Research Article |
Corresponding author: Feiyang Liang ( feiyang_sco@foxmail.com ) Corresponding author: Xingyue Liu ( xingyue_liu@yahoo.com ) Academic editor: André Nel
© 2024 Feiyang Liang, Xingyue Liu.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Psocomorpha, as one of the suborders of the order Psocodea, represents a large group of free-living bark lice. The phylogenetic relationships among/within the major groups, such as families, subfamilies, tribes, etc., of the bark lice need extensive investigation. The family Stenopsocidae belongs to the infraorder Caeciliusetae, which is a large group of Psocomorpha and is one of the common groups in Asia. Here we infer the intergeneric relationships of Stenopsocidae based on the mitochondrial genes and the nuclear 18S rRNA. The result supports the monophyly of Stenopsocidae and suggests the paraphyly of the genus Stenopsocus. Combining the morphological evidence, we propose a new genus, namely Neostenopsocus gen. n., which includes a number of species previously placed in Stenopsocus. The checklist of this new genus is provided. This new genus can be distinguished from Stenopsocus by the glabrous forewing CuP. In the revised classification system, Stenopsocidae includes four genera: Graphopsocus, Malostenopsocus, Neostenopsocus gen. n., and Stenopsocus. A key to the genera of Stenopsocidae is provided. We also present the diagnostic characters for each of these genera.
mitochondrial genome, taxonomy, new genus, Neostenopsocus gen. n.
The paraneopteran insect order Psocodea is an inconspicuous group, containing more than 10000 valid species, and is divided into three suborders: Trogiomorpha, Troctomorpha, and Psocomorpha (Lienhard & Smithers, 2002;
The bark louse family Stenopsocidae is a large group of the suborder Psocomorpha, currently comprising over 190 valid species in the world (
The genus Stenopsocus Hagen, 1866 was proposed by
In the present study, we investigate the phylogenetic relationships within the family Stenopsocidae using DNA sequences from the mitochondrial genes and nuclear 18s rRNA gene. Combining the molecular phylogeny and morphological characters, we propose a new genus Neostenopsocus gen. n., and present a revised generic classification of Stenopsocidae.
The genomic DNA was extracted from the thoracic muscle tissues of a single specimen using TIANamp Micro DNA Kit (TIANGEN, Beijing, China). For the sequencing library for each specimen and used the Illumina NovaSeq platform to obtain 4 GB sequence data. Raw reads were trimmed of adapters using BWA (Burrows-Wheeler Aligner) software (Li & Durbin, 2010). All sequences were assembled mapping to the mitogenome as the reference sequence using Geneious Prime 2023 (
Taxa included in this study with their number of Genbank accession number.
Family | Species | Mitogenome | 18S | Reference | |
Stenopsocidae | Graphopsocus cruciatus | OR608384 | AY630490 | This study | |
Malostenopsocus sp._HN | OR608385 | OR544596 | This study | ||
Malostenopsocus yunnanicus | OR608386 | OR544597 | This study | ||
Neostenopsocus brevicapitus | OR608387 | OR544598 | This study | ||
Neostenopsocus capacimacularus | OR608388 | OR544599 | This study | ||
Neostenopsocus dactylinus | OR608389 | OR544600 | This study | ||
Neostenopsocus externus | OR608390 | OR544601 | This study | ||
Neostenopsocus genostictus | OR608391 | OR544602 | This study | ||
Neostenopsocus longitudinalis | OR608392 | OR544603 | This study | ||
Neostenopsocus maximalis | OR608393 | OR544604 | This study | ||
Neostenopsocus metastictus | OR608394 | OR544605 | This study | ||
Neostenopsocus nepalensis | OR608395 | OR544606 | This study | ||
Neostenopsocus obscurus | OR608396 | OR544607 | This study | ||
Neostenopsocus periostictus | OR608397 | OR544608 | This study | ||
Neostenopsocus pygmaeus | OR608398 | OR544609 | This study | ||
Neostenopsocus tripartibilis | OR608403 | OR544610 | This study | ||
Neostenopsocus wangi | OR608399 | OR544611 | This study | ||
Stenopsocus aphidiformis | OR608400 | OR544612 | This study | ||
Stenopsocus lachali | OR608401 | OR544613 | This study | ||
