 2019, light trap, leg. H. Song (NJAU).
Paratype: P.R. China, Hainan Province: one male, Ledong Li Autonomous County, Jianfeng Town, Jianfengling National Forest Park, bridge over a tributary, 0.5 km east of Jianfeng Parking Lot, 18.7087° N, 108.8844° E, alt. 920 m, 27 Jul 2022, light trap, leg. L. Peng & H. Zang (NJAU).
Distribution: China (Hainan).
Etymology: The name is derived from that of the locality type: Jianfengling mountain, Hainan Province.
3.1.2. Eoneureclipsis foraminulatus Peng, Zang & Sun, sp. nov. (Figure 4A–C and Figure 7B)
Description: Length of each forewing 8.1–8.2 mm (n = 1). Specimen in alcohol with compound eyes black; antennae, palpi, legs, spurs, and wings pale yellow; other parts of body yellow. Venation typical for genus, similar to that of E. jianfenglingensis sp. nov. Wings uniformly brown, without transparent patches (Figure 7B).
Male genitalia: Tergite IX nearly vertical, boomerang-shaped in lateral view (Figure 4C); posterior margin strongly expanded into blunt apicodorsal lobe, with several long ossifying spines on each side in dorsal view (Figure 4B). Sternite IX well developed, trapezoidal in ventral view (Figure 4A) with anterolateral angles produced; nearly rectangular, with anterodorsal angles markedly enlarged in lateral view (Figure 4C). Segment X membranous, extending beyond apex of tergite IX, with apex shallowly incised in dorsal view (Figure 4B); bordering tergite IX and extending beyond it in lateral view (Figure 4C). Preanal appendages lanceolate, setose in dorsal view (Figure 4B); each curved and clavate with narrow base in lateral view (Figure 4C). Intermediate appendages originating from anterolateral bases of sternite IX, about same length as preanal appendages, tilted downward in lateral view (Figure 4C); clamp-shaped (basally divergent and apically curved mesad) in dorsal view (Figure 4B). Inferior appendages sturdy, pilose in lateral view (Figure 4C). Coxopodites each subtriangular in ventral view (Figure 4A), with cluster of apicomesal teeth; subtriangular in lateral view (Figure 4C), each with anterodorsal angle produced into long apodeme. Harpagones elongate, each with apex rounded, bent slightly upward in lateral view (Figure 4C); inner surface with row of dense short spines and bent apicomesad in ventral view (Figure 4A). Phallic apparatus in lateral view long, arched, basal and apical portions wider than middle, apical portion divided into dorsal tubular aedeagus and ventral trapezoidal plate with round hole between them basally (Figure 4C); spindle-shaped in dorsal view, aedeagus slightly constricted subapically, ventral plate triangular (Figure 4B); and V-shaped in caudal view.
Diagnosis: This species is similar to Eoneureclipsis afonini Arefina-Armitage & Armitage, 2015 from Vietnam. However, the new species can be easily distinguished from the latter by the following: (1) the apex of the apicodorsal lobe of tergite IX is rounded rather than incised as in E. afonini; (2) the paired intermediate appendages are curved in the shape of a clamp in dorsal view but are straight in E. afonini; and (3) the phallic apparatus is spindle-shaped in dorsal view rather than tapering from base to apex in E. afonini.
Holotype: Male, P.R. China, Guangxi Province: Fangchenggang City, Shangsi County, Shiwandashan National Forest Park, a fourth tributary of Shitou He, 3.8 km southwest of main entrance to Park, 21.8914° N, 107.9047° E, alt. 420 m, 06 Jun 2004, light trap, leg. J.C. Morse & C. Sun (NJAU).
Distribution: China (Guangxi).
Etymology: The Latin adjective foraminulatus means foraminate, referring to the small circular aperture formed between the aedeagus and ventral plate at their bases in lateral view.
3.1.3. Eoneureclipsis spinosus Peng, Zang & Sun, sp. nov. (Figure 5A–F and Figure 7C)
Description: Length of each forewing 7.4–9.4 mm (n = 3). Specimens in alcohol with compound eyes black; antennae, palpi, and wings pale yellow; legs, spurs, and abdomen yellowish brown; thorax dark brown. Venation typical for genus, similar to that of E. jianfenglingensis sp. nov. Wing surface uniformly brown, without transparent patches (Figure 7C).
