﻿Seven new species and four new records of Psychomyiidae (Insecta, Trichoptera) from China

﻿Abstract Seven new species of the family Psychomyiidae Walker, 1852 are described and illustrated from China; they are Psychomyiashunisp. nov., Ps.mangshanensissp. nov., Ps.capricornissp. nov., Lypesagittalissp. nov., Paduniellafasciariasp. nov., Pa.sanyaensissp. nov., and Tinodesaviformissp. nov. The genus Lype is reported for the first time from mainland China. In addition, four psychomyiids are found to be new to the Chinese caddis fauna: Psychomyiaindra Malicky & Chantaramongkol, 1993; Paduniellaandamanensis Malicky, 1979; Pa.dendrobia Malicky & Chantaramongkol, 1993; and Tinodesgapbona Johanson & Oláh, 2008. Moreover, Psychomyiapolyacantha Li, Qiu & Morse, 2021 is reviewed and synonymized with Psychomyiaimamiah Malicky, 2020.

In this study, we describe three new species of the genus Psychomyia (Psychomyia shuni sp.nov., Ps. mangshanensis sp.nov., and Ps.capricornis sp.nov.), two of Paduniella (Paduniella fasciaria sp.nov.and Pa.sanyaensis sp.nov.), and one each in Lype (Lype sagittalis sp.nov.) and Tinodes (Tinodes aviformis sp.nov).The genus Lype is newly reported from mainland China.In addition, the present study provides four new geographical records for China

Morphological study
Male abdomens used for illustrations were cleared with 10% NaOH solution and heated to 90 °C for 10 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.Genitalia 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 v. 19.0 (Adobe Inc 2018) software and inked digitally with a Wacom CTL-671 tablet to produce final illustrations.Then each abdomen was stored in a microvial together with the remainder of the specimen in 95% ethanol.All specimens are deposited in the Insect Collection, Nanjing Agricultural University, Nanjing, Jiangsu Province, P.R.China (NJAU).

Terminology
The terminology of the male genitalia for the genus Psychomyia mainly follows Schmid (1998), with the adoption of the term "mesal ridge" from Qiu and Morse (2021) to refer to the piece along the inner side of each superior appendage; the term "basoventral process" refers to a single or paired protruding structure at the base of the phallotheca, and "basal process" refers to the paired protruding structure at the base of the superior appendages.Terminology for the genus Lype largely follows Schmid (1998), with the adoption of the term "subapical projection of aedeagus" from Arefina (2002) to refer to the dorsal process near the distal end of the aedeagus.Terminology for the genus Paduniella follows Li and Morse (1997), and that for the genus Tinodes follows Peng et al. (2022).To ensure coherency, "superior appendages" are used to refer to the "preanal appendages", and "coxopodite" and "harpago" are used to refer to the first and second segments of the inferior appendages, respectively.Diagnosis.This species is unique among Psychomyia in that the coxopodites and the harpagones are completely fused and together form an S or Z shape in ventral view.
Etymology.Latin noun in genitive singular.The new species is named after Shun, a leader of tribal alliances in ancient China, who is considered an important founder of the Chinese civilization.Moreover, the holotype and one of the paratypes were collected at Mt Shun-huang, the mountain named after Shun.
Distribution.China (Hunan).(Qiu and Morse 2021).However, P. mangshanensis sp.nov.can be easily distinguished by the following characteristics: (1) each superior appendage has a small triangular protrusion in the middle of the ventral margin in lateral view, which is missing in P. cuspidata; (2) the basal process of each superior appendage is unbranched and with thick spine apically, whereas each superior appendage is two-branched, each branch has a thick spine apically in P. cuspidata; and (3) the phallotheca is slightly wavy at midlength in lateral view but with an obtuse angle in P. cuspidata.Description.Male.Length of each forewing 2.8-3.1 mm (n = 10), holotype forewing 3.0 mm.Specimens in alcohol with compound eyes black; body dark brown dorsally and light brown ventrally.Forewings each with forks II-V present, and hind wings each with forks II and V present.Genitalia.Sternum IX subrectangular in ventral, dorsal, and lateral views (Fig. 2A, C).Tergite IX short and triangular in dorsal and lateral views (Fig. 2B, C).Division between tergite IX and segment X indiscernible in dorsal view (Fig. 2B) but distinguished by membrane in lateral view (Fig. 2C).Segment X parallel-sided, same width as apex of tergite IX, apically truncate in dorsal view, digitate in lateral view (Fig. 2C); with several long thick apical setae in dorsal and lateral views (Fig. 2B, C).Superior appendages well developed in lateral view, each tapering from base towards apex, divided into one narrow dorsomesal branch and one broad ventrolateral branch subapically; dorsomesal branch sclerotized and bare, acute in lateral view, ventrolateral branch setose about twice as wide as upper branch in lateral view (Fig. 2C); in ventral and dorsal views (Fig. 2A, B), dorsomesal branches of superior appendages angled mesad, ventrolateral branches curved slightly mesad; paired basal processes of superior appendages tubular, and slender, each with thick spine apically; in lateral view each with base directed dorsad, then recurved posteroventrad and evenly curved caudad (Fig. 2C); in ventral and dorsal views each slightly curving outwards (Fig. 2A, B).Phallobase slender, lanceolate in lateral view (Fig. 2C).Phallotheca tubular, with base produced caudad in lateral view (Fig. 2C), main portion sinuate, with apex hooked dorsad; phallotheca stick-like in ventral view (Fig. 2A), basoventral process plate-shaped, three times wider than main portion of phallotheca.Inferior appendages extending posterolaterad; coxopodites subtriangular, with their bases fused in ventral view (Fig. 2A); subrectangular, about 3 times as long as tall with middle of dorsal margin concave in lateral view (Fig. 2C); harpagones setose, arising from apices of coxopodites, fingerlike (Fig. 2A, C).

