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Life Sciences Contribution 72
Royal Ontario Museum

Contributions to the

Systematics of the
Caddisfly Family Molannidae
in Asia (Trichoptera)

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GLENN B. WIGGINS

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ROYAL ONTARIO MUSEUM

Contribution No. 72
LIFE SCIENCES

ROYAL ONTARIO MUSEUM

Contributions to the Systematics
of the Caddisfly Family Molannidae
in Asia (Trichoptera)

Publication date: 11 July, 1968
Suggested citation: Life Sci. Contr., R. Ont. Mus.
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GLENN B. WIGGINS is Curator of the Department of Entomology and Invertebrate
Zoology, Royal Ontario Museum, and Professor in the Department of Zoology,
University of Toronto.

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PRINTED AT THE UNIVERSITY OF TORONTO PRESS

Contributions to the Systematics
of the Caddisfly Family Molannidae
in Asia (Trichoptera)

Abstract

Study of a collection of several hundred adult caddisflies of
the family Molannidae from India has yielded three new
species of Molanna and five new species representing a new
genus, Indomolannodes. For the new genus Indomolannodes,
which is closest to Molannodes, a key to males and females
of the five new species is presented. The genitalia of the male
holotype of Molanna moesta Banks from Japan are illus-
trated and this name shown to be the senior synonym of the
widespread Asian species known hitherto as M. falcata
Ulmer. Molanna coreana Tsuda is designated a junior syno-
nym of M. submarginalis McL. In the genus Molannodes
variation within Eurasian material of M. tinctus Zett. is found
to include the form described as M. bergi Ross from Alaska,
and this latter name is designated a junior synonym. A key to
the three Holarctic genera of the Molannidae is given.

Introduction

Between the years 1959 and 1962 Dr. Fernand Schmid, now of the
Entomology Research Institute, Ottawa, collected Trichoptera in India.
Included among these exceedingly rich research materials are several
hundred specimens of the family Molannidae. Dr. Schmid has made the
molannid material available to me and this paper presents the results of my
study of it. A few specimens from other sources have been added. Dr.
Schmid’s specimens are divided between his own collection, where all of
the holotypes are deposited, and that of the Royal Ontario Museum.

This molannid material comprises only adult specimens, with no imma-
ture stages represented. Specimens were collected by net during the day, or
by attracting them to a light at night. Those taken by the latter method are
so identified by the words at light. The collection is of considerable interest
because of the relatively large number of new species it adds to this small
family. There are three new species in the genus Molanna, and five in a
genus previously unknown. During the course of the study it has become
evident that several existing names are actually synonyms for other Asian
species, and the appropriate changes are here proposed.

Genus Molanna Curtis
Molanna moesta Banks

Molanna moesta Banks 1906, p. 110, pl. III, figs. 5, 6.
Molanna falcata Ulmer 1908, p. 347, figs. 8-12. NEW SYNONYMY.
Molanna falcata Martynov 1910, p. 367, figs. 15-18.

This species was originally based on a single male specimen from Gifu
(Honshu), Japan, but is unidentifiable from the original description and
illustrations. In order to clarify its status, this specimen (Type 11839,
Museum of Comparative Zoology, Harvard) has been examined and the
genitalia illustrated here (Fig. 2). This specimen lacks a head, but is
otherwise complete and adequate for comparative study. The genitalic
structures had been cleared previously and some of the appendages showed
slight damage.

In describing Molanna falcata from Japan, Ulmer stated it was dif-
ferentiated from M. moesta by darker colouring, by the presence of apical
fork no. 5 (i.e. separation of Cu;, and Cuj,) in the hind wing, and by
different genital appendages. I have not seen the holotype of M. falcata,
but Ulmer’s illustrations and description seem clear, if somewhat dia-
grammatic.

Following comparison between the holotype of M. moesta and Ulmer’s
description of M. falcata, it is my conclusion that valid reasons do not now
exist for the separation of the two, and therefore I am regarding M. falcata
Ulmer as a junior synonym of M. moesta Banks. Evidence in support of
this is as follows.

1) The genitalia of the holotype of M. falcata illustrated by Ulmer are
almost identical with the holotype of M. moesta, even to the arched and
rather narrow sclerotized plates of the tenth segment, as seen in lateral
view. Although this same structure in dorsal view is shown by Ulmer
(1908, fig. 10) to be a single plate with a deep apical incision, in the
holotype of M. moesta it is composed of two separate plates. A similar
situation is shown by Martynov (1910, fig. 17) for Siberian specimens
which he assigns to M. falcata, and it seems likely that Ulmer’s drawings
are in error on this point. The degree of separation of these two plates
appears to be variable in different figures, and it seems likely, too, that
these differences are not significant at the species level. These Siberian
specimens described by Martynov do in fact differ from the holotype of
M. moesta by the somewhat enlarged tip of the sclerotized plates of the
tenth segment, which bears enlarged spines. I concur with Martynov’s
assessment that these are but regional variants of a single species ranging
through Japan and much of Siberia and the eastern U.S.S.R.