Male genitalia: Tergite IX slender, pilous, nearly vertical in lateral view (Figure 5C); posterior margin produced into blunt apicodorsal lobe in dorsal view (Figure 5B). Sternite IX well developed, nearly square in ventral view (Figure 5A); in lateral view (Figure 5C), subrectangular, with each anterodorsal angle produced into irregular process with which each intermediate appendage is articulated. Segment X membranous, extends beyond apex of tergite IX in dorsal view (Figure 5B); irregular in lateral view (Figure 5C). Preanal appendages clavate, long, setose, slightly curved near base in dorsal view (Figure 3B); each with base narrow and apex truncate and with strong spines apicodorally and apicoventrally in lateral view (Figure 5C). Intermediate appendages straight, clavate in dorsal view (Figure 5B); arched in lateral view (Figure 5C), each with distal half having strong spines of varied sizes and number (13–14 in holotype; 10–15 in paratypes) randomly arranged on dorsal and lateral surfaces (Figure 5B). Inferior appendage stout, hairy, extending beyond phallic apparatus in lateral view (Figure 5C). Coxopodites with basal inner halves almost touching in ventral view (Figure 5A), with several small teeth apicomesally; spoon-shaped in lateral view, with anterodorsal angles each produced into apodeme extending deeply into sternite IX (Figure 5C). Harpagones elongate, each with basal half fused with its coxopodite in lateral view and with middle portion slightly constricted (Figure 5C); bent apicomesally in ventral view (Figure 5A); each with inner margin having a row of spines (Figure 5A,C). Phallic apparatus well developed, slightly curved, gradually enlarged from base to apex, apex sinuate in lateral view (Figure 5C), in dorsal view slightly constricted at middle, apex with sloped opening exposing aedeagus (Figure 5B).
Diagnosis: The species is similar to Eoneureclipsis okinawaensis Torii & Nishimoto, 2011 from Japan in the shape of the phallic apparatus when viewed laterally but differs from the latter in the following: (1) the preanal appendages are straight, each with its basal half slender in lateral view (clavate and curved in the middle, almost of equal width in E. okinawaensis); (2) the distal half of each preanal appendage has strong spines (without spines in E. okinawaensis); and (3) the intermediate appendages are unbranched (branched subapically in E. okinawaensis).
Holotype: Male, P.R. China, Guangxi Province: Hechi City, Huanjiang County, Jiuwandashan Provincial Nature Preserve, an unnamed tributary of Yangmeiao Xi, 100 m upstream of County Road 5309 marker 124.9 km, 25.1976° N, 108.6494° E, alt. 1155 m, 15 June 2004, light trap, leg. J.C. Morse & C.J. Geraci (NJAU).
Paratypes: P.R. China, Guangxi Province: one male, same data as holotype. P.R. China, Guangdong Province: one male, Shaoguan City, Ruyuan County, Nanling National Nature Preserve, an unnamed tributary of Laopengkeng, Route X327, marker 17.45 km, 24.9128° N, 113.0342° E, alt. 935 m, 21–22 May 2004, light trap, leg. J.C. Morse & C. Sun (NJAU).
Distribution: China (Guangxi, Guangdong).
Etymology: The Latin adjective spinosus means spiny, referring to the intermediate appendages with many spines.
3.1.4. Eoneureclipsis gei Peng, Zang & Sun, sp. nov. (Figure 6A–D and Figure 7D)
Description: Length of each forewing 7.4–7.5 mm (n = 1). Specimen in alcohol with compound eyes black; antennae and palpi pale yellow; legs, spurs, and abdomen yellowish brown; thorax and wings dark brown. Venation typical for genus, similar to that of E. jianfenglingensis sp. nov. Forewings with irregular hyaline area across r-m, m, m-cu, and middle of M and small transparent spots irregularly scattered on anterior portions of fore- and hind wings (Figure 7D).
