Zootaxa 4121 (4): 485–494
http://www.mapress.com/j/zt/
Copyright © 2016 Magnolia Press

ArticleISSN 1175-5326 (print edition)ZOOTAXAISSN 1175-5334 (online edition)

http://doi.org/10.11646/zootaxa.4121.4.8
http://zoobank.org/urn:lsid:zoobank.org:pub:F756E510-30F8-4250-87A3-78474AFE65AA

Larvae and females of two Stenopsyche species in Taiwan with redescription of
the male of S. formosana (Insecta: Trichoptera)
TAKAO NOZAKI1,6, RIE SAITO2, NOBORU NISHIMURA3, LI-PENG, HSU4 & KOJI TOJO5
1

3-16-15, Midorigaoka, Ninomiya-machi, Naka-gun, Kanagawa, 259-0132 Japan
Division of Mountain and Environmental Science, Interdisciplinary Graduate School of Science and Technology, Shinshu University,
Asahi 3-1-1, Matsumoto, Nagano 390-8621, Japan3
1841, Sekinomiya, Yabu-shi, Hyogo, 667-1105, Japan
4
Department of Bioresource, Dayeh University, No. 168, Univ. Rd., Datsun, Changhua, 515, Taiwan
5Department of Biology, Faculty of Science, Shinshu University, Matsumoto, Nagano, Japan
6
Corresponding author. E-mail: takao.nozaki@nifty.com
2

Abstract
Larvae and females of two Stenopsyche species, S. formosana Kobayashi and S. drakon Weaver, are described for the firsttime. They were associated with the adult male through mitochondrial DNA sequence comparisons. The male of S. formosana, which has been recorded as Stenopsyche banksi Mosely from Taiwan, is redescribed. The males of S. formosana
and S. banksi can be distinguished from each other by the shape of segments IX and X, and endothecal spines. In larvae,
S. formosana and S. drakon are distinguishable from each other and from other known species by the combination of different dark marking patterns and shapes of no. 18 setae of the head, and by the shapes of forecoxae. The larval morphologyof S. formosana and S. drakon suggests that the phylogenetic relationships based mainly on male genitalia in the genus
Stenopsyche should be reconsidered.
Key words: Stenopsychidae, Stenospyche formosana, Stenospyche drakon, mitochondrial DNA, association

Introduction
The genus Stenopsyche McLachlan is a genus composed of more than 90 species distributed in the East Palaearctic,
Oriental and Afrotropical regions (Morse 2016). Larvae of this genus are a distinct element of the fauna of Asianrivers and streams (Dudgeon 1999), however, the larvae of most species are not yet known (Xu et al. 2015).
Knowledge about females of this genus is also poor (Xu et al. 2015). In Taiwan, Yang et al. (2005) listed seven
Stenospyche species: S. banksi Mosely 1942, S. drakon Weaver 1987, S. formosana Kobayashi 1987, S.
pallidipennis Martynov 1926, S. pjasetzkyi Martynov 1914, S. sauteri Ulmer 1907, and S. taiwanensis Weaver
1987. Furthermore, Malicky (2014) recorded three additional species: S. arvadit Malicky 2008, S. remihel Malicky
2014 and S. schmidi Weaver 1987. However, larvae and females of these species have not yet been described.
In 2015, adults and larvae of various aquatic insects were collected from rivers in Nantou County in Taiwan(Nishimura et al. 2015). In the collection, we found males of two Stenopsyche species, S. formosana and S. drakon,
and also two discrete morphotypes of larvae and females, respectively. Each morphotype of these larvae andfemales was associated through DNA sequence comparisons with the males. In this paper, we describe larvae andfemales of these two Stenopsyche species, and redescribe the male of S. formosana which has been recorded as acontinental species S. banksi (Mosely 1942; Biodiversity Research Center, Academia Sinica 2015).