Psychomyia mangshanensis
Etymology.Latin feminine adjective mangshanensis, referring to the type locality.Distribution.China (Hunan).Diagnosis.This species is similar to Psychomyia shuni sp.nov.from China in having the superior appendages with dense spines mesally and in having well-developed subapical processes on the phallic apparatus.However, P. capricornis sp.nov.can be easily distinguished by the long, slender processes arising from the bases of the coxopodites, which are absent in P. shuni.
Etymology.The Latin feminine adjective capricornis means "goat's horn", referring to the shape of the pair of subapical processes on the phallic apparatus.
Distribution.China (Fujian).Diagnosis.This species is similar to Lype lubaretsi Arefina, 2005 from Russia.However, the new species can be easily distinguished by the following characteristics: (1) longitudinally, sternum IX of L. sagittalis sp.nov. is subtriangular in lateral view, rather than subrectangular in L. lubaretsi; (2) each of the coxopodites of the new species is subcircular in lateral view, but elliptical in L. lubaretsi; (3) the fused coxopodites in ventral view have a narrow mesal notch in the new species, rather than with a wide mesal notch in L. lubaretsi; (4) the aedeagus is sagittal, with its apex truncate in dorsal view and pipe-shaped in lateral view in L. sagittalis sp.nov., but nearly triangular with apex broad in ventral view, triangular with a sharp apex in lateral view in L. lubaretsi.
Etymology.The Latin feminine adjective sagittalis, meaning "arrow-shaped", and refers to the shape of the aedeagus in dorsal and ventral views.
Distribution.China (Hunan).Diagnosis.This species is similar to Paduniella sampati Malicky & Chantaramongkol, 1993 from Thailand in having the superior appendages furcated at their bases in dorsal view and in the shape of the phallic apparatus in the lateral view.However, P. fasciaria sp.nov.can be easily distinguished by the possession of a slender median process.