2) The separation of Cu;, and Cu, in the hind wing illustrated by Banks
is based upon an incomplete and somewhat irregular development of
Cu, in the type specimen. This vein is evidently not separated in the type
of M. falcata, but this does occur infrequently in Japanese specimens other-
wise identifiable as M. moesta or M. falcata. The beginning’ of Cu, is
shown, in fact, in the hind wing of the Siberian specimens (Martynov 1910,

Z

fig. 16). This venational difference is therefore not significant at the species
level. The other venational features of the holotype are in close agreement
with the illustrations of M. falcata given by both Ulmer and Martynov.
3) In colour M. falcata is stated by Ulmer to be darker than M. moesta,
and judging by the total wing expanse measurements, the type of M. moesta
(length of fore wing 12 mm.) is somewhat larger than M. falcata (total
wingspread 21-23 mm.). The type specimen of M. moesta is light brown in
colour. In view of the evidence already advanced, however, I would be
inclined to interpret these differences of colour and size as variants within
a single species—M. moesta. Lack of any means of distinguishing between
the two has also been noted by Dr. Kuwayama (pers. comm.) in Japan.

Specimens available to me of the probable female of M. moesta have
been examined, and there is a slight range of variation among them. One
of these (Tokyo) is illustrated (Fig. 4) and its diagnostic characters com-
pared with the following species under that heading.

Records available for this species under the two names M. moesta and
falcata show it to be widely distributed in Japan and China and over much
of eastern Siberia (Fischer 1964).

Molanna paramoesta sp. n.

Represented by many specimens from a wide assortment of localities and
habitats in the collections from India is a Molanna, which although very
close to M. moesta in structure throughout, does show consistent differences
in the male genitalia from M. moesta. In view of the fact, too, that morpho-
logical differences between the well-known species of Molanna are not
great, it seems best to regard these specimens as representing a new species,
at least until intermediates become known.

ADULTS. Length of fore wing: male 9-10 mm.; female 11-12 mm. Overall
structure as in M. moesta and typical for the genus; overali colour medium
brown. Apical segments of first legs medium brown, second and third legs
and femora of first lighter. Fore wings medium brown with some lighter
patches, more pronounced in the females. Venation as in M. moesta
(Ulmer 1908, figs. 8, 9); hind wing with the groove in the anal part,
regarded by Martynov (1910) as a character diagnostic for the subgenus
Molanneria.

Male Genitalia (Fig. 1). In general structure similar to M. moesta
(Fig. 2), but distinguished from that species by the inflated, hammer-like
form of the paired sclerotized plates of the tenth segment in lateral view.
In M. moesta this structure appears deflated and pendant. Comparison of
the genitalic structures of the holotype specimens for these two species will
indicate certain other differences, such as those in the superior (lateral)
appendages of the tenth segment. These appear to be somewhat variable
within each species and are not reliable diagnostic characters. The same is
true of the degree of separation of the two sclerotized plates of the tenth
segment in dorsal view, which more likely is determined by their position
at death.

M. moesta

Figs. 1-2—Male genitalia in Molanna spp. 1. M. paramoesta sp. n. (holotype).
2. M. moesta Banks (holotype). a, lateral view; 6, ventral view;
c, dorsal view; d, caudal view.

Female Genitalia (Fig. 3). Essentially similar to females assumed to
represent M. moesta (Fig. 4; Tokyo, 24 May 1931; MCZ), but separable
by a number of characters of degree. These appear to be subject to con-
siderable variation in M. paramoesta, at least, where many specimens are
available for study. Following is a summary of all of the characters which
have been found to have some diagnostic significance. The specimens
illustrated present all of these, but others do not. It appears that distinctions
between the females of these two species have to be based on a consensus
of the majority of the characters involved.

Aah ter
Wy eT

\ My ‘
RONCHI 2
aa aN | ee ees
CS rey,

WISE Ad | Ly [i 3b

Figs. 3-4—Female genitalia in Molanna spp. 3. M. paramoesta sp. n.
4. M. moesta Banks. a, lateral view; b, dorsal view; c, ventral view.

=)

Segment IX of M. paramoesta in ventral view with a wide lateral collar
or trough at the junction with the tenth segment; in M. moesta this trough
is barely represented.

Segment X of M. paramoesta in ventral view with a pair of blunt
posterolateral lobes and with a thinly sclerotized ridge running the length
of the rectangular sclerite lying at each side of the genital opening; in dorsal
view a pair of very low lobes lies mesad to the posterolateral lobes, and a
short triangular median ridge has its base at the junction of segments IX
and X. Segment X of M. moesta in ventral view bears a pair of acute
posterolateral lobes, and a heavily sclerotized ridge running the length of
the rectangular sclerite lying at each side of the genital opening; in dorsal
view a pair of fairly prominent lobes lies mesad to the posterolateral lobes,
and a long triangular median ridge has its base at the junction of segments
IX and X.