Male genitalia: Tergite IX slender, setose, nearly vertical in lateral view (Figure 6C); somewhat T-shaped in dorsal view, with produced apicodorsal lobe fused with segment X (Figure 6B). Sternite IX well developed, trapezoid in ventral view (Figure 6A); subrectangular, with each anterodorsal angle produced into irregular projection in lateral view (Figure 6C). Segment X membranous, in dorsal view, with apex slightly incised mesally and setose laterally (Figure 6B); two sides extending downward and enclosing phallic apparatus in lateral view (Figure 6C). Preanal appendages clavate, setose in dorsal view (Figure 6B); elongate-rectangular in lateral view, each with apex truncate (Figure 6C). Intermediate appendages originating from produced projections of sternite IX, in lateral view longer than preanal appendages, each with basal 2/3 narrow and apical 1/3 enlarged, distal end sharp in lateral view (Figure 6C); in dorsal view (Figure 6B) slightly bent mesad, with apices crossed. Inferior appendages stout, hairy in lateral view (Figure 6C). Coxopodites approximately semicircular, their inner margins almost touching along full length in ventral view, each with single spine apicomesally (Figure 6A); subtriangular, long, and upcurved apically in lateral view (Figure 6C). Harpagones elongate-clavate, basal two-fifths fused with coxopodites in lateral view (Figure 6C); somewhat parentheses-shaped in ventral view, with several small spines scattered randomly on inner surfaces in both ventral and lateral views (Figure 6A,C). Phallic apparatus long, slightly curved in lateral view (Figure 6C), lower margin with hook-like incision near 1/3 from apex; in dorsal view (Figure 6D) middle portion swollen; ventral plate protruding upward to acute apex-like scimitar in lateral view (Figure 6C); aedeagus in dorsal view straight and slender, issuing dorsally at 1/3 from apex of phallic apparatus, with its apex slightly swollen, parallel to apicoventral plate in lateral view (Figure 6C); narrower in dorsal view (Figure 6D).
Diagnosis: The species is similar to Eoneureclipsis varsikiyja Schmid, 1972 from India in the shape of the intermediate appendages in lateral view but differs from the latter in that (1) the preanal appendages are elongate-rectangular in lateral view (slightly enlarged basally in E. varsikiyja); (2) the dorsal margin of the phallic apparatus is slightly arched in lateral view (strongly sinuate in E. varsikiyja); and (3) the ventral margin of the phallic apparatus has a semicircular incision in lateral view (nearly straight in E. varsikiyja).
Holotype: Male, P.R. China, Fujian Province: Wuyishan City, Xingcun Town, Wuyishan National Park, the upper tributary of Dazhulan 27.6985° N, 117.6521° E, alt. 884 m, 16 July 2022, light trap, leg. L. Peng, H. Zang, X. Ge & C. Sun (NJAU).
Distribution: China (Fujian).
Etymology: The new species is named after Mr. Xinyu Ge in honor of his unprecedented contributions to the study of mitochondrial genomics and phylogenetic research in Trichoptera.
3.2. Key to Male Chinese Eoneureclipsis Species
With four new species in this study, a total of seven Eoneureclipsis species occur in China. A concise key to these seven species is presented below.
- 1
- -
- Preanal appendages acute or narrow in lateral view ……………………………………4
- 2
- Intermediate appendages with stout spines (Figure 5B,C,E) ……E. spinosus sp. nov.
- -
- 3
- Intermediate appendages longer than phallic apparatus, slender, each curved upward subapically in lateral view [16] ……………………………………………E. sarach
- -
- Intermediate appendages of the same length as phallic apparatus, each broad subapically then abruptly tapered to acute apex in lateral view (Figure 6C) …E. gei sp. nov.
- 4
- -
- Phallic apparatus with ventral plate; intermediate appendages simple ………………5
- 5
- Phallic apparatus with dorsal process basally in lateral view [6].…………E. malchidael
- -
- 6
- Tergite IX posterior margin strongly expanded, with several long ossifying spines on each side in dorsal view (Figure 4B) …………………………….E. foraminulatus sp. nov.
