Lacewings of Vietnam: new species and distributional records of Osmylidae and Sisyridae (Neuroptera: Osmyloidea)

ABSTRACT

Despite ongoing taxonomic efforts, species diversity and geographic distributions remain poorly documented for many groups of Osmyloidea, including in parts of Asia. In this study, we contribute to the taxonomic knowledge of Osmyloidea, the sole group of lacewings (Insecta: Neuroptera) with both freshwater and terrestrial representatives, while also reducing distributional gaps for the group. We conducted detailed examination of lacewing specimens from Vietnam from worldwide entomological museums. We described two new species of Osmylidae, Osmylus hagiangensis sp. nov. and Thaumatosmylus solitarius sp. nov. This is the first record of the genera Osmylus Latreille and Thaumatosmylus Krüger from Vietnam, totalling new records of four species of Osmylidae. We also provide the first record of three species of Sisyridae in the country Sisyra nikkoana (Navás), S. nobilia Yang, Zheng & Liu and Sisyrina qiong Yang & Gao. Previous studies recorded only five species and four genera in Vietnam, and our research nearly doubles this diversity: 12 species in six different genera.

Keywords:
Identification guide; Lance lacewings; Neuropterida; Oriental region; Spongillaflies

Introduction

Insects, one of the most incredibly abundant and diverse groups of organisms on Earth, play an essential role in all aspects of environmental functioning. A particularly intriguing, yet underrepresented, group of insects is the lacewings (Insecta: Neuroptera). These ecologically diverse insects have been recorded on every continent, except for Antarctica (Oswald and Machado, 2018). Approximately 5,800 extant species of lacewings are divided into seven superfamilies (Oswald and Machado, 2018; Winterton et al., 2018).

Osmyloidea, one of the superfamilies of lacewings, encompass three families: Sisyridae, Nevrorthidae, and Osmylidae. They comprise the only group of lacewings with aquatic, semi-aquatic and terrestrial species (Oswald and Machado, 2018; Winterton et al., 2018). Osmyloidea display distinct feeding behaviors among its larvae, including generalist predation or specialized feeding on freshwater sponges and bryozoans (Oswald and Machado, 2018). The adults present wider feeding habits, including herbivory, omnivory or predation, and they can be found both in the vicinity of freshwater habitats or, in the case of terrestrial species, far from aquatic ecosystems (Aspöck et al., 2017; Oswald and Machado, 2018; Winterton et al., 2019; Assmar et al., 2022).

The relationships among Osmyloidea families have been extensively discussed over the past decade, with competing hypotheses proposed to explain their origin and evolution (Winterton et al., 2018; Vasilikopoulos et al., 2020). Molecular evidence, associated with morphological features such as the enlarged semi-articulated gonocoxite 9 of the females, provides support for the Osmyloidea clade (Winterton et al., 2018; Vasilikopoulos et al., 2020; Lai et al., 2024; Zhang et al., 2025; Assmar et al., 2026). This superfamily comprises more than 340 extant species (Oswald, 2024), with Osmylidae accounting for approximately 240 species (Winterton et al., 2019; Li et al., 2025). Sisyridae are the second richest family, containing ~80 species, followed by Nevrorthidae with 20 species (Aspöck et al., 2017; Yang and Liu, 2023; Oswald, 2024). The origin of Osmyloidea is estimated to date to the Late Permian (~250 Ma) (Winterton et al., 2018), but their biogeographic history remains uncertain (Engel et al., 2018; Yang and Liu, 2023).

While Osmyloidea are distributed worldwide, individual families have distinct geographic ranges. Osmylidae have been recorded in all zoogeographic regions except the Nearctic, where only Tertiary-aged fossils of the family have been discovered (Winterton et al., 2017, 2019). Sisyridae have a cosmopolitan distribution, while Nevrorthidae are restricted to the Mediterranean region, China, Japan, and East Australia (Aspöck et al., 2017; Assmar et al., 2022). In the Oriental zoogeographic region, a considerably high number of species of Osmyloidea are known to occur (~90 spp.) (Aspöck et al., 2017; Yang et al., 2018, 2024; Winterton et al., 2019; Oswald, 2024; Li et al., 2025). Most of this known diversity is found in India, Southern China and Indonesia, but Vietnam, for example, has only five species recorded (Winterton et al., 2019; Yang and Liu, 2023; Oswald, 2024). Among these, two belong to Osmylidae, Gryposmylus pennyi Winterton & Wang and Spilosmylus inclytus (Navás), and three to Sisyridae, Sisyra bowlesi Yang & Liu, Sisyra indica Needham and Sisyrina vietnamica Yang & Liu (Navás, 1917; Winterton and Wang, 2016; Yang and Liu, 2021, 2023; Oswald, 2024).

The known diversity of Osmyloidea in Vietnam is considerably low, despite the country's rich environmental landscapes, including tropical forests and numerous freshwater habitats (Phuong, 2007). Moreover, the northern neighbor country, China, has a higher number of Osmyloidea species documented, with at least 35 species recorded from the Southern region only (Yang et al., 2018; Winterton et al., 2019; Yang and Liu, 2021, 2023; Li et al., 2025). This discrepancy suggests a significant knowledge gap regarding the Vietnamese fauna. Therefore, the primary goal of this paper is to address the distributional shortfall by providing information on Osmyloidea diversity in Vietnam. More specifically, we provide the first record of three species of Sisyridae from Vietnam. Additionally, we provide the first record of two genera, Osmylus Latreille and Thaumatosmylus Krüger, and four species of lance lacewings in Vietnam, including the description of two new species, Osmylus hagiangensis sp. nov. and Thaumatosmylus solitarius sp. nov.

