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Accepted by J. Weaver: 20 Jun. 2009; published: 3 Aug 2009 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 2174: 1–17 (2009)
www.mapress.com/zootaxa/Article
The larvae of Chinese Hydropsychidae (Insecta: Trichoptera), Part I:
Arctopsyche shimianensis, Parapsyche sp. A, and Diplectrona obscura
XIN ZHOU
Canadian Centre for DNA Barcoding, Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario, Canada N1G 2W1.
E-mail: xinzhou@uoguelph.ca
Abstract
DNA sequence libraries (mitochondrial COI barcode and nuclear 28S D2) are built for Chinese caddisflies, a fauna
largely under-studied. These independent DNA sequences are used to associate larvae and adults of Chinese
Hydropsychidae. As the first part of the result of this work, Arctopsyche shimianensis Gui and Yang, 2000 and
Parapsyche sp. A are associated with their adults across both gene markers. Probable association is also made for
Diplectrona obscura Ulmer, 1930. Larval descriptions and illustrations are provided for all three species for the first
time.
Key words: Trichoptera, Hydropsychidae, Arctopsyche, Parapsyche, Diplectrona, larva, China, DNA barcoding
Introduction
The fauna of Chinese hydropsychid larvae has been poorly studied. Only 13 larvae out of 136 known
hydropsychid species distributed in China have been associated with adults, among which only a few were
fully described or illustrated (Zhou 2007). Conventional approaches to associating larvae with adult
caddisflies include larval rearing and examination of metamorphotypes (Wiggins 1996). Recently, a
complementary larval association approach that integrates morphology and independent DNA sequences
(mitochondrial COI gene and nuclear 28S D2 fragment) was proposed to facilitate and accelerate larval
association in caddisflies when rearing is not possible and metamorphotypes are not available (Zhou et al.
2007). This method is followed in this paper.
This series larval association and description work is part of the U.S. National Sciences Foundation grant
“Phylogeny and Classification of World Hydropsychidae, with a Revision of Chinese Species and Description
of Their Larvae”. During the course of this investigation, many new species have been discovered from China
and will be described in independent papers by correspondence researchers. As a result, many Chinese
hydropsychid materials used in this larval association study remain unidentified. Potentially new species, such
as the associated Parapsyche species in this paper, are given provisional identities. Along with this association
work, DNA reference libraries of the mitochondrial COI DNA barcode and nuclear 28S D2 have been built
for Chinese caddisflies. These libraries will continuously grow as part of the on-going ‘Trichoptera Barcode
of Life’ project (www.trichopterabol.org) coordinated at the University of Guelph, Canada.
In this work, the larval descriptions and illustrations of three Chinese hydropsychid taxa, Arctopsyche
shimianensis Gui and Yang, 2000, Parapsyche sp. A, and Diplectrona obscura Ulmer, 1930, are provided.
Additional larval descriptions will be published as series manuscripts, consisting of a total of 23 species. For
each associated species, a general introduction to each genus is provided first, followed by generic
characteristics. A list of specimens used in the larval association is given, with GenBank accession numbers
for both genes (where available) linked to each individual (Table 1). Association trees constructed from both
genes are provided. Finally, larval descriptions and illustrations are given for each species.
ZHOU2 · Zootaxa 2174 © 2009 Magnolia Press
TABLE 1. Specimens used in larval-adult association of Chinese Arctopsyche, Parapsyche, and Diplectrona, with
GenBank accession numbers for D2 and COI. SampleID codes are in alphabetic order and are corresponding to those in
Fig. 1 and in BOLD project.
BOLD
sampleID Sex/life
stage Genbank
accession
(COI)
Genbank
accession
(D2)
Locality
CNCAD_0007 larva FJ663191 FJ664670 China,Guangdong,Long-men Co.,Nan-kun-shan Prov. Nature Preserve
CNCAD_0008 adult FJ663192 FJ664671 China,Guangdong,Long-men Co.,Nan-kun-shan Prov. Nature Preserve
CNCAD_0013 adult FJ663187 FJ664672 China,Guangdong,Long-men Co.,Nan-kun-shan Prov. Nature Preserve
CNCAD_0023 larva FJ663193 FJ664673 China,Hunan,Yi-zhang Co.,Mang-shan Natl. Nature Preserve
CNCAD_0024 larva FJ663188 FJ664674 China,Hunan,Yi-zhang Co.,Mang-shan Natl. Nature Preserve
CNCAD_0025 larva FJ663184 FJ664675 China,Hunan,Yi-zhang Co.,Mang-shan Natl. Nature Preserve
CNCAD_0026 adult FJ663185 FJ664676 China,Hunan,Yi-zhang Co.,Mang-shan Natl. Nature Preserve
CNCAD_0037 larva FJ663194 FJ664677 China,Guangdong,Ru-yuan Co.,Nan-ling Natl. Nature Preserve
CNCAD_0038 larva FJ663186 FJ664678 China,Guangdong,Ru-yuan Co.,Nan-ling Natl. Nature Preserve
CNCAD_0039 larva FJ663195 FJ664679 China,Guangdong,Ru-yuan Co.,Nan-ling Natl. Nature Preserve
CNCAD_0055 adult FJ663157 FJ664713 China,Guangdong,Ru-yuan Co.,Nan-ling Natl. Nature Preserve
CNCAD_0073 larva FJ663189 FJ664680 China,Guangdong,Yang-chun Co.,Xin-he village
CNCAD_0074 larva FJ663183 FJ664681 China,Guangxi,Shang-si Co.,Shi-wan-da-shan Natl. Forest Park
CNCAD_0078 male FJ663143 FJ664633 China,Guangdong,Bo-luo Co.,Luo-fu-shan
CNCAD_0080 adult FJ663158 FJ664682 China,Guangdong,Bo-luo Co.,Luo-fu-shan
CNCAD_0091 adult FJ663159 FJ664683 China,Guangxi,Tian-lin Co.,Cen-wang-lao-shan Prov. Forest Preserve
CNCAD_0105 adult FJ663160 FJ664684 China,Guangxi,Long-lin Co.,Jin-zhong-shan Prov. Forest Preserve
CNCAD_0109 male FJ663151 FJ664685 China,Guangxi,Long-lin Co.,Jin-zhong-shan Prov. Forest Preserve
CNCAD_0110 male FJ663152 FJ664686 China,Guangxi,Long-lin Co.,Jin-zhong-shan Prov. Forest Preserve
CNCAD_0113 male FJ663161 FJ664687 China,Guangdong,Ru-yuan Co.,Nan-ling Natl. Nature Preserve
CNCAD_0121 male FJ663162 FJ664688 China,Guangdong,Jiao-ling Co.,Huang-you-bi Nature Preserve
CNCAD_0123 male FJ663163 FJ664689 China,Guangdong,Jiao-ling Co.,Huang-you-bi Nature Preserve
CNCAD_0131b male FJ663198 FJ664634 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0138 male FJ663164 FJ664690 China,Guangxi,Tian-lin Co.,Cen-wang-lao-shan Prov. Forest Preserve
CNCAD_0171 larva FJ663144 FJ664635 China,Guangxi,Hua-jiang Co.,Jiu-wan-da-shan Prov. Nature Preserve
CNCAD_0172 larva FJ663145 FJ664636 China,Guangxi,Le-ye Co.,Bu-liu-he
CNCAD_0173 larva FJ663146 FJ664637 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0174b larva FJ663206 FJ664638 China,Beijing,Bei-jing,Song-shan Natl. Nature Preserve
CNCAD_0175b larva FJ663199 FJ664639 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0176b larva FJ663200 FJ664640 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0177 larva FJ663165 FJ664691 China,Guangxi,Long-lin Co.,Jin-zhong-shan Prov. Forest Preserve
CNCAD_0178 pharate
male FJ663166 FJ664692 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0179 larva FJ663167 FJ664693 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0181 pharate
male FJ663201 FJ664641 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0187 male FJ663138 FJ664642 China,Sichuan,Zhao-jue Co.,Jie-fang-cun Village
CNCAD_0188 male FJ663139 FJ664643 China,Sichuan,Mei-gu Co.,Te-xi Village
CNCAD_0189 male FJ663134 FJ664644 China,Sichuan,Mei-gu Co.,Mei-gu Da-feng-ding Natl. Nature Preserve
CNCAD_0190 male FJ663150 FJ664645 China,Sichuan,Mei-gu Co.,Mei-gu Da-feng-ding Natl. Nature Preserve
CNCAD_0191 male FJ663147 FJ664646 China,Sichuan,Ma-bian Co.,Tian-xing Village
continued next page.