TYPES (all collected by F. Schmid, unless otherwise indicated). Holotype,
male: INDIA, Madhya Pradesh: Satanwara, 1308 ft., 28 Nov. 1961. Para-
types: INDIA, Assam, Manipur: Huiahu, 4300 ft., 1 July 1960, 1 4, at
light; Khaiyang, 3200 ft., 18 June 1960, 1 ¢, at light; Khamassom, 3900
ft., 24 June 1960, 19, at light; Khangairim, 4145 ft., 29 June 1960, 1°,
at light; Khanggoi, 4828 ft., 16 July 1960, 44 29, at light, Mapum,
3300 ft., 12 June 1960, 1 ¢, at light; Sirohi, 4300 ft., 26 June 1960, 4 ¢ ;
Ukhrul, 6000 ft., 4 June 1960, 14, at light. Assam, United Jaintia and
Khasi Hills: Demthring, 3800 ft., 16 April 1960, 1¢, at light; Mynso,
3500 ft., 20 April 1960, 26 19, at light; Nongjni, 3750 ft., 19 April
1960, 54 59, at light; Shilliang Myntang, 3500 ft., 20 April 1960, 3
29, at light: Wadlaskem, 4250 it.; 17 April 1960.2 ° bid] Ese hor eat
light; Thangrain, 3000 ft., 22 April 1960, 14; ibid., 14 19, at light;
Umlangshor, 4100 ft., 18 April 1960, 146 19; ibid, 1¢é 49, at light.
Kerala: Maukut, 200 ft., 15 Jan. 1959, 22, at light. Madhya Pradesh:
Guna, 2000 it., 7 Feb. 1959, 24 1¢, at light; Phalghat, 2000 ft., 6 Feb.
1959, 8¢ 72, at light; Satanwara, 1308 ft., 28 Nov. 1961, 213 239
(type series): ibjd., 62 Feb. 1962, 33° 19, vat light (topotypes); ibid.,
8 Feb. 1962, 256 529, at light (topotypes). Madras: Avalanche, 7200 ft.,
28230 Dec. 1958; 1-9 Kodaikanal, 7400 ft, 5-l2.Dec 1955.23 6 =
Maraiyur, 3000 ft, 19 Dee, 1958,56 19; at light; Perumalmalai, 5000 ft.,
7 Dec. 1961, 14, at light. Mysore: Subrahmanya, 400 ft., 20 Jan. 1959,
1¢, at light. NEPAL: Bharatpur, 8 April 1957, 36 19, EI. Coher;
Rapti Tal, Megouli, 300 m., 29 March-4 April 1962, 1 ¢, G. Ebert und
H. Falkner (coll. Staatslg. Munchen); Shawani, 19 March L957... 2.6; Eek
Coher.

The name given to this species is intended to emphasize its close
similarity to M. moesta Banks.

DISTRIBUTION. Known only from India and Nepal, but very widely distri-
buted in India, from the Himalayas south to Kerala and Madras.

HABITAT AND BIOLOGY. The very wide variety of aquatic situations at which
this species was collected will be treated in more detail by Dr. Schmid in a

6

later publication. At this point it is sufficient to summarize these habitats
as follows: artificial lakes with turbid water and marshy shores; large rivers
with turbid water, little current, scattered rapid sections, gravel bed with
algae; small streams in dense jungle; turbulent streams with clear water,
sand and gravel beds; large slow rivers in bamboo jungle, clear water, sand
bed. Collections of adults were made at elevations ranging from 200 to
7400 feet and throughout most of the year. This appears to be a widespread
and common species of very broad ecological tolerance.

Molanna crinita sp. n.

ADULT. Length of fore wing: male 11 mm. Overall colour medium brown.
Head of male (Fig. 6) with maxillary palpi very much elongated and
covered with very thick and very long dark brown hairs; dorsum of head
recessed around bases of antennae, remaining narrow area between the
antennae with a dense pile of coarse hairs. Pronotum with two patches of
extremely dense, coarse hairs; similar coarse hairs covering the median
area of the mesonotum (Fig. 7). First legs darker than the others, but
second and third tarsi slightly darker apically. Other structures typical for
the genus. Wings medium brown, no distinctive markings evident; venation
(Fig. 8) of fore wings as in M. moesta; hind wing narrow and elliptical,
venation and anal area reduced more so than in M. moesta, a darkened fold
extending the length of the wing, just in advance of the anal margin; anal
margin of hind wing with a very densely packed band of thick hairs.

Male Genitalia (Fig. 5). Without close similarity to the genitalia of any
known species. Ninth segment typical for the genus. Clasper scoop-like with
the posterodorsal and posteroventral corners extended into prominent
processes; a clavate mesal process arises from the base. Tenth segment
with two flat sclerotized plates vertically appressed, the posteroventral
corner of each extended into a linear process with an apical point turned
sharply dorsad. Phallus with four stout spines in the invaginated endotheca.