Material and methods

The specimens examined here are deposited in the Texas A&M University Insect Collection (TAMUIC), the American Museum of Natural History (AMNH) and the California Department of Food and Agriculture (CDFA), all in the United States of America. We dissected the last segments of the abdomen of each specimen, prepared the genitalia using heated KOH 10% for 5-10 minutes, and followed the protocol by Blahnik and Holzenthal (2004) to neutralize the reaction. All genitalia are stored in microvials containing glycerin, which are attached to the specimen. Stacked photographs of the specimens were made with a Canon EOS 70D Macro Photo Lens 65mm, attached to the Cognisys Stack Shot, with support of Helicon Remote^©, and merged with Helicon Focus^©. Illustrations and image plates were made using Adobe Illustrator^© 2023 and we prepared the photos using Adobe Photoshop^© CC 2024.

We developed distributional maps using QGIS3 3.10 A Coruña, and the geographic location of the specimens were based on primary literature and material examined. Names in bold in the distribution sections represent new records, and we follow Holt et al. (2013) for the zoogeographic regions. Terminology for the genitalia follows Aspöck and Aspöck (2008) and for the wing venation follows Breitkreuz et al. (2017). Abbreviations: C – Costa, Sc – Subcosta, R – Radius, RA – Radius anterior, RP – Radius posterior, M – Media, MA – Media anterior, MP – Media posterior, Cu – Cubitus, CuA – Cubitus anterior, CuP – Cubitus posterior, A – Anal.

For the Thaumatosmylus specimen identification, we conducted a phylogenetic analysis and calculated genetic distances between this specimen and other specimens of Thaumatosmylus. We included a total of 11 Cytochrome C Oxidase subunit I (COX1) gene sequences, seven ingroups and four outgroups. From these sequences, five were newly sequenced and six obtained from GenBank. The list of taxa included in this analysis is as follows: Thaumatosmylus solitarius sp. nov. (newly sequenced), Thaumatosmylus hainanus Yang (GenBank MK408756.1), Thaumatosmylus aff. hainanus (newly sequenced), Thaumatosmylus punctulosus Yang (GenBank KY491698), Thaumatosmylus sp.1 (newly sequenced), Thaumatosmylus umbratus New (newly sequenced), Thaumatosmylus zheanus Yang & Liu (GenBank KY491699), Thyridosmylus langii McLachlan (newly sequenced, outgroup), Thyridosmylus paralangii Wang et al. (GenBank SAMN07526209, outgroup), Thyridosmylus qianus Yang (GenBank KY491705, outgroup), Thyridosmylus triypsiloneurus Yang et al. (GenBank KY491706, outgroup).

The newly sampled COX1 were obtained by capturing mitochondrial genes from whole genome sequences using MitoFinder (Allio et al., 2020), as well as from anchored hybrid enrichment (AHE) sequences, both using metaSPAdes (Nurk et al., 2017) as the assembly option. The DNA of these specimens were extracted using thoracic tissues or by submerging the entire specimen in the buffer. We aligned the sequences as nucleotides using MAFFT L-INS-I strategy (Katoh and Standley, 2013). We estimated a maximum likelihood phylogenetic tree using IQ-TREE (Minh et al., 2020), with random seed, two runs, ultrafast bootstrap (Hoang et al. 2018) with 2000 iterations and substitution model estimated using ModelFinder (Kalyaanamoorthy et al., 2017). The phylogenetic tree was edited using FigTree. Genetic distance was calculated with the Kimura 2-parameter (K2P) model (Kimura, 1980) in MEGA 12.1 (Stecher et al., 2020, 2025), including transitions and transversions substitutions, with gamma rates among sites and pairwise-deletion options. Final alignment and log files are available in the Supplementary Material.

Results

Osmylidae

Gryposmylus pennyi Winterton & Wang

Gryposmylus pennyi Winterton & Wang, 2016: 38 (description) – Winterton et al., 2019: 49 (checklist).

Distribution

China, Vietnam (Winterton and Wang, 2016).

Remarks

Winterton and Wang (2016) designated a male holotype for this species, which is deposited at the California State Collection of Arthropods, USA. It is currently recorded to Northern Vietnam, in the Ninh Binh province, and Southern China, in the Yunnan province (Winterton and Wang, 2016).

Osmylus angustimarginatus Xu et al.

(Figures 1a-b)

Figure 1
Osmylus angustimarginatusXu et al. (2016) a) Male habitus, forewing and hind wing in evidence; b) male genitalia in lateral view.

Osmylus angustimarginatus Xu et al., 2016: 117 (description) – Winterton et al., 2019: 38 (checklist).

Distribution

China, Vietnam.

Remarks

Xu et al. (2016) designated a male holotype for this species, which is deposited at the Entomological Museum of China Agricultural University (CAU), Beijing. The holotype and paratypes were collected in the Chongqing municipality, central China, North of Guizhou province (Xu et al., 2016). The specimens we examined were identified as O. angustimarginatus based not only on the characteristic wing venation (Figure 1a), but also on genitalia, especially the ventral margin of the gonocoxite 9, which is very sclerotized (Figure 1b). This is the first record of this species to Vietnam.

