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_ LEESGIENCES

Revision of the Caddisfly Genus Psilotreta
(Trichoptera: Odontoceridae)

C.R. Parker
and
G. B. Wiggins

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Revision of the Caddisfly Genus Psilotreta
(Trichoptera: Odontoceridae)

Ls
-
fi

LIFE SCIENCES CONTRIBUTIONS 144

Revision of the Caddisfly Genus Psilotreta
(Trichoptera: Odontoceridae)

C. R. Parker
and
G. B. Wiggins

ROM

ROYAL ONTARIO MUSEUM

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LIFE SCIENCES EDITORIAL BOARD
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Editor: Judith L. Eger

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Production editor: J. E. Hawken

Charles R. Parker was a postdoctoral associate, Department of Entomology, Royal Ontario
Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 and Department of Zoology,
University of Toronto, Toronto, Canada; current address is National Park Service, Great
Smoky Mountains National Park, Rt. 2, Box 260, Gatlinburg, Tennessee 37738, USA.

Glenn B. Wiggins is Curator-in-charge, Department of Entomology, Royal Ontario
Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6; and Professor, Department of
Zoology, University of Toronto, Toronto, Canada.

Canadian Cataloguing in Publication Data
Parker, C. R. (Charles R.), 1948—

Revision of the caddisfly genus Psilotreta
(Trichoptera: Odontoceridae)

(Life sciences contributions, ISSN 0384-8159 ; 144)
Bibliography: p.
ISBN 0-88854-332-8

1. Caddis-flies — North America — Identification.
2. Caddis-flies — Asia — Identification.

I. Wiggins, Glenn B. II. Royal Ontario Museum.
III. Title. IV. Series.

QL518.04P37 1987 595.7'45 C87-095015-0

Publication date: 15 November 1987

ISBN 0-88854-332-8

ISSN 0384-8159

100 Queen’s Park, Toronto, Canada M5S 2C6
PRINTED AND BOUND IN CANADA AT THE ALGER PRESS

Contents

Abstract l

Introduction I

Materials and Methods l

Locations of Specimens Examined Z

Genus Psilotreta Banks 3
North American Psilotreta 2)
Key to Adults of North American Species
Key to Larvae of North American Species

Psilotreta indecisa Group 1]
Psilotreta indecisa (Walker) 1]
Psilotreta frontalis Banks 13
Psilotreta labida Ross 18
Psilotreta rufa (Hagen) aA
Psilotreta rossi Wallace 3
Psilotreta amera (Ross) WS)

Asian Psilotreta 32
Key to Males of Asian Psilotreta 32
Psilotreta japonica (Banks) 34
Psilotreta chinensis Banks 34
Psilotreta kisoensis lwata 32)
Psilotreta locumtenens Botosaneanu
Psilotreta falcula Botosaneanu 40
Psilotreta quadrata Schmid 40
Psilotreta schmidi sp. nov. 4]
Psilotreta assamensis sp. nov. 4]
Psilotreta sp. 43
Psilotreta kwantungensis Ulmer 45
Psilotreta trispinosa Schmid 47
Species Not Examined 47
Phylogenetic and Biogeographic Considerations
List of Species 50
Acknowledgements ay

Literature Cited 53

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Revision of the Caddisfly Genus Psilotreta
(Trichoptera: Odontoceridae)

Abstract

Twenty species of Psilotreta are recognized, 6 from eastern North America and 14 from
Asia. Adults and larvae are described and illustrated for the North American species, and
keys are provided. For the Asian species, males and females of P. japonica (Banks), P.
chinensis Banks, and P. kwantungensis Ulmer, and larvae of P. kisoensis Iwata are
described and illustrated; observations are made on adults of P. falcula Botosaneanu, P.
locumtenens Botosaneanu, P. quadrata Schmid, and P. trispinosa Schmid, and on larvae
and pupae of P. locumtenens. Two new species are described from eastern India: P.
schmidi sp. nov. and P. assamensis sp. nov. A key to the males of the Asian species is
presented. The larva and pupa of an undetermined species from India are described.
Phylogenetic analysis indicates that the North American species form a monophyletic
group, and a cladogram based on characters of adults and larvae is presented for them.
Possible relationships among the Asian species are suggested based on characters of the
males.

Introduction

Caddisflies of the genus Psilotreta Banks, 1899, occur in
cool running-water habitats in Asia and eastern North
America. Adults are cryptic and infrequently collected, but
the larvae and pupae are often found in large numbers
because of the gregarious habit in some species of fastening
the cases for pupation in layers to the undersides of rocks.
Twenty species of Psilotreta are recognized, 6 from North
America and 14 from Asia. The larva of only 1 North
American species, P. labida Ross, has been described (by
Lloyd, 1921, as P. frontalis Banks), and that of another
species, P. rufa (Hagen), has been illustrated (by Wiggins,
1977, as P. sp.). Ross (1944) provided keys to males of 5
North American species, but the females and larvae have
not yet been distinguished. The purpose of this study was
to make North American species of Psilotreta identifiable

in larval and other stages and to place them in a phylogene-
tic context in the world fauna.

Males, females, and larvae are available for all 6 North
American species; we have studied also males of 10,
females of 6, and larvae of 3 species from Asia. Iwata
(1928) described as new 2 species from Japan on the basis
of larvae and cases. Adding to this base the published
descriptions of the other Asian species, we have made
limited comparisons of the known world fauna. Characters
of the male genitalia indicate that the 6 North American
species form a monophyletic group, and that this group can
be subdivided further into two subgroups of 3 species each
based on adult and larval characters. Insufficient informa-
tion is available to permit an adequate phylogenetic analy-
sis of the relationships among the Asian species.

Materials and Methods

Associated adult and larval material in the Royal Ontario
Museum forms the basis of this study. Specimens from
other collections, including types, have been widely used;
source and deposition of all specimens examined are noted
at the end of this section. Distribution maps include all
peripheral localities known to us, but only representative
localities are plotted in the central part of the ranges of

more common species; thus, not all records listed appear
on the maps.

Identification of adult males of the Nearctic species can
be accomplished without reference to genitalia, by the
presence of eversible lobes on the second segment of the
maxillary palpi of two species and on the dorsum of the
head of one species, but these characters may be difficult to

detect in a retracted condition. Clearing the head or the
genitalia in KOH solution will confirm identification.
Females are usually identifiable only by examination of
cleared genitalia. Characters used in larval identification
rely heavily on the standardized larval chaetotaxy
developed by Williams and Wiggins (1981), and work
most reliably with instar V larvae, although earlier instars
can be identified in some species. Counts of notal setae
were made at X 80 magnification and included all setae.
To avoid obscuring other details of the illustrations, not all
setae are shown in the figures. Colour patterns are helpful

in larval identification, but should not be relied upon
wholly because of the variability within species. No pupal
characters were found to be diagnostic at the species level,
but pupae can be identified by reference to the larval
exuviae which remain in the case at pupation. The criteria
of Wiggins (1977) were used for the distinctions made
between immature stages listed in the Material Examined
sections; these stages are indicated by the following abbre-
viations: L—larva; PP—prepupa; P—pupa; MMT—
metamorphotype.

Locations of Specimens Examined

ANSP Academy of Natural Sciences, Philadelphia,
Pennsylvania.

CLEM Insect Museum, Department of Entomology and
Zoology, Clemson University, Clemson, South
Carolina, per J. C. Morse.

CNC Canadian National Collection, Ottawa, Ontario,
per F. Schmid.

CU Cornell University, Department of Entomology,
Ithaca, New York, per Q. D. Wheeler.

pGp___D. G. Denning, Moraga, California.

Gas’ G. A. Schuster, Department of Biological Sci-
ences, Eastern Kentucky University, Richmond.

INHS _ Illinois Natural History Survey, Natural Resources
Building, Urbana, per J. D. Unzicker.

JCH J. C. Hodges, Roswell, Georgia.

KSBS-_ Kansas State Biological Survey, Lawrence.

LB L. Botosaneanu, Institut voor Taxonomische Zoo-
logie, Universiteit van Amsterdam.

mMcz Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts.

OSF O. S. Flint, Jr, Department of Entomology,
National Museum of Natural History, Smithsonian
Institution, Washington, D.C.

ROM Department of Entomology, Royal Ontario
Museum, Toronto.

UL Université Laval, Provancher Collection, Sainte-
Foy, Québec, per J.-M. Perron.

USNM United States National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C.,
perO7S=Elint Jn

UT University of Tennessee, Department of Zoology,
Knoxville, per D. A. Etnier.
VPI Virginia Polytechnic Institute and State University,

Blacksburg, per J. R. Voshell, Jr.
zim Zoologisches Institut und Museum, Hamburg, per
H. Strumpel.

Genus Psilotreta Banks

Psilotreta Banks, 1899:213, type species P. frontalis
Banks, by monotypy.

Astoplectron Banks, 1914:264—265, type species Hetero-
plectron borealis Provancher, by original designation.
Synonymy by Betten, 1934:240.

The adult is distinguished by having the separation of R,
and R, markedly basad of the separation of R, and R, in
both wings (Figs. 4-7, 20, 52). The larva is distinguished
by a pointed anterolateral corner of the pronotum, and
genae contiguous along most of the median ventral
ecdysial line of the head.

ADULT :
Medium sized (forewing 6—17 mm in length), black, grey,
reddish or yellowish brown. Head (Figs. 1-3, 50) short and

broad; antennae longer than forewing, slightly longer in
male than in female, scape subequal to length of head, first
two flagellar segments fused, with or without a suture line;
eyes glabrous, large, slightly larger in male. Setal warts
with long hairlike setae and/or scales; frons with ante-
rolateral setal wart along anterior margin of eye below
antennal socket, with or without a large median wart or pair
of warts on frontoclypeus; vertex with anteromesal warts
absent or situated on raised prominences separated by
narrow, deep fissure extending below base of antennal
sockets; narrow transverse anterior warts posterad of
antennae present or absent, larger transverse to oval pos-
terior warts near posterior margin of head, and elongate,
narrow posterolateral warts extending ventrally along pos-
terior margin of eye, or anterior and posterior warts appar-
ently combined into one large wart (Fig. 3), or posterior

Fics. 1-3. Psilotreta spp., adults. 1. P. indecisa: a,b, head of d—dorsal, lateral. 2. P. frontalis: a,b, head
of 6—dorsal, lateral; c, maxillary palpi of d, lobe everted. 3. P. amera, head of 6 —dorsal.

pair replaced by large eversible scale-covered lobes (Fig.
2). In some species vertex of male excavated (Fig. 50).
Maxillary palpi five-segmented in both sexes, long, with
apex of third segment reaching past base of scape, very
setose, sometimes with scales intermingled, and
unmodified or with dorsal eversible lobe on second seg-
ment of male of some species (Fig. 2c). Labial palpi with
first segment short, second and third long, subequal.

Tibial spurs 2, 4, 4. Wings (Figs. 4-7) elongate, fore-
wings longer and narrower than hindwings, densely hairy,
with or without scattered scales; discal cell usually present,
but absent in some species; Cu,, and Cu,, separate in
forewings of both sexes and in hindwings of female, but
undivided in hindwings of male; male with Cu,, fused with
M,,, and stems of Cu, and M fused in forewings; M, , »
divided in both wings of female in some species; stigmatic
region whitish opaque.

Male genitalia. Segment VII of some species with a
small eversible lobe at posterior corner of pleuron, usually

> — A

withdrawn and indistinct, or with narrow dorsoventral
invagination near posterior margin. Segment VIII
unmodified. Segment IX with anterolateral margin invagi-
nated into VIII, heavily sclerotized, much broader and
thicker below midline, with a few long setae postero-
laterally and ventromesally, dorsum subtriangular or pro-
duced into a long slender process extending posteriorly
beyond rest of segment, often finely granular, with sharp
steep sides, or produced laterally into a small overhanging
ledge, and with lateral borders fused with body of segment
X (these segments together forming an elongate ‘phallus
guide,’ open ventrally, and into which the phallus normally
is retracted); anal aperture in membranous roof of phallus
guide; inferior appendages two segmented—basal segment
elongate, cylindrical, slightly tapered, either somewhat
curved in ventral aspect and with or without a low median
lobe basally, or thick and quadrate in lateral aspect
(Schmid, 1959, pl. 4, fig. 8; Botosaneanu, 1970, pls. 36—
38), and apical segment short, rounded, tapered or foot
shaped (Mosely, 1942, fig. 8; Hwang, 1957, figs. 94, 95),

Fics. 4-7. Wings of Psilotreta spp. 4. P. indecisa 3.5. P. indecisa 2.6. P. rufa 3.7. P. rufa 2.

and with from two to many stout black teeth apically—both
segments with numerous long setae. Segment X consisting
of several pieces and processes variously developed in
different species; pair of rounded or elongate setal warts at
base; in dorsal aspect a bifid or entire, narrow median
process extending posteriorly, with a few setae apically;
thin, sclerotized lateral processes arising from median
process, each fused partially or completely along its dorsal
edge to the median process; lateral processes large,
rounded or pointed, and with or without anterior projec-
tions or processes, or very narrow and sinuous, or absent;
ventrally from the lateral processes arises a pair of stout,
curved sclerotized intermediate appendages (sensu
Schmid, 1980), which may be short and inconspicuous
(Fig. 5la) or, more often, large and recurved into a
ramshorn shape (Fig. 11a), and occasionally bifid; preanal
appendages either large and thin, somewhat foliaceous in
lateral aspect, widest just beyond base, and pointed api-
cally, or small and subcircular with long setae.

Phallus with phallotheca sclerotized, long and cylindri-
cal, curved, with one or both ends flared ventrally; endo-
theca short and membranous; parameres absent in North
American species (Fig. 11d,e), present in Asian species,
varying from long, slender, acute rods (Fig. 48d,e) to
broad, flat, bladelike structures (Fig. 55d,e); endotheca of
one species possessing a pair of small endothecal processes
arising dorsad of parameres (Fig. 51d,e); aedeagus mem-
branous with three or more eversible lobes, some species
having a ventral sclerite (Fig. 11d,e); phallotremal sclerite
long and delicate, not prominent.

Female genitalia (Figs. 12, 47). Sternum VIII rec-
tangular in ventral aspect, with a row of fine setae along
posterior edge. Sternum IX with conspicuous sclerotized
triangular or hemispherical posterior portion, divided long-
itudinally in some species; anterior portion rectangular and
usually clearly distinct from the posterior portion by less
sclerotization, less pigmentation, or different texture;
tergum IX fused laterally with anterior portion of sternum
IX, extruded posteriorly on each side as broad, rounded
lobes meeting in a posteromedian cleft, and flexibly joined
to posterior edge of sternum IX; apex of posterior portion
of sternum IX delineates narrow vaginal aperture. Tergum
X fused indistinguishably with tergum IX dorsally and
bearing two large, low, oval setal warts (appendages of
Schmid, 1980). Vaginal chamber elongate and extending
into segment VIII, with prominent posterior and anterior
vaginal sclerites (sensu Unzicker, 1968), the posterior
vaginal sclerites heavily sclerotized in some species and
less sclerotized to membranous in others, and the anterior
vaginal sclerites broadly rectangular or oval and connected
to the posterior by complexly folded sclerites.

LARVA (Figs. 8, 9)

Head without secondary setae, sclerites smooth and shiny,
or textured with pebbling or fissures; a prominent dorsal
ridge extending posteriorly from anterolateral corner of
head capsule at dorsal mandibular articulation, along inner
margin of eye to about two-thirds length of head; antenna
short and inconspicuous, situated near dorsal mandibular
articulation, bearing an apical papilla and seta; ventral
apotome short, genae contiguous for much of their length;
two heavily sclerotized plates situated on maxillolabial
complex at anterior border of ventral apotome between
cardines (Fig. 8c). Mandibles (Fig. 8d) very broad and
scoop shaped, each with apex having one or two points,
dorsal margin thin, bladelike, and extending farther mes-
ally than ventral margin; mandibles asymmetrical, with left
mandible having less pronounced apex and broader dorsal
mesal margin; both mandibles lacking mesal brush of
setae.

Thorax and legs with numerous secondary setae. Ante-
rolateral corner of pronotum produced into point that
ranges from long and acute (Fig. 25) to short or absent
(Fig. 30). Mesonotum with two plates, each with a low,
sharply defined ridge along posterior margin extending
from midline laterally to posteroventral corner of sclerite
and ending there abruptly at a narrow excision of the
posterior margin (Fig. 25) or at a light spot (weakness) in
the same area (Fig. 30). Metanotum with two transverse
sclerites—anterior sclerite broadly rectangular, much
larger, and with many more secondary setae than the
narrow posterior sclerite, which has essentially a single
row of setae—and an oval lateral sclerite with numerous
setae on each side of metanotum above pleurites; metanotal
pleurites with primary setae 10 and 11 long and prominent,
setae 19 and 20 fine and inconspicuous, secondary setae
absent. Each thoracic segment with sternum sclerotized,
the sternal sclerite extending completely across segment,
articulating behind coxae with posterior corner of notum,
and extending forwards to near anterior margin of segment;
prosternite of one solid piece, mesosternites and metaster-
nites subdivided into four plates that are readily discernible
only in very dark specimens. Trochanters composed of
four sclerites (Fig. 8e,f), protrochanters and mesotrochan-
ters with secondary setae (trochanteral brush).

Abdomen with segment I lacking gills and secondary
setae, a pale oval sclerite bearing paired setae 12, and a
median dorsal and 2 lateral spacing humps, the lateral
humps with setae 7 and 8 posteriorly and with a patch of
fish-scale-like sculpturing anteriorly (Fig. 9), each scale
consisting of a low crescentic mound armed with a row of
approximately 30 spines. Branched gills present dorsally
on segments II to VI, VII, or VIII, ventrally on II to VII or
VIII, and sometimes laterally on II to III. Lateral fringe

or SN a’
S et

Fic. 8. Larva of Psilotreta rufa: a, habitus—lateral, with detail of anal proleg; b, head and thorax—dorsal;
c, head—ventral; d, mandibles—ventral; e,f, right mesotrochanter—posterior, dorsal; g, larval case—lateral
with details of construction and posterior end.

Fic. 9. Spacing hump of first abdominal segment of Psilotreta labida, dorsal aspect, right side: a, * 50; b,
x 940.

Fic. 10. Pupal structures of Psilotreta spp. a, Asian species, anal processes—dorsal; b, Asian species,
mandible—frontal; c, North American species (P. indecisa), anal processes—dorsal.

present on III to VII. Lateral tubercles present on VIII,
variable in number. Dorsal sclerite of segment IX with 4
pairs of setae (2, 3, 1, 5 in dorsal to lateral sequence;
Williams, 1983), setae 2 and 5 large and dark, setae 3 and |
fine and pale; seta 7 laterad of dorsal sclerite. Anal proleg
with lateral sclerite large and rounded dorsally, with a
prominent ventral extension; seta | present dorsally, seta 6
on membrane between lateral sclerite and ventral sole
plate, setae 2, 5, and 7 on small dorsal plate, and seta 12 on
ventral sole plate; no secondary setae or spines on lateral
plate, dorsal plate, or ventral sole plate; anal claw slender
in lateral aspect with apex broadly rounded, with 8 primary
setae present and usually a brush of small secondary setae
present ventrally just beyond ventral sulcus, and with 2 to 3
accessory hooks present dorsomesally.

CASE (Fig. 8g)

Cylindrical, curved, and usually tapered, although cases of
mature larvae of some species have little or no taper; made
of coarse sand grains closely fitted, and on the inside, of
smaller sand grains filling interstices between the larger
grains and firmly attached with silken bracebands between
adjacent rock pieces (Fig. 8g, detail), resulting in a
remarkably sturdy case resistant to crushing. Posterior end
of case with (usually) a single sand grain attached over
opening and small spaces around edges allowing passage
of water.