Female unknown.

TYPE. Holotype, male: INDIA, Assam, North East Frontier Agency,
Kameng Frontier Division: Chug, 7200 ft., 16 April 1961, F. Schmid. This
single male holotype is the only specimen known.

The name given to this species refers to the remarkable development of
long hair on the maxillary palpi and of thick hairs on areas of the head
and thorax.

HABITAT AND BIOLOGY. This specimen was collected with a net along a small
quiet stream in dense jungle.

Molanna saetigera sp. n.

ADULT. Length of fore wing: male 9 mm. Body medium brown overall;
structure typical for the genus. Fore wings clothed with golden brown
hairs; venation as in Fig. 9e; hind wing with a darkened fold running along
the mid-line of the wing, and a patch of very long hairs immediately behind
this line.

M. crinita

Figs. 5-8—Molanna crinita sp. n. 5. Male genitalia (holotype): a, lateral view;
b, ventral view; c, dorsal view; d, caudal view. 6. Head of male.
7. Thorax of male, dorsal view. 8. Wings of male.

Fig. 9—Molanna saetigera sp. n. 9. Male (holotype): a, genitalia, lateral view;
b, ventral view; c, dorsal view; d, caudal view; e, wings.

Male Genitalia (Fig. 9a—d). Without close similarity to the genitalia of
any known species. Ninth segment generally typical for the genus; in dorsal
view with a pair of short sharp points arising on the posterodorsal margin.
Claspers scoop-shaped, the posterodorsal corner extended into a long
process; a finger-like mesal process arising from the base. Tenth segment
with two flat mesal plates vertically appressed; in lateral view each plate is
rounded posteriorly, bearing many long stout spines, and terminating
ventrally in a beak-like incision; in dorsal view each plate bears a short,
thick mesal knob. Phallus strongly arched and bearing four stout spines in
the invaginated endotheca.

Female unknown.

TYPES. Holotype, male: INDIA, Assam, United Jaintia and Khasi Hills:
Mawlang, 4000 ft., 12 April 1960, F. Schmid. Paratypes: same data as
above, 2 ¢.

The name given to this species refers to the long spines on the tenth
segment.

HABIT AND BIOLOGY. These specimens were collected along two medium-
sized, clear rivers with falls, the headwaters of the Wahrashi; two during
the day and one at a light after dark.

Molanna submarginalis McLachlan

Molanna coreana Tsuda 1942, p. 231, figs. 5-7. NEW SYNONYMY.

Although no new material of this species is at hand, it has become apparent
in the course of studying the Asian molannids that M. coreana, by the
published description and illustrations, is not separable from M. submargi-
nalis. In view of the fact that M. submarginalis has been recorded from the
Amur River area (Martynov 1937), its presence in Korea is not surprising.

Genus Molannodes McLachlan

Up to the present time this has been the only other genus known to be
represented in the molannid fauna of the northern hemisphere. The
Eurasian representatives of this genus are now regarded as belonging to a
single species, and study of material which has come recently to hand
requires that the populations in extreme northwestern North America also
be assigned to this species.

Molannodes tinctus Zett.

Molannodes bergi Ross 1952, p. 85, figs. 1-4. NEW SYNONYMY.

Comparison of both sexes in material of Molannodes from Alaska with
specimens of M. tinctus from Finland and from the Amur River area of the
U.S.S.R. shows that there is little, if any, difference among them. Examina-
tion of a series of specimens shows that the differences in the clasper of the
male attributed to the Alaskan specimens are not concordant. The Finnish
and Alaskan specimens are virtually identical, and exhibit some slight
differences in size and male genitalia from the Amur specimens. The
concept of a single species with minor regional variations over its enor-
mous range seems to fit the available evidence.

Although the traditional spelling of this species name has been tincta,
the International Code of Zoological Nomenclature (1961) takes the
definitive position (Art. 30) that generic names ending in -odes are mascu-
line, necessitating this emendation.

10

Genus [ndomolannodes gen. n.

Type-species Indomolannodes decurvatus sp. n.