Material examined

VIETNAM: Ha Giang, IX-9 2000, 1210m, 22 46.15N 104 49.38E, sweep, col. Johnson; det. Alice Assmar, 1 ♀ (AMNH); Ha Giang, IX-23 2000, 1210m, 22 46.15N 104 49.38E, Malaise, col. Johson and Nguyen, det. Alice Assmar, 1 ♂ (AMNH).

Osmylus hagiangensis sp. nov.

(Figures 2ak)

Figure 2
Osmylus hagiangensissp. nov. Habitus, in dorsal view, with forewing and hind wing in evidence of a) male holotype and b) female paratype. Lateral view of genital sclerites of the c) male and d) female. Digital illustration of genital sclerites e-j: e) male, lateral view; f) female, lateral view; g) female egg sac; h) male ventral view; Gonostyli 10 in i) lateral and j) ventral view. k) Proximal view of the head. (Abbreviations: t = tergite, dp = dorsal processus, cc = callus cerci, st = sternite).

LSID urn:lsid:zoobank.org:act:461F2A5C-0ABA-4632-A29A-EC214CC4EA14

Holotype

A male specimen collected in Ha Giang, Vietnam (AMNH). Specimen preserved in ethanol, genitalia in glycerin.

Distribution

Vietnam.

Diagnosis

Head with yellow transversal streak on vertex. Apical margin of the femur infuscate, forming a narrow ring-like mark at the joint with the tibia. Tibia darkened along the outer margin. Forewing with scattered small brown streaks. Long dorsal projection of the male tergite 9, the apex is rounded and the dorsal margin is slender.

Description

Body. Size varied from 16.4–19 mm.

Head. Brown; vertex with a yellow transversal band. Three distinct ocelli present, with long light brown setae. Scape light brown, pedicel and flagellum dark brown. Presence of a small tubercle below each eye.

Thorax. Brown with dark brown marks on sutures. Long brown and light brown setae spread across the pro, meso and metanotum. Legs. Yellow to light brown, covered with short brown setae; femur with darker color near the joint with the tibia, tibia dark along outer margin, last tarsomere darker than the previous ones; pretarsal claws curved.

Wings. Forewing length varied from 24.8–28 mm, broad and slightly lanceolate; membrane hyaline with five small dot-like streaks along distal margin, on the outer series of gradate crossveins, and one at the wing apex; veins covered with setae; costal space basally broad with approximately 45 subcostal veinlets before pterostigma, several forked, interlinking veinlets present; Pterostigma slightly marked; Presence of one subcostal crossvein, approximately 30 RA-RP crossveins, and at least two recognizable series of gradate veins; Crossveins are numerous; presence of two darkened nigmata; RP with 10 or more branches; M forked close to the first RP fork; Cu forked near wing base, CuA and CuP seems to fuse near distal margin, CuP highly pectinated; Anal veins pectinated. Hindwing length varied from 22–24.4 mm, slightly lanceolate; membrane hyaline; presence of hyaline nigmata; veins covered with fine setae; costal space with approximately 37 crossveins before pterostigma, basal ones not forked; Pterostigma slightly marked; Presence of approximately 30 RA-RP crossveins, and at least two recognizable series of gradate veins; sigmoid rp-m present; RP with 12 or more branches; M forked near wing base before RA-RP fork; Cu forked near wing base, CuP highly pectinated; Anal veins pectinated, presence of two to three crossveins in anal area.

Abdomen. Brown, paler in the male specimen.

Male abdomen. Scent glands rod-shaped. Tergite 9 long and slightly rectangular with a very long, dorsal process, curving upwards (Figures 2c, e). Sternite 9 subtriangular in lateral view, covered by long, thick setae. Ectoproct triangular in lateral view, callus cerci round (Figures 2e, h). Posterior part of gonocoxite 9 well sclerotized, postero-ventral margin with small protuberance in lateral view. Endoprocessus beneath gonocoxite 9, apex bent upward. Gonostyli 10 slender basally with a long, sharp backward end, thin apically in lateral view and distally curved; parameres slender behind gonostyli 10, posterior end sharp (Figures 2i, j).

Female abdomen. (Figures 2d, f-g). Tergite 8 large and sub-quadrangular. Sternite 8 small and triangular. Tergite 9 long, sharp ventrally. Ectoproct quadrangular, with minor projection at dorsal margin apex, callus cerci round. Gonocoxite 9 approximately fusiform, larger apically, with small medial projection on ventral margin; Gonapophysis 9 triangular; spermatheca simple, rod-shaped.

Etymology

The specific name “hagiangensis” derives from the Latin and refers to the type locality, Ha Giang.

Remarks

The new species is similar to Osmylus punctipennis Walker in regard to the wing coloration, the femur with darker color near the joint with the tibia, and the tibia has a darker band dorsally (Walker, 1860; Martins and Price, 2020). However, the new species can be differentiated from O. punctipennis by the forewing MP fused with the CuA vein, which is absent in the new species. Osmylus hagiangensis sp. nov. also has the yellow transverse space above the antennae, similar to other species in Osmylus hyalinatus species group (Li et al., 2025). It can be easily differentiated from its congeners by the shape of the male genitalia, as the dorsal processus of the tergite 9 curves upwards and is longer than in other species that have similar projections.