PUPA (Fig. 10)
Reference to the Indian pupa in the following description is
to specimens of one undetermined species from India.
Mandibles (Fig. 10b; Wiggins, 1978, fig. 14.84) termi-
nating in a long, curved, whiplike style, the apex of which
may be hooked; mesal edge of both mandibles with 9 to 10
small teeth in Nearctic species and in P. locumtenens
Botosaneanu and with 4 to 7 larger pointed teeth in the
Indian species (Fig. 10b). Head with vertex lacking setae in
Nearctic species but bearing 2 pairs of setae in P. locum-
tenens and in the Indian pupa; Nearctic species with eye

bearing a seta posteriorly, P. locumtenens and the Indian
pupa lacking ocular seta; antennal scape with a basolateral
tuft of setae, the Indian pupa also having a dorsal tuft.
Pronotum bearing 2 to several setae dorsally on each side.
Mesonotum bearing 5 to 8 setae anterolaterally and 2 to 6
setae posteriorly between wing base and mesoscutellum on
each side in Nearctic species and in P. Jocumtenens, these
areas bearing 20 to 27 setae and 5 to 7 setae, respectively,
in Indian pupa. Metanotum with an anterolateral and a
posterolateral tuft of setae on each side.

Abdomen (Wiggins, 1978, fig. 14.85) with paired dor-
sal hook-plates anteriorly on segments III to VII, each
having a single hook, and posteriorly on segment V, each
having two hooks, hooks raised and projecting posteriorly
from anterior plates and anteriorly from posterior plates,
with apex of each hook blunt in dorsal aspect. Branched
gills distributed on segments as in larvae. Lateral fringe
arising on posterior corner of segment VI and extending
posteriorly along segment VII to end of segment VIII, the
filaments fine and silky, long on VI, short anteriorly on VII
and becoming longer posteriorly, very long on VIII.
Antennae of most species very long and extending ven-
trally to segment VIII, coiling around anal processes up to
two complete turns; antennae of some species entangled in
the long lateral fringe of segment VIII. Anal processes
undivided in Nearctic species (Fig. 10c) and in P. locum-
tenens, approximately Xx 10 longer than wide at mid-
length, tapering to a point and curved laterally at tips,
dorsoventrally flattened and bearing numerous small spines
and setae along lateral margins; in Indian pupa (Fig. 10a)
anal processes bifid near midlength, with mesal branch
short, straight, and bearing a few spines along mesal
margin, and with lateral branch longer, slightly curved
laterally at tip, and bearing small spines and setae along
lateral margin.

Pupal cases closed at each end by a single capstone but
with a small peripheral space for water to circulate through
case. Pupal cases are usually attached to a solid substrate,
and in some species large aggregations of pupae are found
attached in layers to the undersides of rocks.

NORTH AMERICAN PSILOTRETA

The six North American species of Psilotreta form a
natural group phylogenetically (see Phylogenetic and Bio-
geographic Considerations), as well as geographically. All
SiX species are restricted to small springs, creeks, and
rivers of the eastern forests.

Key to Adults of North American Species

1 Genitalia with elongate paired appendages
CBieee li) ivtale sass fs SSE el ees Ase Pepe: 2
1’ Genitalia lacking elongate paired appendages
(hign)s Bemaleses 250) 226 Sy. tke oi a
2(1) Forewing with R, straight or with a slight kink

at junction of crossvein r (Fig. 4); median dor-
sal process of segment X reduced, mem-
branous, and fused throughout length to lateral
processes (Figs. 11, 13, 15). indecisa subgroup

Jas Forewing with distinct kink in R, at junction of
crossvein r (Fig. 6); median dorsal process of
segment X prominent, elongate, and largely
free from lateral processes (Figs. 17, 19, 22).
FUG SUDETOUDRY awacrae Jomo eee se ag a 5

3(2) Maxillary palp with a membranous eversible
dorsal lobe on second segment, set with brush
of long silky setae; terminal segment mem-
branous, flexible, and subequal to combined
length of preceding segments (Fig. 2b,c) ....

a Maxillary palp unmodified; terminal segment
not membranous and flexible, not more than 2

x length of preceding segment (Fig. 1b) ....

I, fechas Sion VR: ND Re indecisa, p. 11

4(3) Head dorsally with a pair of large eversible
lobes (Fig. 2b), appearing as scale-covered
warts when retracted (Fig. 2a); segment IX with
pair of long, thin, pointed styles arising beneath
preanal appendages and extending posteriorly
beneath segment X (Fig. 13a); inferior appen-
dages with terminal segment bearing two to six
black teeth (Fig: 113¢)2 24. 2¢:.. frontalis, p. 13

4’ Head lacking dorsal eversible lobes (as in Fig.
1); segment IX without styles (Fig. 15a);
inferior appendages with second segment bear-
ing three to six black teeth, one (rarely two) of
which is much longer than the others (Fig. 15c)
RY eels, AR SOBs ie iatataen labida, p. 18

6(5)

6’

8(7)

98)

Head with anterior and posterior warts distinct
(as in Fig. la); intermediate appendages
C-shaped, pointed posterodorsally; lateral pro-
cesses broad and rounded, each with short or
long projection ventrally (Figs. 17a, 19a) . . .6
Head with one pair of large warts dorsally
occupying most of vertex, covered with black
scales laterally and golden scales mesally (Fig.
3); intermediate appendages comma shaped,
directed ventrally; lateral processes of segment
X long, thin, and sinuate, each with apex acute
and directed posteroventrally (Fig. 22a)......
NS etal MER TIRES Le atten Cate amera, p. 25

Lateral processes of segment X broad and
rounded, often with a short ventral projection
(Fig. 17a); dark reddish brown species ......

Lateral processes of segment X rounded api-
cally, each with a long, narrow ventral projec-
tion (Fig. 19a); light grayish brown species
i Rae, Std, Soil) eee rossi, p. 23

Forewing with R, straight or with a slight bend
at attachment of crossvein r (Fig. 5); M three-
branchedtin bothuwin@sS A224 2% ees 5 =a ac 8
Forewing with distinct kink in R, at junction of
crossvein r (Fig. 7); M two-branched in both
WHITES 75 eet SL.. r N 8 ok A ock 10

Sternum IX with anterior portion lightly scle-
rotized (apIXs, Fig. 12c), or with scattered
flecks of sclerotization resulting in a granular or
reticulate appearance (apIXs, Fig. 14c); pos-
terior vaginal sclerites lightly sclerotized (Fig.
P2D FARE UE SAIS RS | sertontays 9
Sternum IX with anterior portion heavily scle-
rotized (apIXs, Fig. 16c); posterior vaginal
sclerites heavily sclerotized (Fig. 16b,d)
ee oC toe ee. labida, p. 18

Sternum IX with anterior portion lacking granu-
lar or reticulate appearance (apIXs, Fig. 12c);
posterior vaginal sclerites broad in ventral
aspect (pvs, Fig. 12d); frontoclypeus with 5 to
TORSELACE: tier). alt cilwe cha chew - indecisa, p. 11
Sternum IX with anterior portion having granu-
lar or reticulate pattern (apIXs, Fig. 14c); pos-
terior vaginal sclerites narrow (pvs, Fig. 14d);
frontoclypeus with 20 to 30 setae...........

11(10’)

Hindwing with discal cell open (Fig. 20); light

grayish brown species .......... rossi, p. 23
Hindwing with discal cell closed (Fig. 7); dark
LEdiShiDTOwWNESPECleSeun aan yer eiey ee 11

Sternum IX with anterolateral corners deeply
invaginated (inv., Fig. 23c), tergum IX with
shallow indentation (ind., Fig. 23a); posterior
vaginal sclerites in ventral aspect with medial
edges nearly, parallel (Big. 23d)aea sae soe.
ET AR Oe ae eT Ae amera, p. 25
Sternum [X with anterolateral corners not
invaginated and tergum IX lacking indentations
(Fig. 18a,c); posterior vaginal sclerites in ven-
tral aspect with medial edges widely separated
anteriorly, converging posteriorly until they
meet or nearly meet at vaginal aperture (Fig.
TS i aay Pee tee of cats Pee rufa, p. 21

Key to Larvae of North American Species

Mesofemur with ventral seta 3 arising in line
with dorsal seta | or distad of it by no more than
approximately the distance between ventral
setae 4 and 5 (Fig. 24); head seta 17 subequal to
or greater in thickness than seta 15 and more
than one-half length of seta 15 (Fig. 25), gener-
ally dark brown; head black with whitish or
yellowish markings forming distinctive pat-
terns of broad dark or light longitudinal bands
that are continued to thoracic nota (Figs. 32-
SW cals Wii: AN go yd tes ei oN eg: p
Mesofemur with ventral seta 3 arising distad of
dorsal seta 1 by more than the distance between
ventral setae 4 and 5 (Fig. 28); head seta 17
much thinner than seta 15 and no longer than
one-half length of seta 15 (Fig. 29), colourless
to light brown; head uniformly reddish brown
or with pale areas laterally; thoracic nota uni-
formly reddish brown to pale yellowish, or with

2(1)

3(2)

indistinct darker markings marginally (Figs.
SOS iy theres (siteknerttans one ee a ee eee 4

Anterolateral corner of pronotum long and
acute (Figs. 25, 27); anteroventral angle of
mesepisternum generally pointed (Figs. 25,
27); head in dorsal aspect with broad black
stripe widest at anterior margin of head capsule
and either narrowing gradually posteriorly (Fig.
32) or of more or less uniform width throughout
or widening posteriorly (Fig. 34); frontocly-
peus uniionmlyblackssaemes eae aso ae 3
Anterolateral corner of pronotum short (Fig.
26); anteroventral angle of mesepisternum short
(Fig. 26); head in dorsal aspect with broad
black stripe constricted anterior to eyes and
narrowing gradually posteriorly; frontoclypeus
black except for light areas at the anterolateral
COmMersM(Fig 33) aes frontalis, p. 13

Mesonotum with posterior margin dark medi-
ally, fading laterally and becoming obscure
before reaching posterior inflection (Figs. 25,
32) scolour pattern, asnmy Fi. 5 Denes ee ee

cyanate: Setins Aa enc eae totes Spy Tet indecisa, p. 11
Mesonotum with posterior margin black at least
to posterior inflection (Figs. 27, 34); colour
pattern)as in: Fis: 34.-eee ee labida, p. 18

Head seta 17 much less than one-half length of
Seta JSiKigse305 36) eaaeoeeeeee rossi, p. 23
Head seta 17 about one-half length of seta 15
(Fige29a) is: tes oeaphelte doers ie Gere 5

Dorsum of head appearing domed between car-
inae (Fig. 31b); head as long as wide (Fig. 37)
. Preaeksgyeunerinn petty cipal amera, p. 25
Dorsum of head relatively flat between carinae
Fig. 29b); head longer than wide (Fig. 35) ...

Psilotreta indecisa Group

This group, characterized by the absence of parameres in
the male, comprises the six North American species and is
divided into the indecisa and rufa subgroups. The dis-
tinguishing characteristics in the indecisa subgroup—P.
indecisa (Walker), P. frontalis Banks, and P. labida
Ross—are as follows: R, is straight or has only a slight kink
at the junction of crossvein r in the forewings of both sexes
(Figs. 4, 5); M is three-branched in both wings of the
female (Fig. 5); the male has the median dorsal process of
segment X reduced, membranous, and fused throughout its
length to the lateral processes (Fig. 11); the larva has head
seta 17 dark brown, thicker than seta 15, and longer than
one-half the length of seta 15 (Fig. 25), and has the
mesofemur with ventral seta 3 more or less in line with
dorsal seta 1, or distad of it by approximately the distance
between ventral setae 4 and 5 (Fig. 24). The rufa sub-
group—P. rufa (Hagen), P. rossi Wallace, and P. amera
(Ross)—is distinguished as follows: R, is distinctly kinked
at the junction of crossvein r in the forewings of both sexes
(Figs. 6, 7); M is two-branched in both wings of the female
(Fig. 7); the median dorsal process of segment X in the
male is a prominent, elongate process largely free of the
lateral processes beyond its base (Fig. 17); the larva has
head seta 17 colourless to clear brown, much thinner than
seta 15, and no longer than one-half the length of seta 15
(Fig. 29), and has the mesofemur with ventral seta 3
situated distad of dorsal seta 1 by a distance clearly greater
than that between ventral setae 4 and 5 (Fig. 28).

Psilotreta indecisa (Walker)

Goera indecisa Walker, 1852:95. Holotype d (British
Museum [Natural History]), St Martin’s Falls, Albany
River, Hudson Bay, Ontario.

Leptocerus indecisus—Hagen, 1861:279.

Heteroplectron borealis Provancher, 1877:263. Holotype
2 (UL), Québec. NEW SYNONYMY.

Heteroplectron indecisus—Betten, 1934:237 (Calamo-
ceratidae).

Heteroplectron indecisum—Milne, 1936:80.

Psilotreta indecisa—Betten and Mosely, 1940:46—49, fig.
22.

Psilotreta indecisa—Ross, 1944:286, fig. 956.

Psilotreta indecisa—Schmid, 1983:46—47, figs. 258, 262—
269, 283, map 21.

This species is readily distinguished from the others in the
indecisa subgroup by the unmodified maxillary palpi of the
male, as well as by details of the genitalia of both sexes as
set out in the key. The larva is distinguished by the
attenuated posterior dark ridge of the mesonotum, the acute

anterolateral corner of the pronotum, and the colour
pattern.

ADULT

Forewing 10-14 mm in length. Antennae dark brown
basally, becoming paler apically, with numerous closely
appressed, short brown setae; antennae long and slender,
about 1.2 x forewing length in female, 1.7 x in male;
scape (Fig. la) 4 x longer than pedicel, first two flagellar
segments fused and 2.5 x length of pedicel. Maxillary
palpi (Fig. 1b) identical in both sexes, unmodified, with
many long erect setae on first three segments and shorter,
recumbent setae on last two segments; labial palpi with
long erect setae, first segment two-thirds length of second
and third segments. Frontoclypeus convex, appearing
slightly swollen ventrally, with a few (S—10) scattered
setae. Thorax dark brown to blackish dorsally, lighter
laterally and ventrally; legs light brown. Forewings brown
with faint irrorations; hindwings lighter with numerous
long, unmodified setae in anal area. Abdominal terga and
sterna light to dark brown; pleura whitish.

Male genitalia (Fig. 11). Segment VII with pleural
membranes forming distinct enlarged lobe posteriorly.
Segment IX with apical segment of inferior appendages
possessing five to nine short, black, mesally curved teeth;
number of teeth often different on right and left sides of
same individual. Segment X with lateral processes project-
ing posteriorly nearly as far as tips of inferior appendages;
lateral processes broad in lateral aspect (Fig. 11a), sub-
parallel in basal half, tapering to sharp, laterally curved
point apically, and with actual shape differing considerably
between specimens.

Phallus (Fig. 11d,e) long, with phallotheca heavily scle-
rotized, cylindrical and narrow basally, and having ventral
lip enlarged apically; endotheca short; aedeagus with large
dark phallotremal sclerite and indistinct ventral sclerite,
and with four membranous apical lobes when fully everted.

Female genitalia (Fig. 12). Genitalia typical for genus.
Anterior portion of sternum IX lightly sclerotized; pos-
terior vaginal sclerites broad and lightly sclerotized.

LARVA (Figs. 24, 25, 32)

Colour pattern with frontoclypeal apotome entirely black;
parietals each with black stripe along frontoclypeal
ecdysial lines, the two stripes meeting at coronal ecdysial
line and continuing posteriorly to margin of head capsule,
the whole appearing as broad, triangular black stripe,
parietals vary from reddish brown to whitish dorsally,
becoming black below eye, and fading to dark reddish
brown along ventral ecdysial line and along posterior

tie

vill

Fics. 11, 12. Psilotreta indecisa. 11. a,b, 3 genitalia—tlateral, dorsal; c, apical segment of inferior
appendage—ventral; d,e, phallus—lateral, ventral (aed.—aedeagus; bsw—basal setal wart; dmpX—median
dorsal process of segment X; end.—endotheca; inf. app.—inferior appendage; int. app.—intermediate
appendage; IXt—tergum segment IX; lat. pr.—lateral process of segment X; phall.—phallotheca; pr. app.—
preanal appendage; vs—ventral sclerite of aedeagus). 12. a,c, 2 genitalia—tateral, ventral; b,d, vaginal
apparatus—lateral, ventral (apIXs—anterior portion of sternum IX; avs—anterior vaginal sclerite; IXt—
tergum segment IX; ppIXs—posterior portion of sternum IX; ppIXt—posterior portion of tergum IX; pys—
posterior vaginal sclerite).

margin of head capsule; pronotum black to dark reddish
brown with pale spot posteromesally on each sclerite;
mesonotal and metanotal sclerites progressively lighter
posteriorly, so that posterior metanotal sclerite is light
yellowish; pleural sclerites dark reddish brown to black,
coxae and trochanters reddish brown with black dorsally,
femora black dorsally and reddish brown ventrally, and
tibiae and tarsi black.

Head (Fig. 32) longer than wide, narrowest anteriorly;
seta 17 dark brown, thicker than seta 15, subequal in
thickness to seta 14 (Fig. 25). Pronotum with projection of
anterolateral corner about 0.2—0.4 x middorsal length of
pronotum. Mesonotum with dark marking of posterior
ridge not extending laterad to posterior inflection; mes-
episternum with anteroventral projection prominent, long
(Fig. 25). Metanotum (Fig. 32) with anterior sclerite bear-
ing 32-47 setae, posterior bearing 17—23, each lateral
bearing 29-38. Abdominal segment VIII bearing 6-24
lateral tubercles on each side. Abdominal gills: dorsal—II
18-27, IIIf 22-38, IV 15-20, V 12-17, VI 0-7, VII 0-5;
ventral—II 12-23, III] 20-34, IV 14-15, V 19-20, VI 17-
18, VII 13—15. Larva up to 12 mm in length; head up to 1.1
mm in width across eyes. Larval case up to 14 mm in
length.

NOTES

We have examined the type of Heteroplectron borealis
Provancher. It is a female identical in all respects with P.
indecisa females; accordingly we designate the name a
junior subjective synonym of Psilotreta indecisa (Walker).

HABITAT AND BIOLOGY

According to Hilsenhoff (1981), P. indecisa lives ‘in
small, sand- and gravel-bottomed creeks, where larvae
feed on algae in the sand.’ Collection records of Rom field
parties indicate that this species also occurs in moderate-
sized rivers with strong current, and in water temperatures
of 14~-23°C.

DISTRIBUTION (Fig. 38)

The northernmost and westernmost of the Nearctic
Psilotreta: New Brunswick, New Hampshire, Nova
Scotia, Ohio, Ontario, Québec, Pennsylvania, Wisconsin.