Stenopsocus niger | OR608402 | OR544614 | This study | ||
Stenopsocus immaculatus | KX187004 | — |
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Dasydemellidae | Teliapsocus conterminus | MZ274210 | AB856951 |
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Amphipsocidae | Polypsocus corruptus | MZ274208 | AY630488 | ||
Paracaeciliidae | Paracaecilius japanus | MZ274207 | AY630501 | ||
Xanthocaecilius sommermanae | MZ274212 | AY630500 | |||
Caeciliusidae | Dypsocus coleoptratus | MZ274186 | AB856955 | ||
Fuelleborniella sp. | MZ274189 | AY630496 | |||
Asiopsocidae | Asiopsocus sonorensis | MZ274183 | AY630481 | ||
Lachesillidae | Graphocaecilius interpretatus | MZ274193 | — | ||
Ectopsocidae | Ectopsocopsis cryptomeriae | MZ274187 | AY630511 | ||
Peripsocidae | Kaestneriella sp. | MZ274196 | AY630506 | ||
Archipsocidae | Archipsocus nomas | MG255135 | — |
In this study, we used mitochondrial genes (exclude the control region, intergenic spacers and tRNA genes, including the protein-coding genes and rRNA genes) and partial nuclear18S rRNA. The alignment of 13 protein-coding genes was based on amino acid sequences translated with the invertebrate mitochondrial genetic codon table using MUSCLE (Edgar, 2004) complied in MEGA 7.0 (
Genitalic preparations were made by clearing the apex of the abdomen in a cold, saturated NaOH solution for 6 h. After rinsing the NaOH with water, the apex of the abdomen was transferred to glycerin for further dissection and examination. Observations were made using an Olympus CX-33 (Olympus Imaging Corporation, Tokyo) light microscope. Photographs and drawings were taken using a Sony Alpha 7II (Sony Corporation, Tokyo) digital camera attached to the Olympus CX-33. The figures were prepared with Adobe Photoshop 24.0.0 (Adobe, San Jose, CA, USA). The terminology follows
A total of 20 nearly complete mitochondrial genomes of Stenopsocidae were assembled. The mt genome of Stenopsocidae was determined to be a double-strand circular DNA molecule, including 37 typical genes: 13 PCGs, 22 tRNAs, and two rRNAs. In the 13 PCGs, ND2, COX1, COX2, ATP8, ATP6, COX3, ND3, ND6 and CytB are encoded on the major strand (J-strand), and ND5, ND4, ND4L and ND1 are encoded on the minor strand (Fig.
The mitogenome of Stenopsocidae shows highly biased AT composition (75.63%) (Table
Dataset | Graphopsocus (1 species) | Malostenopsocus (2 species) | Neostenopsocus (14 species) | Stenopsocus (4 species) | Stenopsocidae (21 species) | |
PCGs | A+T% | 72.74 | 75.19 | 72.55 | 75.77 | 73.43 |
AT-skew | –0.13 | –0.15 | –0.16 | –0.15 | –0.15 | |
GC-skew | 0.017 | 0.027 | 0.021 | 0.055 | 0.026 | |
Size | 11070 | 11106 | 11102 | 11108 | 11102 | |
tRNAs | A+T% | 77.75 | 79.57 | 78.26 | 79.67 | 78.63 |
AT-skew | 0.017 | 0.0036 | 0.0051 | –0.0027 | 0.0041 | |
GC-skew | 0.13 | 0.15 | 0.14 | 0.17 | 0.15 | |
Size | 1411 | 1410 | 1413 | 1414 | 1413 | |
rRNAs | A+T% | 80.55 | 81.44 | 80.39 | 82.11 | 80.83 |
AT-skew | –0.033 | –0.044 | –0.061 | –0.047 | –0.056 | |
GC-skew | 0.25 | 0.22 | 0.23 | 0.23 | 0.23 | |
Size | 1959 | 1937 | 1970 | 1961 | 1965 | |
PCGs+RNAs | A+T% | 74.29 | 76.50 | 74.18 | 77.01 | 74.94 |
AT-skew | –0.10 | –0.12 | –0.13 | –0.12 | –0.12 | |
GC-skew | 0.023 | 0.058 | 0.053 | 0.083 | 0.058 | |
Size | 1440 | 14453 | 14485 | 14483 | 14480 | |
Full | A+T% | 75.30 | 77.09 | 74.76 | 78.07 | 75.63 |
AT-skew | 0.026 | -0.017 | -0.014 | -0.016 | -0.013 | |
GC-skew | -0.18 | -0.087 | -0.10 | -0.095 | -0.10 | |
Size | 16215 | 15369 | 16602 | 16164 | 16367 |
All species of Stenopsocidae share a similar mitochondrial genetic arrangement with the infraorder Caeciliusetae, and show the trnE-trnS1 rearrangement which was considered to be an autapomorphy for Caeciliusetae by
In the present study, we use the mitogenomes and 18S rRNA of 21 species of Stenopsocidae with additional 11 mitogenomes of other bark lice as outgroup to infer the phylogeny of the family Stenopsocidae. The aligned and cleaned full data matrix was 14738 bp (mitogenomic data: 13006 bp, 18S rRNA data: 1732 bp) in length (File S1). Phylogenetic analyses under ML and BI analyses showed the same topology and overall high nodal supports (Fig.