Materials and methods
Specimens. DNA sequences of eight specimens of Stenopsyche caddisflies from two localities in Taiwan were

Accepted by J. Morse: 11 Apr. 2016; published: 10 Jun. 2016

485

analyzed for the DNA sequences at the mitochondrial COI region (636-bp.). We analyzed males of Stenopsyche
formosana (specimen No. 1: Table 1) and Stenopsyche drakon (specimen Nos. 2–3: Table 1) identified by theirmorphological characteristics as the reference specimens. We also analyzed two females (specimen Nos. 4–5:
Table 1) and three larvae (specimen Nos. 6–8: Table 1) of Stenopsyche caddisflies which have not been associatedwith their adult males. Some additional specimens were used for morphological study.
TABLE 1. Specimen information for Stenopsyche spp. used for molecular phylogenetic analyses.
SampleNo.

Locality(Ren'ai Xiang, Nantou County)

LatitudeLongitude

Altitude
(m)

Stage

GenBank
Accession No.

1

Beigang River, Huisun Forest Area

24°4.992'

121°0.651'

490

Male

LC1096742

Beigang River, Huisun Forest Area

24°4.992'

121°0.651'

490

Male

LC109678

3

Meiyuan River

24°5.09'

120°57.22'

504

Male

LC109679

4

Meiyuan River

24°5.09'

120°57.22'

504

Female

LC1096755

Beigang River, Huisun Forest Area

24°4.992'

121°0.651'

490

Female

LC109680

6

Meiyuan River

24°5.09'

120°57.22'

504

Larva

LC109676

7

Meiyuan River

24°5.09'

120°57.22'

504

Larva

LC109677

8

Beigang River, Huisun Forest Area

24°4.992'

121°0.651'

490

Larva

LC109681

Morphological study. Male and female genitalia were figured after being cleared in a 10% solution of KOH.
Photographs of larval characters were taken using Nikon Coolpix P340 through Olympus SZX 12
stereomicroscope, and combined by the image stacking software Combine ZP. Larval characters described are
based on final instar larvae. Morphological terms mostly follow Xu et al. (2015). Setal nomenclature is based onthat of Williams & Wiggins (1981) and Nozaki & Shimura (2015). Depositories of specimens are abbreviated asfollows: Natural History Museum and Institute, Chiba, Japan (CBM); Shinshu University, Matsumoto, Japan (SU)and T. Nozaki, Ninomiya, Kanagawa, Japan (TN).
DNA extraction, amplification, sequencing and alignment. Total genomic DNA was extracted from ethanol
preserved tissues of specimens and purified using the DNeasy Blood and Tissue Kit (QIAGEN, Hilden), according
to the manufacturers’ instructions. Total genomic DNA was used to amplify DNA fragments [the mitochondrial
DNA (mtDNA) cytochrome c oxidase submit I (COI) by polymerase chain reaction (PCR) with sets of primers[LCO1490: 5'-GGTCAACAAATCATAAAGATATTGG-3', and HCO2198: 5'-TAAACTTCAGGGTGACCAAAAAATCA-3' (Folmer et al. 1994)]. The PCR protocol was: 94°C for 1 min; 35× (94°C for 1 min, 45°C for 2 min,
72°C for 1 min); 72°C for 7 min. PCR products were purified using the illustra ExoProStar (GE Healthcare,
Buckinghamshire). Purified DNA fragments were sequenced directly by an automated method using a BigDye
Terminator v1.1 Cycle Sequencing Kit (Applied Biosystems, California) on an automated DNA Sequencer (ABI
3130 or 3130xl DNA Analyzer; Perkin Elmer/Applied Biosystems, California). All of the sequence data have been
submitted to the DNA data bank of Japan (DDBJ database), and the accession numbers are given in Table 1. As
outgroups, we have utilized the sequence data of the two philopotamid caddisflies: Chimarra marginata (GenBank
accession No. FM998346) and Chimarra social (GU114726). In addition, we have also used the sequence data of
East Asian Stenopsyche species listed below, which have been registered in GenBnak: S. marmorata, S. schmidi
and S. sauteri (GenBank accession No. LC109682–109695).
Sequence alignment and editing were performed for each gene separately using MEGA version 5.02 (Tamura
et al. 2011) and CLC Workbench software (CLC bio, Aarhus). All sequence data were aligned automatically with
Clustal W (Thompson et al. 1994) and MEGA version 5.02. The alignments were determined for unique
haplotypes using the software DnaSP v4.0 (Rozas et al. 2003) prior to subsequent analysis.
Phylogenetic analysis. Phylogenetic analyses were performed by neighbor joining (NJ: Saitou & Nei 1987)
and maximum likelihood (ML: Felsenstein 1981) using MEGA version 5.02 with 1,000 bootstrap replications.
Prior to the ML phylogenetic estimations, Best-fit ML models were chosen using MEGA version 5.02; Best-fit ML
models were chosen as follows: GTR + Γ+ I. In addition, phylogenetic analyses were performed using Bayesian
approaches in MrBayes v3.1 (Huelsenbeck & Ronquist 2001). The program Kakusan4 (Tanabe 2007) was used to
select appropriate models based on Schwarz’s Bayesian Information (BIC) and the selected sites model: HKY85 +486 · Zootaxa 4121 (4) © 2016 Magnolia Press