Zootaxa 2174 © 2009 Magnolia Press · 3
CHINESE HYDROPSYCHIDAE
TABLE 1. (continued)
BOLD
sampleID Sex/life
stage Genbank
accession
(COI)
Genbank
accession
(D2)
Locality
CNCAD_0192 male -FJ664647 China,Sichuan,Mian-ning Co.,Da-jia-cun Village
CNCAD_0193 male -FJ664648 China,Sichuan,Zhao-jue Co.,Jie-fang-cun Village
CNCAD_0194 larva FJ663135 FJ664649 China,Sichuan,Zhao-jue Co.,Jie-fang-cun Village
CNCAD_0195 male -FJ664650 China,Jiangxi,Wu-yi-shan Natl. Nature Preserve
CNCAD_0196 male FJ663207 FJ664651 China,Beijing,Bei-jing,Songshan Mountain Natl. Nature Reserve
CNCAD_0197 male FJ663202 FJ664652 China,Sichuan,Du-jiang-yan Co.,Qing-cheng-hou-shan Scenic Area
CNCAD_0280 larva FJ663208 FJ664653 China,Beijing,Bei-jing,Songshan Mountain Natl. Nature Reserve
CNCAD_0281 larva -FJ664654 China,Sichuan,Zhao-jue Co.,Jie-fang-cun Village
CNCAD_0282 larva -FJ664655 China,Sichuan,Zhao-jue Co.,Jie-fang-cun Village
CNCAD_0283 larva FJ663142 FJ664656 China,Sichuan,Wen-chuan Co.,San-jiang Scenic Area
CNCAD_0289 male FJ663168 FJ664695 China,Guangxi,Tian-lin Co.,Cen-wang-lao-shan Prov. Forest Preserve
CNCAD_0290 male FJ663169 FJ664696 China,Guangdong,Zhao-qing City,Ding-hu Shan Forest Ecosystem,
Research Station
CNCAD_0291 male FJ663170 FJ664712 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0292 male FJ663171 FJ664697 China,Guangxi,Hua-jiang Co.,Jiu-wan-da-shan Prov. Nature Preserve
CNCAD_0293 male FJ663172 FJ664698 China,Guangxi,Xing-an Co.,Mao-er-shan Natl. Nature Preserve
CNCAD_0294 male FJ663173 FJ664699 China,Guangxi,Tian-lin Co.,Cen-wang-lao-shan Prov. Forest Preserve
CNCAD_0295 male FJ663174 FJ664700 China,Guangxi,Nan-dan Co.,Qing-shui-he
CNCAD_0299 male FJ663175 FJ664701 China,Jiangxi,Wu-yi-shan Natl. Nature Preserve
CNCAD_0300 male FJ663176 FJ664702 China,Jiangxi,Wu-yi-shan Natl. Nature Preserve
CNCAD_0301 male FJ663177 FJ664703 China,Jiangxi,Jiu-lian-shan Natl. Nature Preserve
CNCAD_0302 male -FJ664704 China,Jiangxi,Jiu-lian-shan Natl. Nature Preserve
CNCAD_0303 male FJ663178 FJ664705 China,Jiangxi,Jiu-lian-shan Natl. Nature Preserve
CNCAD_0304 male FJ663179 FJ664706 China,Jiangxi,Jiu-lian-shan Natl. Nature Preserve
CNCAD_0305 male FJ663180 FJ664707 China,Sichuan,Tian-quan Co.,Xiao-yu-xi Stream
CNCAD_0306 male FJ663181 FJ664708 China,Sichuan,Shi-mian Co.,Liziping Nature Preserve
CNCAD_0307 male FJ663182 FJ664709 China,Jiangxi,Wu-yi-shan Natl. Nature Preserve
CNCAD_0308 male FJ663153 FJ664710 China,Sichuan,Ma-bian Co.,Tian-xing Village
CNCAD_0309 male FJ663154 FJ664711 China,Jiangxi,Jiu-lian-shan Natl. Nature Preserve
CNCAD_AT01 larva FJ663203 -China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_AT02 larva -FJ664657 China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_AT03 larva FJ663204 FJ664658 China,Yunnan,Li-jiang Co.,Yu-shui-zhai Park
CNCAD_AT04 larva FJ663205 FJ664659 China,Yunnan,Da-li Co.,Cang-shan Park
CNCAD_AT09 larva FJ663140 FJ664660 China,Sichuan,Bao-xing Co.,Feng-tong-zhai Natl. Preserve
CNCAD_AT10 larva FJ663141 FJ664661 China,Sichuan,Bao-xing Co.,Feng-tong-zhai Natl. Preserve
CNCAD_AT11 male FJ663148 FJ664662 China,Yunnan,Li-jiang Co.,Yu-shui-zhai Park
CNCAD_DP02 larva FJ663190 FJ664694 China,Yunnan,Da-li Co.,Cang-shan Park
CNCAD_PR01 larva -FJ664663 China,Yunnan,Da-li Co.,Cang-shan Park
CNCAD_PR02 larva -FJ664664 China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_PR03 larva FJ663196 FJ664665 China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_PR04 larva -FJ664666 China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_PR05 larva -FJ664667 China,Yunnan,Zhong-dian Co.,Zhong-dian
CNCAD_PR07 larva -FJ664668 China,Beijing,Bei-jing,Song-shan Natl. Nature Preserve
CNCAD_PR08 larva FJ663197 FJ664669 China,Yunnan,Li-jiang Co.,Yu-shui-zhai Park
ZHOU4 · Zootaxa 2174 © 2009 Magnolia Press
Material and methods
Larval and adult specimens were collected from a series of collection trips in six provinces of China:
Guangdong, Guangxi, Jiangxi, Sichuan, Yunnan, and Beijing, from 2001 to 2005. All specimens were
collected in 95% ethanol. Hydropsychid larvae were collected using kick-nets and D-nets or by examining the
substratum. Adults were collected by light-trap and sweep-net. Both larval and adult specimens were cursorily
sorted after collecting and preserved in 95% ethanol. The ethanol was replaced within a few days after
collecting and changed again within approximately 30 days. Larvae and adults were sorted to morpho-species
as soon as possible (typically within 6 weeks) and individuals were preserved in separate vials. Vouchered
specimens are deposited at Nanjing Agriculture University (NAU), China.