Material examined

Holotype: VIETNAM: Ha Giang, IX-9 2000, 1210m, 22 46.15N 104 49.38E, sweep, col. Johnson, det. Alice Assmar, 1 ♂ (AMNH); Paratype: VIETNAM: Ha Giang, IX-23 2000, 1210m, 22 46.15N 104 49.38E, Malaise, col. Johson and Nguyen, det. Alice Assmar, 1 ♀ (AMNH).

Spilosmylus inclytus (Navás)

Centrolysmus inclytus Navás, 1917: 18.

Spilosmylus inclytus – Winterton et al., 2019: 64 (checklist).

Distribution

“Indochina: Chapa” [Vietnam] (Navás, 1917).

Remarks

Navás (1917) described this species based on a female specimen, but it remains uncertain where the type material is deposited. Winterton et al. (2019) synonymized Centrolysmus Navás with Thyridosmylus Krüger but transferred S. inclytus (Navás) to Spilosmylus Kolbe. Regarding its geographic distribution, the type locality of this species is “Indochina: Chapa”. Indochina was a large territory encompassing the modern countries of Vietnam, Laos, and Cambodia before their independence in the mid 1950’s. According to Michaud and Turner (2000), “Chapa” was how the French called Sa Pa district, in the province of Lào Cai, Vietnam. Therefore, S. inclytus is also recorded from Vietnam.

Spilosmylus inthanonensis New

(Figures 3a-b)

Figure 3
Spilosmylus inthanonensis New. a) Male habitus, forewing and hind wing in evidence; b) male genitalia in lateral view.

Spilosmylus inthanonensis New, 1991: 18, figs 60–64 (description) – New, 2003: 152 (list) – Winterton et al., 2019: 64 (checklist) – Martins and Price, 2020: 42 (type material).

Distribution

Thailand (Winterton et al., 2019), Vietnam.

Remarks

New (1991) described this species based on a male specimen from Doi Inthanon National Park in Thailand. Here, we expand the known geographic distribution of S. inthanonensis by recording this species for the first time in Vietnam. Most likely, S. inthanonensis also occurs in Laos, and New (2003) suggested that it might occur in Malaysia as well.

We identified this species based on characteristics of the wings, such as the absence of an embossed spot in the forewing, the presence of two brownish transversal bands in the basal half of the forewing, and the darkened sides of the pterostigma with a hyaline center. Additionally, the specimen examined herein displays the same shape of the male terminal segments as described by New (1991).

Material examined

VIETNAM: Ha Giang, IX-9-23 2000, 1210m, 22 46.15N 104 49.38E, Yellow pan trap, col. Johnson, det. Alice Assmar, 2 ♀ 1 ♂ (AMNH); Ha Giang, IX-9-16 2000, 1210m. 22 46.15N 104 49.38E, black light/mercury vapor lamp, col. Johson and Nguyen, det. Alice Assmar, 1 ♀ (AMNH).

Thaumatosmylus solitarius sp. nov.

(Figures 4a-f)

Figure 4
Thaumatosmylus solitariussp. nov. Female holotype a) forewing and b) hind wing. c) Habitus in lateral view, d) head and prothorax in lateral view, e) and in dorsal view, f) genitalia in lateral view. (Abbreviations: tb = tubercle).

LSID urn:lsid:zoobank.org:act:DA1BD614-CD56-4D84-BC49-2B232F5B2ADF

Holotype

A female specimen collected in Cuc Phuong National Park, Vietnam (CDFA). Specimen preserved in ethanol, genitalia is in glycerin, thorax detached, wings damaged and half of the antennae are missing.

Distribution

Vietnam.

Diagnosis

Pronotum with two lateral tubercles near the joint with the head. Wings sub-triangular with relatively dense coloration pattern. Presence of conspicuous streaks in the radial area, also on the second and fourth fork of RP of the wing. Wing veins are brown with scattered pale regions. Forewing with 2 m-cu before fork of M.

Description

Body. Size approximately 10.7 mm.

Head. Pale yellow to light brown. Vertex pale near pronotum, with two dark brown longitudinal streaks, ocelli area dark brown. Three distinct ocelli present, with long dark brown setae. Scape and pedicel light brown, and flagellum yellow.

Thorax. Dorsally brown and ventrally pale yellow. Long dark brown and smaller yellowish to light brown setae spread across the pro, meso and metanotum. Pronotum with antero-lateral tubercles, near the joint with the head. Legs yellow, covered with short light brown setae; pretarsal claws curved.

Wings. Forewing length approximately 20 mm, broad and slightly lanceolate; membrane hyaline with distinct coloration pattern; presence of more conspicuous transversally aligned streaks near the wing base in the radial area, also on the second fork of RP and right after the middle of the wing; veins darkened especially in the basal half; costal space relatively broad with 48 simple subcostal veinlets before pterostigma; Pterostigma medially hyaline, with brown streak before and after; Presence of one subcostal crossvein, 14 RA-RP crossveins, and at least one recognizable series of gradate veins; Crossveins are numerous; basal nigma darkened; RP with 14 branches; M forked anteriorly to the first RP fork; Cu forked near wing base, most of CuP branches are simple; Branches of anal veins simple. Hindwing length approximately 18 mm, sub-triangular; membrane hyaline, with four streaks in similar position to those in forewing, two near the second RP fork and two right after the middle of the wing; costal space with 41 simple subcostal veinlets before pterostigma; Pterostigma slightly marked; Presence of 16 RA-RP crossveins, and at least one recognizable series of gradate veins; sigmoid rp-m incomplete; RP with 12 branches; M forked near wing base; Cu forked near wing base, CuP simple; Anal veins simple.