- -
- Tergite IX tapering to end, without spines in dorsal view [15] …………E. hainanensis
3.3. Molecular Analysis
DNA sequences were aligned for 56 extracts of 32 psychomyiid species with a full barcode length of 658 base pairs. For the genus Eoneureclipsis, a single clade was well supported in the NJ tree (Figure 8). The interspecific distances (K2P p value) between specimens of genus Eoneureclipsis and species of other genera were more than 17.60% (Table S1). Based on existing DNA barcodes, the minimum interspecific divergence was 12.60% between E. okinawaensis and E. montanus, and the maximum interspecific divergence was 20.60% between E. montanus and E. hainanensis. The mean interspecific divergence was 17.10% in the genus Eoneureclipsis. Among them, the interspecific divergence of the Eoneureclipsis species distributed in Japan ranged from 14.30% to 16.70%, with a mean divergence of 15.91%; Eoneureclipsis species distributed in China ranged from 12.60% to 15.90%, with a mean divergence of 14.33%. The mean intraspecific divergence of the Eoneureclipsis species was 5.47% with a minimum of 0.5% and a maximum of 8.20% in E. montanus; it seems that E. montanus perhaps has diverged into two geographic populations.
4. Discussion
The genus Eoneureclipsis was established by Kimmins within Polycentropodidae to accommodate new species Eoneureclipsis limax Kimmins, 1955 from Borneo [1]. Schmid [17] added three Indian species to Eoneureclipsis and transferred the genus to Psychomyiidae based on primitive wing venation, the primitive inferior appendages of the male genitalia, and the female ovipositor and regarded the group as a primitive lineage of the family Psychomyiidae [15]. In 1997, Li and Morse [27] performed a phylogenetic analysis on the relationships of six genera of Psychomyiidae based on adult and larval morphology; they suggested that the family Psychomyiidae consists of two subfamilies, i.e., Psychomyiinae and Tinodinae, and treated the subfamily Paduniellinae as a synonym of Psychomyiinae. However, they failed to include the genus Eoneureclipsis in their analysis, and the taxonomic position of the genus Eoneureclipsis has remained unsolved [28]. In 2013, based on the large body size and other features, Mey [15] established a new monotypic subfamily Eoneureclipsinae to include the genus Eoneureclipsis and its members. Thamsenanupap et al. [19] and Torii and Nakamura [20] proved that the larvae also have larger body sizes than those of other psychomyiid genera, clearly different from Psychomyiinae and Tinodinae [29]. Silva’s analysis on the relationship of Xiphocentronidae and Psychomyiidae indicated that the subfamily Eoneureclipsinae was the first clade of the two families with Lype placed as a clade right after Eoneureclipsis [30]. Our neighbor-joining cladogram based on the 658 bp long mtCOI sequence of six Eoneureclipsis species is a little bit different from the result of Silva (Figure 8) in that Eoneureclipis is a sister genus to Lype, and these two genera are the sister group to the remaining Psychomyiidae genera rather than the sister genus to all other Psychomyiidae and Xiphocentronidae genera as Silva suggested. However, discussing the relationship among psychomyiid genera based only on mtCOI data is not reliable enough. In this study, the additional data of three species support the monophyly of each species. Unfortunately, we failed to obtain molecular data from E. foraminulatus sp. nov. and E. spinosus sp. nov.; therefore, we are unable to conduct a complete phylogenetic analysis on the relationships of the psychomyiid genera. More samples are needed to complete the analysis.
Supplementary Materials
The following Supplementary Materials can be downloaded at: https://www.mdpi.com/article/10.3390/insects14020158/s1. Table S1: Kimura 2-parameter pairwise genetic distances based on mtCOI barcodes of Psychomyiidae.
Author Contributions
Conceptualization, all authors; methodology, all authors; software, L.P. and C.S.; validation, L.P. and H.Z.; formal analysis, L.P. and H.Z.; investigation, L.P., H.Z., C.S., and L.W.; resources, all authors; data curation, L.P.; writing—original draft preparation, L.P.; writing—review and editing, C.S. and B.W.; visualization, L.P.; supervision, C.S. and B.W.; project administration, B.W.; funding acquisition, C.S., L.W., and B.W. All authors have read and agreed to the published version of the manuscript.
Funding
This research was supported by the National Natural Science Foundation of China (NSFC, No. 41771052; No. 32200364) and The Project of Biological Resources Survey in Wuyishan National Park.
Data Availability Statement
The voucher specimens from this research were deposited in the Insect Classification and Aquatic Insect Laboratory, College of Plant Protection, Nanjing Agricultural University, Nanjing, China.