Abdominal segments III–VI or legs of larvae and adults were used for DNA extraction. Larval intestines
and gut contents were removed carefully to reduce the potential for contaminants. The rest of the specimen
was preserved in ethanol for morphological study. Each extracted individual was given a unique sample ID
linked with two GenBank accession numbers (for both D2 and COI where available) (Table 1). DNA barcode
sequences and detailed specimen information are accessible in the project ‘Larvae of Chinese Hydropsychidae
Part I’, in the Barcode of Life Data System (BOLD, www.boldsystems.org).
General molecular protocols were used for genome DNA extraction, PCR amplification, and sequencing.
The sequencing of D2 gene was conducted at Rutgers University, USA. Full-length DNA barcodes (658bps)
were acquired at the Canadian Centre for DNA Barcoding, University of Guelph, Canada. D2 sequences were
aligned manually according to the secondary structure, following Kjer (1995). The COI tree was constructed
in the BOLD system using Neighor-joining and K-2-P parameter. The D2 tree was constructed in
PAUP*4.0b10 (Swofford 2003) using the same method and parameter. Both association trees were exported
to and annotated in Adobe Illustrator®. Association criteria described by Zhou et al. (2007) are followed in
this paper. Briefly, provisional species boundaries were recognized by morphological examination of male
specimens and confirmed by two genes. Two conditions were used to make an association: if both D2 and
COI sequences were identical, the association was assumed; if this condition was not met, then the unknown
larva had to nest within a clade of identified adults in order for an association to be made.
Wiggins (1996) and Williams and Wiggins (1981) are followed for general larval morphology and setal
nomenclature. Larval illustrations were performed on a Nikon SMZ-U Zoom 1:10 compound microscope and
camera lucida. Final ink illustrations were scanned, arranged and annotated in Adobe Photoshop® and Adobe
Illustrator®.
The lengths of the membranous body parts, such as abdominal segments, are affected by natural variation
and distortions from the ways the animals were preserved. Thus only the length and width of the hard,
sclerotized head capsule are measured. The measurements were performed using a 10 x 10 ocular grid reticle
installed in the eyepiece of the microscope. The grid lines were calibrated with a stage micrometer for various
magnifications.
Among many larval features, the characteristics of the head (such as the shape of the frontoclypeal
apotome, color patterns, texture of the cuticle and setal characters), thoracic nota (shape of borders and marks,
setation), foretrochantin, prosternal plate and posterior prosternal sclerites, abdominal hairs and tracheal gills
have been emphasized.
Specimens were handled with extreme care because some characters, such as hairs and tracheal gills, are
easily broken and lost during specimen transfer and dissection. All characters were examined in standardized
aspects. In most cases, these are the exactly dorsal, ventral or lateral views of a particular structure. For some
characters, such as the foretrochantin, the lateral parallel plane of the structure was observed, not the lateral
plane of the animal’s body. Inaccurate placement of the structure may result in inaccurate measurements, such
as the relative length and width of the dorsal and ventral branches of the foretrochantin in some caddis larvae.
Hairs, tracheal gills, and anal gills were variable among individuals of the same species because of
intraspecific variation or abnormal development in some individuals. In this case, multiple individuals for a
particular species were examined.
Zootaxa 2174 © 2009 Magnolia Press · 5
CHINESE HYDROPSYCHIDAE
FIGURE 1a–b. Larval-adult association of Chinese Arctopsyche, Parapsyche, and Diplectrona— 1a. COI neighbor-
joining phylogram; 1b. D2 neighbor-joining phylogram. Specimens are labeled with taxonomy, followed by sample ID
(corresponding to Table 1) and province location. Specimens of immature life stage are noted at the end (pp: pharate
pupa; lv: larva). COI tree is constructed in the BOLD system using Neighbor-joining and K-2-P parameter. D2 tree is
constructed in PAUP*4.0b10 using Neighbor-joining and K-2-P parameter. Numbers on the internodes represent the
bootstrap values using Neighbor-joining/1000 replicates. Species boundaries recognized in morphology and two DNA
sequences are marked in brackets on the right of each DNA tree and noted as haplogroups (HPGP) 1-34 respectively.
ZHOU6 · Zootaxa 2174 © 2009 Magnolia Press
Genus Arctopsyche McLachlan, 1868
A total of eight Arctopsyche species were recorded previously from China (Yang et al. 2005), of which A.
shimianensis Gui and Yang, 2000, A. trispinosa Schmid, 1968, and two possibly new species, as well as
various of larval morphs were collected and sequenced in this study (Table 1; Fig. 1). Lepneva (1964)
provided comprehensive larval and pupal descriptions and illustrations for A. palpata Martynov, 1934, a
species with wide distribution including the Russian Far East and the Chinese northeastern provinces and
Inner Mongolia. The larval ecology of this species was studied by Kocharina (1997). This is the only Chinese
Arctopsyche species whose larvae have been associated.
LARVA—HEAD. Frontoclypeal apotome conspicuously constricted opposite eyes. Genae completely
separated by ventral apotome. Ventral apotome narrowed posteriorly [but also tapered in at least one North
American Parapsyche species (Wiggins 1996)]. Submentum trapezoidal, anterior margin not cleft.
Stridulating files consisting of series of short, thick bars, very conspicuous. Seta 22 on anterolateral corners of
pronotum not conspicuous. Foretrochantin triangular, pointed, not forked. Prosternal plate single-pieced,
posterior prosternal sclerites absent. Sagital section of mesonotum more or less triangular. Meso- and
metanota bearing transverse ecdysial lines. Abdominal segments covered by regular hair-like setae and
elongated club-hairs. Setal areas, Sa2 and Sa3 in most abdominal segments each bearing only one single long
seta, not cluster of club-hairs. Lateral gills on abdominal pleura similar to ventral gills, with numerous
filaments distributed mostly at apex of each stalk. Abdominal segment VII bearing ventral and lateral gills.
Ventral plates on abdominal segment IX large.