Abdomen. Dorsally brown and ventrally pale yellow.

Male abdomen. Unknown.

Female abdomen. Tergite 8 subquadrate, posteroventral corner ventrally projected, rounded. Presence of an invagination ventral to the tergite 8, with a short projection directed anteriorly, adjacent to the “joint” of tergite 9 and gonocoxite 9. Tergite 9 narrow, with light brown setae specially on dorsal and ventral regions; ventral margin is subtriangular; posterior margin is sinuous. Ectoproct subtriangular covered with long yellowish to light brown setae; callus cerci near the anterior margin. Gonocoxite 9 brown, setose and with subcylindrical format. Spermatheca is clavate.

Etymology

The specific name “solitarius” is a Latin word that means “alone”, “only”, or “single”. It refers to the single female specimen used to describe the species.

Remarks

The results we obtained by calculating genetic distances indicate this species is distinct from its congeners (Table 1). The genetic distance within the genus ranged from 0–0.16, with the intraspecific rate around 0.01 between specimens identified as T. hainanus Yang. Interspecific genetic distances ranged from 0.01 to 0.16, and the new species presented distances ranging from 0.08 to 0.16. Li et al. (2025), in their work with Osmylus hyalinatus species group, recovered intraspecific genetic distances between 0–0.034, while the interspecific was between 0.05–0.13. Although some species in our taxon sampling share very low genetic distances, the distance between T. solitarius sp. nov. and the other species are consistent with interspecific genetic distances.

Table 1
Pairwise genetic distance among COX1 DNA barcodes calculated using Kimura 2-parameter model. Values ranged from 0 to 0.17. The higher the value, the greater the genetic distance.

	Thaumatosmylus aff. hainanus	Thaumatosmylus hainanus mtDNA NCBI	Thaumatosmylus punctulosus KY491698	Thaumatosmylus sp1	Thaumatosmylus solitarius sp. nov.	Thaumatosmylus umbratus	Thaumatosmylus zheanus KY491699	Thyridosmylus langii	Thyridosmylus paralangii ahe_NCBI	Thyridosmylus qianus KY491705	Thyridosmylus triypsiloneurus KY491706
Thaumatosmylus aff. hainanus
Thaumatosmylus hainanus mtDNA_NCBI	0.017
Thaumatosmylus punctulosus KY491698	0.011	0.011
Thaumatosmylus sp1	0.144	0.135	0.141
Thaumatosmylus solitarius sp. nov .	0.102	0.089	0.089	0.162
Thaumatosmylus umbratus	0.143	0.132	0.138	0.000	0.158
Thaumatosmylus zheanus KY491699	0.036	0.030	0.022	0.139	0.089	0.134
Thyridosmylus langii	0.146	0.134	0.146	0.161	0.153	0.157	0.130
Thyridosmylus paralangii ahe_NCBI	0.139	0.126	0.119	0.159	0.143	0.160	0.123	0.096
Thyridosmylus qianus KY491705	0.123	0.110	0.114	0.155	0.139	0.151	0.110	0.130	0.129
Thyridosmylus triypsiloneurus KY491706	0.140	0.134	0.132	0.166	0.170	0.165	0.130	0.133	0.123	0.110

The phylogenetic tree estimation recovered two clades, one including species occurring in Malaysia (T. umbratus and Thaumatosmylus sp.) and another one including species known to occur in Vietnam (T. solitarius sp. nov.) and China (T. hainanus, T. aff. hainanus, T. zheanus, T. punctulosus) (Figure 5). The new species was recovered as sister to all other species in this latter clade, and it was expected that T. solitarius sp. nov. would be closer related to species from China rather than Malaysia due to its geographical distribution.

Figure 5
ML phylogenetic tree reconstructed using seven COX1 sequences of Thaumatosmylus Krüger and four outgroup sequences. Statistical support values are represented by the number at the node, including ultrafast bootstrap (left) and approximate likelihood ratio test (aLRT, right).

The wings’ coloration of T. solitarius sp. nov. is similar to those of Thaumatosmylus ornatus Nakahara, however, they differ by the coloration of the veins, which is brown in the new species and, according to Nakahara (1955), pale in T. ornatus. Additionally, T. ornatus is from Taiwan, which is very far from where the new species was collected (Nakahara, 1955; New, 1991). The wing coloration pattern differs from all other congeners. Additionally, this species has a tubercle-like structure in each side of the prothorax, in the pleura (Figure 4).

Material examined

Holotype: VIETNAM: Ninh Binh Province, Cuc Phuong National Park, 24-28.iii.2012, 20°21'03" N 105°35'36" E [12-09], 390m, mercury vapor light, col. SD Gaimari, M Hauser, Pham HT, det. Alice Assmar, 1 ♀ (CDFA).

Sisyridae

Sisyra bowlesi Yang & Liu

Sisyra bowlesi Yang & Liu, 2023: 3, figs 1, 2 (description).

Distribution

China, Laos, Vietnam (Yang and Liu, 2023).