Acknowledgments
We are very thankful to Haitian Song from the Fujian Academy of Forestry for donating caddis specimens selflessly. We sincerely thank Hongying Sun of Nanjing Normal University for providing us with a precious opportunity to visit Hainan for collection. Thanks also to John Morse from Clemson University, USA, and Christy Jo Geraci for contributions to the Trichoptera fauna of China.
Conflicts of Interest
The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.
References
- Kimmins, D.E. Results of the Oxford University expedition to Sarawak, 1932. Order Trichoptera. Sarawak Mus. J. 1955, 6, 374–442. [Google Scholar]
- Walker, F. Cataloque of the Specimens of Neuropterous Insects in the Collection of the British Museum, Part I: Phryganides-Perlides; British Museum: London, UK, 1852; pp. 1–192. [Google Scholar]
- Holzenthal, R.W.; Blahnik, R.J.; Prather, A.L.; Kjer, K.M. Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. Zootaxa 2007, 1668, 639–698. [Google Scholar] [CrossRef]
- Morse, J.C. Trichoptera World Checklist. Available online: http://entweb.clemson.edu/database/trichopt/index.htm (accessed on 4 November 2022).
- Suwannarat, N.; Malicky, H.; Laudee, P. Four new species of caddisflies (Trichoptera: Polycentropodidae, Psychomyiidae, Hydropsychidae, Odontoceridae) from Khao Nan and Tai Rom Yen National Parks, southern Thailand. Zootaxa 2020, 4801, 577–583. [Google Scholar] [CrossRef] [PubMed]
- Malicky, H. Beiträge zur Kenntnis chinesischer Köcherfliegen (Insecta, Trichoptera). Linz. Biol. Beiträge 2020, 52, 417–455. [Google Scholar]
- Torii, T.; Nishimoto, H. Discovery of the genus Eoneureclipsis Kimmins (Trichoptera: Psychomyiidae) from Japan. Zoosymposia 2011, 5, 453–464. [Google Scholar] [CrossRef]
- Malicky, H. Beschreibung von neuen Kocherfliegen (Trichoptera) aus Thailand und Burma. Entomol. Ber. Luzern 1989, 22, 117–126. [Google Scholar]
- Malicky, H.; Chantaramongkol, P. Weitere neue Köcherfliegen (Trichoptera) aus Thailand. Arbeit Nr. 20 über thailändische Köcherfliegen. Linz. Biol. Beiträge 1997, 29, 203–215. [Google Scholar]
- Malicky, H.; Chantaramongkol, P.; Bunlue, P.; Changthong, N.; Nawvong, J.; Nuntakwang, A.; Prommi, T.; Thamsenanupap, P.; Thapanya, D. 27 neue Köcherfliegen aus Thailand (Insecta, Trichoptera) (36. Arbeit über thailändische Köcherfliegen). Linz. Biol. Beiträge 2004, 36, 287–304. [Google Scholar]
- Thapanya, D.; Chantaramongkol, P.; Malicky, H. An updated survey of caddisflies (Trichoptera, Insecta) from Doi Suthep-Pui and Doi Inthanon National Parks, Chiang Mai Province, Thailand. Trop. Nat. Hist. 2004, 1, 21–40. [Google Scholar]
- Malicky, H. Neue Kocherfliegen (Trichoptera, Insecta) aus Vietnam. Linz. Biol. Beiträge 1995, 27, 851–885. [Google Scholar]
- Arefina-Armitage, T.I.; Armitage, B.J. New species and new country records for Vietnamese caddisflies (Insecta: Trichoptera). Insecta Mundi 2015, 0438, 1–19. [Google Scholar]
- Armitage, B.J.; Mey, W.; Arefina, T.I.; Schefter, P.W. The caddisfly fauna (Insecta: Trichoptera) of Vietnam. In Proceedings of the 11th International Symposium on Trichoptera, Shiga, Japan, 12–19 June 2003; Tanida, K., Rossiter, A., Eds.; Tokai University Press: Kanagawa, Japan, 2005; pp. 25–37. [Google Scholar]
- Mey, W. Eoneureclipsis hainanensis spec. nov. from China and its systematic position (Insecta, Trichoptera, Psychomyiidae) (pl. 14). Esperiana Band 2013, 18, 255–258. [Google Scholar]