LARVAL-ADULT ASSOCIATION. For this study 15 individuals of Arctopsyche, representing 6 provisional
species (HPGP 1–6, Fig. 1), were extracted for DNA amplifications, including two possibly new species
(CNCAD_0078 and CNCAD_0191/CNCAD_AT11, Table 1 and Fig. 1). Species delimitations are congruent
in two genes in 5 of the 6 species. HPGP 4 is split into two subgroups (HPGP 4a and 4b) in the COI tree, with
a mean divergence of 8.7% between them, and 0.5% and 0% divergence within HPGP 4a and 4b, respectively.
However, members of these two subgroups have essentially identical D2 sequences. Likely, D2 has not
evolved as fast as COI gene in this recent speciation. Additional adult materials are needed to clarify the
species boundary because HPGP 4b is only represented by larval specimens, which are indistinguishable from
that of HPGP 4a. A. shimianensis was associated across both D2 and COI makers. Its larva is described and
illustrated.
Arctopsyche shimianensis Gui and Yang, 2000
Figures 2a–l
LARVA—HEAD. Head oval, 3.1 mm long, 2.8 mm wide, dorsoventrally flattened. Frontoclypeal apotome
strongly constricted opposite eyes, maximum width at anterior end. Anterior margin of frontoclypeus curved,
convex in middle, smooth, without flanges or notches. Areas near tentorial pits depressed, but not connected
by complete mediotransversal crease as in some species of Hydropsychinae.
Primary setae mostly long, thick and dark except seta 17 short, yellowish, and clear. Secondary setae
small, fine, tapered, blackish with yellowish basal ends. Setal pits of these small hairs often large and
relatively deep, posterior edge elevated and dark, anterior edge depressed and pale, giving head capsule rough
appearance. No blunt-setae present on dorsum of head. Several secondary setae on anterolateral corners of
frontoclypeal apotome long, curved, yellowish, and clear. Setae on posterior half of frontoclypeus sparser.
Thickened spiky setae distributed on ventrolateral sides of head, short to moderately long, dark, clearly visible
from dorsal, lateral, and ventral aspects.
In dorsal aspect, head mostly dark brownish, with some inconspicuous pale marks: anterior half of
frontoclypeus with pair of elongated marks on lateral portions immediately anterior to depressed regions near
tentorial pits; series of four round blackish dots transversely arranged on middle portion connecting lateral
Zootaxa 2174 © 2009 Magnolia Press · 7
CHINESE HYDROPSYCHIDAE
FIGURE 2a–l. Arctopsyche shimianensis. Larva— a. Head, dorsal; b. head, ventral; c. head, left lateral; d. pro-, meso-,
and metanota, dorsal; e. pro-, meso-, and metanota, left lateral (with setae on anterolateral corners of pro- and
mesonotum further enlarged: hl: hair-like setae; ch: club-hairs; pl: peg-like setae); f. prosternal plate, ventral; g. base of
left fore leg, showing lateral view of prosternal plate and position of foretrochantin, left lateral (with left foretrochantin
further enlarged); h. ventral plates on sterna VIII and IX, ventral; i. abdominal hairs on tergum of segment II (hl: hair-like
setae; ch: club-hairs); j. ventral gill on right side of mesosternum, anterior aspect with ventral side facing upward; k.
ventral and lateral gills on right side of abdominal segment II, anterior aspect with ventral side facing upward; l. lateral
(pleural) gills on left side of abdominal segment V, left lateral.
ZHOU8 · Zootaxa 2174 © 2009 Magnolia Press
pale marks; bases around setae 9 yellowish; posterior half of frontoclypeus with V-shaped pale mark along
posterior half of frontoclypeal suture; longitudinal pale stripe visible along mesal line of frontoclypeus and
coronal suture, although often very weak; small, round, bright mark located posterior to apex of expanded
portion of frontoclypeus, followed by longitudinal pale stripe along proprioceptors 19, 20, and 21 on posterior
parietal sclerite of each side. Muscle scars mostly confined on posterior portion of head, often slightly paler
than background, with dark borders. Lateral sides of head generally pale, yellowish, eyes each surround by
whitish area. Muscle scars pale, not conspicuous. Stout spiky setae pronounced.
Parietal sclerites separated completely on ventral side by ventral apotome. Anterior end of ventral
apotome enlarged; anterior margin slightly curved; both anterolateral corners acute; lateral borders narrowed
gradually toward posterior end and almost parallel to each other on posterior half; posterior end of ventral
apotome with small brownish triangular projection. Submentum trapezoidal, anterior and posterior borders
slightly curved; anterior margin not cleft; area adjacent to posterior margin with curved transverse ridge.
Outline of stridulating files wide; striae on anterior portion of stridulating files thicker than those on posterior
portion; each stria consisting of series of dark brownish, horizontally arranged short bars. Muscle scars mostly
confined to posterior portion of venter of head, about as dark as background, not conspicuous. Stridulating
files and following area dark; ventral apotome and areas on parietal sclerites mesal of stridulating files
generally pale; ridges along occipital margins pale. Setae on ventrolateral sides of head consisting of short,
fine hair-like setae and moderately long, stout, spiky, dark brownish setae, with setal pits pronounced.
THORAX. Thoracic nota covered mainly by moderately long, tapered, blackish, thick and hair-like setae.
Hairs on meso- and metanota generally sparser than those on pronotum. Pronotum subdivided longitudinally.
Majority of pronotum dark brownish, lateral fourth of each side of notum pale, yellowish (not visible in dorsal
aspect). Posterior margin bearing transverse blackish border, followed by wide yellowish area and brownish
narrow sclerotized stripe. Posterior three-fourths of lateral borders blackish, anterior fourth of lateral borders
and anterior margin without blackish mark. Anterior fourth of lateral borders bearing series of moderately
long, thick, blunt, brownish peg-like setae; anterolateral corners of pronotum also with few similar setae (Fig.
2e, with hairs on anterolateral corner of left side of pronotum further enlarged). Hairs on anterior margin of
pronotum similar to those on dorsum of notum, but longer and thicker.
Foretrochantin triangular, not forked, distal end gradually tapered into sharp apex. Dorsal surface of
trochantin nearly straight, bearing two large, pointed, brownish spurs and series of blackish bristles (Fig. 2g,
with foretrochantin further enlarged). Prosternal plate large, folding transversely at posterior third into almost
right angle (Figs. 2f, g). Anterior margin protrusive in middle, forming acute angle. Anterior margin blackish;
posterior border with wide, oblong, blackish mark in middle third. No posterior prosternal sclerites present. At
least in some individuals (probably mature larvae before becoming prepupae), intermediate space between
prosternal plate and ventrolateral borders of pronotum appearing sclerotized and yellowish; these sclerites
appressing each other tightly and forming integrated piece (as shown in Fig. 2f).
Mesonotum paler than pronotum, mostly brownish, with longitudinal pale stripe along mid-dorsal line.