NOZAKI ET AL.

Γ. Bayesian MCMC simulations were run for 100 million generations, sampling every 2,000 generations. The
resulting tree was visualized in FigTree v1.3.1 (Rambaut 2009).

Results
Phylogenetic analysis. Based on our genetic analysis of S. formosana and S. drakon, it was clearly revealed thatthey form two separate monophyletic clades (Clade I vs II: Fig. 1). This trend was supported as robust results ofstrong reliability in all analyzed methods (i.e., ML, NJ, and Bayesian analyses: Fig. 1). The Bayesian posterior
probability support, NJ bootstrap support, and ML bootstrap support of each clade are 1.00/100/100, respectively(Fig. 1). From the positioning of the reference specimens, clade I corresponds to S. formosana, and clade II
corresponds to S. drakon.

FIGURE 1. Bayesian phylogenetic relationships based on the sequence data of the mitochondrial DNA COI region (636-bp.)of Stenopsyche caddisflies. The numbers at major nodes indicate the Bayesian posterior probabilities, NJ, and ML bootstrap
support, respectively. Numbers of the OTUs of clades I and II indicated specimen numbers in Table 1.

Species description
Stenopsyche formosana Kobayashi 1987
(Figs. 2, 3)
Stenopsyche formosana Kobayashi 1987, 37–38, 43, male, Taiwan; Malicky 2014, 1624, Taiwan.
Stenopsyche banksi: Mosely 1942 (in part), 359, Taiwan; Nishimura et al. 2015, 78, Taiwan.

Adult. Forewings of male each 21–22 mm long (n=2) and of female 19–23 (n=3), brown with many pale spots,
irregularly reticulated; hind wings white or slightly pale brown, subhyaline. Antennae more than 1.5 times as longas forewings in male, slightly longer than forewings in female. Legs yellowish brown, with dark markings on foreand middle tibiae and on 1st to 2nd (often 3rd) segments of middle tarsi. Tibial spurs 3, 4, 4.
Male genitalia. Segment IX (IX) short dorsally, long ventrally, anteroventral margins round in lateral aspect,
posterolateral margins each with long triangular projection in middle. Segment X (X) forming isosceles trapezoid