Phylogenetic tree inferred from maximum likelihood analyses of the mitochondrial genes (the protein-coding sequences and rRNA genes) and the 18S rRNA gene. Numbers associated with each branch indicate ML bootstrap/Bayesian posterior probabilities. Side bars on the right are classification according to
Order Psocodea Hennig, 1966
Suborder Psocomorpha Badonnel, 1951
Infraorder Caeciliusetae Pearman, 1936
Family Stenopsocidae Koble, 1880
Graphopsocus Kolbe, 1880:185. Type species: Hemerobius cruciatus Linnaeus, 1768: 225, original designation.
Body length 2–3 mm, from postclypeus to wing tip length 3–5 mm. Antenna slightly longer than body length, not longer than wing length. Labrum with distal styli. Head with brown vertex. Forewing with several dark markings, proximal half with one or two brown marking between CuP and A, and distal half with brown markings at middle and also long distal margin, forming a large and a small V-shaped patterns, pterostigma general with brown markings on basal region. Forewing anterior margin rather sparsely setose, costal vein without seta from base to pterostigma; CuP glabrous, all other veins with single-row setae; pterostigma broad, with a distinct posterior angle. Abdomen with two ventral vesicles. Male genitalia: tip of aedeagal arch not exceeding parameres (aedeagal arch exceeding parameres in G. vietnamensis); endophallus with one pair of weakly sclerotized lobes. Female genital: spermathecal sac with a small pouch.
The genus is widely distributed in all main zoogeographical regions of the world.
Malostenopsocus Li, 1992: 250. Type species: Malostenopsocus yunnanicus Li, 1992: 250, original designation.
Male body shorter than 3 mm, female body longer than 3 mm, from postclypeus to wing tip length 5–7 mm. Antenna slightly longer than body length, not longer than wing length. Labrum without distal styli. Head with several dark brown markings. Wings transparent, without distinct markings (M. plurifasciatus with markings on wings), veins darker than wing margin. Forewing costal vein with setae from base of anterior margin to pterostigma, all veins with single-row setae at least, but R, Rs+M, CuA+M, and CuA with two rows of setae, and basal membranous part of forewing with setae. Pterostigma broad, with a distinct posterior angle. Abdomen with three ventral vesicles. Abdomen with segments 8–9 indistinctly sclerotized. Male phallosome strongly sclerotized, endophallus with three lobes. Female spermathecal sac with a lateral pouch connecting to spermathecal duct.
Oriental region.
Stenopsocus externus Banks, 1937: 259.
Body length from postclypeus to wing tip 4–7 mm. Antenna distinctly longer than body and forewing in length. Labrum with distal styli. Forewing with setose anterior margin and single-row setae on all veins except for CuP, CuP glabrous; pterostigma elongate, with a distinct posterior angle; Rsm slightly curved. Forewing pterostigma with variable brown markings, other regions transparent or yellowish. Genitalia from yellowish to dark brown. Male genitalia: tip of aedeagal arch not exceeding parameres. Female subgenital plate posteriorly convex; dorsal valve and ventral valve sclerotized, narrow; external valve reduced; spermathecal sac generally pear-shaped without pouch.
Oriental, Palaearctic and Australian regions.
The generic name is a combination of Greek “neo-” (meaning “new”) and “Stenopsocus”, meaning the new genus of Stenopsocidae. Gender masculine.
Based on morphological characters, we transferred 115 species previously placed in Stenopsocus to Neostenopsocus gen. n.. The checklist is provided..
Stenopsocus Hagen, 1866: 203. Type species: Psocus immaculatus Stephens, 1836: 125, original designation.
Cubipilis Li, 1993: 350. Type species: Cubipilis hamaocaulis Li, 1993.