Area following blackish posterior median mark pale, yellowish. Lateral half of each side of mesonotum
generally pale. Transverse ecdysial line extending from posterior third of median part of notum nearly to
middle of lateral borders. Anterior margin bearing heavy blackish mark, disconnected from lateral marks near
anterolateral corners. In lateral aspect, mesonotum appearing triangular in sagital section, with dorsal border
convex (Fig. 2e). Posterior border with blackish crescent mark on middle two-fifths. Pale areas following
median mark triangular, followed by pair of brownish short bars with crenulated posterior borders. Muscle
scars mostly confined on posterior portion of notum, typically paler than background. Areas near anterolateral
corners of notum without peg-like setae, but with few scattered club-hairs.
Metanotum lightest among thoracic nota, dorsoventrally depressed, almost quadrate, posterior margin
bearing wide, deep excision. Mostly light brownish, with longitudinal yellowish stripe along mid-dorsal line.
Transverse ecdysial line bearing sharp angle pointing caudad at position immediately anterior to Sa2 of each
side. Anterior margin with blackish border, not as broad as that on mesonotum, disconnected from lateral
borders near anterolateral corners. Posterior border bearing narrow, wide, blackish mark on middle three-
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fourths, curving caudad. Mesosternum with one pair of ventral tracheal gills; metasternum bearing two pairs
(ventrolateral and ventromesal) (Table 2). All thoracic gills with single, globular gill stems; each gill stem
bearing 25–40 gill filaments distributed evenly on upper hemi-sphere (Fig. 2j).
TABLE 2. Number and arrangement of gills in mature larva of Arctopsyche shimianensis. Numbers in the table represent
the number of gills present on one side of a particular body segment. Gill types: a: simple, finger-like gill (note: no type
a gill present in Arctopsyche shimianensis, term being kept to be compatible with other hydropsychid species); b: bifid-
stemmed gill; c: single-stemmed gill.
ABDOMEN. Abdominal terga and pleura densely covered by long, pointed, curved hair-like setae and long,
thick, dark brownish club-hairs, each with apex slightly enlarged and truncated (Fig. 2i). Club-hairs typically
longer and stouter than hair-like setae. Club-hairs not forming clusters. Abdominal pleura II–VII with large
lateral (pleural) gills posteroventrally on each pleuron: II and VII with one gill; III–VI with two gills on each
side; I, VIII and IX without lateral gills (Table 2; Fig. 2l). Lateral gills similar to ventral gills, only smaller;
each gill with single cylindrical stalk; numerous gill filaments arising mostly from apex of each stalk. Lateral
gills not covered by abdominal hairs.
Abdominal sterna bearing two types of ventral gills: single-stemmed gills and bifid-stemmed gills, with
single-stemmed gills typically situated anteromesal of bifid- stemmed gills when both present on particular
segment. Gill filaments arising from apex of each gill stem or each branch of bifid gill stem. Abdominal
segment I with pair of single-stemmed ventral gills (ventrolateral and ventromesal) on each side, isolated from
each other; II–V each with one single-stemmed ventromesal gill and one bifid-stemmed ventrolateral gill on
each side (Fig. 2k); VI with only one bifid-stemmed ventrolateral gill on each side; VII with two single-
stemmed gills on each side; VIII and IX without ventral gills (Table 2).
Sterna VIII and IX each bearing strongly sclerotized yellowish plates: ventral plate VIII relatively small,
fused into one single piece, nearly trapezoidal, posterior border longer, bearing small median notch; ventral
plates IX large, separated, each nearly trapezoidal, with mesal border longer, posteromesal corner slightly
acute. Anterior half of plate VIII bearing almost no hairs; posterior half with series of short, thick, peg-like,
golden setae, whose setal sockets pronounced; posterior border with row of thick (about as thick as peg-like
setae), long, blackish bristles. Plates IX large, lateral portions folding dorsad toward pleura; anterolateral
corner of each plate IX bearing almost no hairs; remaider of each plate with several short (longer than those
on plate VIII), thick, golden, peg-like setae and short, thin, tapered, hair-like setae; posterior border of each
plate IX bearing dense row of long, thick, blackish bristles pointing caudad; longitudinal area between plates
IX bearing series of short, fine, tapered, blackish hair-like setae (Fig. 2h).
Diagnosis. The mature larva of A. shimianensis is a very large caddis. The head is typically wide and
dorsoventrally depressed. Most of the dorsum of the head is dark, with slightly pale marks, which can be very
inconspicuous in some individuals. The stridulating files on the ventral side of the head are pronounced. The
thick, spiky setae on the ventrolateral side of the head and the hairs on the anterolateral corners of both the
pro- and mesonota appear to be diagnostic for the species. Each of the bifid-stemmed ventrolateral gills on
abdominal pleura VI can be two separate single-stemmed gills that are very close together.
Material examined. Sichuan: Mian-ning County, Jia-wu-cun Village. Yan-jian-gou stream, 100–150 m
upstream of S215 at 410.4 km stone marker. N28.36409, E101.99700. 2379 m. 3-July-2005. X. Zhou, C-F
Zhou, 13 specimens. Zhao-jue County, Jie-fang-cun Village. Jie-fang-gou stream, S307 at 553.0 km stone
marker. N27.88159, E102.55734. 2925 m. 4-Jul-2005. X. Zhou, C-F Zhou, 36 specimens.
Thorax Abdominal segments
Mesothorax Metathorax III III IV VVI VII VIII IX
Lateral (pleural) - - - 1c2c2c2c2c1c- -
Ventro-lateral 1c1c1c1b1b1b1b1b1c- -
Ventro-mesal -1c1c1c1c1c1c-1c- -
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Genus Parapsyche Betten, 1934
A total of eight Parapsyche species were previously recorded from China (Yang et al. 2005). At the moment,
no larval descriptions or illustrations are available for Chinese Parapsyche. In this work, 22 individuals of
Parapsyche, representing 5 provisional species (HPGP 7–12, Fig. 1), were extracted for DNA amplifications,
among which nuclear D2 of all were sequenced while amplification of mitochondrial sequences were
unsuccessful in some individuals, especially that of HPGP 9. This is presumably due to some significant
nucleotide changes in the primer regions of the COI fragment. Two species (corresponding to HPGP 8 and 11)
are associated across two DNA markers (Fig. 1). However, neither of these two provisional species were
identified to any known Chinese Parapsyche species. Larva of HPGP 8 is described and illustrated in this
paper as Parapsyche sp. A.
LARVA—HEAD. Frontoclypeal apotome greatly constricted opposite eyes and significantly expanded
laterally posterior to constriction. Genae completely separated by ventral apotome. Ventral apotome very
broad, lateral borders mostly parallel, not narrowed posteriorly. Submentum trapezoidal, anterior margin not
cleft. Stridulating files consisting of series of conspicuous, short, thin bars. Sagital section of mesonotum
more or less triangular. Setae 22 on anterolateral corners of pronotum not conspicuous. Each foretrochantin
triangular, not forked. Prosternal plate single, posterior prosternal sclerites absent. Meso- and metanota
bearing transverse ecdysial lines. Abdominal segments covered by regular hair-like setae and thickened club-
hairs. Sa2 and Sa3 on most abdominal segments bearing cluster of elongated club-hairs in addition to long,
tapered hairs. Lateral gills on abdominal pleura simple, finger-like or bearing few short filaments. Abdominal
segment VII bearing ventral and lateral gills. Ventral plates on abdominal segment IX large.