MATERIAL EXAMINED AND ADDITIONAL RECORDS

NEW BRUNSWICK—Nipisquit R., 12.vili.1980, L
(ROM). NEW HAMPSHIRE—Coos Co., Colebrook,
19.vi-20.vii (fide Morse and Blickle, 1953). NOVA
SCOTIA—Baddeck Forks, 1.vii.1936, 1 2 (ROM).
OHIO—Portage Co., W. Branch, Mahoning R., v—vi (fide
McElravy et al., 1977). ONTARIO—Algonquin Prov. Pk:
str. crossing Hwy 60, 4.4 mi. e. Opeongo L. Rd,
11.v.1972, L (Rom 720118); Madawaska R. crossing Hwy
60, 29-31.v.1972 (preserved 6.vii.1972), P (ROM

720203); same, 27—28.1x.1972, L (Rom 720244).
Cochrane Dist.: Ft Albany, 52° N, 29.vi.1942, 1 2 (Rom);
Pitopiko R. between Longlac and Hearst on Rt. 11,
25.vi.1971 (emerged and preserved 27.vi—18. viii. 1971), L
P, 135 2 (Rom 710472); St Martin’s Falls, Albany R.,
holotype ¢d (BMNH, not examined). Grey Co., North Spey
R., Hwy 10s. Rockford, 21.v.1976, 2 P (Rom). Hastings
Co., Thurlow Twp, Latta, Moira R. below bridge, Conc.
VI, 15.x.1968, L (Rom). Nipissing Dist., Samuel de
Champlain Prov. Pk ca 9 mi. w. Mattawa, Pautois Cr. at
Hwy 17, 22.v.1977 (preserved 1.viii.1977), P (ROM
770036). Carleton Co., Richmond, Jock R., 20.v.1948, 3
3 (ROM). Sudbury Dist., 34.2 mi. w. Timmins, Eastman
Cr. crossing Hwy 101, 23.v.1972 (preserved 3. viii. 1972),
PP (RoM 720181). Thunder Bay Dist.: Geraldton, Creel-
man’s Cr., shallow rapids, 13.vi.1966, P (Rom); 13.6 mi.
sw. Geraldton crossing Rt. 11, 22—23.vi.1971, L P (Rom
710451); 11.1 mi. ne. Jellicoe, Sturgeon R. at Rt. 11,
22.vi.1971 (emerged and preserved 23.vi—26. vii. 1971), L
P, 2 6 4 2 (Rom 710450); Kakabeka Falls Prov. Pk,
Kaministikwia R., 19.vi.1971, L (Rom 710434). PENN-
SYLVANIA—Crawford Co.: Linesville Cr., 2.x.1965, L;
same, 5.vi.1968, 2 d 1 9; same, 22.v.1971, 1 5 1 9;
Same, 22:v21973,, 10"6% ‘same, -22:-v 1977, 10" 5re:
same, 22.x.1977, L; all USNM. QUEBEC—C .(henal?) Ste
Catherine, 15.vi.1955, 7 do (Rom). Gatineau Co.,
Wakefield, 4.vi.1935, 1 6 4 @ (Rom). Ile Ste Héléne,
5.vi.1965, P; same, 9.vi.1965, P; same, 10.v1.1965, P;
same, 18.x.1965, L (Rom). (St Helen’s Island) Montréal,
St Lawrence R., L (fide Corbet et al., 1966). Lake St John
Co., Mistassini, 10.vii.1954, 1 ¢ 2 @ (fide Harper et al.,
1975). Mistassini Post, 28.vii.1956, 1 d (ROM). Pare du
Mont Tremblant: 16—17.vi, 2 d (fide Roy and Harper,
1975); Riviére du Diable, 9.vi, adults (fide Robert, 1960).
Riviére Nabisipi, 11.vii.1962, 1 d (fide Harper et al.,
1975). Saguenay Co., 32 kme. Sept-Iles, Matamek R. nr
Matamek Res. Stn, 50°17’ N x 5°57’ W, 15,20. vii. 1974,
L PP P, 3 @ (fide Williams and Williams, 1979). Sher-
brooke Co., Sherbrooke, vi.1894, 2 (mcz). WISCON-
SIN—10,20.vi (fide Longridge and Hilsenhoff, 1973;
Hilsenhoff, 1981). Florence Co.: Pine R., 10.vi, adults;
same, 16.xi, L (fide Longridge and Hilsenhoff, 1972).

Psilotreta frontalis Banks

Psilotreta frontalis Banks, 1899:213. Holotype d (mcz),
Sea Cliff, New York.

Heteroplectron indecisum—Milne, 1936:80.

Heteroplectron gameta Ross, 1939:69, fig. 10.

Psilotreta frontalis—Betten and Mosely, 1940:49; as syn-
onym of P. indecisa.

Psilotreta frontalis—Ross, 1944:286—287, figs. 86, 9SSA;
H. gameta synonym of P. frontalis.

Psilotreta hansoni?—Morse et al., 1980, table 9; status
uncertain, probably synonym of P. frontalis.

Psilotreta sp.—Morse et al., 1980, table 9.

Psilotreta frontalis—Schmid, 1983:47, figs. 259, 274—
277, map 12.

This species is readily distinguished by eversible lobes on
the maxillary palpi and head of the male, and by details of
the genitalia of both sexes as set out in the key. The larva
can be distinguished by the short projections of the ante-
rolateral corner of the pronotum and of the mesepisternum,
the large number of setae on the metanotal sclerites, and
the colour pattern.

ADULT

In general appearance similar to P. indecisa; maxillary
palpi of female identical with P. indecisa, but maxillary
palpi of male (Fig. 2b,c) with eversible gland dorsally on
second segment and with numerous long silky setae form-
ing brush when lobe is retracted; first, second, and fourth
segments subequal and short, third segment subequal to
first two combined, fifth subequal to first four combined
and flexible, with dark appressed setae. Frontoclypeus of
female identical in shape with those of P. indecisa and P.
labida females, but with numerous (20-30) yellowish
setae; male with frontoclypeus a little swollen and with
thick patch of scales covering setal warts on either side next
to eyes, these patches extending dorsally between anten-
nae. Male with dorsum of head bearing two large, evers-
ible, scale-covered lobes (Fig. 2a,b) that appear as large
setal warts when retracted.

Male genitalia (Fig. 13). Abdominal segment VII with
pleural membranes forming enlarged lobe posteriorly. Seg-
ment IX with long, thin, acute style projecting posteriorly
from below insertion of preanal appendages, and with
apical segment of inferior appendage having two to five
black teeth. Segment X similar in all respects to P.
indecisa.

Female genitalia (Fig. 14). Anterior portion of sternum
IX with granular or reticulate appearance (Fig. 14c); pos-
terior vaginal sclerites narrow (Fig. 14d).

LARVA (Figs. 26, 33)

Coloration highly variable, most specimens as illustrated
(Fig. 33), but some very dark with light areas reduced, and
some very light with median dark stripe. A broad black
stripe extending dorsally over head capsule, widest at level
of eyes where it extends full width of head capsule between
eyes, gradually narrowing posteriorly; parietals with lateral
light areas extending to anterolateral corners of fron-
toclypeus in all but darkest specimens, and widening grad-
ually posteriorly; postgenae and genae very dark, becom-

ing lighter along midventral ecdysial line; pronotal and
mesonotal sclerites with lateral and middorsal black stripes
becoming narrower posteriorly as longitudinal light stripes
become larger posteriorly; mesonotum with black marking
of posterior ridge extending past inflection to ventral mar-
gin; metanotum with anterior and lateral sclerites brown,
posterior sclerite yellow; legs with coxae light brown, each
successive segment darker than the preceding, claws dark.
Pronotum with projection of anterolateral corner short
(Fig. 26), about 0.2 x middorsal length of tergite, and
usually slightly upcurved. Mesepisternum with antero-
ventral corner short. Metanotum (Fig. 33) with anterior
sclerite bearing 56—90 setae, posterior bearing 19-32, each
lateral bearing 37-50. Abdominal segment VIII bearing 4—
13 lateral tubercles on each side. Abdominal gills: dorsal—
I] 21-26, I 25-31, IV 17-23, V 12-16, VI 1-12, VII 0-
10, VIII O—9; lateral—II O—S, III 0-8; ventral—II 15-23,
II 21-31, IV 18-23, V 15-22, VI 15-22, VII 12-21, VIII
0-16. Length of larva up to 14 mm; width of head across
eyes up to 1.4 mm. Larval case up to 16 mm in length.

NOTES

Psilotreta frontalis was described in 1899 and P. labida in
1944; much confusion exists in the literature prior to 1944
concerning the identity of P. frontalis, and many of the
earlier records should be referred to P. labida. Unfor-
tunately, since the ranges of these two species are largely
sympatric (Fig. 38), it is not always possible to ascertain
which species an author actually dealt with, and in fact it is
clear that some authors dealt with both. Lloyd (1921),
Sibley (1926), and Betten (1934) all worked with material
collected at McLean Bog, New York. We have examined
specimens from the Cornell University collection used by
these authors and have found both P. frontalis and P.
labida represented but all specimens identified as P. fron-
talis. The description and illustration of larvae by Lloyd
lack the details concerning structural features used here to
distinguish these species, but the colour pattern of the larva
he illustrated is much more like that of P. labida than that
of P. frontalis. Sibley described only pupae, which cannot
be distinguished at the species level. Illustrations of the
male by Betten (1934, pl. 27, figs. 9,10; pl. 28, figs. 7,8)
are clearly of P. /abida rather than P. frontalis; the apical
segment of the inferior appendage has an obvious mesal
spine much larger than the other spines, and the lateral
aspect of segment IX shows no indication of the style found
in P. frontalis. We conclude that these three authors dealt
with a mixture of both species, but primariiy with P.
labida, which Lloyd and Betten, at least, illustrated as P.
frontalis.

Denning (1948) described Psilotreta hansoni on the
basis of a single male from Massachusetts; P. hansoni
differs from P. frontalis in lacking the style on segment IX.
Our examination of the holotype of P. hansoni revealed

Fics. 13, 14. Psilotreta frontalis. 13. a,b, 3 genitalia—tateral, dorsal (with detail of variation in lateral
processes of segment X); c, apical segment of inferior appendage—ventral. 14. a,c, 2 genitalia—lateral,
ventral; b,d, vaginal apparatus—lateral, ventral (apI[Xs—anterior portion of sternum IX; pvs—posterior
vaginal sclerite).

that it has the eversible lobes on the second segment of the
maxillary palpi and on the head as found in P. frontalis. In
addition, we examined the two male specimens from South
Carolina listed by Morse et al. (1980) as P. hansoni?; one
is clearly a specimen of P. frontalis in which the style of
segment IX is present; the other specimen lacks the style,
but has eversible lobes on the head and maxillary palpi and
in all other respects is indistinguishable from P. frontalis.
No other specimens of P. frontalis that we examined lack
the style. Although we cannot rule out the possibility that
P. hansoni is a valid species differing from P. frontalis
only by the absence of the style from segment IX, we
believe that it is more likely that P. hansoni represents an
unusual variant of P. frontalis; we therefore consider P.
hansoni to be a species of doubtful validity, probably a
synonym of P. frontalis.

HABITAT AND BIOLOGY

As discussed above, much of the published information on
the biology of P. frontalis probably refers to P. labida, or
to both species. Our observations indicate that P. frontalis
is found primarily in well-shaded springs and spring-fed
streams. Larvae can be observed crawling about in sandy-
bottomed areas, and pupae are found attached to the under-
surfaces of rocks in the stream or buried in the gravel up to
6 cm deep in marginal areas where no surface water is
present. In South Carolina instar III larvae were observed
(by C. R. Parker) feeding on Goniobasis snails; larvae
attacked the snails through the shell opening. In one
instance, two larvae were observed attacking the same
snail.

DISTRIBUTION (Fig. 38)

P. frontalis reaches the easternmost extent of North Amer-
ica and extends farther south than any other Psilotreta:
Connecticut, Delaware, Florida, Georgia, Massachusetts,
New Brunswick, Newfoundland, New Hampshire, New
Jersey, New York, North Carolina, Pennsylvania, Québec,
South Carolina, Tennessee, Vermont, Virginia.

MATERIAL EXAMINED AND ADDITIONAL RECORDS

CONNECTICUT—Hartford Co., Granby, McLean Wild-
life Refuge, 7.vi.1962, 2 ¢ (USNM). Litchfield Co., W.
Woodbury, 21.111.1952, L (osF). DELAWARE—Kent
Co., New Castle Co., Sussex Co. (fide Lake, 1984).
FLORIDA—Gadsden Co., Bear Cr. 1 mi. n. Hwy 65C,
13.v.1970, 2 6 1 2 (INHS). GEORGIA—Dawson Co.,
Amicalola Cr. at St. Pk, 17.v.1982, L, 1 6P 1 2P (Jcn).
Greene Co., Cable Branch, 8.i1.1982, L (cH). Lumpkin
Co.: Jakes Branch, 8.ii.1982, L; same, 14.v.1982, 2 dP
(JCH). Rabun Co.: Chattahoochee Nat. For. s. Tate City,

Tate Branch at Tate Br. Cpgrd, 16—-17.v.1970, P (osF);
same, 17.v.1970, P (Rom 700343). Union Co.: Vogel St.
Pk, Wolf Cr., 7.i11.1981, L (CLEM); same, 30—31.v.1980,
14 3d (cLeEmM). MAINE—Isle au Haut Twp, off Stonington
Hbr, Laundry Brook, 15.vi.1981, L (Rom 811098). MAS-
SACHUSETTS—1 ¢ (rom). Franklin Co.: Colrain, sm.
str. on Rt. 33, 796-V, 20.v.1972, L P (INHs); Leverett,
Roaring Brook, 6 6 (fide Neves, 1979); Sutherland, North
Mountain Pond Rd, 64 (fide Neves, 1979). Hampshire
Co.: Amherst, 11.iv.1953, L PP (osF); nr Chesterfield,
6.vi1.1958, L (osF); Cummington, 22.111.1954, L; same,
30.iv.1954, PP (osF); Middlefield, Factory Brook, 66
(fide Neves, 1979); Westhampton, str. nr South and Stage
Rds, 9.vi.1968, 2 2 (Rom). Suffolk Co., Jamaica PI.,
7.vi, 1 2 (Rom). NEW BRUNSWICK—Charlotte Co.,
Waweig R. at Hwy 765, 45°14’ N x 67°08’ W,
14.viii.1980, L (Rom 801048). NEWFOUNDLAND—
Avalon Peninsula: nr Bay Bulls, 21.vi.1961, 1 d 1 @ (fide
Denning, in litt.); same, 27.vi.1961, 1 2 (usNm); Long
Cove, 2.vi.1961, 1 3 (fide Denning, in litt.); Loon Bay,
12.vii.1961, 1 ¢& (fide Denning, in litt.). NEW
HAMPSHIRE—Coos Co.: Base Rd nr Ammonoosuc R..,
sm. Str... 9.v1i.1977,, L. PP PB, 1.6 (Grem); Cutlenyke:
Pinkham’s Notch, 8—9.vii.1977, 1 d (CLEM); Fabyan,
3. vill. 1958, L (osF); Jefferson Notch Rd, Abenaki Brook,
7-8.vii.1977, 3 5 1 2 (cLEM). Grafton Co., Plymouth,
Fox Inlet #3, 25.iv.1969, L (UsNM). Stratford Co.: Lee,
7.vi.1965, 1 3 (INHS); same, 30.vi (fide Morse and
Blickle, 1953). NEW JERSEY—Bamber Lake:
31.11.1962, L; same, 22.iv.1962, L (USNM). Mt Misery,
16.1x.1962, L (USNM). Ocean Co.: Lakehurst, Wrangle
Brook, 16.v.1962, PP P (USNM); sphagnum bog,
16.v.1962, P (ROM); Wrangle Brook Rd, 20.vi.1955, 1 d;
same, 23.vi.1955, 1 2 (cu); Old Hurricane Brook on
Beckerville Rd, 19.vili. 1975, L (GAs); trib. to Middle Br.
at bridge on Dover Rd, 17.vili.1975, L (KsBs). NEW
YORK—Herkimer Co., Old Forge, 31.vii.1905, 1 9;
same, 18.viii.1905, L (cu). Nassau Co., Sea Cliff, vi,
lectotype 6 (fide Banks, 1899). Sullivan Co., Roscoe, #5,
4.iv.1959, PP (osF). Tompkins Co.: Ithaca, Mud Cr. nr
Freeville (fide Lloyd, 1921); McLean, vi, 2 2P, 1 2 (cu);
1 mi. e. McLean, Sphaerium Brook, Lloyd-Cornell Reser-
vation, 16—23.vi, adults (fide Sibley, 1926); Termonch L.,
Trout Branch, #4, 4.iv.1954, PP (osF). NORTH CAR-
OLINA—Avery Co., Stacked Rock Cr. at F. S. Rd 192,
3.5 kms. F. S. Rd 1514, xii.1975, L (cLEM). Blue Ridge
Parkway: milepost 250.7, 17.1x.1958, L (OsF and ROM);
milepost 251, 22.v.1959, L PP P, 1 6 4 2 (osF); spring,
2.1x.1959, L (OSF); sm. str. above falls, 2.ix.1959; L
(ROM). Buncombe Co.: Bent Cr. nr Asheville, 4 d; same,
20.v. 1950.1, 1 «dizvsame, 21-v21950- 1 9: same;
22.v.1950, 2 5d; same, 23.v.1950, 1 3; same, 24.v.1950,
36 1 2; same, 26.v.1950, 8 6 6 2; same, 29.v.1950, 2
6; same, 10.v.1951, OP; same, 13.v—2.vi.1951, 36;

same, 14.v.1951, 1 d; same, 15.v—2.vi.1951, 2 2; same,
[7ve1951> 1" 2 same, 20:v 1951, 1 oP; 5 6; ‘cases;
same, 2l.v.1951, 1 O="same, 22.v.1951, 2d: same,
13:vi (951 ISG: sames 6.vi. 1951, 1 2s same,
25.v.1953, 1 2 (cu); Black Mtn, 13.v.1952, 1 2; same,
seve l952e 1S Se e= "same, L7-v.1952> 1 3; same,
19.v.1952, 2 2: same, 20.v.1952, 1 6; same, 21.v.1952,
lS same, 24%v. 19525 1% "same, 25-v.1952, L (eu):
Jackson Co.: Cashiers, Greens Cr., 11.1x.1958, L; same,
14.1x.1958, L (Rom). Macon Co.: Highlands, 3-
8.vi.1961, 5 6 2 2 (USNM); unnamed cr. 4 rd mi. n.
Highlands on U.S. Hwy 64, NC #14, 17.v.1979, P
(CLEM); Clear Cr., 10.1x.1958, L (OsF); same, 6.vi.1961,
1 2 (UsNM); Green’s Cr., 19.v.1959, P (OSF); Whiteside
Cove, 11,14.1x.1958, L (osF); Highlands Biol. Stn,
1.vii.1958, PP P (OsF); same, 21.11.1981, PP (CLEM); at
Highlands inflow str. s. end L. Revenel, #12, 17.v.1979,
P (CLEM); str. entering reservoir, 26.iv.1980, L (GAs);
Nantahala Nat. For., cpgrd, 20.vi.1981, P (CLEM); Skitty
Cr., nw. Highlands, 20.vi.1981, L P (CLEM); ca 5 mi. w.
Highlands’ on’ U:S. 64, NC ‘#11, 16.v.1979, L PP P
(KSBS); same, 31.viii. 1980, L (CLEM); same, 20.vi.1981,
L P (cLEM). McDowell Co.: Crabtree Meadows, sm. trib.
above falls, 2.ix.1959, L (Rom); Blue Ridge Pkwy, 5—
6.vi.1962, 2 d 2 2 (USNM). Orange Co., Chapel Hill, 1 ¢
(cu). Swain Co., Great Smoky Mtns Nat. Pk: Couches Cr.
at Rt, 4415 2x. 1967; L (Rom); Deep’ ‘Cr: ‘Cperd;
Juneywhank Falls, 21.v.1970, | 2 (Rom 700366); New
Found Gap along Little Pigeon R., 13.vi.1935, holotype ¢
Heteroplectron gameta Ross (fide Ross, 1939);* Smoke-
mont Cpgrd, 11—14.v.1970, 3 2 (osF); Smokemont Cpgrd
at Bradley Fk of Oconaluftee R., 2.x.1967, L (ROM).
Transylvania Co.: Davidson R. nr Ecusta, 3 ¢; same,
7.v.1952, 1 3d; same, 14.v.1952, 1 3 (cu); White Water
R. at Rt. 171 n. Salem, 18.v.1970, 2 2 (Rom 700353);
same, 2 2 (USNM). Yancey Co.: Black Mtns, Hemphill
Spring, 4760 ft, 1—-2.vii.1979, 1 d (CLEM); South Toe R.,
Black Mtn Cpgrd, 8.viii. 1980, L (CLEM); sm. feeder str. ca
3 mi. n. Blue Ridge Pkwy Cperd (se. foot of Mt Mitchell),
NC #4, 14.v.1979, PP P (KsBs); sm. trib., Black Mtn
Cpegrd, 8.vili.1980, L P (cLEmM). PENNSYLVANIA—
Carbon Co.: e. Hickory Run St. Pk, Laurel Run crossing
Rt. 534, 20.iv.1968, PP (Rom); Swamp Run crossing Rt.
534, 20.iv.1968, L (Rom). Chester Co., Atglen, Glen
Run, 2—3.vi.1977, 3 3 (CLEM). Lancaster Co., Welsh
Mtn, New Holland Watershed Area, 19—20.vi.1978, 3 ¢ 2

* Ross’s designation for this locality appears to be in error.
Newfound Gap is on the border between North Carolina and
Tennessee, but Little Pigeon River is in Tennessee about 10 km
downhill from the Gap. In Tennessee, the nearest named creek
to the Gap is Walker Camp Prong; in North Carolina, the
nearest named creek is Beech Flats Prong.