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in dorsal aspect, semimembranous in apical half; apex slightly concave, with pair of stout lateral lobes. Pair ofintermediate appendages (int) dorsolaterally near base, each with apex round, directed posterad. Preanal
appendages (pr) long, more than 3 times as long as segment X. Inferior appendages (inf) long triangular in ventral
aspect, each with round apex; superior arms (sup) about half as long as preanal appendages, evenly curved
outward, apically semicircular in dorsal aspect. Phallus long, endotheca with numerous stout spines along its entire
length.
Female genitalia. Segment IX (IX) rectangular in lateral aspect, mostly sclerotized but membranous
posterodorsally and bearing 1 to 3 setae on each anterodorsal side, with long apodemal extension on eachanteroventral corner. Segment X (X) (segment XI of Schmid 1969) membranous dorsally, with pair of setoselongitudinal sclerites dorsolaterally; ventral setose area weakly sclerotized, but membranous apicomesally; withpair of setose distal lobes, each bearing cercus. Vulvar scale (vs) large, each side sclerotized, thumb-like in lateral
aspect, with long stout setae apicolaterally; posterolateral margins round in ventral aspect, semimembranous
mesally, projecting slightly beyond lateral corners. Vaginal chamber (vc) sclerotized, expanded anterad and dorsadin lateral aspect; ventral part wide, vase-shaped in ventral view.
Final instar larva. General appearance as in genus. Length up to 35 mm. Head 4.4–5.0 mm long (n=5);
anterior part about 1.6 mm wide (eye level), posterior widest part about 1.8 mm; ground color yellowish to reddish
brown, with many small dark spots; frontoclypeus with dark brown longitudinal band on mid-line, crossing three
black transverse stripes; primary setae 1, 4, 10, 11, 13 not dark pigmented; setae 8, 15, 18 located ventrally; eachseta 18 (18) bifurcated, about 0.2 mm long, located near postoccipital ridge, each surrounded by rugged
semicircular area. Pronotum 2.3–2.5 mm long, about 2.3 mm wide; ground color yellowish brown, with many dark
spots except for anterodorsal area, posterior margin black. Forecoxae pentagonal in mesal view; each apicodorsal
process (ap) finger-like, straight, directed anterad, with short semitransparent spine-like seta apically; eachbasodorsal process (bp) elongate triangular, slightly longer than apicodorsal process, with long mesal seta in
middle; each central process (cp) on mesal face small, with short stout seta apicodorsally.

FIGURE 2. Male genitalia of Stenopsyche formosana. A, left, lateral; B, phallus, left lateral; C, dorsal; D, ventral.
Abbreviations: IX = segment IX, X = segment X, inf = inferior appendage, int =intermediate appendage, pr = preanal
appendage, sup = superior arm of an inferior appendage.

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FIGURE 3. Female and larva of Stenopsyche formosana. A–C, female genitalia: A, left lateral; B, ventral; C, dorsal. D–G,
larva: D, head, dorsal; E, head, ventral; F, prothorax, dorsal; G, mesal surface of right forecoxa, left lateral. Abbrebiations: 18 =primary seta no. 18 of larval head, VIIIs = sternum VIII, VIIIt = tergum VIII, IX = segment IX, X = segment X, ap =apicodorsal process of a forecoxa, bp = basodorsal process of a forecoxa, cp = central process of mesal face of a forecoxa, vs =vulvar scale; vc = vaginal chamber.

Specimens examined. Holotype male (with handwritten label “7143 ♂”), [TAIWAN] Nangzunkei, wulia,
16.iv.1982, S. Uchida (CBM). [TAIWAN] Beigang River, Huisun Forest Area, Ren’ai Xiang, Nantou, 24°4.992’N,
121°0.651’E, alt. 490 m, 3.iii.2015, N. Nishimura, 1 male (SU); same locality, 3.iii.2015, L.P. Hsu, 1 male (TN);
Huisun Forest Area, Ren'ai Xiang, Nantou, 24°5.628’N, 121°1.843’E, alt. 910 m, 30.iv.2010, K. Tojo, 2 larvae(SU); Meiyuan River, Ren'ai Xiang, Nantou, 24°5.09’N, 120°57.22’E, alt. 504 m, 3.iii.2015, N. Nishimura, 2larvae (SU); same locality, 4.iii.2015, N. Nishimura, 3 females (1 female: SU, 2 females: TN); Sankeng Ecology
Park, Dahan River, Longtan Xiang, Taoyuan, 24°50.169’N, 121°15.329’E, alt. 127 m, 26.xi.2010, K. Tojo, 1 larva(SU).