ADULTS. Basically similar to Molannodes, but distinguished by the structure
of the male genitalia. Dorsum of head (Fig. 10) as in Molannodes, and
distinct from Molanna, with a pair of elongate warts and a second smaller
pair posterolaterad from these; a large triangular wart between the bases of
the antennae. Maxillary palpi stout and long, with a dense hair covering,
5-segmented in both sexes; the basal segment very short, the second seg-
ment twice the length of the basal, the three remaining segments each
approximately twice the length of the second segment. Antennae stout, with
a dense hair covering, slightly longer than the insect itself. Pronotum
(Fig. 10) different in the sexes, as in Molannodes, the females with two
pairs of rounded warts, the males with one; the median warts in the male
replaced by a pair of membranous pouches. Mesonotum and metanotum
typical for the family, without conspicuous warts, the mesoscutellum
swollen and uniformly covered with short hairs. Legs with a dense covering
of short hairs; spurs well developed 2, 4, 4; spines black and well developed.
Wing shape (Figs. 16, 17) as in Molannodes, and distinct from Molanna
in not being curled around the body and in having R»+3 separate from R,
in the fore wing; a row of coupling hooks along the costal margin of the
hind wing, typical for the family. Venation generally as in Molannodes,
but distinct in that M, and Mz, are united in the fore wing of the male, but

OS

©

Figs. 10-—12—10. Indomolannodes excavatus sp. n.: a, head and thorax of male,
dorsal view; 6, head and pronotum of female, dorsal view.
11. Molannodes tinctus Zett., pronotum and part of mesonotum,
male, dorsal view. 12. Molanna flavicornis Banks, head, pronotum
and part of mesonotum, male, dorsal view.

I]

Separate in the female, with the result that the venation of the sexes is
different in Indomolannodes (although in one species, I. excavatus, the
venation is the same in the two sexes as it is in Molannodes).

Male Genitalia (Fig. 13). Characterized by a highly complex develop-
ment of all the component parts represented in Molannodes. Ninth segment
rather narrow throughout, the dorsum exposed in the normal way, but the
entire ventral area, along with approximately one-half the width of the
lateral area, recessed into the eighth segment; the posteroventral margin of
the sternum of segment VIII rolled dorsad and developed into prominent
points or lobes. Claspers composed of two distinct branches: a ventral
branch which in lateral view is distinctly hook-shaped; and a dorsal branch
which usually takes the form of a slender finger-like appendage. Tenth
segment composed of three parts: a dorsomedian roof-like plate; a lateral
pair of prominent superior appendages; and a pair of mesal appendages,
usually extended into conspicuous lobes or processes, which may be
homologous with the intermediate appendages (sensu Nielsen 1957).
Phallus composed of a short, tubular, sclerotized phallotheca, from within
which there can be everted a membranous endotheca bearing a dorsal
extensile arm terminating in one or two bristle-covered lobes, and also
other patches of setae; the ejaculatory duct and phallotremal sclerite can be
seen within the endotheca in certain preparations (Figs. 14e, 15e). A
heavily sclerotized, median piece lies ventrad of the endotheca.

Female Genitalia (Fig. 22). Of the same basic structure as in Molan-
nodes. Sternum of the eighth segment forming a prominent posteroventral
lip which is variously excised in different species. A semi-membranous
plate with many fine linear striations lies at each side of the genital opening.
Tenth segment forms a prominent roof-like plate extending above the
genital opening; the apical margin of this plate is variously developed in
different species.

IMMATURE STAGES. No material is available for this genus.

HABITAT AND BIOLOGY. Adults have been collected in large numbers, largely
at lights but also by net, in the immediate vicinity of rivers and streams of
mountainous, wooded country.

DISTRIBUTION. This genus is known from a long segment of the Himalayan
range, the hills situated south of the Brahmaputra valley in Assam and
from northeastern Burma.

REMARKS. The reason for the pronotal warts in Indomolannodes (Fig. 10)
and Molannodes (Fig. 11), but not Molanna (Fig. 12), comprising two
pairs in the females and one pair in the males, appears to be that in the
males the mesal pair is modified into a pair of whitish membranous
pouches. These appear to be capable of sufficient extension that as two
ribbon-like strands they can be passed into a slit-like concavity in the
cleavage depression at the anterior-most point of the mesonotum. In view
of the fact that these structures are totally absent in the females, it seems
clear that they are associated in some way with sexual attraction by the
males. The structure of the membranous pouches is not clear in specimens

12

ww We

he Ea
a ee clasper, ventral branch ~~
ef | NS oe

io SS ee aS Y ES
a ae Saee seg> VAllli =e eae a

13¢

5 seg. X, superior appendage

~—>~ seg. X, mesal appendage --~-4

I. incurvatus

Figs. 13-14—Male genitalia in Indomolannodes spp. 13. I. decurvatus sp. n.
(holotype). 14. J. incurvatus sp. n. (holotype). a, lateral view;
b, ventral view; c, dorsal view; d, caudal view; e, phallus (from

paratypes).
13

I. excavatus

17

I. decurvatus fe)

Figs. 15—17—15. Indomolannodes excavatus sp. n., male genitalia (holotype):
a, lateral view; b, ventral view; c, dorsal view; d, caudal view;
e, phallus (from paratype). 16. J. decurvatus sp. n., wings of
male. 17. 7. decurvatus sp. n., wings of female.

14

of Indomolannodes available to me, which are dried, but in Molannodes,
where alcoholic specimens are available, the hard parts of the pronotum and
mesonotum are identical, and the membranous pouches (Fig. 11) can be
seen in detail.