Body length from postclypeus to wing tip 4–7 mm. Antenna longer than body and forewing. Head with variable brown markings. Labrum without distal styli. Pterostigma transparent, partly brown or wholly brown. Forewing with setose anterior margin, all veins with single-row setae; pterostigma narrowly elongate, with indistinct or distinct posterior angle; Rsm slightly curved. Genitalia generally yellowish white, weakly sclerotized. Male genitalia: tip of aedeagal arch generally exceeding parameres; endophallus with complex sclerotized papillae regions. Female subgenital plate posteriorly convex; dorsal valve and ventral valve sclerotized; proximal portion of dorsal valve broad; external valve reduced; spermathecal sac generally pear-shaped.
Oriental and Palaearctic regions.
1 | Forewing with glabrous CuP; labrum with distal styli | 2 |
1’ | Forewing with setose CuP; labrum without distal styli | 3 |
2 | Forewing: one or two markings between CuP and A; abdomen with two ventral vesicles | Graphopsocus Kolbe |
2’ | Forewing: no marking between CuP and A; abdomen with three ventral vesicles | Neostenopsocus gen. n. |
3 | Forewing: base membrane with setae, Rsm obviously curved | Malostenopsocus Li |
3’ | Forewing: membrane without seta, Rsm slightly curved, nearly straight | Stenopsocus Hagen |
Neostenopsocus abnormis (Liang, Li & Liu, 2017) comb. n.
Neostenopsocus adisoemartoi (Cole, New & Thornton, 1989) comb. n.
Neostenopsocus albipileus (Smithers, 1974) comb. n.
Neostenopsocus albus (Li, 1992) comb. n.
Neostenopsocus angustifurcus (Li, 2002) comb. n.
Neostenopsocus angustistriatus (Li, 2002) comb. n.
Neostenopsocus anthracinus (Li, 1989) comb. n.
Neostenopsocus aureus (Li, 2002) comb. n.
Neostenopsocus bellatulus (Li, 1989) comb. n.
Neostenopsocus betulus (Li, 2002) comb. n.
Neostenopsocus biconicus (Li, 2002) comb. n.
Neostenopsocus biconvexus (Li, 1997) comb. n.
Neostenopsocus bimaculatus (Li, 2002) comb. n.
Neostenopsocus bipunctatus (Li, 2002) comb. n.
Neostenopsocus bombusus (Li, 2002) comb. n.
Neostenopsocus brachychelus (Li & Yang, 1988) comb. n.
Neostenopsocus brachycladus (Li, 1989) comb. n.
Neostenopsocus brachyodicrus (Li, 2002) comb. n.
Neostenopsocus brevicapitus (Li, 1997) comb. n.
Neostenopsocus brevivalvaris (Li, 2002) comb. n.
Neostenopsocus capacimacularus (Li, 1993) comb. n.
Neostenopsocus cassideus (Li, 1992) comb. n.
Neostenopsocus ceuthozibrinus (Li, 2002) comb. n.
Neostenopsocus changbaishanicus (Li, 2002) comb. n.
Neostenopsocus concisus (Li, 2002) comb. n.
Neostenopsocus daozheniensis (Li, 2005) comb. n.
Neostenopsocus dichospilus (Li, 2002) comb. n.
Neostenopsocus dictyodromus (Li, 1993) comb. n.
Neostenopsocus disphaeroides (Li, 2002) comb. n.
Neostenopsocus emeishanicus (Li, 2002) comb. n.
Neostenopsocus eucallus (Li & Yang, 1988) comb. n.
Neostenopsocus externus (Banks, 1937) comb. n.
Neostenopsocus fanjingshanicus (Li & Yang, 1988) comb. n.
Neostenopsocus faungi (Li, 1999) comb. n.
Neostenopsocus flavicaudatus (Li, 2002) comb. n.
Neostenopsocus flavifrons (Li, 1989) comb. n.
Neostenopsocus flavinigrus (Li, 2002) comb. n.
Neostenopsocus floralis (Li, 2002) comb. n.
Neostenopsocus foliaceus (Li, 1997) comb. n.
Neostenopsocus formosanus (Banks, 1937) comb. n.
Neostenopsocus frontalis (Li, 1989) comb. n.
Neostenopsocus frontimaculatus (Li, 1992) comb. n.
Neostenopsocus fulivertex (Li, 2002) comb. n.
Neostenopsocus gannanensis (Li, 2002) comb. n.
Neostenopsocus gansuensis (Li, 2002) comb. n.