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Remarks. We examined the holotype of S. formosana deposited in the CBM, and confirmed it to be identicalwith males used in this study. The type locality of this species was recorded as “Nangzunkei, wulia” by Kobayashi(1987). Although we could not understand the meaning of “wulia” but “Nangzunkei” must be a Japanese reading ofa Chinese locality name “Nanshan stream.” The collector of this specimen visited this stream in Ren'ai Xiang,
Nantou County on 16 April 1982 (Uchida personal communication).
On the other hand, Mosely (1942) described S. banksi, a species similar to S. formosana, from Foochow(Fuzhou, P.R. China: the holotype) in China, and also recorded it from Taiwan based on a male specimen
designated as a paratype. Stenopsyche banksi has been recorded from north to central Taiwan (Biodiversity
Research Center, Academia Sinica 2015). For confirming the taxonomic status of S. formosana, we comparedmales of S. formosana with photographs of the holotype and paratype males of S. banksi kindly provided by Dr. B.
Price, The Natural History Museum, London. The paratype male of S. banksi from Taiwan is identical to our malesof S. formosana. However the S. banksi holotype from continental China is slightly different from the paratype.
Segment IX in lateral aspect is longer in S. formosana than that in the holotype of S. banksi (Figs. 2A, 4B); a pair ofintermediate appendages is directed posterad in S. formosana (Fig. 2A) but they are large and strongly curved
ventrad in the holotype of S. banksi (Fig. 4B; Mosely 1942, fig. 51); spines arising from the base of the endotheca
are short in S. formosana (Figs. 2B, 2D), but very long in the S. banksi holotype (Fig. 4B; Mosely 1942, figs. 51,
53). Furthermore, in S. formosana a pair of intermediate appendages of the segment X arise from the dorsolateralsides of segment X (Fig. 2C), but they arise dorsally and close to each other in the S. banksi holotype (Mosely
1942, 358, fig. 52). Although S. formosana may be a subspecies level taxon of S. banksi, we treat them as distinct
species. Further study with molecular data is needed to confirm the taxonomic status of this/these species.
The larva of this species is very similar to those of S. angustata Martynov 1930 (Hoang & Bae 2007, Xu et al.
2015) and S. schmidi Weaver 1987 (Tanida 2003, Nozaki & Shimura 2015), but is distinguishable from them by theshape or length of the apicodorsal processes of the forecoxae. In this species, the apicodorsal process of each
forecoxa is straight and slightly shorter than the basodorsal process, but it is curved ventrad in S. angustata and lessthan half as long as the dorsal process in S. schmidi.

FIGURE 4. Holotype male of Stenopsyche banksi (Photos B. Price). A, body and genitalia (mounted), B, genitalia, left lateral.
Abbreviations: IX = segment IX, int =intermediate appendage. Arrows indicate long spines arising on basal part of endotheca.

Stenopsyche drakon Weaver 1987
(Fig. 5)
Stenopsyche drakon Weaver 1987, 165–167, male, Taiwan; Malicky 2014, 1624, Taiwan; Nishimura et al. 2015, 78, 81,
Taiwan.

Adult. Forewings of male each 19 mm long (n=2), of female 20–22 (n=3), brown with many pale spots, irregularly

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reticulated; hind wings white or slightly pale brown, subhyaline. Antennae about 1.5 times as long as forewings inmale, slightly longer than forewings in female. Legs yellowish brown, middle legs with dark markings on tibiae
and 1st to 2nd segments of tarsi, markings on foretibiae indistinct. Tibial spurs 3, 4, 4.

FIGURE 5. Female and larva of Stenopsyche drakon. A–C, female genitalia: A, left lateral; B, ventral; C, dorsal. D–G, larva:
D, head, dorsal; E, head, ventral; F, prothorax, dorsal; G, mesal surface of right forecoxa, left lateral.