- Malicky, H. Köcherfliegen (Trichoptera) von Taiwan, mit Neubeschreibungen. Linz. Biol. Beiträge 2014, 46, 1607–1646. [Google Scholar]
- Schmid, F. Sur quelques nouvelles psychomyiines tropicales (Trichoptera: Psychomyiidae). Le Nat. Can. 1972, 99, 143–172. [Google Scholar]
- Malicky, H. Beiträge zur Kenntnis asiatischer Trichopteren. Braueria 2009, 36, 11–58. [Google Scholar]
- Thamsenanupap, P.; Chantaramongkol, P.; Malicky, H. Description of caddis larvae (Trichoptera) from northern Thailand of the genera Himalopsyche (Rhyacophilidae), Arctopsyche (Arctopsychidae), cf. Eoneureclipsis (Psychomyiidae) and Inthanopsyche (Odontoceridae). Braueria 2005, 32, 7–11. [Google Scholar]
- Torii, T.; Nakamura, M. DNA identification and morphological description of the larva of Eoneureclipsis montanus (Trichoptera, Psychomyiidae). Zoosymposia 2016, 10, 424–431. [Google Scholar] [CrossRef]
- Xu, J.H.; Wang, B.X.; Sun, C.H. The Stenopsyche simplex species group from China with descriptions of three new species (Trichoptera: Stenopsychidae). Zootaxa 2015, 3785, 217–230. [Google Scholar] [CrossRef]
- Zang, H.; Ge, X.; Peng, L.; Sun, C.; Wang, B. A New Species of Limnephilus (Insecta: Trichoptera: Limnephilidae) from China, with Revision of the Genus Limnephilus on the Chinese Mainland. Insects 2022, 13, 653. [Google Scholar] [CrossRef]
- Folmer, O.; Black, M.; Hoeh, W.; Lutz, R.; Vrijenhoek, R. DNA primers for amplification of mitochondrial cytochrome coxidase subunit I from diverse metazoan invertebrates. Mol. Mar. Biol. Biotechnol. 1994, 3, 294–299. [Google Scholar]
- Ge, X.Y.; Peng, L.; Du, J.; Sun, C.H.; Wang, B.X. New species of the genus Molanna Curtis, 1834 (Trichoptera, Molannidae) in China inferred from morphology and DNA barcodes. ZooKeys 2022, 1112, 161–178. [Google Scholar] [CrossRef]
- Kumar, S.; Stecher, G.; Li, M.; Knyaz, C.; Tamura, K. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Mol. Biol. Evol. 2018, 35, 1547–1549. [Google Scholar] [CrossRef] [PubMed]
- Kimura, M. A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. J. Mol. Evol. 1980, 16, 111–120. [Google Scholar] [CrossRef] [PubMed]
- Li, Y.L.; Morse, J.C. Phylogeny and classification of Psychomyiidae (Trichoptera) genera. In Proceedings of the 8th International Symposium on Trichoptera, Minneapolis and Lake Itasca, MI, USA, 9–15 July 1995; Holzenthal, R.W., Flint, O.S., Jr., Eds.; Ohio Biological Survey: Columbus, OH, USA, 1997; pp. 271–276. [Google Scholar]
- Sharma, R.M.; Chandra, K. Checklist of Indian caddisflies (Insecta: Trichoptera). Zool. Surv. India 2009, 1–33. Available online: https://www.researchgate.net/publication/237460449_CHECKLIST_OF_INDIAN_CADDISFLIES_INSECTA_TRICHOPTERA (accessed on 15 January 2023).
- Tachet, H.; Coppa, G.; Forcellini, M. A comparative description of the larvae of Psychomyia pusilla (Fabricius 1781), Metalype fragilis (Pictet 1834), and Paduniella vandeli Décamps 1965 (Trichoptera: Psychomyiidae) and comments on the larvae of other species belonging to these three genera. Zootaxa 2018, 4402, 91–112. [Google Scholar] [CrossRef]
- Silva, A.V.S.D. Phylogeny of Xiphocentronidae Ross, 1949 and Taxonomy of Neotropical Xiphocentroninae (Trichoptera: Annulipalpia: Psychomyioidea). Ph.D. Thesis, Universidade de São Paulo, São Paulo, Brazil, 2020. [Google Scholar]
Figure 1.