Remarks

Yang and Liu (2023) designated a male holotype for this species, which is deposited at CAU. This species has a characteristic striped coloration pattern of the antennae, which differentiate it from most of the other species from the region.

Sisyra indica Needham

(Figure 6)

Figure 6
Sisyra indica Needham. Habitus of a male specimen in lateral view.

Sisyra indica Needham, 1909: 206, pl. 21, fig 1 (description) – Banks, 1934: 568 (record) – Navás, 1935: 58 (re-description), fig 31 – Ghosh and Sen, 1977: 287 (record) – Monserrat, 1981: 166 (re-description), figs 1–3 – Yang and Liu, 2023: 12 (re-description), figs 5, 6.

Sisyra vigana Navás, 1923: p. 8 (description) – Navás, 1931: 16 (re-description) – Navás, 1935: 61 (re-description), fig 32 – Monserrat, 1981: 166 (synonym) – Monserrat, 1985: 243 (list) – Baltazar, 1990: 16 (record).

Sisyra aquavivai Navás, 1929: 52, fig 28 (description) – Navás, 1931: 16 (re-description) – Navás, 1935: 56 (re-description), fig 29 – Ghosh and Sen, 1977: 287 (record) – Monserrat, 1981: 166 (synonym) – Monserrat, 1985: 243 (list).

Sisyra hainana Yang & Gao, 2002: 286 (description) – Yang and Liu, 2023: 14 (synonym).

Sisyra nervata Yang & Gao, 2002: 287 (description) – Yang and Liu, 2023: 14 (synonym).

Distribution

China, India, Malaysia, Philippines, Thailand, Vietnam (Yang and Liu, 2023).

Remarks

Yang and Liu (2023) have recently conducted a comprehensive revision of this species. The authors redescribed the species and placed S. hainana Yang and Gao and S. nervata Yang and Gao as junior synonyms of S. indica Needham. They have also expanded the known distributional range of the species. Here, we further contribute to the known geographic distribution of S. indica, by recording it to the Quang Tri province, in Vietnam. Sisyra indica is widespread throughout the Oriental region.

Material examined

VIETNAM: Quang Tri Prov., 1 mi N Quang Tri, VII-11-1970, at light, Coll. A. R. Gillogly, det. Alice Assmar, 1 ♂ (TAMUIC-ENTO X0155026).

Sisyranikkoana (Navás)

(Figures 7a-b)

Figure 7
Sisyra nikkoana (Navás). a) Male habitus, in lateral view; b) male genitalia in lateral view.

Nopia nikkoanaNavás (1910): 398 (description) – Banks, 1913: 218 (checklist) – Nakahara, 1915: 18 (checklist) – Nakahara, 1920: 163 (synonym) – Navás, 1935: 71 (checklist) – Kuwayama, 1962: 339 (checklist) – Zakharenko, 1988: 767 (checklist).

Sisyrella nikkoana– Banks, 1913: 218 (new genus) – Nakahara, 1915: 18 (re-description) – Nakahara, 1920: 163 (synonym); Navás, 1935: 71 (re-description) – Kuwayama, 1962: 339 (checklist).

Sisyra japonica Nakahara, 1914: 493 (description) – Nakahara, 1915: 99 (record) – Nakahara, 1920: 163 (synonym) – Navás, 1935: 73 (checklist) – Baba, 1955: 12 (checklist) Nakahara, 1960: 34 (record) – Kuwayama, 1962: 341 (checklist) – Zakharenko, 1988: 767 (checklist).

Sisyra ozememana Nakahara, 1914: 495 (description) – Nakahara, 1915: 99 (record) – Nakahara, 1920: 163 (synonym) Navás, 1935: 53 (re-description) – Kuwayama, 1962: 341 (checklist) – Zakharenko, 1988: 767 (checklist).

Sisyra yamamurae Nakahara, 1914: 496 (description) – Nakahara, 1915: 99 (record) – Nakahara, 1920: 163 (synonym) – Navás, 1935: 54 (re-description) – Kuwayama, 1962: 341 (checklist) – Zakharenko, 1988: 767 (checklist).

Sisyra esakii Nakahara, 1915: 99 (description) – Kuwayama, 1962: 341 (checklist) – Zakharenko, 1988: 767 (checklist).

Sisyra nikkoana– Nakahara, 1920: 163 (synonym, record) – Kuwayama, 1962: 339 (record) – Zakharenko, 1988: 767 (record) – Makarkin, 1990: 38 (record) – Makarkin, 1995: 46 (record) – Makarkin, 2000: 625 (record) – Hayashi, 2005: 387 (re-description) – Yang and Liu, 2023: 17 (re-description), figs 7–9.

Sisyra aurorae Navás, 1933: 13 (description) – Navás, 1935: 55 (re-description) – Yang and Liu, 2023: 20 (synonym).

Sisyrella japonica– Navás, 1935: 73 (re-description) – Kuwayama, 1962: 341 (checklist).

Distribution

China, Japan, Korea, Russia (Yang and Liu, 2023), Vietnam.

Remarks

Yang and Liu (2023) redescribed S. nikkoana Navás and recorded it in several locations in China. The authors also placed Sisyra aurorae Navás as junior synonym of S. nikkoana (Yang and Liu, 2023). This is the first record of this species outside the Palearctic region. We identified this species based on characteristics of the male genitalia and wing venation following its redescription by Yang and Liu (2023).