2 (CLEM). Lycoming Co., 5 mi. n. Trout Run, str. beside
Rt. 15, 7.v.1968, PP (Rom). Tioga Co., Liberty,
29.v.1939 (fide Hyland, 1948). QUEBEC Argenteuil
Co., Harrington, str. in Can. Ind. Pulp For., 21.vii. 1966,
L (ROM). Parc du Mont Tremblant: 16—17.vi (fide Roy and
Harper, 1975); Riviere du Diable, 9.vi (fide Robert, 1960);
Tremblant, str. 2 mi. nw. Riviere du Diable, 16.vii.1969,
L (Rom). Rouville Co., Mont St-Hilaire, West Creek,
1966-1968, L, 3 d 2 2 (Rom). St-Laurent (fide Roy and
Harper, 1979). Systeme du St-Laurent, des Outaouais (fide
Roy and Harper, 1979). SOUTH CAROLINA—Barnwell
Co.: Savannah R. Plant, Upper Three Runs Cr. at SRP Rd
8-1, 4.x1i.1976, L; same, 29.ii1.1977, 2 6; same,
IQMiwaO77e 1-2] same. Suv 197t 2) 2 same, “12—
13.iv.1979, 2 d; same, 27.x.1979, L; same, 8.1.1980, PP;
same, 7.11.1980, PP; same, 24.iv.1980, P, 1 2; same,
4.x.1980, L; same, 18.x.1980, L (CLEM). Oconee Co.::
Chattooga R., e. fk, U.S. Fish Hatchery, 28.111.1980, PP;
same, 2.iv.1981, L; same, 29.v.1981, 1 2 MMT (cLEM);
Chattooga R. at Ellicott’s Rock, 27.ix.1978, L (CLEM);
Crane Cr. at Tamassee Rd 3.8 mi. e. SC #107, 7.1x. 1980,
L (CLEM); n. Salem on Rt. 171, glades area, 18.v.1970, L
(ROM 700352); Stumphouse Tunnel, 7 mi. w. on SC #107
from SC #28, 22.xi.1976, L (CLEM); 6 mi. s. Walhalla,
Crane Cr. n. Tamassee, U.S. Fish Hatchery, 26—
27.v.1980, 2 d (CLEM); s. Walhalla Fish Hatchery and Rt.
109, str. crossing rd to Burrell’s Ford Cpgrd, 19.v.1970, L
PP P (Rom 700351). Pickens Co.: Clemson, trib. Camp
Cr., 20.i1i. 1963, P (CLEM); 9 km nw. Clemson, Clemson
Univ. For., Wildcat Cr., 1968-1969, 1 5 1 2; same,
28.111.1973, L P; same, 22.ix.1976, L; same, 22.1x.1976,
L; same, 21.1x.1978, L; same, 21.1x. 1978, es same,
5.x.1978, L; same, 11.iv.1979, PP; same, 5—6.v.1979, 5
6;same, 5.x.1979, L; same, 13.iv.1981, LPP P,767 2
MMT (cLEM); Fox Cr., Six Mile, 18.x.1967, L; same, 18—
19.x.1977, L; same, 19.x.1977, L (CLEM); Indian Cr.,
Issaqueena For., 13.iv.1979, P (cLEM); Rock Quarry,
28.11.1968, L (CLEM); Six Mile Cr., 8.ix.1965, L (CLEM);
Wildcat Cr., 24.1x.1975, L; same, 5.5 mi. nw. Clemson,
28.i11.1973, L P (cLEM). Sumter Nat. For.: str. in Yellow
Branch Picnic Area, ca 1500 ft, 19.v.1970, L P, 1 6
(USNM); same, L P, 1 6 (Rom 700356). TENNESSEE—
Carter Co., Roan Mtn St. Pk, outflow of seepage area nr
trib. Dave Miller Hollow Branch, 7.v.1977, L P (ut).
Davidson Co.: Little Marrowbone Cr., 14.iv.1954, L;
same. 14.iv.1955, 1°92: same, 2.v. 1955, 12 (fide
Edwards, 1966). Dickson Co.: Montgomery Bell St. Pk,
str. at entr., 17.iv.1955, L (fide Edwards, 1966); Green-
briar Cove, 4.iv.1941, L P (osF). Monroe Co., str. at
Betsy Coker restaurant, Rt. 68, 22.iv.1977, P (uT).
Sequatchie Co., Sequatchie R., 26.vii.1953, L (fide
Edwards, 1966). Sevier Co., e. slope of Webb Mtn,
George Cox property, 17.iv-17.v.1975, 2 5 (GaAs). Great
Smoky Mtns Nat. Pk: Cole Branch of Little R. at Rt. 441,

1.x.1967, L (ROM); Jakes Branch nr Elkmont Cpegrd,
LOsiv21 981,14, (GAs); Little, Regat) Elkmont’€perd:
10.iv.1981, L (GAs); spring 0.5 mi. n. Greenbriar Ranger
Stn; J-iv: 1977, J, (ksss),., Unicoi |Co,, ponds2. mi up
Rocky Fork Cr., U.S. 23, 7.v.1977, L P (ut). Williamson
Co., Leipers Fork Cr., 24.vii.1953, L (fide Edwards,
1966). VERMONT—L (cu). Windsor Co., just s. West
Bridgewater, sm. str. crossing Rt. 100, 26.vii.1969, L
(ROM 690376). VIRGINIA—Str. on Rt. 104, 0.4 mi. e.
junction with 22, 2.v.1970, 1 ¢ (INHs). Fairfax Co.,
George Mason Univ., I1.vi.1973, | d (INHS). Fauquier
Co.: Catharpin Cr., Jackson Hollow, 8.v.1962, PP P;
spring, 10.v.1974, L PP P; upper springs and creek, Rt.
629, 9.vi.1962, LP, 1 6 1 2 (USNM). Frederick Co., 8 mi.
nw. Winchester, vi. 1976, L (USNM). Giles Co., Hunter Cr.
at Mountain L. Biol. Stn, 18.i1v.1968, L (Rom). Grayson
Cos, Laurel Cr., Rte 603, 22civ1979 (emerged
14.v.1979), P,2 d 1 2 (vet). Rockbridge Co., Cave Mtn
Lake Area #12, 29.111.1957, PP (osF). Shenandoah Nat.
Pk, Skyline Dr. milepost 79, 24.vi.1961, P (USNM). Smyth
Co.: Hungry Mother St. Pk, Marion, 26.iv.1979, PP P
(UT); n. side White Top Mtn, 21 mi. n. Grayson Co. Line
on s. side Virginia Co. Rd 600, sm. str., Va #2,
11.v.1979, L (ksBs); Jefferson Nat. For., Mt Rogers, sm.
str. and seepages 1.4 mi. n. Grayson Co. Line on Rt. 600,
Va #1, 10.v.1979, L(KsBs); sm. str. nr trail to Mt Rogers,
Grindstone Cpgrd, Va #4, 11.v.1979, PP (KsBs). Spot-
sylvania Co., unnamed trib. Massaponax Cr., | mi. s.
4-mile Fork, Rt. 208, P (USNM). Washington Co., Straight
Branch, Beartree Camp, 12.vi.1979, | d 1 2 (USNM).

Psilotreta labida Ross

Psilotreta labida Ross, 1944:287, fig. 954A,C. Holotype
S (INHS), Cedar River near Indian Lake, Adirondack
State Park, New York.

Adults of this species are separated from P. indecisa and P.
frontalis by the modified maxillary palpi and unmodified
posterior warts on the head of the male, and by details of
the genitilia of both sexes as set out in the key. The larva
can be distinguished by the acute projection of the ante-
rolateral corner of the pronotum, the long, blunt mes-
episternal projection, the extension of the black marking of
the posterior ridge of the mesonotum at least as far as the
posterior inflection, and the colour pattern.

ADULT

Frontoclypeus of female as in P. indecisa; frontoclypeus of
male similar to that of P. frontalis but with patches of
scales less dense. Maxillary palpi identical with those of P.
frontalis in both sexes. Head of male lacking large, scale-

covered eversible lobes, instead with pair of lunate setal
warts (as in Fig. la). Male with setae on undersurface of
anal area of hindwing slightly flattened.

Male genitalia (Fig. 15). Segment VII with pleural
membranes bearing a distinct enlarged lobe posteriorly.
Segment IX lacking posteriorly projecting style; inferior
appendage with apical segment bearing two to five short,
stout black teeth laterally and apically and one long,
pointed, mesally directed tooth, which (rarely) may be
paired. Segment X similar to P. indecisa and P. frontalis.

Female genitalia (Fig. 16). Anterior portion of sternum
IX heavily and uniformly sclerotized; posterior vaginal
sclerites heavily sclerotized and broad.

LARVA (Figs. 27, 34)
Colour pattern similar to P. indecisa, but dorsal black
stripe of head much broader posteriorly with little or no
taper from front to rear; extent of light and dark areas
variable among populations, some having light areas
reduced as to appear almost completely black, others with
light areas very large so that nota are predominantly light.
Pronotum with projection of anterolateral corner acute,
shorter in most specimens of P. labida than in most
specimens of P. indecisa but with the same range of size
overall. Mesonotum with posterior ridge black as far as
posterior inflection, black stripe extending ventrally past
inflection to ventral edge of sclerite; mesepisternum with
anteroventral projection long and narrow. Metanotum
(Fig. 34) with anterior sclerite bearing 29-66 setae, pos-
terior bearing 17—30, each lateral bearing 24-43. Abdomi-
nal segment VIII bearing 8-18 lateral tubercles on each
side. Abdominal gills: dorsal—II 20-32, III 28-46, IV 18—
28, V.14-29, VI 10-29, VII 5—16, VIII 0-10; lateral—II
0-6, III 0-9; ventral—II 13-28, III 24-34, IV 18-33, V
18-30, VI 16-27, VII 16-25, VIII 13-16. Larva up to 13
mm in length; head up to 1.2 mm in width across eyes.
Larval case up to 16 mm in length.

HABITAT AND BIOLOGY

As discussed under P. frontalis, many of the published
observations on the habitat and biology of that species
actually refer to P. labida, or to mixtures of P. frontalis
and P. labida. We have found that larvae of P. labida
occur in small spring-fed streams 1—3 m wide and in large
rivers at least 15 m wide. Larvae are found on the sides and
lower surfaces of rocks and are notable for their markedly
gregarious habits at pupation. In large populations it is not
uncommon to find a single rock with several layers of pupal
cases attached one atop another with old cases from pre-
vious years still attached on the lower layers.

Fics. 15, 16. Psilotreta labida. 15. a,b, 3 genitalia—lateral (with detail of variation in lateral process of
segment X), dorsal; c, apical segment of inferior appendage—ventral. 16. a,c, 2 genitalia—lateral, ventral;

b,d, vaginal apparatus—lateral, ventral (apI[Xs—anterior portion of sternum IX; pvs
sclerite).

posterior vaginal

DISTRIBUTION (Fig. 38)

P. labida has a more restricted range than P. indecisa or P.
frontalis, extending neither as far north nor as far south:
Connecticut, Georgia, Kentucky, Maine, Maryland, Mas-
sachusetts, New Hampshire, New Jersey, New York,
North Carolina, Pennsylvania, Tennessee, Virginia, West
Virginia.

MATERIAL EXAMINED AND ADDITIONAL RECORDS

CONNECTICUT—Tolland Co., Storrs, 10.vi.1954, 1 @
(INHS). GEORGIA—Rabun Co., Tate City, Tate Br.
Cperd, 17.v.1970, P (preserved 22.vi.1970) (ROM). KEN-
TUCK Y—McCreary-Wayne Co. Line, Little S. Fork at
Cumberland R., mi. 12.5 at Corder Cr., 1.xi.1980, L
(GAS). MAINE—Penobscot Co., Passadumkeag,
29.vi.1956, 1 2 (cu). MARYLAND—Garrett Co.:
6.vi.1931, paratype d (USNM); ca 1.5 mi. e. Friendsville,
river beside rd, 9.v.1968, P (Rom). MASSACHU-
SETTS—Hampshire Co.: N. Amherst, Cushman Br.,
4.v.1954, PP (osF); Middlefield, Factory Brook, v, vi,
3 6 (fide Neves, 1979); Pelham, 10.vi.1939, 1 d (DGpb);
Williamsburg, Rt. 143, 16.vi.1958, L PP P (osF). NEW
HAMPSHIRE—Coos Co.: Ammonoosuc R., Zealand
Camp, #117 112vi. 1957, PP: same; 25..vii0. 1957, L (Osr);
Jefferson, 28.vi.1965, 8 d6 2 2 (iINHS); Dundee, #26,
14.vi.1957, P (osF). Grafton Co., Whitcheville Brook nr
Benton, 21.vi.1941, 2 d paratypes (fide Ross, 1944).
Stratford Co.: Durham (fide Morse and Blickle, 1953); Lee
(fide Morse and Blickle, 1953). NEW JERSEY—Passaic
Co., Passaic, 8.vi, 1 d (mMcz). Sussex Co., Stokes St.
For., beaver pond at Big Flatbrook Cr. by Steam Mill
Camp, 3.ix.1975, L (ksBs). NEW YORK—Adirondack
St. Pk: Cedar R. nr Indian L., 20.vi.1941, holotype ¢,
allotype 2,2 ¢ 1 2 paratypes (fide Ross, 1944); small cr.
nr Tahawus, 20.vi.1941, paratype d (fide Ross, 1944).
Albany Co., Nassau, 2.vi.1906, 2 6 1 2 (cu). Beaverkill
R., e. branch, 6.vii.1959, L (osF). Chemung Co., Van
Etten, McCorn Cr. CYL-5, 27.vii.1975, L (cu). Dela-
ware Co.: Beaverkill R. at Rt. 17 (exit 92) nr Simcoe Stn,
19.v.1979, L PP (Rom 790002); same, 22.v.1979, P (ROM
790003); Horton, emerged 8.v.1976, 2 6 1 @; same,
5.vi.1966, 4 6 4 2; 2 2 (usNM). Hamilton Co., Wells,
17 -viel92Z5, 1 6 (cu): Sacadaga’ RiS Sport: Isies
19.vi.1900, 1 5 (mcz). St Lawrence Co.: Ogdensburg,
11.viii. 1906, L; same, 21.viii, empty cases (cu). Steuben
Co.: 4 mi. wnw. Avoca (just w. Bloomerville), Castle Cr..,
#14, 17. vii. 1978, L (kKsBs); same, 29.x1i. 1977, L (KsBs).
Sullivan Co.: Rockland, 1,2,4.iv.1959, L; same,
6.vi. 1959, P; same, 6.xi1.1959, L PP (osF); Roscoe, Little
Beaverkill R., 13.11.1977, L 13 6 2 MMT (usNm); same,
6.vi. 1959, P (OsF). Tompkins Co.: Danby, Michigan Hol-
low Str., 3.vii.1959, L; same, 23.i1x.1956, L (OSF); Free-
ville, 7.11.1915, 1 2; same, 6.vii.1913, 1 6 1 2 (cu);

Ithaca, 8.vi, 1 2° (eu); Mekean, 1°9ssame, 23vil LP:
same, 28.vii, d P (cu); same, 13.vi.1935, paratype 6 (fide
Ross, 1944); same, 1.v.1959, PP (osF); McLean Bogs
Reserve, 3.vi.1939, 2 2 (cu); McLean Reserve,
Sphaerium Brook, 22.vi.1956, P (OSF); same, 29.v.1959
(preserved 23.vi. 1959), L P (ROM); same, PP (osF); Mich.
Swamp, 6.x.1914, 1 2 (cu); Slaterville Springs, Wild-
flower Preserve, 7.vi.1958, P (OsF). Warren Co., North
Cr., 4.vii.1918, 1 2 (cu). NORTH CAROLINA—Bun-
combe Co.: Black Mtns, 31.v-—4.vi.1912, 1 2; same,
vi.1912, 1 3d (cu); Swannanoa, 26.v.1933 (fide Denning,
1950). Swain Co.: Great Smoky Mtns Nat. Pk, 7.vii.1961,
2 6 2 2 MMT (usNM); Bryson City, Deep Cr. at Deep Cr.
Cperd, 131x19585 9 gsamen 2 lv. 1970) Ee Pees (Rom
700365). ONTARIO—Grey Co., North Spey R., Hwy 10
just s. Rockford, 21.v.1976, P (Rom). PENNSYL-
VANIA—3 @ (cu). Monroe Co.: Analomink, 6.vi. 1907,
1 3 (cu); Ole Bull Park, 1 ¢ (cu); same, 11.vi.1947,6 3
6.2 (cv); nr Swiftwater (Lot 258), 1928, 2 ¢ 1 92
paratypes (fide Ross, 1944). Potter Co., 10 mi. n. Cou-
dersport, str. crossing rd 0.5 mi. w. Rt. 44, 15.1v.1968, L
(ROM). Union Co., Penn’s Cr., 16.v.1938, paratype d
(fide Ross, 1944). Wayne Co., Gouldsboro, 19.1x.1958, L
(ROM). TENNESSEE—Blount Co., trib. to Little R. at
Townsend, 19.11.1973, L P (ut). Cheatham Co., Big
Marrowbone Cr., 2.v.1954, 2 d (fide Edwards, 1966).
Davidson Co., Little Marrowbone Cr., 15.iv.1954, adults
(fide Edwards, 1966). Dickson Co.: str. at entr. to
Montgomery Bell St. Pk across U.S. 70, 18.iv.1955, 1 3
(fide Edwards, 1966); same, 2.v.1975, L P (ut); same, L P
(GAS). Franklin Co., 4 mi. w. Winchester, Wellington
Mills, Owl Hollow Cr. 1.5 mi. n. Rt. 50, 14-15.v.1970, L
PP P (Rom 700338). Grundy Co., Monteagle, 12.vi, 1 @
(cu). Hamilton Co., Chattanooga, Rt. 11, Sta. 8, Tenn.
#490a, 17.xii.1956, L (UsNM). Marion Co., Martin
Spring, |.viii.1955, L, 3 ¢ 6 @ (fide Edwards, 1966).
Overton Co.: 0.5 mi. w. Alpine on Rt. 52, Means Cr.,
30.1x.1967, L; same, 13.1v.1978, L (ROM 780063). Pickett
Co., 7.2 air mi. n. Jamestown, Wolf R., second bridge on
county rd offiUeS. 1275 22:xs1977,, (um), Sevier Cox
Elkmont, fk Little Pigeon R., 27.v.1934, paratype d (fide
Ross, 1944); e. slope Webb Mtn, George Cox property,
17.iv-17.v.1975, 2 3 (uT). Wayne Co., Eagle Cr. at dirt
rd 6.1 mi. from Lookout Tower off Clifton Turnpike
(between Tenn. #114 and U.S. 64), 15.i11.1972, L (ur).
VIRGINIA—Bath Co., ca 5 mi. n. Mountain Grove, Back
Cr. on Rt. 600, 2.vi.1972, 1 o (UsSNM); Curles Neck
Bridge, 19.iv.1938, paratype ¢ (fide Ross, 1944). Giles
Co.2 Little Walker. Cr. sateRt: + [00% Pearisbure:
10.viii. 1968, L (Rom); White Gate, Walker Cr.,
29.i1x.1977, L (USNM). Hyland Co.: e. fk Potomac R., Rt.
220, 18-20.v.1963, P, 1 5 (USNM); 5 mi. sw. Monterey,
Jackson R. beside Rt. 84, 5.x.1967, L (ROM). Madison
Co., Graves Mill, Rapidan R., 800 ft, 18.iv.1975, L

(USNM). Montgomery Co.: Mill Cr., Rt. 785, 29.111.1977,
P (emerged 15.iv.1977), 3 6 1 9; same, 11.iv.1978
(emerged and preserved 28.iv.1978), L PP P, 7 6d 2 9
(USNM); same, | 6 1 2 (ROM). Rockbridge Co.: nr Cave
Mtn Area, #25, 11.ix.1957, L (osF); Greys Run, Goshen
Wildnerness Mgmt Area, 11.vi.1977, 1 3 (ver). Shenan-
doah Nat. Pk: Pass Run, 11.vi.1961, L PP P, 1 6 MMT
(USNM); Thornton R., 8.vii. 1961, PP P; same, 27.v.1962,
P (USNM). Smyth Co., Crewey Br., Rt. 610, 30.i11.1979,
10 3d 11 & (ver). Wythe Co., 1.8 mi. w. Wytheville, Reed
Cr. at Rt. 11, 4.x.1967, L (Rom). WEST VIRGINIA—
Pendleton Co., Gandy Cr., Spruce Knob, 22.vi.1973, 1 3
(USNM).