Neostenopsocus genostictus (Li, 2002) comb. n.
Neostenopsocus gibbulosus (Li, 1995) comb. n.
Neostenopsocus gracilimaculatus (Li, 2002) comb. n.
Neostenopsocus gracillimus (Li & Yang, 1988) comb. n.
Neostenopsocus guizhouiensis (Li, 2002) comb. n.
Neostenopsocus hemiostictus (Li, 2002) comb. n.
Neostenopsocus hexagonus (Li, 2002) comb. n.
Neostenopsocus huangshanicus (Li, 2002) comb. n.
Neostenopsocus isotomus (Li, 2002) comb. n.
Neostenopsocus jocosus (Banks, 1939) comb. n.
Neostenopsocus lacteus (Li, 1997) comb. n.
Neostenopsocus laterimaculatus (Li, 2002) comb. n.
Neostenopsocus lemniscsingulaeis (Li, 2005) comb. n.
Neostenopsocus lifashengi (Mockford, 2003) comb. n.
Neostenopsocus liuae (Li, 2002) comb. n.
Neostenopsocus liupanshanensis (Li, 2002) comb. n.
Neostenopsocus longicuspis (Li, 1997) comb. n.
Neostenopsocus longitudinalis (Li, 2002) comb. n.
Neostenopsocus macrocheirus (Li, 2002) comb. n.
Neostenopsocus maculosus (Li & Yang, 1988) comb. n.
Neostenopsocus makii (Takahashi, 1938) comb. n.
Neostenopsocus maximalis (Li, 1997) comb. n.
Neostenopsocus melanocephalus (Li, 1997) comb. n.
Neostenopsocus metastictus (Li, 2002) comb. n.
Neostenopsocus naevicapitatus (Li, 2002) comb. n.
Neostenopsocus nepalensis (New, 1971) comb. n.
Neostenopsocus nigricellus (Okamoto, 1907) comb. n.
Neostenopsocus obscurus (Li, 1997) comb. n.
Neostenopsocus oculimaculatus (Li, 1992) comb. n.
Neostenopsocus pallidus (Thornton & Wong, 1966) comb. n.
Neostenopsocus parviforficatus (Li, 2002) comb. n.
Neostenopsocus pavonicus (Li, 2002) comb. n.
Neostenopsocus paxillivalvaris (Li, 2002) comb. n.
Neostenopsocus percussus (Li, 1995) comb. n.
Neostenopsocus periostictus (Li, 2002) comb. n.
Neostenopsocus perspicuus (Li, 1997) comb. n.
Neostenopsocus phaeostigmus (Li, 1992) comb. n.
Neostenopsocus phaneostriatus (Li, 2002) comb. n.
Neostenopsocus platynotus (Li, 1995) comb. n.
Neostenopsocus platyocephalus (Li, 2002) comb. n.
Neostenopsocus podorphus (Li, 1997) comb. n.
Neostenopsocus polyceratus (Li, 2002) comb. n.
Neostenopsocus pygmaeus (Enderlein, 1906) comb. n.
Neostenopsocus qianipullus (Li, 2005) comb. n.
Neostenopsocus radimaculatus (Li, 1989) comb. n.
Neostenopsocus revolutus (Li, 1993) comb. n.
Neostenopsocus rubellus (Thornton, 1984) comb. n.
Neostenopsocus shennongjiaensis (Li, 2002) comb. n.
Neostenopsocus sichuanicus (Li, 2002) comb. n.
Neostenopsocus signatipennis (New, 1978) comb. n.
Neostenopsocus silvaticus (Li, 2002) comb. n.
Neostenopsocus spongiosus (Li, 2002) comb. n.
Neostenopsocus stigmaticus (Imhoff & Labram, 1842) comb. n.
Neostenopsocus striolatus (Li, 1992) comb. n.
Neostenopsocus symipsarous (Li, 2002) comb. n.
Neostenopsocus thermophilus (Li, 2002) comb. n.
Neostenopsocus tibialis (Banks, 1937) comb. n.
Neostenopsocus tribulbus (Li, 1993) comb. n.
Neostenopsocus tripartibilis (Li, 2002) comb. n.
Neostenopsocus trisetus (Li, 2002) comb. n.
Neostenopsocus turgidus (Li, 1997) comb. n.
Neostenopsocus uniformis (Hagen, 1859) comb. n.
Neostenopsocus valvilacteus (Li, 2005) comb. n.