Distribution map of Eoneureclipsis species.
Figure 2.
Photograph of a mobile light trap (A) and photographs of ecosystems (B–D): (A) light trap with mobile power and a high-pressure mercury lamp, in Hainan; (B) a tributary in Jianfengling National Forest Park, Hainan Province; (C,D) the upper tributary of Dazhulan, Wuyishan National Park, Fujian (photographs by Haoming Zang and Lang Peng).
Figure 2.
Photograph of a mobile light trap (A) and photographs of ecosystems (B–D): (A) light trap with mobile power and a high-pressure mercury lamp, in Hainan; (B) a tributary in Jianfengling National Forest Park, Hainan Province; (C,D) the upper tributary of Dazhulan, Wuyishan National Park, Fujian (photographs by Haoming Zang and Lang Peng).
Figure 3.
Male genitalia of Eoneureclipsis jianfenglingensis sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) intermediate appendages of paratype, dorsal; (E) one intermediate appendage of paratype, left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus. Scale bar refers to A–E.
Figure 3.
Male genitalia of Eoneureclipsis jianfenglingensis sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) intermediate appendages of paratype, dorsal; (E) one intermediate appendage of paratype, left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus. Scale bar refers to A–E.
Figure 4.
Male genitalia of Eoneureclipsis foraminulatus sp. nov.: (A) ventral; (B) dorsal; (C) left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus; ven. pla. = ventral plate. Scale bar refers to A–C.
Figure 4.
Male genitalia of Eoneureclipsis foraminulatus sp. nov.: (A) ventral; (B) dorsal; (C) left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus; ven. pla. = ventral plate. Scale bar refers to A–C.
Figure 5.
Male genitalia of Eoneureclipsis spinosus sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) phallic apparatus, left lateral; (E) intermediate appendages of paratype (Guangxi), left lateral; (F) intermediate appendages of paratype (Guangdong), left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus. Scale bar refers to A–F.
Figure 5.
Male genitalia of Eoneureclipsis spinosus sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) phallic apparatus, left lateral; (E) intermediate appendages of paratype (Guangxi), left lateral; (F) intermediate appendages of paratype (Guangdong), left lateral. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus. Scale bar refers to A–F.
Figure 6.
Male genitalia of Eoneureclipsis gei sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) phallic apparatus, dorsa. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus; ven. pla. = ventral plate. Scale bar refers to A–D.
Figure 6.
Male genitalia of Eoneureclipsis gei sp. nov.: (A) ventral; (B) dorsal; (C) left lateral; (D) phallic apparatus, dorsa. Abbreviations: ste. IX = sternum IX; ter. IX = tergum IX; seg. X = segment X; pre. app. = preanal appendages (paired); int. app. = intermediate appendages (paired); cox. = coxopodite (paired); har. = harpago (paired); pha. = phallic apparatus; aed. = aedeagus; ven. pla. = ventral plate. Scale bar refers to A–D.
Figure 7.
Wing venations of Eoneureclipsis species: (A) Eoneureclipsis jianfenglingensis sp. nov.; (B) E. foraminulatus sp. nov.; (C) E. spinosus sp. nov.; (D) E. gei sp. nov. Abbreviations: Sc = subcosta; R = radius; M = media; Cu = cunitus; A = anal; DC = discoidal cell; MC = medial cell; TC = thyridial cell. Scale bar refers to A–D.
Figure 7.
Wing venations of Eoneureclipsis species: (A) Eoneureclipsis jianfenglingensis sp. nov.; (B) E. foraminulatus sp. nov.; (C) E. spinosus sp. nov.; (D) E. gei sp. nov. Abbreviations: Sc = subcosta; R = radius; M = media; Cu = cunitus; A = anal; DC = discoidal cell; MC = medial cell; TC = thyridial cell. Scale bar refers to A–D.
Figure 8.
Neighbor-joining cladogram based on the 658 bp long mtCOI sequence of six Eoneureclipsis species. Numbers on branches represent the bootstrap values based on 1000 replicates; scale bar indicates Kimura’s 2-parameter genetic distance of 0.02.
Figure 8.