Psilotreta rufa (Hagen)

Molanna rufa Hagen, 1861:276. Lectotype 3d (mcz),
Trenton Falls, New York, Osten Sacken, 1858.

Heteroplectron? dissimilis Banks, 1897:30. Lectotype @
(mcz), Sea Cliff, Long Island, New York, vi. NEW
SYNONYMY.

Astoplectron connexa Banks, 1914:265.

Molannodes rufa—Milne, 1934:9, 11.

Heteroplectron indecisum—Milne, 1936:57, 80.

Heteroplectron rufa—Ross, 1938:20, as synonym of P.
indecisum.

Psilotreta rufa—Ross, 1944:286—287, 300, fig. 9S2A—C;
A. connexa synonym of P. rufa.

Psilotreta rufa—Wallace, 1970:244, fig. 4.

Psilotreta rufa—Schmid, 1983:48, figs. 278-282.

This species is distinguished from others in the rufa sub-
group by the details of genitalia given in the key. The larva
is distinguished by the shape of the head and of the
anterolateral corner of the pronotum and by the condition
of the setae as outlined in the key.

ADULT

Forewing 9-11 mm in length. Coloration almost uniformly
dark reddish brown with antennae, legs, and abdomen
paler. Antennae long and slender, |.2-1.7 x forewing
length; scape 3.5-4.0 x pedicel length; first two fused
segments of flagellum 2.3—2.7 x pedicel length. Maxil-
lary palpi with first two segments subequal in length,
approximately three-quarters length of third and fourth
segments, and fifth segment about twice length of first.
Labial palpi short, with many short setae, first segment
two-thirds length of second, and one-half length of third.
Frontoclypeus with large convex wart medially, covered
with long setae projecting dorsally. Vertex with ante-
romesal setal warts very small or absent; anterior warts
divided into numerous small areas arranged transversely

behind anteromesal warts, with largest contiguous areas
laterad; posterior setal warts large, discrete, and trans-
versely lunate.

Male genitalia (Fig. 17). Abdominal segment VII with
pleural membrane unmodified. Segment [X with apical
segment of each inferior appendage bearing three to six
sharp black teeth apicomesally. Segment X with dorsome-
dian lobe rarely extending beyond midlength of lateral
processes in dorsal aspect, and with apex entire or bifid and
bearing a few to several setae; lateral processes triangular
in dorsal aspect, highly variable in lateral aspect, ranging
from simple broad, bluntly rounded plates to narrow,
sinuous plates with a pronounced apicoventral projection
(Fig. 17a).

Female genitalia (Fig. 18). Anterolateral region of ster-
num IX lacking indentations, but this area may appear
slightly granular. Posterior vaginal sclerites with mesal
edges approximate posteriorly, diverging rapidly, and
widely separated anteriorly.

LARVA (Figs. 8, 28, 29, 35)

Sclerites uniformly reddish brown, although posterior scle-
rites may be lighter than more anterior sclerites. Head
longer than wide (Fig. 35), and in lateral aspect posterodor-
sal angle more or less squared; dorsum relatively flat
between carinae (Fig. 29b); seta 17 thinner than, and
usually about one-half as long as, seta 15 (Fig. 29a).
Pronotum with projection of anterolateral corner short,
about 0.2 x middorsal length of pronotum. Mesonotum
with black marking along posterior ridge extending laterad
to posterior inflection; mesepisternum lacking projection of
anteroventral corner. Metanotum (Fig. 35) with anterior
sclerite bearing 60—67 setae, posterior bearing 21—30, each
lateral bearing 35-43. Abdominal segment VIII bearing 0—
10 lateral tubercles on each side. Abdominal gills: dorsal—
II 12-16, Ill 13-17, IV 9-13, V 7-10, VI 4~7, VII 3-6,
VIII 0-3; ventral—II 9-11, Ill 16-20, IV 12-16, V 8-12,
VI 9-11, VITO—11, VIII O04. Larva up to 12 mm in length;
head up to 1.1 mm in width across eyes. Larval case up to
11 mm in length.

NOTES

We examined the female lectotype of Heteroplectron?
dissimilis Banks, 1897, and found it to be identical with
females of P. rufa. We therefore place H. dissimilis as a
junior subjective synonym of Psilotreta rufa (Hagen).

HABITAT AND BIOLOGY

P. rufa larvae occur in small spring seeps and spring-fed
streams. Graham (1982) studied a small population in a
spring seep in southeastern Pennsylvania; larvae required

Fics. 17, 18. Psilotreta rufa. 17. a,b, 3 genitalia—lateral (with detail of variation in lateral process of
segment X), dorsal; c, apical segment of inferior appendage—ventral; d,e. phallus—lateral, ventral (dmpX—
dorsal median process of segment X; int. app.—intermediate appendage; [Xt—tergum of segment IX; lat.
pr.—lateral process). 18. a,c, 2 genitalia—lateral, ventral; b,d, vaginal apparatus—lateral, ventral.

WN
N

two years to complete development, growth was tempera-
ture-dependent, occurring from May to October, and
optimum growth was at 15°C which was reached during
early spring and late fall.

DISTRIBUTION (Fig. 39)

This is the most widely distributed of the three species of
the rufa subgroup: Alabama, Delaware, Kentucky, Mas-
sachusetts, Mississippi, New York, North Carolina, Penn-
sylvania, Tennessee, Vermont, Virginia.

MATERIAL EXAMINED AND ADDITIONAL RECORDS

ALABAMA—Cleburne Co., trib. Shoal Cr. 5.8 mi. from
Ala #73, R9E-T155-28NW, 22.iv.1973, 1 6 (ur). DEL-
AWARE—New Castle Co. (fide Lake, 1984). KEN-
TUCKY—Mammoth Cave Nat. Pk, Good Spring,
20.v.1957, 5 do (INHS). MASSACHUSETTS—Berkshire
Co., Mount Greylock, 15.vi, 1 d (Rom). MISSISSIPPI—
Lafayette Co., 6 mi. e. Oxford, Hopewell L. (fide
Holzenthal et al., 1982). Marshall Co.: Wall Doxey St. Pk,
Rt. 7s. Holly Springs, sm. spring str., 12.v.1970, L (Rom
700332); Spring Lake, v (fide Holzenthal et al., 1982); w.
side Spring Lake, spring seep, 15.v.1980, L (CLEM); same,
16.v.1980, L (Paul Lago, University of Mississippi).
Tishomingo Co., Iuka (fide Holzenthal et al., 1982). NEW
YORK—Tompkins Co.: Ithaca, Dryden, Ellis Hollow,
spring, 4x. 1956, “Escsame,’ Sci. 1957, 1: “same,
PeeS57 Pe: same, 1250-19575 LC@sr); same,
11.vi.1956, 1 d (ROM); Ellis, spring, 5.v.1956, L; same,
6.vii. 1956, 5 3d 1 2; same, 14.vii.1956, P, 13 ¢; same,
17.ix.1956, L (osrF); Ellis Hollow Rd, spring seepage,
29.v.1959, PP (ROM); same, 23.vi.1956, P (osF). NORTH
CAROLINA—Macon Co.: trib. Chattooga R. e. High-
lands, #2, 6.vii.1958, L (osF); str. feeding Cliffside L.
(Skitty Cr.) at cpgrd ca 6 mi. nw. Highlands, 27.iv.1980,
L PP (GAs). Transylvania Co., n. Salem, White Water R.
at Rt. 171, 18.v.1970, P (Rom 700353). PENNSYL-
VANIA—Chester Co.: Atglen, Glen Run, 2.iv.1977, P;
same, 24.iv.1977, L PP P, d MMT; same, 2-3.vi.1977,
14 3d 1 @ (cLEM). Fulton Co.: Chestnut Bridge Hollow,
21.vi.1976, L P (cLEM); Crystal Spring, 29.vi.1976, 1 3 1
2 MMT; same, 31.vii.1976, 1 6 (CLEM). TEN-
NESSEE—Chester Co., at spring ne. Henderson,
21.vi.1959, 1 d (INHS). Franklin Co.: Wellington Mills,
I9hiv.. 1955,° 1 Sesame, 26-1v-1955, 2) S:. same,
30.iv.1955, 1 3 (fide Edwards, 1966). Hardin Co., Cave
Hollow Br. of Turkey Cr., 15.v.1978, P (ur). Sevier Co.,
Great Smoky Mtns Nat. Pk, str. 10.2 mi. e. Gatlinburg
entr. on Rt. 441, 20.v.1970, PP (Rom 700361). VER-
MONT—Windham Co., Brattleboro, 14.vii, 1 ¢d (Rom).
VIRGINIA—Albemarle Co., s. fk Mooremans R. above
Charlottesville Reservoir, 13.iv.1974, PP (UusNM). She-
nandoah Nat. Pk: Ivy Creek, 2700 ft, 18.ix.1958, L (Rom);
Piney R., 1100 ft, 28.v.1977, 6 d 1 2 (UsNM). Skyline

Drive, milepost 79, 18.ix.1958, L; same, milepost 79.5,
23.v.1959, PP P; same, milepost 79.6, #27, 12.ix.1957,
L (CLEM).

Psilotreta rossi Wallace

Psilotreta rossi Wallace, 1970:243-245, figs. 1-3.
Holotype d (INHS), Coweeta Hydrological Laboratory,
Macon County, North Carolina.

Adults of this species are distinguished from other North
American Psilotreta by their pale grayish coloration, the
open discal cell in the hindwing of the female, and details
of the genitalia of both sexes as outlined in the key. The
larva is distinguished by the very small projection of the
anterolateral corner of the pronotum and the short head seta
17.

ADULT

Forewing 9—10 mm in length. Coloration pale yellowish to
grayish brown, abdomen purplish. Antennae 1.4—-1.5 x
forewing length; scape 4.5-5.0 x pedicel length; first two
fused flagellar segments 3.0-3.3 x pedicel length. Palpi
identical with those of P. rufa except male maxillary palpi
with a few dark, scalelike setae ventrally along length of
first two and base of third segments. Frontoclypeus with
median wart narrower than in P. rufa and constricted below
middle; vertex with anteromesal warts larger, anterior
warts smaller and narrower, and posterior warts more
triangular than in P. rufa. Hindwing of female with discal
cell open apically.

Male genitalia (Fig. 19). Abdominal segment VII with
pleural membrane unmodified. Segment IX with apical
segment of inferior appendage not truncate in lateral or
ventral aspects (Fig. 19c), and with mesal surface concave
and possessing a sharp black tooth on the basomesal corner
and one to three teeth apically. Segment X with dorsome-
dian lobe almost always extending beyond midlength of,
and often entirely beyond apex of, lateral process in dorsal
aspect, apex bifid and bearing several setae (Fig. 19b);
lateral processes semicircular or triangular in dorsal aspect,
short and broadly rounded apically, with a narrow posteri-
orly curved projection from the apicoventral border in
lateral aspect (Fig. 19a), this projection. moderately long
and blunt; intermediate appendages as in P. rufa.

Female genitalia (Fig. 21). Anterolateral corner of ster-
num IX with a small depression; posterior vaginal sclerites
broadly oval and lightly sclerotized.

LARVA (Figs. 30, 36)
Head and pronotum dark reddish brown, mesothoracic and

_int. app.

Fics. 19-21. Psilotreta rossi. 19. a,b, 3 genitalia—lateral, dorsal; c, apical segment of inferior appen-
dage—ventral (int. app.—intermediate appendage; lat. pr.—lateral process). 20. 2 hindwing. 21. a,c, @
genitalia—lateral, ventral; b,d, vaginal apparatus—lateral, ventral (ind.—indentation).

metathoracic sclerites paler; legs light brown basally, dark
apically, with dark borders. Head subcircular (Fig. 36),
with area between lateral carinae slightly domed (Fig.
30b), and appearing more wrinkled and less shiny than on
other Psilotreta; seta 17 much shorter than seta 15, clear
and thin. Pronotum with projection of anterolateral corner
short. Mesonotal sclerites with dark marking of posterior
ridge extending to posterior inflection, which is repre-
sented as a light spot (Fig. 30a). Metanotum (Fig. 36) with
anterior sclerite bearing 86—90 setae, posterior bearing 37—
40, each lateral bearing 43-50. Abdominal segment VIII
bearing 34 lateral tubercles on each side. Abdominal gills:
dorsal—II 10, III 16, TV 11, V 7-8, VI 4-5, VII 2;
ventral—II 7-9, III 16-18, IV 8-12, V 11, VI. 10, VII 9-
10. Larva up to 11 mm in length; head up to 1.0 mm in
width across eyes. Larval case up to 12 mm in length.

HABITAT
P. rossi larvae have been collected from small spring
seeps.

DISTRIBUTION (Fig. 39)
P. rossi is known only from the mountains of North
Carolina, Virginia, and West Virginia.

MATERIAL EXAMINED AND ADDITIONAL RECORDS
NORTH CAROLINA—Macon Co.: Coweeta Hydro-
biological Lab., 15.vi.1970, holotype ¢ (fide Wallace,
1970); Cullasaja R. 5 mi. from Highlands on U.S. 64,
18.vi.1976, 1 d (GAs); Wayah Bald, 8.vi.1961, 1 6 MMT
(USNM); sm. spring str. 2.7 mi. from Wayah Bald Rd on
For. Serv. Rd 69, 20.v.1970, L (Rom 700363). Tran-
sylvania Co.: 4 mi. s. Blue Ridge Pkwy on NC #215, sm.
roadside seepage, #8, 15.v.1979, 1 5 (KsBs); same, L PP
P, 2 3d 1 9 MMT (cLEM). VIRGINIA—Grayson Co.:
series of spring-fed strs off Mt Rogers Trail into Lewis
Fork, 20.vi.1981, 4 d (ROM); spring seep-bog along For-
est Rd 89, 13.viii.1979, 1 5 2 2 (ver). Shenandoah Co.:
nr Liberty Furnace, Little Sluice Mtn trail, springs,
21.vii.1974, 1 3d; same, 27.x.1974, L (UsNM). WEST
VIRGINIA—Pocahontas Co.: Right Fork Tea Cr., red
spruce forest, small spring with sedge, Monongahela Nat.
For., 4.vii.1984, 18 3d (USNM); small seep to Hills Cr.,
Hills Creek Scenic Area, Rt. 39, Monongahela Nat. For.,
20.x1.1982, L (USNM).

Psilotreta amera (Ross)
Heteroplectron amera Ross, 1939:68, fig. 8. Holotype d ,
allotype 2 (INHS), Parksville, Tennessee.

Psilotreta amera—Ross, 1944:286, 300, fig. 953A,B.

The adult of this species are distinguished by the very large

setal warts on the head of the male, and by the distinctive
genitalia of both sexes. The larva is distinguished by the
circular domed head.

ADULT

Forewing 10-11 mm in length. Antennae 1.2—1.5 x fore-
wing length; scape 3.5-4.1 x pedicel length; first two
fused flagellar segments 2.0—-2.6 < pedicel length. Maxil-
lary palpi as in P. rossi. Frontoclypeus with median wart
large; vertex of female with warts identical to P. rufa;
vertex of male (Fig. 3) with very small anteromesal warts,
anterior pair of warts either absent or fused with posterior
pair, and posterior pair greatly enlarged, occupying entire
posterior half of head, each one having raised prominence
near midline of head thickly covered with erect brown setae
and dense pile of erect blackish setae on rest of wart,
resulting in distinct two-toned appearance.

Male genitalia (Fig. 22). Segment VII with narrow
dorsoventral invagination near posterior edge of pleural
membranes. Segment IX with apical segment of inferior
appendages small, and bearing one basal and one apical
long black curved tooth. Segment X with apexes of dor-
somedian process bifid and widely separated (Fig. 22b),
extending considerably beyond bases of lateral processes
in some specimens (Fig. 22a); lateral processes narrow
throughout, sinuate and directed posteroventrally, and end-
ing in sharp points; intermediate appendages short, arcuate
but not curled, and ending in sharp ventrally directed
points.

Female genitalia (Fig. 23). Segment IX with median
plate deeply invaginated at anterolateral corner (Fig. 23a);
tergum IX with shallow indentation laterally. Posterior
vaginal sclerites narrow, parallel sided, and approximately
uniformly separated throughout their length (Fig. 23d).

LARVA (Figs. 31, 37)

Coloration and shape of head similar to P. rossi. Head
setation and shape of pronotum similar to P. rufa. Head
strongly domed (Fig. 31b), subcircular in dorsal aspect
(Fig. 37); in lateral aspect, posterodorsal angle of head
rounded (Fig. 31a). Metanotum (Fig. 37) with anterior
sclerite bearing 60—63 setae, posterior bearing 20—24, each
lateral bearing 41-47. Abdominal segment VIII bearing 2—
3 lateral tubercles on each side. Abdominal gills: dorsal—
II 7, Ill 15-16, IV 6-9, V 6-7, VI 4-6, VII 3, VIII 2;
ventral—II 34, I11 17, 1V 8, V 6, VI 7, VII 7-8. Larva up
to 9 mm in length; head up to 1.1 mm in width across eyes.
Larval case up to 10 mm in length.

HABITAT
Like P. rossi, P. amera larvae apparently inhabit small
spring seeps.

MN
nN

DISTRIBUTION (Fig. 39)

P. amera has the most restricted range known for any
North American species, a relatively small area in the
vicinity of Great Smoky Mountains National Park:
Georgia, North Carolina, South Carolina, Tennessee.