Neostenopsocus wangi (Liang, Li & Liu, 2017) comb. n.
Neostenopsocus wuxiaensis (Li, 1997) comb. n.
Neostenopsocus xanthophaeus (Li, 2002) comb. n.
Neostenopsocus xanthostigmus (Li, 2002) comb. n.
Neostenopsocus xiangxiensis (Li, 1992) comb. n.
Neostenopsocus xilingxianicus (Li, 1997) comb. n.
Neostenopsocus zonatus (Li, 1989) comb. n.
Shape of the vertex (Fig.
Head of stenopsocid species in frontal view. A Graphopsocus cruciatus, male; B Graphopsocus cruciatus, female; C Malostenopsocus yunnanicus, male; D Malostenopsocus yunnanicus, Female; E Neostenopsocus externus, male; F Neostenopsocus externus, female; G Stenopsocus immaculatus, male; H Stenopsocus immaculatus, female. Scar bars = 0.5 mm.
Presence or absence of labral styli (Fig.
The markings on forewing (Fig.
Forewing venation (Fig.
Ciliation of forewing (Fig.
Number of ventral abdominal vesicles. In Graphopsocus, two ventral vesicles are present. In Neostenopsocus gen. n., Malostenopsocus and Stenopsocus, three ventral vesicles are present. There can be no question that the condition of two ventral abdominal vesicles is plesiomorphic for Stenopsocidae.
Phallosome (Fig.
As discussed above, several plesiomorphic features in Stenopsocidae are revealed, i.e., round vertex, presence of labral styli, broad pterostigma with a distinct posterior angle, straight Rsm on the forewing, glabrous anterior margin and CuP on the forewing, presence of two abdominal vesicles and the bilobed endophallus with weakly sclerotized portion. The new genus Neostenopsocus gen. n. is supported by several apomorphies: (i) forewing mostly transparent, (ii) abdomen with three ventral vesicles; (iii) endophallus with partly divided upper and base parts of minute sclerotized papillae.
The present phylogenetic analyses confirm the monophyly of the family Stenopsocidae, which has been strongly supported based on the morphological characters and multi-loci phylogenetic studies (Yoshizawa, 2002;
The phylogeny recovered a sister relationship between Neostenopsocus gen. n. and Malostenopsocus + Stenopsocus. This clade is supported by several apomorphies: (i) labrum with styli, (ii) abdomen with three ventral vesicles; (iii) endophallus with several lobes of minute sclerotized papillae. The genus Graphopsocus is estimated to be the sister group of all other stenopsocid taxa, as the stem branch of Stenopsocidae in the present study.
Herein, we propose a new classification scheme for the family Stenopsocidae based on our phylogenetic analysis of molecular data and re-evaluation of morphological characters. In this scheme, Stenopsocidae is classified into four genera: Graphopsocus, Malostenopsocus, Neostenopsocus gen. n., and Stenopsocus.
We sequenced 20 new nearly complete mitochondrial genomes of Stenopsocidae, and reconstructed the intergeneric phylogeny of this family. A new genus, namely Neostenopsocus gen. n., is proposed based on the molecular phylogeny herein inferred. Most species of Stenopsocidae were described from the Oriental realm, and it is likely that unknown genera and species are still to be discovered in this region. Future fieldwork and taxonomic studies based on morphological and molecular data could lead to further insight into the diversity and phylogeny of this family.
The authors declare no conflicts of interest.
We appreciate Prof. André Nel and the two anonymous reviewers for their valuable and critical comments on this paper. We are grateful to Mr. Fasheng Li for his great attribution of the study of Stenopsocidae. We thank all the people who collected the specimens examined in the present paper. We thank Prof. Kazunori Yoshizawa for the information on the Stenopsocidae from Japan. Many thanks to Mr. Yuchen Zheng for his kind help in the phylogenetic analyses. We thank Mr. Ruiyang Wang, Mr. Qianle Lu and Mr. Fan Gao for sharing their photographs of living stenopsocids. We thank Dr. Phillip Perkins for his kind help when LFY visited the collection of ‘Psocoptera’ in the Museum of Comparative Zoology, Harvard University. This research was supported by the National Natural Science Foundation of China (32100362), the Scientific Research Fund of Hunan Provincial Education Department (23B0490), the Natural Science Foundation of Hunan Province (2021JJ40194) and the National Animal Collection Resource Centre, China.
File S1
Data type: .dat
Explanation notes: Aligned and cleaned full data matrix.