Paduniella fasciaria
Description.Male.Length of each forewing 2.6-3.0 mm (n = 10), holotype 2.9 mm.Specimens with compound eyes black, antennae approximately of same length as forewings; body brown; head, bases of antennae, thorax covered with brown, short hair; wings mostly covered with brown, short hair; each forewing with transversal white band at middle (Fig. 5D, E).Genitalia.Sternum IX subrectangular with anterodorsal angle produced into subrectangular process in lateral view (Fig. 5C); and transversely elongate-rectangular with anterior margin having deep U-shaped incision in ventral view (Fig. 5A).Tergum IX membranous, with base fused with superior appendages, somewhat clavate in lateral view (Fig. 5C), directed posterodorsad; transversely subrectangular in dorsal view, with anterior margin slightly convex and posterior margin undulated (Fig. 5B).Superior appendages large, forming a parallelogram shape in lateral view (Fig. 5C); in dorsal view (Fig. 5B) each with basal portion furcate, basolateral lobe slightly longer than inner one, tapering distally with apex curved mesad and crossing apex of opposing superior appendage, setose subapically.Sclerotized strips slightly clavate in lateral view (Fig. 5C) and somewhat V-shaped in dorsal view (Fig. 5B).Median process mostly slender with sharp apex, insertion between sclerotized strips broad in dorsal view (Fig. 5B).Inferior appendages each with basal half broad, then abruptly narrowed at mid length and tapered towards apex in lateral view (Fig. 5C); basal half broad, then abruptly narrowed, with apex slightly enlarged and curved mesad in ventral view (Fig. 5A), mesal branch lamellar, setose, arising from middle part of inner surface (Fig. 5B, C).Phallobase well developed, basally clavate in lateral view (Fig. 5C); with upper margin having deep incision in ventral and lateral views (Fig. 5A, C), tapering from base to apex, phallicata tubular, curved slightly upwards in lateral view (Fig. 5C); about same length as phallobase (Fig. 5A, C).
Etymology.The Latin feminine adjective fasciaria, meaning "banded," refers to the band of white hairs across each wing in dorsal view.
Etymology.Latin feminine adjective sanyaensis, the name referring to the location of the type locality in Sanya City.
Distribution.China (Hainan).Diagnosis.This species is similar to Tinodes igok Kimmins, 1955 from Malaysia in the composition and morphology of the male genitalia, but it can be distinguished by: (1) two unpaired inner branches of the phallic sheath processes, of which one is short and straight and the other one curved, whereas both branches are curved in T. igok; (2) the mesal digitate process of each coxopodite is shorter than the harpago, curved, and with a sharp apex in ventral view, but is undulated and almost the same length as the harpago in T. igok; and (3) the phallic guide is divided into two branches with the dorsal one extending backward beyond the coxopodites in lateral view, rather than having only one uncinate branch that is not longer than the coxopodites in T. igok.

Tinodes aviformis
Description.Male.Length of each forewing 3.1 mm (n = 1).Specimen in alcohol with compound eyes black, antennae yellowish white; thorax and legs brown, wings light brown without any distinctive markings; abdomen dark brown dorsally, pale yellow ventrally.Genitalia.Sternum IX transversely subrectangular in ventral view (Fig. 7A); subtriangular in lateral view (Fig. 7C).Tergum IX covered with microchaetae, with anterior margin slightly sinuate and middle of posterior margin produced posterad in dorsal view (Fig. 7B); broader and subapically angled slightly caudad in lateral view (Fig. 7C).Segment X membranous, closely fused with tergum IX, its posterior margin nearly truncate in dorsal view (Fig. 7B); subrhomboid in lateral view (Fig. 7C).Superior appendages each with distal half setose, clavate, and with apex rounded in lateral view (Fig. 7C); parallel-sided in dorsal view (Fig. 7B).Phallic sheath process consisting of paired lateral branches and unpaired inner branches; paired lateral branches compressed and spoon-like, semicircular, and resembling nesting bird in lateral view (Fig. 7C, E), each with several strong spines at middle near ventral edge and distal half setose; fused basally, resembling pair of clam shells in ventral view (Fig. 7D); two unpaired inner branches strongly sclerotized (Fig. 7D, E); one of them straight, one curved; phallus slightly extending beyond tip of phallic sheath process, with distal end membranous in ventral view (Fig. 7D), base and apex swollen in lateral view (Fig. 7E); ejaculatory duct slender, with subapex S-shaped in lateral view (Fig. 7C).Phallic guide with subapex wider and then divided into two branches in lateral view (Fig. 7C), dorsal branch slender, curved downwards distally and gradually narrowed to sharp apex; ventral branch short.Coxopodites elliptical in lateral view (Fig. 7C); fused with each other basally in ventral view (Fig. 7A), each with posterodorsal angle digitate and posterior margin having tiny submesal digitate process; harpago setose, small, and simple (Fig. 7A, C).
Etymology.The Latin masculine adjective aviformis, meaning "bird-shaped," and refers to the shape of the pair of lobes of the lateral phallic sheath process in lateral view.