Female genitalia. Segment IX rectangular in lateral aspect, with long apodemal extension on eachanteroventral corner; in dorsal view mostly sclerotized but separated by membrane along dorsal midline,
posterodorsal area membranous, each sclerotized plate with transverse row of setae about 1/3rd distance fromanterior margin, 1 pair of setae near posterior margin. Segment X (segment XI of Schmid 1969) membranous
dorsally, with pair of setose longitudinal sclerites dorsolaterally; ventral setose area weakly sclerotized withapicomesal incision; apex divided into pair of distal lobes, each with cercus. Vulvar scale large; each side
sclerotized, subrectangular in lateral aspect, with long stout setae apicolaterally; posterior margin sinuous in ventral
aspect, membranous mesally. Vaginal chamber sclerotized, projecting anterodorsad in lateral aspect; ventral partpentagonal posteriorly in ventral aspect, tongue-shaped anteriorly in lateral and ventral aspects.

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Final instar larva. Length up to 35 mm. Head 3.6–3.8 mm long (n=2); anterior part about 1.6 mm wide (eyelevel), posterior widest part about 1.8 mm; ground color yellowish brown, with many dark distinct spots andmarkings, frontoclypeus without dark longitudinal band on mid-line; with irregular dark spots and two small dots
anteriorly, big dark spot and two transverse stripes on middle and posterior position; other parts of head withvarious dark spots; primary setae 1, 4, 10, 11, 13 not dark pigmented; setae 8, 15, 18 located ventrally; setae 18 notbifurcated, about 0.25 mm, located near postoccipital ridge, each surrounded by rugged, triangular area. Pronotum
2.4–2.6 mm long, about 2.2 mm wide, ground color yellowish brown, with many small dark spots except foranterodorsal area, posterior margin black. Forecoxae pentagonal in mesal view, each apicodorsal process very long,horn-like, directed anterad, curved slightly ventrad; each basodorsal process about 1/4 as long as apicodorsal
process, close to apicodorsal process, with long dorsal seta near apex; each central process on mesal face with shortstout seta dorsally in middle.
Specimens examined. [TAIWAN] Beigang River, Huisun Forest Area, Ren’ai Xiang, Nantou, 24°4.992’N,
121°0.651’E, alt. 490 m, 3.iii.2015, N. Nishimura, 1 male, 2 larvae (SU); same locality, 3.iii.2015, L.P. Hsu, 2females (1 female: SU, 1 female: TN); Meiyuan River, Ren'ai Xiang, Nantou, 24°5.09’N, 120°57.22’E, alt. 504 m,
4.iii.2015, N. Nishimura, 1 male, 1 female (1 male: SU, 1 female: TN).
Remarks. The female of this species has dorsum IX mostly sclerotized but separated by membrane along thedorsal mid-line. A continental species, S. huangshanensis Xu et al. 2015, also has this character (Xu et al. 2015),
but S. huangshanensis can be distinguishable from the former by the lack of a row of setae on each sclerotizedplate. The larva of this species can be easily distinguished from known Stenopsyche larvae by the combination offollowing characteristics: Large distinct markings of the head but lacking dark longitudinal band on mid-line offrontoclypeus, unbifurcated no. 18 setae of the head, and long apicodorsal process of each forecoxa.

Discussion
Xu et al. (2015) associated identifiable males of four Chinese Stenospyche species with their females and larvae by
mitochondrial DNA COI sequences. In this study, we also analyzed two Stenopsyche species collected from
Taiwan, and their monophyly at the species level was strongly supported by the mitochondrial DNA COI region(Fig. 1). Thus, we were able to identify the species for Stenopsyche caddisflies using the DNA barcoding method.
We suggest that the effectiveness of the DNA barcoding method is demonstrated clearly with regard to the
Stenopsyche caddisflies.
Schmid (1969) and Weaver (1987) recognized six species groups in the genus Stenopsyche based on male
genitalia, and Xu et al. (2014) assigned S. formosana to the S. simplex Group mainly based on the short segment Xof the male genitalia illustrated by Kobayashi (1987). However, the S. formosana specimens examined in this study
have segment X longer than the intermediate appendages (Figs. 2A, C), although segment X of the holotype male
was over-cleared. The shape of segment X in dorsal view and the numerous spines dispersed along entire length ofthe endotheca in this species are rather similar to those of males belonging to the S. marmorata Group. The larva of
S. formosana is also similar to those of S. angustata Martynov 1930 (Hoang & Bae 2007, Xu et al. 2015) and S.
schmidi Weaver 1987 (Tanida 2003, Nozaki & Shimura 2015), both belonging to the S. marmorata Group as
indicated by the presence of the dark longitudinal band on the mid-line of the frontoclypeus, the shape and lengthof no. 18 setae on the head, and the shape of the forecoxae (Figs. 3D, E, G).
On the other hand, Weaver (1987) assigned S. drakon to the S. marmorata Group, despite the fact that the larva
is very different from known larvae of the S. marmorata Group in the large dark markings and unbifurcated no. 18
setae on the head, and the shape of the forecoxae. Further studies including molecular study are needed to solve thephylogenetic relationships in the genus Stenopsyche.