The name given to this genus is intended to convey the concept of a

group related to Molannodes, with distribution centred in India. As indi-
cated under the preceding genus, the name is of masculine gender.

Key to the Species of Indomolannodes

iy

in)

Males .. I ar SAEED ota rel PEt i 8 ent anne A olen ate Soe te D
Isic UIIeelll Spee RM © uD Rin rari ce py te Niet wl OB ad oa a Fa nth a PE sere tan 6
Mesal appendages of tenth segment in the form of broad, leaf-
Hike mes) AMS TOMS o CheU OS Sea AE A he Sees a kan eel Ala ere ag ONE ee: g
Mesal appendages of tenth segment not expanded and leaf-like
CEigs. 18, 19) See aE I ics tte enn I ie gmter red are eect etn 4
. Dorsal branch of the clasper with its apex curved strongly ven-

trad; mesal appendage of the tenth segment with a very long,

slender, pointed, ventral process and a small dorsal protu-

Benamcem (liga)! Woke. Aol Ne ReaD ise eich eee I. decurvatus
Dorsal branch of the clasper with its apex curved strongly

mesad; mesal appendage of the tenth segment with a broader,

pointed, ventral process and a finger-like dorsal process

(IRIS V1 EO ee RU nee are Were Ae ce at eer ee eee ee I. incurvatus
Mesal appendages of tenth segment with long and very slender

processes; posterior margin of the eighth sternum without a

Prominent mediamenotclh wigs. 1S 1D een 5 tere et es at 5
Mesal appendages of tenth segment without long, slender

processes; posterior margin of the eighth sternum with a

PLrommentemedian noten (Pigs WD) as I. excavatus

. Posterior margin of the eighth sternum extended dorsad into a

lateral pair of prominent vertical lobes, each with an apical

horizontal lip; dorsal branch of the clasper finger-like in

lavenaligvicwe (tees) eee Ce Pie orc ie: eee ee ee I. falcifer
Posterior margin of the eighth sternum extended caudad into

a lateral pair of pointed lobes; dorsal branch of the clasper

widened in lateral view, with a slender terminal process

LS OA Rok es oe Ml AS. aeteat cy te Rea Aa den) en se aoe Se I. comans
Tenth segment extended into an elongate, compressed median

RO CCSSIAM UCSB. SNE AIA Bote ee ANE started a ball San dees Ganado’ dxgh
Tenth segment without a compressed median process (Figs.
20-22) ee eRe ERE Ee a ee ar nC ee coe 8
Median process of tenth segment of uniform depth in lateral
SUR emt a sg) ch Mie Mee eo Ry 9 2g ts Pecan eo SE aL shown _I. incurvatus
Median process of tenth segment with a pronounced knob at
tie taper Ai ratetal wiew UG” 23)! oy cco cada crtacc ase radsesic I. comans
. Posterior margin of eighth sternum in ventral view entire;
apical margin of tenth segment in ventral view rounded and
SOME WU thNCAte (EUR a 22)) over cc sncscnsie. seis srs sasavutpemenaenls': I. excavatus
Posterior margin of eighth sternum with a prominent mesal
notch (Figs. 20, 21) et ee hile Fd en nae Se 9

iS

9. Posterior margin of eighth sternum with mesal notch wide;
apical margin of tenth segment evenly rounded in ventral
VIEW TCE IIE) eter se oe orci be cat iced ee ee, Scene te ee I. decurvatus
Posterior margin of eighth sternum with mesal notch much
narrower and somewhat V-shaped; apical margin of tenth
segment roughly trilobate in ventral view (Fig. 20)... I. falcifer

Indomolannodes decurvatus sp. n.

ADULTS. Length of fore wing: male 6 mm.; female 7 mm. Overall structure
typical for the genus; medium brown in colour. Antennae with very
narrow light alternating bands. Fore wings uniform medium brown,
thickly covered with hairs. Venation (Figs. 16, 17) different in the sexes,
with M, and Mz united (usually) in the fore wing of the male but separated
in the female.

This species is most readily distinguished by characters of the male
and female genitalia.

Male Genitalia (Fig. 13). Most closely resembling J. incurvatus, but
clearly distinguished by characters given in the key. Ninth segment narrow
on the extreme dorsum, but of fairly uniform width elsewhere. Posterior
margin of the eighth sternum rounded dorsad and extended into a pair of
small pointed lobes. Ventral branch of the clasper shaped like a large hook
in lateral view; dorsal branch of the clasper in the form of a long slender
process which curves ventrad. Tenth segment with dorsomedian plate
short and parallel-sided; superior appendages simple lobes; mesal appen-
dages enlarged and leaf-like with a very long and slender ventral process
and a small dorsal protuberance. Phallus with a sclerotized, bifurcate
ventral piece and an extensile, bristle-tipped dorsal appendage (illustrated
with the endotheca invaginated because efforts to evert it in several speci-
mens met with no success).