Material examined

VIETNAM: Quang Tri Prov., 1 mi N Quang Tri, VII-11-1970, at light, Coll. A. R. Gillogly, 1 ♀ 5 ♂ (TAMUIC-ENTO X0156900, X0155153, X0154165, X0152216, X0156912, X0154623); same data, but XII-25-1969, 1 ♂ (TAMUIC-ENTO X0155352); same data, but IV-30-1970, 1 ♀ 1 ♂ (TAMUIC-ENTO X0156930, X0156865); same data, but III-9-1970, (TAMUIC-ENTO X0156955).

Sisyranobilia Yang, Zheng & Liu

(Figures 8a-i)

Figure 8
Sisyra nobilia Yang, Zheng & Liu. Habitus, in lateral view of a a) male and b) female specimens. Forewings and hind wings of a c) male and d) female specimens. Lateral view of genital sclerites of the e) male and f) female specimens. Digital illustration of male genital sclerites g) in lateral view; h) same, but in dorsal and i) ventral view. (Abbreviations: t = tergite, st = sternite, ect = ectoproct, gx = gonocoxite, gs = gonostyli).

Sisyra nobilia Yang, Zheng & Liu, 2024: 133, figs 3, 4 (description).

Distribution

China (Yang et al., 2024), Vietnam.

Remarks

This species was recently described by Yang et al. (2024) and recorded for Guangxi and Hainan, China. Here, we further contribute to the known geographic distribution of S. nobilia by recording it for Vietnam for the first time. Sisyra nobilia can be easily differentiated by its congeners by the shape of the male genital sclerite appendages. Sternite 9 is long, projected distally, with dorsal margin covered by numerous setae, resembling a comb. Gonocoxite 9 is long, thin, and cylindrical, with a thick seta on the distal apex. Additionally, this species has striped coloration in the antennae, and a radial streak is present on the forewing. It is similar to S. arcuata Yang, Zheng & Liu, but the male gonocoxite 9 differs in these two species.

Material examined

VIETNAM: Quang Tri Prov., 1 mi N Quang Tri, VII-06-1970, at light, Coll. A. R. Gillogly, det. Alice Assmar, 1 ♂ (TAMUIC-ENTO X0155507); same data, but III-09-1970, 1 ♂ (TAMUIC-ENTO X0157000); same data, but VII-11-1970, 1 ♀ (TAMUIC-ENTO X0155231).

Sisyrina qiong Yang & Gao

(Figure 9)

Figure 9
Sisyrina qiong Yang & Gao. Habitus of a male specimen in lateral view.

Sisyrina qiong Yang & Gao, 2002: p. 288 – Yang et al., 2018: 19 (checklist) – Yang and Liu, 2021: 560 (re-description), figs 6–8.

Distribution

China (Yang and Liu, 2021), Vietnam.

Remarks

Yang and Liu (2021), in their work about the spongillaflies from the Oriental region, redescribed Sisyrina qiong Yang & Gao. Prior to our work, S. qiong had only been collected in Hainan, China. This study expands the known distributional range of S. qiong to central Vietnam. The identification of this species was based on wing venation and the shape of the male gonocoxite 9.

Material examined

VIETNAM: Quang Tri Prov., 1 mi N Quang Tri, VII-11-1970, at light, Coll. A. R. Gillogly, det. Alice Asmar, 2 ♀ (TAMUIC-ENTO X0155033, X0154993); same data, but VIII-24-1970, 1 ♀ 1 ♂ (TAMUIC-ENTO X0156942, X0154828); same data, but VIII-24-1970, 1 ♂ (TAMUIC-ENTO X0154998).

Sisyrina vietnamica Yang & Liu

Sisyrina vietnamica Yang & Liu, 2021: 559, fig 5.

Distribution

Vietnam (Yang and Liu, 2021).

Remarks

Yang and Liu (2021) recently described this species based on a single female holotype, which is deposited at CAU. The male of this species remains unknown.

Identifying adults Osmyloidea known from Vietnam

Key to extant Osmyloidea families (modified from Winterton et al. (2019)).

• - Wings medium to large (> 10 mm), crossveins often numerous; FW veins CuA, CuP and A1 strongly pectinately branched; FW with more than four RP branches ……… Osmylidae

• - Wings relatively small (< 10 mm), crossveins largely absent or few in number; FW veins CuA, CuP and A1 weakly pectinately to dichotomously branched; FW with four RP branches ……… Sisyridae

Osmylidae

A key to osmylid genera can be found in Winterton et al. (2019). Currently, only one species of Gryposmylus and one of Thaumatosmylus occur in Vietnam. The following characters can be used to differentiate species of Osmylus and Spilosmylus recorded in Vietnam:

Osmylus Latreille

• - Forewing with several brown marks throughout the wing; male dorsal processus (dp) thin and small, nearly two times smaller than tergite 9; Male ectoproct subquadrate ………………… O. angustimarginatusXu et al. (2016)

• - Forewing hyaline with scattered small dots near the posterior margin; male dorsal processus (dp) large and long, nearly the same size as tergite 9; Male ectoproct subtriangular …………………. O. hagiangensis sp. nov.