Parapsyche species A
Figure 3a–m
LARVA— HEAD. Head oval, almost round, 2.6 mm long, 2.5 mm wide, slightly constricted laterally near
posterior third, dorsoventrally thickened. Frontoclypeal apotome strongly constricted opposite eyes;
constriction point forming sharp angle, after which frontoclypeus greatly expanded laterally, reaching
maximum width; anterior margin smoothly curved. In dorsal view, head mostly dark, yellowish brown, with
yellowish pale marks and number of dark muscle scars. Anterior margin and anterolateral corners of
frontoclypeus, dark brownish. Eyes circled by yellowish pale areas. Frontoclypeus with longitudinal
yellowish stripe along posterior two-thirds of mid-dorsal line. Pale stripe on frontoclypeus broken at its
posterior third by number of dark muscle scars, but posteriorly connected with pale stripe along coronal stem.
Tentorial pits dark, each pit followed by small oval pale mark with anterolateral margin bordered by lateral
border of frontoclypeus after constriction point. Posterior portion of each parietal sclerite bearing wide,
longitudinal pale stripe, along which situated anteriorly small, round yellowish mark and large, round
yellowish mark in middle, immediately anterior to proprioceptor 19 (Fig. 3a).
Primary setae mostly long, thick and brownish; seta 17 short, yellowish and clear. Secondary setae small,
fine, tapered, mostly dark with yellowish basal ends. Setal pits of these small hairs often large and relatively
deep, posterior edge elevated and dark, anterior edge depressed and pale, giving head capsule rough
appearance. No blunt-setae present on dorsum of head. Some secondary setae on anterolateral corners of
frontoclypeal apotome long, curved, yellowish and clear. Several thickened spiky setae on ventrolateral sides
of head, short to moderately long, dark or pale. In ventral view, genae completely separated by ventral
apotome. Ventral apotome wide; anterior margin slightly concave; anterior eighth expanded laterally, forming
acute anterolateral angles; lateral borders mostly parallel; posterior border slightly excised in middle.
Submentum trapezoidal, anterior margin concave and posterior margin convex. Stridulating files conspicuous,
each stria consisting of series of short, thin bars. Muscle scars mostly confined to posterior halves of parietal
sclerites, slightly darker than background. Posteromesal areas of parietal sclerites contiguous with posterior
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CHINESE HYDROPSYCHIDAE
border of ventral apotome, lightly sclerotized. Venter of head mostly brownish, areas surrounding stridulating
files slightly paler (Fig. 3b).
FIGURE 3a–m. Parapsyche sp. A. Larva— a. Head, dorsal (with secondary setae on dorsum of head further enlarged);
b. head, ventral (with secondary setae on ventrolateral head further enlarged); c. head, left lateral (setae and color
patterns on frontoclypeus not shown); d. pro-, meso-, and metanota, dorsal (with secondary setae on dorsum of
mesonotum further enlarged); e. pro-, meso-, and metanota, left, lateral; f. prosternal plate, ventral; g. left foretrochantin,
left lateral; h. ventral plates on abdominal sterna VIII and IX; i. abdominal hairs on tergum of segment II (hl: hair-like
setae; ch: club-hairs); j. lateral (pleural) gills on left side of abdominal segment V, left lateral; k. ventral gills on right side
of abdominal segments I and II, anterior aspect with ventral side facing upward; l. ventral gills on right side of abdominal
segment III, anterior aspect with ventral side facing upward; m. ventral gills on right side of abdominal segments VII and
VIII, ventral aspect with caudal orientation placed upward.
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THORAX. Thoracic nota densely covered by moderately long, thick, tapered, blackish hairs and small, fine,
tapered, dark hairs (Fig. 3d, with hairs on dorsum of mesonotum enlarged). Pronotum subdivided
longitudinally, but mid-dorsal ecdysial line weak. Majority of pronotum pale yellowish, with pair of
ambiguous dark areas on middle of dorsum and series of dark muscle scars distributed mainly on lateral areas
of each side. Posterior margin of pronotum with wide blackish border. Lateral blackish border not reaching
anterolateral corners (Figs. 3d, e). Foretrochantins triangular, not forked, each very broad at base; two thick,
long, blackish spurs and series of short, thick bristles located on dorsal surface (Fig. 3g). Prosternal plate
large, folding transversely at posterior third (Fig. 3f). Anterior margin protrusive in middle, forming acute
angle. Anterior margin blackish except in middle; posterior border with wide, oblong, blackish mark in
middle half. No posterior prosternal sclerites present. Meso- and metanota each with transverse ecdysial line
near posterior third, extending anterolaterad toward anterior third of lateral borders.
Meso- and metanota each with pair of large dark areas on lateral portions of notum; longitudinal area
along mid-dorsal line pale, yellowish; dark areas on mesonotum darker than those on metanotum. All of
anterior margin of mesonotum with wide, transverse blackish border; posterior border with wide, U-shaped
blackish mark on middle three-fifths, with lateral corners angled posterolaterad; this posterior middle mark
not connected with lateral blackish borders. Anterior margin of metanotum also bearing blackish border, but
border thinner than on mesonotum. Posterior margin bearing wide, shallow notch. Posterior middle mark
straight, connected laterally with lateral borders. Tracheal gills on meso- and metasterna single-stemmed; gill
stalks short, globular, each with numerous gill filaments mainly distributed on upper half of sphere.
Mesosternum with one ventrolateral (coxal) gill on each side; metasternum with one ventrolateral and one
ventromesal gill on each side (Table 3).
TABLE 3. Number and arrangement of gills in mature larva of Parapsyche sp. A. Numbers in the table represent the
number of gills present on one side of a particular body segment. Gill types: a: simple, finger-like gill (note: no type a gill
present in Parapsyche spp., term being kept to be compatible with other species); b: bifid-stemmed gill; c: single-
stemmed gill.
ABDOMEN. Abdominal terga and pleura densely covered by small, fine, tapered hair-like setae and large,
thickened, blackish club-hairs (Figs. 3i, j). Club-hairs typically truncated at apex and broadened near middle
or base. Sa2 and Sa3 of most abdominal segments with cluster of elongated club-hairs in addition to long hair-
like setae.
Segment I with one finger-like lateral (pleural) gill on each side. II–VII each with one laterodorsal and one
lateroventral gill on each side: dorsal one simple, finger-like; ventral one typically with short, bulging stalk,
with one longer filament located on inner side at apex of stalk, and one short filament on outer side;
lateroventral gill on V with two short filaments on outer side (Fig. 3j). Lateral gills not covered by hairs.