Female Genitalia (Fig. 21). Most closely resembling J. falcifer, but
distinguished by characters given in the key. Posterior margin of the eighth
sternum with a wide, rounded excision in ventral view. Apical margin of
the tenth segment extended into an evenly rounded lobe.

TYPES (all collected by F. Schmid, unless otherwise indicated). Holotype,
male: INDIA, Assam, Manipur: Chingsao, 4500 ft., 15 June 1960. Para-
types: INDIA, Assam, Manipur: Chingsao, 4500 ft., 15 June 1960, 7 3
11 ¢ (type series); ibid., 5400 ft., 13 June 1960, 3 ¢, at light; ibid., 3800
ft., 14 June 1960, 36 119, at light; Langdang, 5300 ft., 5 June 1960,
28 19, at light; Mattiyang, 2800 ft., 17 June 1960, 1 ¢, at light; Sirohi
Kashong, 6200 ft., 10 June 1960, 24 19, at light. BURMA: Kambaiti
(N.E. Burma), 7000 ft., 1 June 1934, 1 6 (Swedish Exp. to Burma; coll.
Riksmuseum, Stockholm).

The name given to this species refers to the ventrally directed dorsal
branches of the claspers.

DISTRIBUTION. Known only from Manipur (Assam) and northeast Burma.

16

HABITAT AND BIOLOGY. All the available material of this species bearing
information concerning habitat was collected at lights, but in every case the
lights were located beside turbulent streams in densely wooded moun-
tainous areas, at elevations ranging from 2800 to 7000 feet. Adults were
collected only in June, occasionally along with /. falcifer.

Indomolannodes incurvatus sp. n.

ADULTS. Length of fore wing: males 6-7 mm.; females 8-9 mm. Overall
structure typical for the genus; somewhat lighter brown in colour than the
other species. Head with light hairs; antennae with light alternating bands.
Fore wings with light hairs mixed with the brown, apparently more so in
the males, but the somewhat rubbed condition of the wings in most of the
specimens make this uncertain. Venation as in J. decurvatus, different in
the sexes with M, and Mz united in the fore wing of the male but separated
in the female.

This species is most readily distinguished by characters of the male
and female genitalia.

Male Genitalia (Fig. 14). Most closely resembling J. decurvatus, but
clearly distinguished by characters given in the key. Ninth segment nar-
rowed at the dorsal and ventral extremities, wider laterally, with a promi-
nent point arising mid-laterally from the posterior margin. Posterior margin
of the eighth sternum rounded dorsad to form a pair of smoothly margined
lobes. Ventral branch of the clasper hook-shaped in lateral view as in
I. decurvatus, but the hook much smaller (there is some variation in the
size of the hook); dorsal branch of the clasper in the form of a long
slender process with a ventral spur, the process curving sharply mesad,
instead of ventrad as in J. decurvatus. Tenth segment with dorsomedian
plate long and narrow, the lateral margins somewhat deflated; superior
appendages rather more expanded than in J. decurvatus, their margins
irregular; mesal appendages enlarged and leaf-like as in J. decurvatus, but
with a broader, pointed ventral process and a finger-like dorsal process; in
some specimens (Pauri Garhwal: Tungnath) this dorsal process is much
longer than illustrated. Phallus basically typical for the genus, distinct
from I. decurvatus in having a stouter ventral piece and in having two arms
on the extensile appendage, each with a bristle-tipped lobe.

Female Genitalia (Fig. 24). Resembling most closely J. comans, but
distinguished by characters given in the key. Posterior margin of the eighth
sternum slightly excavate in ventral view. Tenth segment developed into a
prominent median process, the lateral margins sharply compressed, the
apex simple.

TYPES (all collected by F. Schmid). Holotype, male: INDIA, Assam,
North East Frontier Agency, Kameng Frontier Division: Shergaon, 6500
ft., 9 May 1961. Paratypes: INDIA, Assam, North East Frontier Agency,
Kameng Frontier Division: Bilo La, 6000 ft., 10 June 1961, 1 ° ; Bomdi
La, 8800 ft., 13-21 June 1961, 1 2; Dirang Dzong, 6500 ft., 21-22 July
1961, 19, at light; Kelang, 6000 ft., 2 July 1961, 24 19, at light; Mosh-

Ly

ing, 6800 ft., 4-7 Sept. 1961, 1 9°, at light; ibid., 7200 ft., 8-10 Sept. 1961,
1, at light; Nafra, 4000 ft., 26 June 1961, 2 4 ; Nakhu, 4500 ft., 3 July
1961, 2, at light; ibid., 4800 ft., 4 July 1961, 22, at light; Nizong,
4800 ft., 27 June 1961, 1 6; Nyukmadong, 6600 ft., 1 Aug. 1961, 1 9,
at light; Rahung, 7000 ft., 16 Aug. 1961, 1¢, at light; ibid., 6500 ft.,
ly Suly 196126 3°, at hehtyabid, 730000. 17 Ame 19Gb isaac
at light; Shergaon, 6800 ft., 1 Sept. 1961, 14, at light; Talung Dzong,
7800 ft., 13-14 Sept. 1961, 1¢, at light. INDIA, Pauri Garhwal: Khum-
yara, 4500 ft., 3-4 May 1961, 1 6; Tungnath, 7500 ft., 1 June 1961, 14.