Acknowledgements
We express our cordial thanks to R.B. Kuranishi, Natural History Museum and Institute, Chiba; for the loan of theholotype of S. formosana; and to B. Price, The Natural History Museum, London, for providing fine photographsof the type series specimens of S. banksi. We also are grateful to S. Uchida, Aichi Institute of Technology, for

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providing the information about the collecting site of the holotype of S. formosana. We are also indebted to K.
Hatta, Nagoya Women’s University; staff of Entomological Department of the National Chung Hsing University;
A.T-S. Lin, the National Central University, Taiwan, for their cooperation with the field research and collection of
specimens in Taiwan. Finally, we are deeply grateful to C.H. Sun, Nanjin Agriculture University, J.C. Morse,
Clemson University, and an anonymous reviewer for their valuable suggestion to improve the manuscript.
The genetic analysis of this study was supported by a Grant-in-Aid for JSPS Fellows (#13J10312, R.S.), andby Grants-in-Aid for Challenging Exploratory Research from the Japan Society for Promotion of Science (JSPS,#23657064, K.T.; #26650160, K.T.).

References
Biodiversity Research Center, Academia Sinica (2015) Taiwan Biodiversity Information Facility. Available from: http://
taibif.tw/zh (accessed 1 December 2015)
Dudgeon, D. (1999) Tropical Asian Streams: Zoobenthos, Ecology and Conservation. Hong Kong University Press, Hong
Kong, 830 pp.
Felsenstein, J. (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular
Evolution, 17, 368–376.
http://dx.doi.org/10.1007/BF01734359
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial
cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3,
294–299.
Hoang, D.H. & Bae, Y.J. (2007) Vietnamese species of Stenopsyche McLachlan (Trichoptera: Stenopsychidae). Zootaxa, 1624,
1–15.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
http://dx.doi.org/10.1093/bioinformatics/17.8.754
Hwang, C. (1957) Descriptions of Chinese caddis flies (Trichoptera). Acta Entomologica Sinica, 8, 373–404.
Kobayashi, M. (1987) Systematic study of the caddisflies from Taiwan, with descriptions of eleven new species (Trichoptera:
Insecta). Bulletin of Kanagawa Prefectural Museum, 17, 37–48.
Malicky, H. (2008) Beschreibungen von neuen Trichopteren aus Asien. Braueria, 35, 45–57.
Malicky, H. (2014) Köcherfliegen (Trichoptera) von Taiwan, mit Neubeschreibungen. Linzer Biologische Beiträge, 46, 1607–
1646.
Martynov, A.V. (1914) Contributions à la faune des Trichoptères de la Chine. Annuaire du Musée Zoologique Académie
Impériale des Sciences de Pétrograd, 19, 323–339.
Martynov, A.V. (1926) On the family Stenopsychidae Mart. With a revision of the genus Stenopsyche McLachl. (Trichopt.).
Eos, 2, 281–308.
Morse, J.C. (2016) Trichoptera World Checklist. Available from: http://entweb.clemson.edu/database/trichopt/index.htm(accessed 2 January 2016)
Mosely, M.E. (1942) Chinese Trichoptera: a collection made by Mr. M. S. Yang in Foochow. Transactions of the Royal
Entomological Society of London, 92, 343–361.
http://dx.doi.org/10.1111/j.1365-2311.1942.tb01210.x
Nishimura, N., Nishida, A., Hatta, K., Kashi, T., Lu, Y.J. & Lien, Y.J. (2015) The records of benthic invertebrates including
caddisflies (Trichoptera) and other species collected in the mountain streams in middle western Taiwan, Hyōgo Freshwater
Biology, 66, 73–81. [in Japanese]
Nozaki, T. & Shimura, N. (2015) Primary seta no. 18 of the larval head of Stenopsyche species (Trichoptera, Stenopsychidae).
Entomological Research Bulletin, 31, 1–6.
Rambaut, A. (2009) FigTree, Version 1.3.1. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 5 January 2016)
Rozas, J., Sánchez-DelBarrio, J.C., Messeguer, X. & Rozas, R. (2003) DnaSP, DNA polymorphism analyses by the coalescentand other methods. Bioinformatics, 19, 2496–2497.
http://dx.doi.org/10.1093/bioinformatics/btg359
Saitou, N. & Nei, M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular
Biology and Evolution, 4, 406–425.
Schmid, F. (1969) La famille des Sténopsychides (Trichoptera). The Canadian Entomologist, 101, 187–224 + pls. 1–7.
http://dx.doi.org/10.4039/ent101187-2
Tamura, K., Peterson, D., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5.02: Molecular evolutionary genetics analysis using
maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28,
2731–2739.
http://dx.doi.org/10.1093/molbev/msr121
Tanabe, A.S. (2007) Kakusan: A computer program to automate the selection of a nucleotide substitution model and theTWO STENOPSYCHE SPECIES IN TAIWAN