Ventral gills on abdominal sterna single-stemmed, with globular to cylindrical gill stalks, gill filaments
distributed mainly at apex of each stalk. Abdominal segments I and II each with one ventrolateral gill and one
ventromesal gill on each side, distinctly separated (Fig. 3k); III–VI each with two closely positioned but
separated ventrolateral gills and one ventromesal gill on each side (Fig. 3l); VII with two ventrolateral gills
but no ventromesal gill; VIII with one simple, finger-like ventrolateral gill on each side (Fig. 3m); VIII and IX
with no lateral or ventral gills (Table 3).
Sterna VIII and IX each bearing strongly sclerotized yellowish plates: ventral plate VIII relatively small,
fused into one single piece, nearly oval; ventral plates IX large, separated, nearly trapezoidal, mesal borders
Thorax Abdominal segments
Mesothorax Metathorax III III IV VVI VII VIII IX
Lateral (pleural) - - - 1c2c2c2c2c1c- -
Ventro-lateral 1c1c1c1b1b1b1b1b1c- -
Ventro-mesal -1c1c1c1c1c1c-1c- -
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CHINESE HYDROPSYCHIDAE
longer, straight. Posterior half of plate VIII with few short, thick, peg-like, golden setae; posterior border with
row of thick, long, blackish bristles. Plates IX large, lateral portions folding dorsad toward pleura; most of
each plate with several short, thick, golden, peg-like setae and few short, thin, tapered, hair-like setae;
posterior border of each plate IX bearing dense row of long, thick, blackish bristles pointing caudad;
longitudinal area between plates IX bearing series of short, fine, tapered, blackish hair-like setae (Fig. 3h).
Diagnosis. Some Parapsyche larvae, such as two unassociated Japanese morpho-species (Figs. 2, 3, p480,
in Tanida, 2005), tend to have similar color patterns on dorsum of the head. Although not many diagnostic
characters are available to differentiate these species, the morphology of tracheal gills seems to provide some
differentiation. For example, the lateral gills on the abdominal segments in the two Japanese morpho-species
are different from the Chinese larva described in this work. However, the gill distribution can be highly
variable among individuals. The validity of this potential diagnostic character is subject to further
examination.
Material examined. Beijing, Song-shan National Nature Reserve: Song-shan Park main scenic route.
N40.52541, E115.82284. 960–1170 m. 2-Aug-2003. X. Zhou. 29 specimens; small stream besides Da-
zhuang-ke Village. 1000 m. 19-Jun-2005. X. Zhou. 29 specimens.
Genus Diplectrona Westwood, 1840
Diplectrona is the only genus of the subfamily Diplectroninae occurring in China (Yang et al. 2005). A total
of six Diplectrona species are recorded from China. No Chinese Diplectorna larvae have been associated with
their adults. Because the original species descriptions were overly simplified for most Chinese Diplectrona
species and a significant portion of Chinese fauna remains unknown, many individuals used in this larval-
adult association work were not identified to named species. However, the male genitalic structures of these
morpho-species are distinctive. Thus, the species boundaries among the included Chinese Diplectrona species
are clear.
LARVA—HEAD. Genae on ventral side of head touching along mid-ventral ecdysial line. Ventral apotome
divided into anterior and posterior parts, posterior ventral apotome at least half as long as mid-ventral ecdysial
line. Frontoclypeal apotome greatly constricted opposite eyes and strongly expanded after eyes.
Foretrochantins not forked (in all Chinese Diplectrona species examined so far). Prosternal plate large,
posterior prosternal sclerites absent. In dorsal view, pronotum slightly constricted at posterior end. Meso- and
metanota each with transverse ecdysial lines. Lateral (pleural) gills on abdominal segments simple, finger-
like. Abdominal terga and pleura densely covered by hair-like setae and elongated club-hairs. Abdominal
segment VII without ventral tracheal gills.
LARVAL-ADULT ASSOCIATION. In this work, 44 individuals of Diplectrona, representing 22 provisional
species (HPGP 13–34, Fig. 1), were extracted for DNA amplifications (Table 1 and Fig. 1). One possible
association was made for Diplectrona obscura Ulmer, 1930 (represented by individuals CNCAD_0109, 0110
and 0177 in Table 1 and Fig. 1). The D2 sequence of the larval specimen (CNCAD_0177) is identical to the
two adult D. obscura (0109 and 0110), while its COI is only 0.8% different from the latter two specimens.
Although this larval form is very likely the larva of D. obscura, additional materials are needed to confirm this
association. The two specimens representing HPGP 31 and 32 share identical D2 sequence but are
distinctively separated in COI tree. This is likely due to the relatively slow evolving rate in D2.
Diplectrona obscura Ulmer, 1930
Figure 4a–k
LARVA (POSSIBLE ASSOCIATION)—Head. Head generally orangish brown, subquadrate, 1.4 mm long, 1.2 mm
wide. Frontoclypeal apotome very broad, narrowed opposite eyes, behind which strongly expanded laterally.
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Middle expansion as wide as anterior margin of frontoclypeus. Anterior margin of frontoclypeus smooth,
slightly convex, without prominent flanges or teeth. Frontoclypeal apotome nearly flat, tentorial pits and
mediotransversal crease not prominent (Fig. 4a). Dorsum of head (including frontoclypeus) densely covered
by two types of secondary setae: long, thick, dark brownish blunt-setae and median to long, fine, tapered, dark
hair-like setae (Fig. 4a). Blunt-setae erect, nearly perpendicular to surface of head capsule; hair-like setae
lying close to surface, pointing cephalad. Posterior sixth of dorsum of head seta-less.
Eyes circled by whitish yellow area. Posterior seta-less area of head whitish yellow. Area on anterior
fourth of frontoclypeus dark brownish, whose posterior border not conspicuous. Series of dark spots posterior
to expansion of frontoclypeus, nearly U-shaped in arrangement. Parietal sclerites along frontoclypeal suture
behind frontoclypeal expansion and along coronal suture with dark brownish thin stripes; contrasting with
undarkened adjacent frontoclypeus. Muscle scars mostly confined to posterior areas of parietal sclerites, dark
brownish, conspicuous (Fig. 4a). Anterior ventral apotome large, subtriangular, with anterior and
posterolateral margins concave, anterior half of anterior ventral apotome dark. Posterior ventral apotome
large, elongated, triangular, posterior border dark brownish. Anterior and posterior ventral apotome about
same length, together longer than mid-ventral ecdysial line. Submentum smaller than anterior ventral
apotome, trapezoidal; posterior margin curved; anterior margin short, not cleft, almost flattened; both
anterolateral corners round.
Venter of head mostly yellowish orange. Stridulating files dark: anterior two-thirds relatively paler,
brownish; posterior third dark brownish and extending laterodorsad to mid-regions of lateral sides of parietal
sclerites. Stridulating files thickened, spaces between striae sparse. Each transverse striae on anterior two-
thirds of stridulating files consisting of series of short, longitudinal small ridges. Striae on posterior third
closely spaced, consisting of short, broken, transverse ridges not clearly defined. Lateroventral sides of
parietal sclerites bearing thick, long, acuminate setae and short, fine, tapered hair-like setae on anterior halves
(Fig. 4b). Series of muscle scars on lateral sides of parietal sclerites dark, very conspicuous (Fig. 4c).