Neighbor-joining cladogram based on the 658 bp long mtCOI sequence of six Eoneureclipsis species. Numbers on branches represent the bootstrap values based on 1000 replicates; scale bar indicates Kimura’s 2-parameter genetic distance of 0.02.
Table 1.
World list of Eoneureclipsis species.
| Number | Species | Distribution |
|---|---|---|
| 1 | E. jianfenglingensis sp. nov. | China: Hainan |
| 2 | E. foraminulatus sp. nov. | China: Guangxi |
| 3 | E. spinosus sp. nov. | China: Guangxi, Guangdong |
| 4 | E. gei sp. nov. | China: Fujian |
| 5 | E. malchidael Malicky, 2020 | China: Guangdong |
| 6 | E. hainanensis Mey, 2013 | China: Hainan |
| 7 | E. sarach Malicky, 2014 | China: Taiwan |
| 8 | E. montanus Torii & Nishimoto, 2011 | Japan: Honshu |
| 9 | E. okinawaensis Torii & Nishimoto, 2011 | Japan: Okinawa |
| 10 | E. shikokuensis Torii & Nishimoto, 2011 | Japan: Shioku |
| 11 | E. yaeyamaensis Torii & Nishimoto, 2011 | Japan: Yaeyama Islands |
| 12 | E. alekto Malicky & Chantaramongkol, 1997 | Thailand |
| 13 | E. chinachotiae Malicky &Laudee, 2020 | Thailand |
| 14 | E. nykteus Malicky & Nawvong, 2004 | Thailand |
| 15 | E. querquobad Malicky & Chantaramongkol, 1989 | Thailand |
| 16 | E. afonini Arefina-Armitage & Armitage, 2015 | Vietnam |
| 17 | E. quangi Malicky, 1995 | Vietnam |
| 18 | E. tieni Malicky, 1995 | Vietnam |
| 19 | E. akrichalakchmi Schmid, 1972 | India: Manipur |
| 20 | E. varsikiyja Schmid, 1972 | India: Assam |
| 21 | E. sebulon Malicky, 2009 | Laos |
| 22 | E. pravrisija Schmid, 1972 | Myanmar |
| 23 | E. limax Kimmins, 1955 | Malaysia: Sarawak |
Table 2.
Information on the collection of 8 specimens.
| Species | Number of Specimens | Collection Site | Collection Date | Molecular Sample |
|---|---|---|---|---|
| E. jianfenglingensis sp. nov. | 1 male | Jianfeng Town, Ledong County, Hainan | 16 Apr 2019 | yes |
| E. jianfenglingensis sp. nov. | 1 male | Jianfeng Town, Ledong County, Hainan | 27 Jul 2022 | no |
| E. foraminulatus sp. nov. | 1 male | Shangsi County, Fangchenggang City, Guangxi | 06 Jun 2004 | no |
| E. spinosus sp. nov. | 2 males | Huanjiang County, Hechi City, Guangxi | 15 Jun 2004 | no |
| E. spinosus sp. nov. | 1 male | Ruyuan County, Shaoguan City, Guangdong | 21–22 May 2004 | no |
| E. gei sp. nov. | 1 male | Xingcun Town, Wuyishan City, Fujian | 16 Jul 2022 | yes |
| E. hainanensis Mey, 2013 | 1 male | Jianfeng Town, Ledong County, Hainan | 28 Jul 2022 | yes |
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Peng, L.; Zang, H.; Sun, C.; Wang, L.; Wang, B. Four New Species of the Genus Eoneureclipsis (Trichoptera: Psychomyiidae) from China Inferred from Morphology and DNA Barcodes. Insects 2023, 14, 158. https://doi.org/10.3390/insects14020158
AMA Style
Peng L, Zang H, Sun C, Wang L, Wang B. Four New Species of the Genus Eoneureclipsis (Trichoptera: Psychomyiidae) from China Inferred from Morphology and DNA Barcodes. Insects. 2023; 14(2):158. https://doi.org/10.3390/insects14020158
Chicago/Turabian StylePeng, Lang, Haoming Zang, Changhai Sun, Le Wang, and Beixin Wang. 2023. "Four New Species of the Genus Eoneureclipsis (Trichoptera: Psychomyiidae) from China Inferred from Morphology and DNA Barcodes" Insects 14, no. 2: 158. https://doi.org/10.3390/insects14020158
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