MATERIAL EXAMINED AND ADDITIONAL RECORDS

GEORGIA—Lumpkin Co., 3 mi. s. Neels Gap, Chestatee
Br., 23.v.1946, 1 2 (INHs). NORTH CAROLINA—Hay-
wood Co., trib. w. fk Pigeon R., | mi. s. Sunburst Cpgrd
on NC #215, NC #6, 15.v.1979, L (KsBs). Macon Co.,
Nantahala Nat. For., Cullasaja R. nr U.S. 64 at entr.
Cliffside L. Rec. Area, 17.v.1979, P (ksBs). Swain Co.,
Great Smoky Mtns Nat. Pk, Smokemont Cpgrd and
nearby, 11—14.v.1970, PP P (usNm). SOUTH CAR-
OLINA—Oconee Co.: Sumter Nat. For., str. in Yellow
Br. Picnic Area, ca 1500 ft, 19.v.1970, 3 6 MMT 3 @
MMT (Rom 700356); 5 mi. nw. Tamasee, sm. spr. brook
above Wash Br. of Townes Cr., 26.v.1980, 2 d; same, 2—
3.vi.1980, 7 6 (CLEM); 6 mi. s. Walhalla, Crane Cr., U.S.

Fish Hatchery n. Tamasee, 26—27.v.1980, 1 3d (CLEM).
Pickens Co., 9 km nw. Clemson, Wildcat Cr., 18—
19.v.1979, 1 3; same, 30.1x.1961, P (cLEM). TEN-
NESSEE—Blount Co.: Tenn. #73 ca 3 mi. w. fk Little
Rat, trib. to Laurel Br. on Cadestone Rd, 11.iv.1981, PP P
(GAS); Great Smoky Mtns Nat. Pk, trib. to Little R. 0.5 mi.
above fk, 4.v.1981, P (uT); Spicewood Br., trib. w. prong
Little R., 11.iv.1981, PP (GAs); same, 4.v.1981, P (uT).
Cocke Co., English Cr. at Carson’s Spring nr Newport, 3—
8.vi. 1946, | d (INHS). Great Smoky Mtns Nat. Pk, 1978,
1 5 (uT). Knox Co., 1.v.1963, 2 MMT (cLEm). Polk Co.:
first rock-face str. below Coker Cr., 5—200 m above
Hiwassee R., 12.v.1977, L PP P, od and 2 MMT (ut);
streams between Coker Cr. and Lars Cr. at Hiwassee R.,
8.iv.1977, PP (uT); trib. Hiwassee R. 2.6 mi. from Hwy
411, Quinn Springs For. Serv. Cpgrd, L PP P, 3 and 2
MMT (ur); Parksville, 25.iv.1938, holotype d, allotype
2, paratype 6 (fide Ross, 1939). Sevier Co.: Great Smoky
Mtns Nat. Pk, Jakes Br. nr Elkmont Cpgrd, 10.iv.1981, PP
(GAS); e. slope Webb Mtn, George Cox property, 17.1v—
L7evalS75. ied (Ur):

Fics. 22, 23. Psilotreta amera. 22. a,b, 3 genitalia—lateral, dorsal (with detail of variations of median
dorsal and lateral processes of segment X); c, apical segment of inferior appendage—ventral (int. app.—
intermediate appendage; lat. pr.—lateral process). 23. a,c, @ genitalia—lateral, ventral; b,d, vaginal
apparatus—lateral, ventral (ind.—indentation; inv.—invagination).

Fics. 24-31. Larvae of Psilotreta spp. 24, 28. Left mesofemur—anterior. 25-27. Head, prothorax, and
mesothorax—lateral. 29-31. Head, prothorax, and mesothorax—lateral (detail of head in frontal aspect).
(Abbreviations: mesep.—mesepisternum, anteroventral angle; post. ang.—posterior angle of head; post.
infl.—posterior inflection.) 24, 25. P. indecisa. 26. P. frontalis. 27. P. labida. 28, 29. P. rufa. 30. P. rossi.
31. P. amera.

Fics. 32-37. Larvae of Psilotreta spp., head and thorax—dorsal. 32. P. indecisa. 33. P. frontalis. 34. P.
labida. 35. P. rufa. 36. P. rossi. 37. P. amera.

Fic. 38. Distribution of Psilotreta spp.: P. indecisa—squares; P. frontalis—circles; P. labida—triangles.

Fic. 39. Distribution of Psilotreta spp.: P. rufa—circles; P. rossi—squares; P. amera—triangles.

ASIAN PSILOTRETA

The 14 species of Psilotreta now known from Asia are a
heterogeneous assemblage. We have examined males of 10
species, females of 6, and larvae of 3, and offer a key to the
males; females and larvae are insufficiently known to
provide keys.

2(1)

sql)

Key to Males of Asian Psilotreta

Inferior appendages with basal segment
C-shaped, apical segment arising mesally and
subapically (Fig. 55); median dorsal process of
segment X indistinguishably fused with median
dorsal process of segment IX, very long and
hood shaped, and free from lateral processes
throughoutlencth (Fig: 55).0.. 2,.5- 47.04." Z
Inferior appendages with basal segment not
C-shaped, apical segment situated apically or
dorsoapically (e.g., Fig. 46a); segment X with
median dorsal process not long and hood
shaped, or if somewhat hood shaped, then
fused throughout length to lateral processes
(Schmid> 1959 plh Vs tiga) 224.5 oe 3

Intermediate appendages long and projecting
posteriorly beneath hood-shaped process;
inferior appendages with ventral lobe of basal
segment subequal in length to apical segment
(ig sS5a)e of Seok kwantungensis, p. 45
Intermediate appendages short and knoblike
apically and not extending far beneath hood-
shaped process; inferior appendages each with
ventral lobe of basal segment extending far
beyond apex of apical segment (Hwang, 1957,
| tds 8) eee Ae oe ae eae lobopennis, p. 50

Segment X with median dorsal process com-
pletely fused to lateral processes, forming hood
over intermediate appendages and lateral pro-
cesses; intermediate appendages straight,
slender, directed anteroventrad (Schmid, 1959,
DU TV Set) coe erie eae quadrata, p. 40

4(3')

5(4’)

5’

6(5)

Segment X with median dorsal process free of
lateral processes (Fig. 46a), or if fused not
forming a hood (Fig. 48a); intermediate appen-
dages of different shapes but not slender,
straight, or directed anteroventrad ......... +

Segment X with preanal appendages short,
median dorsal process fused with lateral pro-
cesses; lateral processes each with a projection
arising from anteroventral corner, with a dis-
tinct dorsal lobe and a prominent horn curved
posteroventrally, and ending in a sharp point;
intermediate appendages arising beneath lateral
processes, curving dorsally around antero-
ventral projections of lateral processes, and
each tapering to a sharp point which is directed
ventrolaterally close to apex of each lateral
process (Fig:/48) 4.2.3. . sta chinensis, p. 34
Segment X with preanal appendages long and
foliose in lateral aspect (Fig. 46a), median dor-
sal process free of lateral processes, and lateral
PIOCESSES NOLAS ADOVEw 4 ee dae 5

Inferior appendages each with apical segment
foot shaped in dorsal aspect (Mosely, 1942, fig.
8), or at least with a prominent lobe along outer
marein (wang, 19577 fis. 94) ee ae ae 6
Inferior appendages each with apical segment
not foot shaped, and without lobes (Figs. 51a,
DSO) Fel oheedg ees eae eee Vi

Intermediate appendages curved, each tapering
to a point directed ventrolaterally; lateral pro-
cesses each with a recurved anteroventral pro-
jection, tapered and pointed (Hwang, 1957, fig.
93); wings lacking discal cell (Hwang, 1957,
fp 9D) sh wake he Sooo orientalis, p. 50
Intermediate appendages much as above, but
each with a dorsal accessory hook (Mosely,
1942, fig. 7); lateral processes apparently lack-
ing; wings with discal cell (Mosely, 1942, fig.
OQ) cae oe, SRE owe echina, p. 50

TO)

8(7)

9(8')

10(7’)

Segment X having lateral processes bean
shaped in lateral aspect and without projections;
inferior appendages each with basal segment
bearing a small apicoventral process (Botosa-
neanu, 1970, pls. XXXVI-XXXVIII)...... 8
Segment X having lateral processes of various
shapes and with (Fig. 46a) or without (Fig. 53a)
projections; inferior appendages each with
basal segment lacking apicoventral process
(Higley ashe tee tele uk eo ae. 10

Segment X in dorsal aspect with median dorsal
process narrow and pointed apically; intermedi-
ate appendages long and narrow, each with a
subapical spine; phallotheca with ventral sur-
face strongly sclerotized to apex; parameres
long, narrow, and pointed (Botosaneanu, 1970,
(6) LO, OG Uae trata Dare Sear he kisoensis, p. 35
Segment X in dorsal aspect with median dorsal
process more robust and apex expanded and
bilobed; intermediate appendages either long
and narrow or more robust but without a subapi-
cal spine; phallotheca with ventral surface
membranous to apex; parameres as above or
short and blunt (Botosaneanu, 1970, pls.
OOOO VLD as SS SY SEY. 9

Parameres short and blunt, strongly decurved
ventrally; inferior appendages each with apico-
ventral projection of basal segment prominent;
head with frontoclypeal wart large and bearing
numerous long, erect setae; vertex with a deep
groove along coronal sulcus (Botosaneanu,
LOT Op) Seana. nee falcula, p. 40
Parameres long, pointed, and only slightly
decurved apically; inferior appendages each
with apicoventral projection of basal segment
weakly developed, not prominent; head with
frontoclypeal warts paired, small, and bearing
normal setae; vertex without groove (Botosa-
Means O70; tpl OOK VIM) ee ae ee,
oe a eg ee Ee oe locumtenens, p. 36

Phallus with parameres long, slender, and acute
(Ris a5 Sdke)) een ee eae tee teeta. 11

10’

11(10)

11’

12(10’)

12’

13(12')

13’

Phallus with parameres variously formed, not
long, slender, and acute (Figs. 46d,e; 51d,e)

Segment X with lateral processes each having
an acute ventral projection, and intermediate
appendages each having two acute, curved pro-
Jections, so that three distinct acute projections
are visible in lateral aspect (Schmid, 1965, pl.
Vib thie. 6 9) ts. Ne ee ree trispinosa, p. 47
Segment X with lateral processes each reduced
to inconspicuous low ridge (Fig. 53a) .......
Daa 9y Sy. See assamensis, p. 41

Segment X with intermediate appendages both
broad and bifid apically in lateral aspect (Kim-
mins, 1964, fig. 40); median dorsal process a
short cordate lobe in dorsal aspect (Kimmins,
1964. eral ee See ae quinlani, p. 50
Segment X with intermediate appendages small
and simple (Fig. 46a); median dorsal process
elongate, bifid, or truncate apically, not cordate
(Rig4 6b) ie Ses Rae Sees pee eee 13

Inferior appendages uniformly cylindrical in
lateral aspect (Fig. 46a) and each with a low,
rounded median lobe in ventral aspect (Fig.
46c); segment X with lateral processes each
having a narrow, acute projection directed pos-
teriorly; intermediate appendages short and
pointed ventrad (Fig. 46a); phallus without pair
of accessory processes above parameres; para-
meres narrow in lateral aspect (Fig. 46d,e)

Inferior appendages each with a ventral projec-
tion in lateral aspect (Fig. 5la) and with a
prominent, angulate basomedian lobe in ventral
aspect (Fig. 5lc); segment X with lateral pro-
cesses each having a relatively broad, rounded
projection directed posteriorly; intermediate
appendages a pair of small, low inconspicuous
lobes (Fig. 51a); phallus with a pair of small
accessory appendages situated dorsad of the
large, broad parameres (Fig. 51d,e).........
be Redes ot arr weer Stas oR Ura A schmidi, p. 41

Psilotreta japonica (Banks)

Odontocerum japonicum Banks, 1906:110. Holotype 2
(mcz, Type 11837), Gifu, Japan.

Odontocerum japonicum—UlImer, 1907a:51-52, figs. 76—
79.

Psilotreta japonica—UI|mer, 1907b:126, figs. 157, 158.

Psilotreta kyotoensis Iwata, 1928:118, 124-125, figs. 213,
214. Synonymy by Tsuda (1942b).

Psilotreta japonica—Tsuda, 1959:143, fig. 256.

This species is most similar to P. schmidi sp. nov., from
which it is distinguished by the narrow, acute lateral
processes, the small, acute, and projecting intermediate
appendages of segment X, and the less prominent develop-
ment of the ventromesal angle of the basal segment of the
inferior appendages.

ADULT

Forewing 10-11 mm in length. Coloration black dorsally;
legs and palpi dark brown; forewings brown with faint
irroration. Each wing with discal cell long and widest at
origin of vein R,, with R, kinked at junction of crossvein r,
and lacking scales. Scape subequal in length to head.
Maxillary palpi broken on males examined, first three
segments cylindrical and unmodified, second segment
shortest and third longest; on females, first three segments
similar, segments four and five subequal, each slightly
longer than segment three. Labial palpi in both sexes with
each segment slightly longer than preceding segment, last
segment thinner than the first two. Setal warts of vertex
similar to those of P. indecisa, anterior warts close to
antennal sockets, posterior warts ovoid.

Male genitalia (Fig. 46). Segment VII with pleural
membranes each bearing a small lobe posteriorly. Segment
IX typical for genus; dorsolateral piece extremely narrow,
dorsum slightly longer than wide and produced laterally
into thin ledge, which overhangs base of segment X and
obscures circular basal setal warts in dorsal aspect; basal
segment of inferior appendage dark in basal fourth and
lacking setae, the remainder light, with low, broad median
basal lobe, and with numerous long setae, which are
subcylindrical and slightly tapered; apical segment dark
and setose, slightly tapered in lateral aspect, obliquely
truncate in ventral aspect, and with numerous small black
spines apically. Segment X with median dorsal process
large, apex flared, bifid, and bearing several setae, and in
lateral aspect deeply excavate basodorsally; lateral pro-
cesses long, acute, posteriorly directed, arising from pos-
terior edge of body of segment X; intermediate appendages
short, stout, pointed, darkly sclerotized, arising beneath
bases of lateral processes, directed ventrolaterad, and
curved slightly caudad; preanal appendages broad, foli-

aceous, broadest in basal fourth and tapering apically,
extending almost to apex of segment X, with numerous
long setae apically and ventrally.

Phallus with two parameres arising close together on
venter of endotheca, rounded apically in lateral aspect,
pointed in ventral aspect; aedeagus long with sclerotized
ventral plate.

Female genitalia (Fig. 47). Females of P. japonica are
distinguished from those of P. kwantungensis by a median
sulcus in the posterior portion of sternum IX, from P.
chinensis by lack of a lateral depression in the tergum of
segment IX and by the anterior vaginal sclerites being
shorter than the posterior, from P. falcula and P. locum-
tenens by the setal warts of the tergum of segment IX being
produced only slightly, and from P.. assamensis sp. nov. by
details of the shape and orientation of the vaginal sclerites.
General structure similar to North American species. Apex
of segment X in lateral aspect angular. Posterior portion of
sternum IX with distinct longitudinal median sulcus.
Vaginal chamber with anterior and posterior portions
approximately equal in length; sclerites of posterior portion
convoluted in lateral aspect, broad and widely separated in
ventral aspect; anterior portion with apical corners pro-
duced as rounded lobes.

LARVA

The larva of P. japonica was described as P. kyotoensis
(Iwata, 1928), a name later synonymized with P. japonica
(Tsuda, 1942b). We have seen no specimens.

MATERIAL EXAMINED
JAPAN. Gifu, holotype 2, paratype 2 (mcz). Kyoto,
8.v.1953, 1 do (P. W. Oman) (USNM).

NOTE

When Banks described P. japonica, he indicated that he
had two specimens, but no holotype was designated at the
time. His material, which we have examined, consists of
the two females listed above, one of which now bears a red
MCZ type label (11837), presumably affixed by Banks
himself at some time after publication of the description.

Psilotreta chinensis Banks

Psilotreta chinensis Banks, 1940:219-—220, pl. 27, fig. 1;
pl. 30, figs. 67, 68, 70. Holotype d (UsNM, Type
53194), China, Szechwan near Washan, 1220—1830 m,
Vil.

The male of P.. chinensis is distinguished from all others in
the genus by the absence of the discal cell of the forewing,
R,,, being fused with R,,, for most of their lengths, and

by the preanal appendages being very short. The forewing
of the male of P. orientalis also appears either to lack the
discal cell or at least to have the discal cell open apically,
but the preanal appendages are long.

ADULT

Forewing 12 mm in length in male, 15 mm in female.
Coloration light brown in male, darker in female. Although
Banks described P.. chinensis as having venation similar to
P. japonica, with a discal cell, the male lacks a discal cell
(Fig. 52). Stem of R,,,,,5 fused throughout length to a-
nastomosis; R, arising about halfway between origins of R,
and R, and kinked at crossvein r; R, and R, stalked; a
narrow fold of membrane overlapping R;,,,,5 dorsally
from behind, this vein bearing numerous stiff setae ven-
trally; forewings covered with setae intermingled ventrally
with scattered scales. Forewing of female with a long,
narrow discal cell; R, arising in basal fourth and kinked at
crossvein r. Hindwings of both sexes each with long discal
cell; R, arising in basal fourth. Maxillary palpi in both
sexes with each segment slightly longer than the preceding
and heavily setose, male with scattered scales on basal
segments. Labial palpi having first segment short, second
and third segments longer and subequal, and long setae.
Face of male with elongate lateral setal warts next to eyes
having long setae and short scales; frontoclypeus with a
narrow scale-filled indentation extending dorsally between
antennae and continuous on vertex (Fig. 50); on vertex
anteromedian pair of warts on raised surfaces on either side
of indentation, anterior pair ovate, and posterior pair large
and transverse, widest laterally and narrowing medially.
Female lacking indentation and scales, but otherwise simi-
lar to male. Pronotum of male with scales intermingled
among long, silky setae; female lacking scales on
pronotum.

Male genitalia (Fig. 48). Segment IX typical for genus;
longer in ventral half, with narrow, subtriangular poste-
romedian lobe dorsally and low anterior lobe laterally;
inferior appendages long; basal segment extending beyond
lateral process and cylindrical with many setae; apical
segment approximately one-third length of basal segment,
cylindrical, and truncate apically with numerous small
black teeth and short setae. Segment X with basal setal
wart circular, distinct; median dorsal process short, pro-
jecting only slightly posterad of segment IX, bifid apically,
and completely fused to lateral processes; lateral processes
each with large anteroventral projection bearing long,
decurved, tapered spine ventrally and short blunt projec-
tion dorsally; intermediate appendages arising on body of
segment X, each beneath anterolateral projection of lateral
process, extending as large spine curving posterodorsally
above lateral process, and ending in sharp ventrally
directed point; in lateral aspect intermediate appendage and

spine of lateral process nearly forming circle with points
meeting posteriorly; preanal appendages short, ovate, and
bearing numerous setae.

Phallus with phallotheca long, narrow, and cylindrical;
endotheca short, with pair of long, sharply pointed para-
meres arising ventrolaterally and extending two-thirds
length of aedeagus; aedeagus large, membranous, in ven-
tral aspect roughly triangular with two pairs of lobes
laterally and single lobe posteromedially, and in lateral
aspect with phallotremal sclerite forming bilobed projec-
tion dorsally.

Female genitalia (Fig. 49). The female of P. chinensis
is distinguished from the others we have examined by the
tergum of segment IX, which has a broad depression
laterally, and by the vaginal chamber, on which the ante-
rior portion is clearly longer than the posterior portion and
the sclerites of the posterior portion are weak and indis-
tinct. Segment IX with broad, shallow depression laterally;
posterior portion of sternum IX with obvious median lon-
gitudinal sulcus, apex of each half produced posteriorly.
Segment X with apex rounded in lateral aspect. Vaginal
chamber with anterior portion longer than posterior por-
tion, anterolateral corners produced laterad, and posterior
portion having weak and indistinct internal sclerites.