THORAX. Thoracic nota densely covered by long, thick, dark blunt-setae and short, fine, tapered hair-like
setae. Pronotum yellowish orange; meso- and metanota dark brownish. Pronotum subdivided longitudinally in
middle, constricted laterally at posterior border. Blackish marks along lateral borders not reaching
anterolateral corners (Fig. 4e). Setae 22 on anterolateral corners inconspicuous, indifferentiable from regular
blunt-setae. Prosternal plate large, folded ventrad at posterior third, with lateral borders strongly incised.
Blackish marks along anterior border thinner near middle of each side. Posterior border of prosternal plate
concave, bearing broad blackish mark, surrounded by reddish brown area. Posterior prosternal sclerites not
present (Fig. 4f). Foretrochantins not forked, each bearing series of blackish, thick bristles (Fig. 4g).
Forefemur with small triangular scraper on middle of dorsomesal surface (Fig. 4h), presumably functioning as
stridulator rubbing against stridulating files on ventral side of head. Mesonotum with transverse ecdysial
suture on posterior third, extending anterolaterad toward anterior third of lateral borders (Fig. 4e). Diagonal
grooves pronounced, extending from anterolateral corners to near transverse suture. Posterior border bearing
wide, flattened blackish U-shaped mark on middle two-thirds; this median mark connected laterally with
lateral borders of mesonotum by thin, curved, blackish lines, with triangular brownish areas behind them (Fig.
4d). Metanotum with transverse ecdysial suture on posterior sixth, extending anterolaterad toward anterior
third of lateral borders. Diagonal grooves not as pronounced as those on mesonotum. Posterior border bearing
wide, thin blackish mark on middle two-thirds, not continuous with blackish marks along lateral borders
(Figs. 4d, e). Mesosternum bearing pair of ventromesal gills; metasternum bearing pair of ventrolateral gills
and single ventromesal gill (Table 4). Each thoracic tracheal gill with single stem.
ABDOMEN. Abdominal terga densely covered by thick, elongated, dark brownish club-hairs, with sparsely
scattered short, fine, hair-like setae among them (Fig. 4i). Abdominal segments III–VII bearing simple, long,
finger-like lateral gills on pleura (Fig. 4j). Abdominal segment III with one lateral gill; IV–VII with three
closely arranged lateral gills on each side (Table 4). Abdominal sternum I bearing pair of single-stemmed
ventral gills on each side (one lateral and one median), stems widely separated; segments II–V each bearing
one bifid ventrolateral gill and one single ventromesal gill, with latter situated slightly anterior of former; VI
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CHINESE HYDROPSYCHIDAE
bearing one ventrolateral and one ventromesal gill on each side, both single; VII–IX with no ventral gills
(Table 4).
FIGURE 4a–k. Diplectrona obscura. Larva (possible association)— a. Head, dorsal (with secondary setae on dorsum of
left parietal sclerite further enlarged, dorsal aspect); b. head, ventral (with secondary setae on lateroventral side of right
parietal sclerite further enlarged, ventral aspect); c. head, left lateral (setae on frontoclypeus not shown); d. pro-, meso-,
and metanota, dorsal (with secondary setae on dorsum of pronotum further enlarged, dorsal aspect); e. pro-, meso-, and
metanota, left lateral; f. prosternal plate, ventral; g. left foretrochantin, left lateral; h. right fore leg, ventromesal aspect,
showing fore femoral process; i. hairs on tergum of abdominal segment II (hl: hair-like setae; ch: club-hairs); j. lateral
gills (pleural gills) on left side of abdominal segment V, left lateral; k. ventral plates on sterna VIII and IX.
Ventral plates on abdominal segment VIII small, triangular, with yellowish pigmentation. Anterior part of
each plate VIII with short, thickened, conical golden setae; posterior border with long, thick, blackish bristles,
pointing posterolaterad. Area between plates VIII sclerotized and pigmented such that ventral plates on
segment VIII appearing as single sclerite. Ventral plates IX large, nearly semi-circular, with lateral borders
convex, mesal borders straight; yellowish. Golden setae on anterior part of each plate IX longer than those on
plates VIII; series of variably long, thick, blackish bristles distributed along almost entire length of lateral
border, pointing posterolaterad or posterad. Posteromesal corner of each plate IX without thickened golden
setae. Thickened golden setae on anterior regions of ventral plates VIII and IX bearing prominent setal
sockets (Fig. 4k).
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Diagnosis. This species can be differentiated from other Chinese Diplectrona species by the presence of a
series of dark muscle scars on the posterior whitish area of each parietal region.
Material examined. Guangxi: Long-lin County, Jin-zhong-shan Provincial Forest Preserve, Nong-heng
Gou, ~1.3 km N of Xi-she Village. N24.57867, E104.91399. 1140 m. 11-June-2004. L-F Yang, X. Zhou, C. J.
Geraci, K. Kjer, 8 specimens.
TABLE 4. Number and arrangement of gills in mature larva of Diplectrona obscura. Numbers in the table represent the
number of gills present on one side of a particular body segment. Gill types: a: simple, long finger-like gill; b: bifid-
stemmed gill; c: single-stemmed gill. * The metathorax only possesses one median ventral tracheal gill on the middle of
metasternum.
Acknowledgements
Dr. John Morse of Clemson University, USA, Dr. Karl Kjer of Rutgers University, USA, and Professor
Lianfang Yang of Nanjing Agricultural University, China, have provided critical advice on this larval
association work and made important comments on the manuscript. Changhai Sun of Nanjing Agricultural
University, China, did crucial work on adult morphology. I especially appreciate the great assistance from the
scientists and staff members of the following Chinese Nature Preserves: Nan-ling, Ding-hu-shan and Nan-
kun-shan Nature Preserves of Guangdong Province; Cen-wang-lao-shan, Jin-zhong-shan, Jiu-wan-da-shan,
Mao-er-shan and Shi-wan-da-shan Nature Preserves of Guangxi Province; Wu-yi-shan Nature Preserves of
Jiangxi Province; and Li-zi-ping and Da-feng-ding Nature Preserves of Sichuan Province. Special thanks to
Dr. Shaoying Liu of Sichuan Forestry Science Institute for his generous help organizing our Sichuan
exploration in 2005. This work was supported by the National Science Foundation (NSF DEB-0316504). COI
sequence analysis is supported by grants from the Natural Sciences and Engineering Research Council of
Canada (NSERC) and from Genome Canada through the Ontario Genomics Institute to Dr. Paul D. N. Hebert,
Biodiversity Institute of Ontario, University of Guelph, Canada.
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Thorax Abdominal segments
Mesothorax Metathorax III III IV VVI VII VIII IX
Lateral (pleural) - - - - 1a3a3a3a3a- -
Ventro-lateral -1c1c1b1b1b1b1c- - -
Ventro-mesal 1c1c* 1c1c1c1c1c1c- - -
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