Psychomyia indra Malicky & Chantaramongkol, 1993
Psychomyia indra Malicky & Chantaramongkol, 1993: 1162 (type  Remarks.The specimens that we examined included topotypes that were collected at the same time as the type specimen of this species (Qiu and Morse 2021).We found that the harpagones each had four apical processes, which is rare in Psychomyia, occurring only in P. polyacantha and P. imamiah.By comparing these specimens and published descriptions and figures of the male genitalia of these two species, we believe that the morphological characteristics of the two species overlap.The type locality of P. polyacantha is in the basin of the Fujiang River, and the Fujiang River flows into the Jialing River, and the paratypes of P. polyacantha and the holotype of P. imamiah are from the basin of the Dadu River, which merges into the Minjiang River.Both the Jialing and Minjiang rivers are tributaries of the Yangtze, and their upstream habitats are similar and geographically adjacent.In summary, we identify P. polyacantha as a synonym of P. imamiah.

Discussion
The Psychomyiidae is a moderately sized family of caddisflies (Holzenthal et al. 2007).However, the number of species has increased rapidly in recent years as new species are discovered and named; in 2008 there were about 400 species (de Moor and Ivanov 2008), in 2019 there were about 600 species (Morse et al. 2019), and now there are more than 650 species (including this study).Though the members of this family are widespread over nearly all the world (Holzenthal et al. 2007), the family's species diversity is uneven across zoogeographical regions.Psychomyiids are found mostly in the Oriental and Palaearctic regions, with the combined total number of species in those two regions constituting more than 90% of the world's species for this family.Of the two regions, the Oriental Region is the only one with all eight psychomyiid genera present, and it has about 65% of the world's recorded species, significantly higher than the number of Palaearctic species, which constitute about 25% of the species among six genera (Psychomyia, Tinodes, Paduniella, Metalype, Eoneureclipsis, and Lype).Additionally, the Afrotropical region has three genera (Tinodes, Paduniella, and Lype) and 6.9% of the world's species, the Nearctic region has four genera (Psychomyia, Tinodes, Paduniella, and Lype) and 2.8% of the world's species, and the Australian region has only one genus (Tinodes) and 1.5% of the world's species.China covers parts of the Oriental and Palaearctic regions and has more than 111 species in six genera (including those in this study), constituting about 17% of psychomyiid species, of which six genera (Psychomyia, Tinodes, Paduniella, Metalype, Eoneureclipsis, and Lype) and 100 species are recorded in China's Oriental region, and four genera (Psychomyia, Tinodes, Paduniella, and Metalype) and 18 species are recorded in its Palaearctic region.
Most of the species in our study were collected from low latitudes, with a few from mid-latitudes (Table 1).Collection sites are near the source of streams or near a reservoir.Headwater streams had high forest cover and were small and cool, which is consistent with most psychomyiid larvae living in cool running water (Fig. 8A-D).While Tinodes aviformis sp.nov.was collected near a reservoir, similar to the habitats of some Tinodes, which have been reported to live in isolated stream pools in western North America and in lake littorals in Europe (Flint 1964;Wiggins 1996).
The male genitalia of the family vary among psychomyiid genera.In some genera, such as Paduniella and Lype, the structures of the male genitalia are simple enough to use common terms for describing them; however, in other genera, for example, in the genus Tinodes, extra structures are present in the genitalia, resulting in the different understanding on their homology, and accordingly, the terminology for the structures has varied among authors.The situation hinders phylogenetic study of the family based on morphology.We sincerely hope that, with phylogenetic studies using DNA sequences, the homology of these extra structures will become more generally understood and consensus for these terms will one day be reached, making interpretation of the evolution of the morphology and functional traits of these interesting and ecologically important animals more reliable.

Table 1 .
Detailed information on Psychomyiidae specimens collected in China.Specific dates are for specimens collected by light trap, while a date range represents specimens collected by Malaise trap over several days.
* indicates the lack of original records for geographic coordinates; the data are based on location information.

Peng & Sun, sp. nov.
Diagnosis.This species is similar to Psychomyia cuspidata Li, Qiu & Morse, 2021 from China