LARVA
Unknown.

MATERIAL EXAMINED

CHINA—Szechwan, near Washan, 4000-6000 ft, vii,
holotype 6, paratype 2 (USNM); paratypes | d 3 2 (Mcz).
Szechwan, Long Ch’i, 17—18.vii.1938, 1 d (USNM).

Psilotreta kisoensis Iwata

Psilotreta kisoensis Iwata, 1928:117, 124-125, figs. 211,
DV

Psilotreta armata Martynov, 1933:144—145, figs. 7-10.

Psilotreta kisoensis—Uéno, 1935:84-85, fig. 59.

Psilotreta kisoensis—Tsuda, 1942a:285 (P. armata Mar-
tynov as synonym).

Psilotreta kisoensis—Tsuda, 1959:143, fig. 255.

Psilotreta kisoensis—Botosaneanu, 1970:313-316, pl. 36,
figs. 1-4.

This species originally was based on larvae and on adults
subsequently described as P. armata by Martynov (1933),
which later was reduced to synonymy with P. kisoensis by
Tsuda (1942a). Botosaneanu (1970) has provided excellent
illustrations of the adults and at the same time described
two closely related species, P. locumtenens and P. falcula.
We have examined two males of P. kisoensis from Japan.

ADULT
The male of P. kisoensis is easily distinguished from those
of P. locumtenens and P. falcula by the characters given in
the key. To published descriptions we add the following
observations: head unmodified, warts on frontoclypeus not
enlarged and not bearing abundant long erect setae, vertex
without groove.

We did not have access to female specimens, but
Botosaneanu (1970) has illustrated the vaginal sclerites.

LARVA (Figs. 40-42)

The larva of P. kisoensis has been described several times
since its original description. Botosaneanu (1970) has sug-
gested that descriptions by Lepneva (1958, 1966) may
apply to P. locumtenens or P. falcula rather than to P.
kisoensis. We have examined larvae in the ROM collection
from Japan that may be P. kisoensis, and we have exam-
ined larvae from North Korea sent to us by Dr Botosaneanu
that we believe to be P. locumtenens on the basis of a
female metamorphotype. The larvae from Japan and North
Korea differ slightly but significantly in colour pattern, and
we concur with Botoganeanu that the larva described by
Lepneva is more likely to be P. locumtenens than P.
kisoensis. Although the associations are not firm,
especially that for P. kisoensis, we treat the larvae tenta-
tively under these names.

The larvae of P. kisoensis and P. locumtenens are
similar in morphology and have been adequately described
and figured by Lepneva (1958, 1966). We describe here
only those features of colour pattern and setation that differ
between the two forms or that our study of the Nearctic
species has indicated are useful in distinguishing larvae.

Head similar to P. Jocumtenens (see Lepneva, 1966, fig.
677—note that the setae are mislabelled in fig. 677, the
seta labelled 15 is actually 13, and that labelled 16 is 15;
seta 16 is not illustrated, but is extremely fine, clear, and
recumbent, positioned near the base of 15 and directed
towards 17). Thorax with dark areas diffuse, much less
distinct than in P. locumtenens, and spread over a wide
area of the nota as blotches rather than stripes. Legs darker
in most specimens of P. kisoensis than in P. locumtenens.

Head longer than wide, narrowest anteriorly, not domed
dorsally; seta 17 dark brown and slightly shorter and
thicker than seta 15. Mesofemur (Fig. 42) with seta 3
situated distad of dorsal seta | by approximately the dis-
tance between setae 4 and 5. Segment VIII with 12-17
tubercles laterally. Abdominal gills: dorsal—II 12-16, III
16-19, IV 13-15, V 11-16, VI 7-9, VII 4-6; lateral—III
3-4; ventral—II 7-12, III 16-18, IV 12-20, V 10-15, VI
8-14, VII 8-14, VIII 0-6. Larva up to 11 mm in length;
head up to 1.1 mm in width across eyes. Larval case typical
for genus, up to 15 mm in length.

MATERIAL EXAMINED

JAPAN—Fukuoka Prefecture: Wakamatsu, 27.vii. 1952, 1
3 (cnc); Sendai, 3.v.1952, 1 3 (cNc). Nara Prefecture:
Yoshino-Kumano Nat. Pk, str. crossing rd 0.3 km above
Mameo, 23.111.1977, L PP (preserved 19.v.1977) (Rom
770029); Omata River 5.5 km upstr. Omata, large side-
pool, 20.111.1977, L (Rom 770023).

NOTE

Tsuda (1942a) synonymized P. armata Martynov, 1933,
under P. kisoensis Iwata, 1928. However, Martynov’s
illustrations of the male genitalia of P. armata do not show
any indication of the large intermediate appendages found
in P. kisoensis, which should be visible in the dorsal
aspect, at least; nor is any mention made of intermediate
appendages in the accompanying description, which is
complete in most other aspects, including the description
of the pleural abdominal lobes of segment VII. It will be
necessary to restudy the type of P. armata to determine its
status.

Psilotreta locumtenens Botosaneanu

Psilotreta locumtenens Botosaneanu, 1970:313-314, pl.
38, figs. 1-5; pl. 39, fig. 2, d 2. Holotype d (Zoologi-
cal Institute, Polish Academy of Sciences, Warsaw),
North Korea, Hamgjong-pukto Province, River Poro-
chon about 2 km northwest of Kjong-song, 4.vi.1965.

This species is most closely related to P. falcula and P.
kisoensis. We have examined uncleared male and female
paratypes, a female metamorphotype, pupae, and larvae.

ADULT

The male can be distinguished by characters given in the
key. To the original description we add the following
observations: head unmodified; frontoclypeus with a pair
of small discrete setal warts on either side of median
groove, these warts bearing long setae; vertex without a
groove, with anteromesal and anterior warts absent and
with posterior warts small, rounded, and separated mesally
by approximately twice the diameter of one wart.

The females may be distinguished from all other species
except P. falcula (and P. kisoensis?) by the elongation of
the setal wart of the tergum of segment X, and from all
species by details of the vaginal sclerites (see Botogsaneanu
1970, pl. 39, fig. 2). To the original description we add the
following observation: tergum of segment X with setal
warts elongated posteriorly on either side as fingerlike
processes.

x Rees TULLE

Fics. 40-45. Larvae of Psilotreta spp. 40, 45. Head and thorax—dorsal. 41, 44. Head, prothorax, and
mesothorax—lateral. 42, 43. Left mesofemur—anterior. 40-42. P. kisoensis. 43-45. Psilotreta sp. (India,
Assam, Kameng).

Fics. 46, 47. Psilotreta japonica. 46. a—c, 3 genitalia—lateral, dorsal, ventral; d,e, phallus—tateral,
ventral. 47. a,c, 2 genitalia—lateral, ventral; b,d, vaginal apparatus—lateral, ventral.

Fics. 48-50. Psilotreta chinensis. 48. a—c, 3 genitalia—lateral, dorsal, ventral; d,e, phallus—lateral,
ventral. 49. a,c, 2 genitalia—lateral, ventral; b,d, vaginal apparatus—lateral, ventral. 50. ¢ head—dorsal.

LARVA

This species may be the one described by Lepneva (1958,
1966) under the name P. kisoensis, as discussed above. To
the description in those works we add the following obser-
vations: colour pattern discrete dark brown streaks on a
yellowish background, as opposed to the more diffuse
blotches of P. kisoensis larvae. Head longer than wide, not
domed dorsally; head seta 17 brown, shorter than seta 15.
Mesofemur with seta 3 situated distad of dorsal seta | by
approximately the distance between setae 4 and 5. Meta-
notum with anterior sclerite bearing 35—40 setae, posterior
bearing 18—23, each lateral bearing 35-40. Segment VIII
with 8-9 tubercles laterally. Abdominal gills: dorsal—II
15-18, I] 21-24, IV 15-19, V 13-18, VI 8-9, VII 5-7;
ventral—II 10—11, II 15-17, IV 16-17, V 14-15, VI 12-
17, VII 11-14. Larva up to 13 mm in length; head up to 1.2
mm in width. Larval case typical for genus, up to 16 mm in
length.

PUPA
Typical for genus. Case up to 19 mm in length.

MATERIAL EXAMINED

NORTH KOREA—Hijangsan District, Mjohjang-san: 16—
22.vi.1965, 3 @ paratypes (LB); Sangvon-am, 17.vi.
1965, L PP P, 2 MMT (rom).

NOTES

The association is based on a female metamorphotype
which is identical with the illustration of P. locumtenens in
Botosaneanu (1970), and on three pupae, one prepupa, and
eight early fifth instar larvae. This collection is from
Botosaneanu’s Station 14, which is one of the paratype
localities of P. falcula. However, the type locality of P.
locumtenens (Station 17) apparently is nearby in the same
river system.

Psilotreta falcula Botosaneanu

Psilotreta falcula Botosaneanu, 1970:314-316, pl. 37,
figs. 1-5; pl. 39, fig. 3, d 2. Holotype ¢ (Zoological
Institute, Polish Academy of Sciences, Warsaw), North
Korea, Hamgjong-pukto Province, Poro-chon River
about 2 km northwest of Kjong-song, 4.vi.1965.

This species is closely related to P. kisoensis and P.
locumtenens and can be distinguished by details of the
genitalia and the structure of the head of the male, as
outlined in the key, and by details of the vaginal sclerites
and the setal wart of the tergum of segment X of the female.
We have examined uncleared male and female paratypes.

To the original description we add the following obser-

vations: frontoclypeus of male having a large median wart
thickly beset with long, erect setae; vertex with deep
groove along coronal sulcus and with anteromesal warts
small and rounded, anterior warts absent, and posterior
warts large. Female with tergum of segment X having setal
warts extended posteriorly on either side as fingerlike
processes, as in P. locumtenens.

LARVA
Unknown.

MATERIAL EXAMINED

NORTH KOREA—Hamgiong-pukto Province, Poro-chon
River about 2 km nw. Kjong-song, 4.vi.1965, ¢ paratype
(LB). Hamhyng-si Province, Hamdzu District, ca 15 km w.
Hamhyng, Hyngpong-ri, 12.vi.1965, 2 paratype (LB).

Psilotreta quadrata Schmid

Psilotreta quadrata Schmid, 1959:327, pl. 4, figs. 8,9.
Holotype 3 (Zoologisches Museum der Humboldt, Uni-
versitat zu Berlin), China, Li-kiang, 22.v.1924.

This species is distinctive in several features, notably the
complete fusion of the body of segment X and of the lateral
processes with the median dorsal extension of segment X,
forming a complete hood over the phallus, and the quadrate
shape of the basal segment of the inferior appendage.

To the original description we add the following obser-
vations. Abdominal segment VII with a posterior lobe on
the pleural membrane. Segment X with basal setal wart
indistinct and elongate, bearing several setae; median dor-
sal process with several short setae apically; lateral pro-
cesses each with a narrow blunt projection arising from the
basoventral border and curving posteriorly; the straight,
more acute intermediate appendage arising from the
posteroventral border of the body of segment X beneath the
lateral process (Schmid, 1959, fig. 8, shows the projection
of the lateral process apparently arising from the body of
segment X beneath the intermediate appendage; the projec-
tion as illustrated also appears to be much longer than in the
specimen we examined). Phallus with parameres short and
talonlike, curved anterodorsally in lateral aspect and later-
ally in ventral aspect when in retracted position (shown in
everted position in fig. 8 of Schmid, 1959).

FEMALE AND LARVA
Unknown.

MATERIAL EXAMINED
CHINA—North Yuennan Province, Li-kiang, 7.vii. 1934,
3 paratype (CNC).

Psilotreta schmidi sp. nov.

This species is similar to both P. japonica and P.. quadrata,
but is distinct from all others of the genus in the inferior
appendages having a ventral projection in lateral aspect and
a prominent basomedian lobe in ventral aspect, and in the
paired accessory phallic appendages being dorsad of the
parameres.

ADULT

Maxillary palpi with first two segments subequal, slightly
shorter than third and fourth, with fifth the longest, and
with segments unmodified, setose. Clypeus convex, with
scattered setae. Anteromedian warts of head slightly
raised, with a moderately deep groove between them;
anterior warts small, oval; posterior warts large, oval,
extending about halfway to coronal suture. Tibial spurs 2,
4, 4. Forewing 9-10 mm in length. Coloration brown;
setae reddish or purplish. Forewing with discal cell present
and approximately one-third length of wing; R, arising in
basal one-third of discal cell and kinked at crossvein r; R,
and R, stalked. Hindwing with discal cell short, R, arising
very close to R,.

Male genitalia (Fig. 51). Segment VII with pleural
membranes lacking posterior lobe. Segment IX with sides
broad, with median dorsal projection long and fused with
segment X throughout; basal segments of inferior appen-
dages each with a median basal flange appearing as a broad
triangle in ventral aspect and as a thin projection in lateral
aspect, apical segments each dorsoventrally depressed and
bearing numerous small black teeth on apical margin, both
segments setose. Segment X with preanal appendages thin,
long, each dorsal edge being smoothly convex, each ven-
tral edge being sinuous and strongly concave beyond base,
apical third of each having dorsal and ventral edges sub-
parallel and apex rounded and bearing scattered, long
setae; median dorsal process in dorsal aspect broad and
enlarged apically, with apex truncate to slightly emarginate
and bearing a row of several long setae laterally on each
side, in lateral aspect thin and free from body of segment X
and lateral processes, extending posteriorly just beyond
apex of each lateral process; body of segment X with apex
bilobed in dorsal or ventral aspect, broad laterally, and
with basal setal wart elongate and poorly defined; lateral
processes long and broad, arising from middle of body of
segment X, in lateral aspect projecting ventrally a short
distance before turning posteriorly at right angles and
ending in a slightly upcurved apex short of end of median
dorsal process, and in dorsal or ventral aspects appearing as
a thick rod tapering to an acute apex; intermediate appen-
dages low, rounded, and inconspicuous, arising beneath
lateral processes and extending anterad half the distance to
the preanal appendages, free from the body of segment X

apically but scarcely visible or not visible in dorsal aspect.

Phallus stocky; phallotheca short and slightly constricted
medially in lateral aspect. Endotheca inflated dorsally and
bearing two pairs of sclerotized processes; parameres
arising ventrally, large, broad, ventrally decurved in lateral
aspect, and thin and forming a narrow V in ventral aspect;
second pair of processes on endotheca short and roughly
triangular, arising laterally, and projecting outwardly.
Aedeagus with a cylindrical base, a prominent ventral lobe
apically, and the phallotremal sclerite C-shaped in lateral
view.

FEMALE AND LARVA
Unknown.

MATERIAL EXAMINED

(All specimens were collected by F. Schmid and are depos-
ited in the CNC, Ottawa.) Holotype ¢: INDIA—Sikkim,
Padamchen, 30.vili.1959. Paratypes: INDIA—Sikkim,
Manu, 5-6.viil.1959, 1 5 (#239); Rongne, 21.viii.1959,
1 3 (#238).

ETYMOLOGY
We take pleasure in naming this species in honour of its
collector, Dr Fernand Schmid.

Psilotreta assamensis sp. nov.

This species is distinguished from other Asian species by
the greatly reduced lateral processes of segment X, by the
long, slender, acute parameres, and by the inferior appen-
dages, which are broad and have a ventromesal lobe.

ADULT

Forewing 9-12 mm in length in male, 14-17 mm in
female. Head and thorax brown; antennae densely clothed
with appressed white setae making them conspicuous in
contrast with the rest of the insect; maxillary palpi dark
brown; labial palpi light brown; forewings light brown with
patches of yellowish setae between the veins in the middle
of the wing and along the apical margin; hindwing uni-
formly light brown; forelegs light brown; middle and hind
legs whitish.

Maxillary palpi with first two segments subequal and
shorter than the rest, third and fifth subequal and about
twice the length of the first or second and slightly longer
than the fourth. Labial palpi with each segment slightly
longer than the preceding. Vertex with anteromedian warts
small, oval, and raised, a deep groove between them;
anterior pair small, transversely oval, and slanting away
from anteromedian pair; posterior warts twice as large as
anterior warts, two pairs of small bumps between them;
warts bearing long yellowish setae, with shorter setae

Fics. 51, 52. Psilotreta spp., adults. 51. P. schmidi: a-c, 3 genitalia—lateral, dorsal, ventral; d,e,
phallus—lateral, ventral (int. app.—intermediate appendage; lat. pr.—lateral process). 52. P. chinensis, 3
wings.

between warts. Face with lateral warts extending from
level of anterior tentorial pits to antennal sockets and
bearing dark brown setae; frons without setae and indented
above level of anterior tentorial pits to form the groove
between the anteromedian warts. Eyes black, large, larger
in male than female.

Venation in both sexes similar except forewing of
female with M, and M, separated and thyridial cell present;
forewing with discal cell present and extending about one-
third length of wing, R, arising in basal fourth of discal cell
and kinked at crossvein r, R, and R, converging near wing
margin but not meeting, R,,, branched beyond end of
discal cell; hindwing with thyridial cell broad near origin of
Cu,, in female, thyridial cell not broadened in male, M, , 5
undivided in either sex.

Male genitalia (Fig. 53). Segment VII without pleural
lobes. Segment IX with median dorsal process long and
triangular and with base wide in dorsal aspect; inferior
appendages each with a broad basal segment having a
mesoventral lobe bearing several short, stiff, thickened
setae at its apex, in addition to the longer, thinner setae
scattered over the rest of the segment, and with a short
apical segment having a broad base and narrow apex in
lateral aspect, the apex broadly rounded in ventral aspect
and bearing numerous short black teeth apically. Segment
X with preanal appendages broadest in basal fourth, taper-
ing gradually to apex, and bearing numerous short, stiff
yellowish setae on basal half and long black setae on apical
half, contrasting sharply in uncleared specimens with
golden yellowish setae of rest of genitalia; median dorsal
process long, acute or truncate in dorsal aspect, slightly
enlarged apically in lateral aspect in some specimens, and
bearing a row of about 10 stiff setae along each side near
apex; body of segment X broad in lateral aspect, largely
membranous, sclerotized along ventral edge, fused dor-
sally to segment IX throughout its length; lateral process(?)
greatly reduced, present as a sclerotized flange arising from
dorsal edge of segment X near midlength, becoming
thicker and flared dorsolaterally, ending far short of apex of
median dorsal process, and bearing dorsally on each side 1
to several setae, which may represent the basal setal
wart(?); intermediate appendages(?) broad, curved,
ramshornlike projections arising from apex of body of
segment X and curved apicodorsally.

Phallus with phallotheca long, cylindrical, slightly
decurved and enlarged ventroapically; endotheca with pair
of long, pointed, slightly curved parameres; aedeagus with
ventral plate sclerotized, with membranous apex bilobed in
ventral aspect, and with phallotremal sclerite elongate and
S-shaped in lateral aspect and having apex flared.

Female genitalia (Fig. 54). The female of P. assa-
mensis is easily distinguished from the other species we

examined by details of the vaginal chamber. Segment IX
lacks depression laterally. Segment X with tergum deeply
incised apically, narrowed and angular in lateral aspect.
Median plate with weak median longitudinal suture. Pos-
terior vaginal sclerites broad in lateral aspect, narrow in
ventral aspect; dorsal edge above posterior sclerites broad
in ventral aspect; anterior portion of vaginal chamber
slightly longer than wide in ventral aspect.

LARVA

The larva of P. assamensis is unknown, but may corte-
spond to Psilotreta sp. collected in Kameng, where the
majority of the specimens of P. assamensis were collected.

MATERIAL EXAMINED

(All specimens were collected by F. Schmid and are depos-
ited in the CNC, Ottawa, except for five paratypes in the
ROM.)