Spilosmylus Kolbe

• - Forewing with a transversal ribbon-like streak near the middle of the wing; Hind wing with posterior margin hyaline……… S. inthanonensis New, 1991

• - Forewing without a transversal ribbon-like streak; Hind wing with posterior margin darkened………S. inclytus (Navás, 1917)

Sisyridae

Key to the genera of Sisyridae of Vietnam (modified from Monserrat (1981)).

• - Outer gradate series of crossveins present in the apical third of forewing and hind wing…. ……………………………………………………………………………….. Sisyrina Banks. (Key to all Sisyrina species can be found in Yang and Liu (2021))

• - Outer gradate series of crossveins absent in the apical third of forewing and hind wing ……………………………………………………………………………… Sisyra Burmeister. (Key to Sisyra species from Asia can be found in Yang and Liu (2023))

Following Yang and Liu’s (2023) identification key, S. nobilia keys out in the third step, together with S. bowlesi Yang & Liu. The following traits can be used to differentiate these two species:

• - Scape, pedicel and antennal basal seven flagellomeres yellow; conspicuous radiomedial streak absent in forewing; male sternite 9 not projected distally …………………………………………….. S. bowlesi Yang & Liu

• - Scape, pedicel and antennal basal twelve flagellomeres yellow; conspicuous radiomedial streak present in forewing; male sternite 9 projected distally (Figure 7e)………. S. nobilia Yang, Zheng & Liu

Discussion

The known diversity of Osmyloidea in the Oriental region has now increased to 89 species. The number of known Nevrorthidae in this region remains the same, totaling four species of Nipponeurorthus Nakahara: N. fasciatus Nakahara (China, Taiwan), N. multilineatus Nakahara (China, Taiwan), N. damingsthanicus Liu, H. Aspöck & U. Aspöck (China, Guangxi) and N. furcatus Liu, H. Aspöck & U. Aspöck (China, Yunnan) (Aspöck et al., 2017).

Previously, 14 species of Sisyra Burmeister were recorded in this region, including recent discoveries: S. mononoke Szőke recorded in India, S. arcuata recorded in Laos, S. stenoloma Yang, Zheng and Liu and S. nobilia recorded in China, as well as S. hainana and S. nervata as junior synonyms of S. indica (Szőke, 2023; Yang and Liu, 2023; Yang et al., 2024). We increased the known number of species of Sisyra in the Oriental region to 15 species by adding one new record, S. nikkoana. In Vietnam, only five species in four genera of Osmyloidea were previously known to occur. Our research significantly increases this diversity, totaling 12 species in six different genera (Figure 10).

Figure 10
Distribution map showing current known geographic distribution of Osmyloidea species in Vietnam.

Despite recent efforts to minimize taxonomic and distributional knowledge gaps for Osmyloidea (Aspöck et al., 2017; Winterton et al., 2019; Yang and Liu, 2021, 2023; Szőke, 2023, 2024; Yang et al., 2024; Li et al., 2025), much work remains to be done. For future steps, we recommend further addressing Linnean and Wallacean shortfalls, as this is essential for answering future questions related to the evolutionary history of the group. We cannot emphasize enough the importance of investing in taxonomic studies to better comprehend Osmyloidea diversity.

Supplementary material

Supplementary material accompanies this paper.

Table S1

Raw MEGA results in CSV (MEGA-result2.csv) showing pairwise genetic distance among COX1 DNA barcodes calculated using Kimura 2-parameter model.

Figure S1

ML phylogenetic tree in TIFF (COX1_aligned-thaumatosmylus.tif) reconstructed using seven COX1 sequences of Thaumatosmylus.

COX1_aligned.fasta

Alignment file including all 11 COX1 sequences used for phylogenetic inference (ingroup and outgroup).

COX1_aligned.fasta.log

Text file containing information of the maximum likelihood statistical analysis.

COX1_aligned.fasta.treefile

Raw tree file with the resulting topology estimated under maximum likelihood framework using IQTREE.

This material is available as part of the online article from https://doi.org/10.1590/1806-9665-RBENT-2025-0108

Acknowledgements

We would like to thank Agnieszka Pierwola and David Grimaldi (AMNH), John Oswald (TAMUIC), and Shaun Winterton (CDFA) for loaning us the specimens examined in this work. Additionally, we are grateful for the funding support received from the Fonds de Recherche du Quebec – Nature et Technologies (FRQNT) for the doctoral scholarship to AA (B2X 2023-2024, file 330130 https://doi.org/10.69777/330130). RJPM thanks the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the grant: Proc.: 441787/2023-1. We also thank to the Entomological Society of Canada (ESC) for a Graduate Research Travel Scholarship, which allowed AA to visit entomological museums to acquire specimens. JPG acknowledges funding from NSERC-DG (NSERC RGPIN-2021-02656). We thank Dr. Yuchen Zheng and another anonymous reviewer for the very important comments.

Data statement

The entire dataset supporting the results of this study was published in the article and in the “Supplementary Materials” section.

• Funding
  This work was supported by the Fonds de Recherche du Quebec – Nature et Technologies (FRQNT B2X 2023-2024, file 330130); the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq Proc.: 402785/2021-5 and Proc.: 441787/2023-1); and the NSERC-DG (NSERC RGPIN-2021-02656).

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Edited by

• Associate Editor:
  Tiago Kütter Krolow

Publication Dates

• Publication in this collection
  29 June 2026
• Date of issue
  2026

History

• Received
  06 Nov 2025
• Accepted
  29 Apr 2026