Zootaxa 4121 (4) © 2016 Magnolia Press · 493

configuration of a mixed model on multilocus data. Molecular Ecology Notes, 7, 962–964.
http://dx.doi.org/10.1111/j.1471-8286.2007.01807.x
Tanida, K. (2003) Trichoptera. In: Nishida, M. Shikatani, N. & Shokita, S. (Eds.), The Flora and Fauna of Inland Waters in the
Ryukyu Islands. Tokai University Press, Tokyo, pp. 370–392. [in Japanese]
Thompson, J.D., Higgins, D.G. & Gibsonv, T.J. (1994) CLUSTAL W: Improving the sensitivity of progressive multiple
sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids
Research, 22, 4673–4680.
http://dx.doi.org/10.1093/nar/22.22.4673
Ulmer, G. (1907) Trichopteren. Collections Zoologiques du Baron Edm. de Selys Longchamps, 6, 1–102.
Weaver, J.S. III. (1987) New species of Stenopsyche from the northeastern Orient (Trichoptera: Stenopsychidae). Aquatic
Insects, 9, 161–168.
http://dx.doi.org/10.1080/01650428709361290
Williams, N.E. & Wiggins, G.B. (1981) A proposed setal nomenclature and homology for larval Trichoptera. In: Moretti, G.P.
(Ed.), Proceedings of the 3rd International Symposium on Trichoptera. Dr. W. Junk Publishers, The Hague. pp. 421–429.
http://dx.doi.org/10.1007/978-94-009-8641-1_52
Xu, J.H., Wang, B.X. & Sun, C.H. (2014) The Stenopsyche simplex Species Group from China with descriptions of three new
species (Trichoptera: Stenopsychidae). Zootaxa, 3785 (2), 217–230.
http://dx.doi.org/10.11646/zootaxa.3785.2.5
Xu, J.H., Sun, C.H. & Wang, B.X. (2015) A new species of Stenopsyche, with descriptions of larvae and females of somespecies associated by gene sequences (Insecta: Trichoptera). Zootaxa, 4057 (1), 63–78.
http://dx.doi.org/10.11646/zootaxa.4057.1.3
Yang, L.F., Sun, C.H., Wang, B.X. & Morse, J.C. (2005) Present status of Chinese Trichoptera, with an annotated checklist. In.
Tanida, K. & Rossiter, A. (Eds.), Proceedings of the 11th International Symposium on Trichoptera. Tokai University Press,
Hadano, Kanagawa, pp. 441–456.

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