Holotype 6: INDIA—Assam, Kameng, Rahung,
17.vii.1961 (#281). Paratypes: INDIA—Assam,
Kameng: Amatulla, 17.v.1961, 3 3d (#192); Bilo La,
10.vi.1961, 1 6 2 2 (#241); Bokhar, 28.v.1961, 1 3
(#218); Chug, 25—-31.vii.1961, 3 d 1 2 (#300); same, 9-
13.viii.1961, 1 3d (#307); Dirang Dzong, 18.vii.1961, 3
3 1 & (#283); same, 21-22.vii, 2 2 (#292); Kujjalong,
28-30.vi.1961, 1 2 (#259), 1 3 (#260); Rahung,
16.vii.1961, 3 2 (#279); same, 17.vii.1961, 3 d 1 2
(#281); Salari, 10.vii.1961, 2 5 1 2 (#275), 2 d (#276).
Sikkim: Dikchu, 9.v.1959, 1 d (#119); Lingtham, 8—
10.viii.1959, 1 o (#243); Mangang, 9.v.1959, 1 3
(#122); Nampung, 8.v.1959, 1 d (#116); Singhik, 7—
10.viii.1959, 1 5d (#242). Non-type specimens: the fol-
lowing four females are not included in the type series
because they were not collected in association with males.
INDIA—Assam, Kameng: Lifakpo, 29.v.1961, 1
(#222); Moshing, 8-10.ix.1961, 1 2 (#333); Jhum La,
15—22.ix.1961, 1 2 (#337); Talung Dzong, 3.vi.1961, 1
2 (#228).

Psilotreta sp.

The larvae and pupae described here may be P. assamen-
sis; they were collected in Kameng, some of them from the
same locality as one of the non-type females listed under
that species. However, since the pupae are insufficiently
developed, except one female pupa which had rotted and
could not be identified, we choose not to assign a name to
this larva until better evidence becomes available.

LARVA (Figs. 43-45)

General appearance typical for genus. Head without dis-
tinct colour pattern but dorsal surface light, with contrast-
ing dark muscle scars and irregular darker areas on

Fics. 53, 54. Psilotreta assamensis. 53. a—c, 3 genitalia—lateral, dorsal, ventral; d,e, phallus—tiateral
ventral (int. app.—intermediate appendage; lat. pr.—lateral process). 54. a,c, 2 genitalia—lateral, ventral.
b,d, vaginal apparatus—lateral, ventral.

parietals along anterior border with frontoclypeus; head
much darker posteriorly and ventrally, with a large light
area around eyes. Pronotum brown, with lighter areas
laterally, and posterior ridge black. Mesonotum brown,
with lighter areas at anterolateral corner and at bases of
some of the larger dorsal setae, and with posterior ridge
black from middorsal line to ventrolateral border. Meta-
notum with anterior sclerite brown, lateral borders yellow-
ish, lateral sclerites brown and having some darkening
posterodorsally, and posterior sclerite light yellowish
brown. Pleurites brown with suture and anterior borders
black; legs light brown basally with distal segments
becoming darker and with proximal articulation of coxae
and femora blackened.

Head longer than wide; seta 17 slightly more than one-
half length of, and similar in appearance to, seta 15.
Pronotum with anterolateral corner of size and shape simi-
lar to P. frontalis, approximately 0.2 x middorsal length
of tergite and slightly upcurved. Mesonotum with posterior
inflection indicated as a short, thin, light streak extending
anteriorly from ventrolateral extent of posterior ridge, and
without an indentation along posterior margin of sclerite;
mesepisternum with anteroventral corner slightly produced
ventrally; mesofemur with | to 2 secondary setae arising
dorsally basad of seta 1 and slightly shorter and thinner
than seta 1, and with position of ventral seta 3 variable
relative to dorsal seta 1 and ventral setae 4 and 5. Meta-
notum (Fig. 45) with anterior sclerite bearing 3446 setae,
posterior bearing 16-20, each lateral bearing 28-37.
Abdominal segment VIII bearing 20 lateral tubercles on
each side. Abdominal gills: dorsal—II 23-25, III 30-33,
IV 24-25, V 17-20, VI 12-13, VII 9-10, VIII 9-12;
lateral—III 5S—7; ventral—II 19-23, III 26-30, IV 26-27,
V 24-25, VI 20-23, VII 16-22, VIII 9-14. Larva up to 12
mm in length; head up to 1.3 mm in width across eyes.
Larval case typical for genus, tapered and curved in early
instars and curved with little or no taper in final instar;
posterior end with a single rock fragment cemented over
opening; case up to 18 mm in length.

Pupa (Fig. 10a,b). The pupa of this species is described
in the generic diagnosis. Pupal case typical for genus, up to
18 mm in length.

MATERIAL EXAMINED

INDIA—Assam, Kameng: Jhum La, 17.1x.1961, many L,
| P (decomposed) (ROM); Kalaktang, 14.v.1961, many L
and PP, 3 P (Rom).

NOTES

The position of ventral seta 3 of the femur (Fig. 43) is
variable relative to the position of dorsal seta | and ventral
setae 4 and 5, being distad of | by less than the distance
between 4 and 5 in most specimens, and by approximately

the same or a greater distance in other specimens. This may
be because of difficulty in interpretation owing to the
presence of the secondary dorsal setae, or it may indicate
that there are two closely similar species present.

The occurrence of early instars and pupae in samples in
May and September may indicate a life cycle requiring at
least two years for completion, as is found in Nearctic
species, coupled with prolonged emergence of adults. It is
also possible that two species are present, as suggested
above.

Psilotreta kwantungensis Ulmer

Psilotreta kwantungensis Ulmer, 1925:66—68, figs. 49-52.
Holotype 6 (zim), China, Lienping, 8.iv.1920.

This species is most closely related to P. lobopennis
Hwang; both have the median dorsal process of segment X
produced as a long narrow hood overhanging the rest of the
segment, and have inferior appendages each with the basal
segment bearing prominent dorsal and ventral lobes and
with the apical segment arising between the lobes. The
male of P. kwantungensis is distinguished from those of P.
lobopennis by the longer apex of the body of segment X
and by the shorter ventral lobe of the basal segment of the
inferior appendage.

ADULT

Forewing 10 mm in length in male, 12 mm in female. Head
and thorax brown; abdomen blackish; antennae whitish;
legs lighter than thorax; wings uniformly brown. Venation
(Ulmer, 1925, figs. 49, 50) without modifications; discal
cell present and short, with R, having a kink at crossvein r.

Male genitalia (Fig. 55). Segment VII with posterior
lobe on pleural membrane. Segment IX with anterolateral
lobe large; small wartlike setal patches present laterally and
ventrally near insertion of inferior appendage; median
dorsal process of segment IX indistinguishably fused with
that of segment X, together forming a long, narrow hood
overhanging body of segment X, in dorsal aspect slightly
constricted at base, broader and parallel sided beyond base
to rounded apex, and bearing several setae; inferior appen-
dages with basal segment C-shaped in lateral aspect,
thumblike dorsal lobe and longer ventral lobe bearing
several short stiff setae at apex, in addition to longer setae
scattered over rest of segment, in ventral aspect with broad
darker area free of setae, extending along mesal margin,
and slightly produced at apex, and with lighter setate
ventral lobe narrowing to apex from this point; inferior
appendages with apical segment arising subapically in
C-shaped basal segment, triangular in lateral aspect, dorso-
ventrally depressed, having apex rounded in dorsal aspect,

Fics. 55, 56. Psilotreta kwantungensis. 55. a—c, 3 genitalia, holotype, Lienping, China—lateral, dorsal,
ventral with detail of apex of inferior appendage; right preanal appendage on type broken as shown, with outline
of left appendage dotted in lateral aspect; d,e, phallus—lateral, ventral (int. app.—intermediate appendage; lat.
pr.—lateral process; pr. app.—preanal appendage). 56. a,c, 2 genitalia, paratype, Mahntsishan, China—
lateral, ventral; b,d, vaginal apparatus—lateral, ventral.

and bearing eight to nine short black spines along apical
margin. Segment X with body fused to segment IX only
near base, completely free from median dorsal process;
lateral processes(?) small, thin, rounded lobes arising near
midlength of body of segment X; intermediate appendages
projecting posteriorly from apex of body of segment X and
extending beneath median dorsal process nearly to apex;
preanal appendages broad basally, with apical half very
narrow and bearing scattered long setae.

Phallus with phallotheca cylindrical and curved in lateral
aspect and both basal and apical ends produced ventrally.
Endotheca bearing large parameres laterally; base broad in
lateral aspect, with a large triangular dorsal projection near
base; ventral margin angled upwards to an acute apex at
about midlength, in ventral aspect appearing as a thin rod.
Aedeagus membranous throughout with two triangular
lobes near midlength and with apex bifid in ventral aspect;
phallotremal sclerite with anterior margin straight, apex
flared in ventral aspect.

Female genitalia (Fig. 56). The female of this species is
easily distinguished from the others we have examined by
the absence of a sulcus on the posterior portion of sternum
IX and by details of the vaginal chamber.

Segment IX with tergum lacking depression laterally;
posterior portion of sternum IX lacking suture and set off
from posteroventral lobes of tergum X by sharp ridge.
Segment X with apex rounded in lateral aspect. Vaginal
chamber with anterior portion bearing low ventromedian
projection; posterior portion with three layers of sclerotiza-

tion, the ventral layer more heavily sclerotized than the
other two, and in ventral aspect with sinuous outer margin.

LARVA
Unknown.

MATERIAL EXAMINED
CHINA—Lienping, 8.iv.1920, holotype 6d (zim).
Mahntsishan, 20.vii.1915, paratype 2 (zim).

Psilotreta trispinosa Schmid

Psilotreta trispinosa Schmid, 1965:148, pl. 7, figs. 9,10.
Holotype ¢ (Alexander Koenig Museum, Bonn),
China, Wenchow, 15 April 1939.

This species is easily recognized by the distinctive three-
spine arrangement of the lateral processes and intermediate
appendages of segment X.

To the original description we add abdominal segment
VII with pleural membranes only slightly produced
posteriorly.

FEMALE AND LARVA
Unknown.

MATERIAL EXAMINED
CHINA—Wenchow (Chekiang), 18.iv.1939, paratype 3
(CNC).

Species Not Examined

Psilotreta lobopennis Hwang, 1957:394-39S, figs. 96-99,
3, China.

Psilotreta ochina Mosely, 1942:345-346, figs. 6-8, d @.
Holotype 3d (BMNH), China, vicinity of Foochow,
1935-1937.

Psilotreta orientalis Hwang, 1957:393-394, figs. 92-95,
3. China.

Psilotreta quinlani Kimmins, 1964:49-SO, figs. 38-43, 3
2. Holotype ¢ (BMNH), Nepal, Ulleri, 1830-2135 m,
19.v.1954.

Phylogenetic and Biogeographic Considerations

Twenty species of Psilotreta are now recognized; 6 are
known from eastern North America, and 14 from Nepal,
India, China, Korea, Siberia, and Japan. These two geo-
graphic groups are also separated phylogenetically, the
North American species forming a monophyletic
assemblage distinct from the Asian species.

Monophyly of the Nearctic species is supported by two
characters in the male genitalia. All lack parameres on the
phallus, whereas all the Asian species possess parameres.
Since the presence of parameres is a plesiomorphic condi-
tion in Trichoptera (Nielsen, 1957, 1981; Schmid, 1970,
1980), absence of these structures in the North American
Psilotreta is a synapomorphy uniting these species in a
monophyletic group. On all North American Psilotreta the
apical segments of the inferior appendages bear a few
relatively stout black teeth (e.g., Figs. 1lc, 17c), whereas
most of the Asian Psilotreta have numerous small spines
(e.g., Figs. 46c, 48c). Numerous small spines is the proba-
ble plesiomorphic condition; certain other genera of o-
dontocerids have numerous very small spines (e.g.,
Odontocerum, Marilia, Nerophilus), and the same condi-
tion occurs in the sister family of the odontocerids, the
Philorheithridae. Thus, we propose that a reduced number
(fewer than 10) of larger teeth on the apical segment of the
inferior appendages in the North American species, and at
least one Asian species, is an apomorphic condition. It is
probable that this condition in the Asian species P. kwan-
tungensis (Fig. 5Sc) (and P.. lobopennis?) is independently
derived, since it is associated with a dorsoventral depres-
sion of the apical segment which does not occur in the
North American species. We have not been able to exam-
ine the females, larvae, or pupae of a sufficient number of
Asian species to detect any characters of phylogenetic
value in these stages.

Further analysis of the phylogenetic relationships among
the North American species was carried out by means of an
outgroup analysis using the Asian Psilotreta, the other
genera of the Odontoceridae, and other limnephiloid fam-
ilies as the outgroups. Characters for which hypotheses of
ancestral and derived states could be developed are listed in
Table 1. In the cladogram (Fig. 57) resulting from the
analysis in Table 1, two clusters of three species each are
proposed—the indecisa and rufa subgroups. The species in
these two groups are readily resolved as three-taxon
statements.

Published descriptions of the Asian species that we have
not examined are inadequate for phylogenetic analysis;
however, we advance the following observations. Certain
groups of closely related species can be recognized among
the Asian Psilotreta. The three species P. kisoensis, P.
falcula, and P. locumtenens share apparently derived
characteristics:” the lateral processes of segment X are bean
shaped and lack projections, and the basal segment of each
of the inferior appendages has an apical projection ven-
trally. P. kwantungensis and P. lobopennis Hwang are
sister species as indicated by the fused condition of the
median dorsal and the lateral processes of segment X and

* While this manuscript was in press, we noted the description of
a new species of Psilotreta from Korea, P. pyonga Olah, 1985
(J. Olah, 1985, Three new Trichoptera from Korea. Folia
Entomologica Hungarica 46(1):137—142). The species as
described is difficult to distinguish from P. falcula Botosa-
neanu, with which it is closely related, if not conspecific.
Comparison of types and of variability in series of the two
species will be required to confirm the validity of P. pyonga.

TABLE |. Character states for cladistic analysis of North American Psilotreta species.

Character

1. Parameres of male present

2. Spines on apical segment of inferior
appendage

3. Median dorsal process of segment X
of male

4. Larval head seta 17

5. Larval head capsule flat dorsally

6. Warts on vertex of male

7. Hindwing of female
8. Maxillary palp of male unmodified

9. Vertex of male

L] Plesiomorph State

@ Apomorph State

numerous, small

well developed, sclerotized
long, thick, dark

small, unmodified

discal cell closed

without eversible lobes

absent

few, large
reduced, membranous

short, thin, clear
domed dorsally

very large, raised medially, with scalelike
setae

discal cell open
second segment with eversible lobe
with paired eversible lobe

57 2

Fic. 57. Cladogram for the North American species of Psilotreta. Characters numbered as in Table 1; open
squares represent plesiomorph character states, closed squares represent apomorph character states.

by the C-shaped basal segment of the inferior appendages.
The new species, P. schmidi, appears most closely related
to P. japonica because of the orientation and shape of the
projection of the lateral processes, the reduced intermedi-
ate appendages of segment X, and the basomesal projec-
tion of the first segment of the inferior appendages. Two
species that we have not examined (P. ochina Mosely and
P. orientalis Hwang) appear from the illustrations and
descriptions to be related on the basis of the unique foot-
shaped apical segment of each inferior appendage, and
perhaps also on the basis of the apex of the median dorsal
process of segment X, which is very wide in lateral aspect.
Kimmins (1964) has suggested that P. orientalis is related
to P. quinlani Kimmins on the basis of the structure of
segment X; however, if the illustrations and descriptions of
the two species are accurate, it is difficult to relate these
two species very closely. In fact, P. quinlani may have
more in common with P. trispinosa, the only other species
having bifurcate intermediate appendages. The remaining
species are rather isolated from the rest of Psilotreta: P.
chinensis because of the very short preanal appendages, the
dorsal projection of the lateral processes of segment X, and
the accessory processes of the endotheca; P. assamensis

because of the reduced lateral appendages; and P. quadrata
because of the fused condition of the median dorsal process
of segment X with the lateral processes to produce a large
hood over the phallus, and the distinctive inferior appen-
dages. The association of larvae and females of more of the
species will improve our understanding of the relationships
within Psilotreta. We also expect that many undescribed
species remain to be discovered in Asia.

The disjunct distribution of the species of Psilotreta in
temperate eastern Asia and eastern North America coin-
cides with a widely recognized global biogeographic pat-
tern (Boufford and Spongberg, 1983). Yet, absence of
Psilotreta from Europe is at variance with the generaliza-
tion, derived from analysis of several arthropod lineages,
that the faunas of Europe and eastern North America have
shared a more recent common biota than either has shared
with Asia (Allen, 1983). The sister genus of Psilotreta
appears to be the European Odontocerum, giving further
support to a Eurasian origin for Psilotreta. The fact that the
Nearctic component of Psilotreta is a derivative group is
consistent with the suggestion (Schmid, 1970:107) that the
more primitive elements of lotic lineages in Trichoptera
have an Oriental origin.

List of Species

The 20 species of Psilotreta are listed with their distribu-
tions and references to figures. New synonymies estab-
lished herein also are listed.

NORTH AMERICAN SPECIES

indecisa Group
indecisa Subgroup
P. indecisa (Walker, 1852). Eastern North America. Figs.
p45. Oem 1242s 52 Oo:
Heteroplectron borealis Provancher, 1877.

P. frontalis Banks, 1899. Eastern North America. Figs. 2,
£3,,14;,.26; 33,38:
P. hansoni Denning, 1948. Status unconfirmed.

P. labida Ross, 1944. Eastern North America. Figs. 9, 15,
16, 27, 34, 38.

rufa Subgroup
P. rufa (Hagen, 1861). Eastern North America. Figs. 6-8,
L718; 28529,.55, 59.
Heteroplectron? dissimilis Banks, 1897.

P. amera Ross, 1939. Georgia, North Carolina, South
Carolina, Tennessee. Figs. 3, 22, 23, 31, 37, 39.

P. rossi Wallace, 1970. North Carolina, Virginia, West
Virginia. Figs. 19-21, 30, 36, 39.

ASIAN SPECIES

P. japonica (Banks, 1906). Japan. Figs. 46, 47.
P. schmidi sp. nov. Eastern India. Fig. 51.

P. falcula Botoganeanu, 1970. North Korea.

. kisoensis Iwata, 1928. Japan. Figs. 40-42.

. locumtenens Botosaneanu, 1970. North Korea.
. kwantungensis Ulmer, 1925. China. Figs. 55, 56.
. lobopennis Hwang, 1957. China.

. ochina Mosely, 1957. China.

. orientalis Hwang, 1957. China.

. quinlani Kimmins, 1964. Nepal.

. trispinosa Schmid, 1965. China.

. chinensis Banks, 1940. China. Figs. 48-50, 52.
. assamensis sp. nov. Eastern India. Figs. 53, 54.
. quadrata Schmid, 1959. China.

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Acknowledgements

This study was conducted under the support of an operating
grant (to G. B. Wiggins) from the Natural Sciences and
Engineering Research Council of Canada. Earlier field-
work was supported by research grants (to G. B. Wiggins)
from the National Science Foundation of the United States
and the Canadian National Sportsmen’s Show. To these
sources and to the individuals and institutions listed pre-
viously as sources of additional material, especially type
specimens, we extend sincere appreciation. O. S. Flint

assisted in providing references and specimens. J. C.
Morse reviewed the manuscript and provided helpful com-
ments on the interpretation of venation; F. Schmid also
reviewed the manuscript. Zile Zichmanis drew all the
illustrations, except for Fig. 8a—c,g, which were drawn by
Anker Odum. Nina Cunniff assisted in compiling locality
data. Susan Pasch and E. R. Fuller helped to organize and
check the manuscript.

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