The name given to this species xefers to the mesally directed dorsal
branches of the claspers.

DISTRIBUTION. Known in the Himalayas from Garhwal (west of Nepal) to
Kameng (northern Assam, east of Bhutan).

HABITAT AND BIOLOGY. The materiai available ail comes irom the vicinity
of turbulent streams in densely wooded mountainous areas, at elevations
ranging from 4500 to 8800 feet. Adults were collected from May to
September.

Indomolannodes excavatus sp. n.

ADULTS. Length of fore wing: male 6-7 mm.; female 7-8 mm. Overall
structure typical for the genus; medium brown in coiour; antennae with
narrow, light, alternating bands; head and thorax as in Fig. 10. Fore wings
uniform medium brown in colour with a dense coating of hairs; venation
unique among the five species of Indomolannodes here described because
in the fore wings of both sexes M,; and Mz are separated, as they are in
Molannodes.

This species is most readily distinguished by characters of the male
and female genitalia.

Male Genitalia (Fig. 15). Generally resembling I. decurvatus and
incurvatus, but abundantly distinct by characters given in the key. Ninth
segment narrowed dorsally and ventrally, of fairly uniform width laterally.
Posteroventral edge of the eighth segment with a rounded median excava-
tion, the two corners of this excavation somewhat enlarged and rolled
dorsad. Dorsal and ventral branches of the clasper separated only near
their apices; dorsal branch forming a curved finger-like appendage in
lateral view, in caudal view a sharp mesal spur is apparent; ventral branch
forming a prominent hook in lateral view. Tenth segment with dorsomedian
plate triangular and the lateral edges convex in dorsal view; superior
appendages with a prominent ventral extension; mesal appendages with
a slender elongate dorsocaudal process. Phallus essentially as in J. incur-
vatus but the median sclerotized piece short.

Female Genitalia (Fig. 22). Most closely resembling I. decurvatus, but
distinguished by characters given in the key. Eighth sternum with the
posterior margin entire. Tenth segment in ventral view evenly rounded
laterally, the apical margin somewhat truncate.

18

TYPES (all collected by F. Schmid). Holotype, male: INDIA, Assam,
Manipur: Poi, 4200 ft., 4 July 1960. Paratypes: INDIA, Assam, Manipur:
Chingai, 4400 ft., 8 July 1960, 1 6, at light; Huiahu, 3800 ft., 3 July 1960,
1 4, at light; Khamassom, 3900 ft., 24 June 1960, 2 ¢, at light; Khanggoi,
4828 ft., 16 July 1960, 1¢, at light; Longbi Khulen, 4500 ft., 30 June
1960, 34 19, at light; Mapum, 3300 ft., 12 June 1960, 1 6 29, at light;
Poi, 4 July 1960, 1 ¢; ibid., 4200 ft., 4 July 1960, 66 49 (type series);
Sirohi, 4300 ft., 26 June 1960, 66 892, at light; ibid., 4100 ft., 9 July
1960, 3 46 99, at light.

The name given to this species refers to the prominent mesal excava-
tion on the eighth sternum of the male.

DISTRIBUTION. Known only from Manipur in Assam.

HABITAT AND BIOLOGY. The data available indicate that this species lives at
somewhat lower elevations, 3300 to 4800 feet, than most of the others. The
collections were made generally in the vicinity of somewhat larger streams,
and often around rice fields. Adults were collected only in June and July.
This species was taken along with Molanna paramoesta Wiggins at several
locations.

Indomolannodes falcifer sp. n.

ADULTS. Length of fore wing: male 7-8 mm.; female 8-9 mm. Overall
structure typical for the genus; medium brown in colour; antennae with
light, narrow, alternating bands. Fore wings with a dense coating of hairs,
medium brown in colour with indefinite light yellowish patches in the apical
area, at the arculus and around the bifurcation of Rs. Venation as in
I. decurvatus.

This species is readily distinguished by characters of the male and
female genitalia.

Male Genitalia (Fig. 18). Most closely resembling /. comans, but dis-
tinguished most readily by characters given in the key. Ninth segment very
narrow dorsally, the lateral portion broadest above the mid-line, and taper-
ing ventrad from that point. Posterior margin of the eighth sternum
extended dorsad into a lateral pair of prominent vertical lobes, each with
an apical horizontal lip; in caudal view a very wide rounded concavity lies
between the two vertical lobes. Ventral branch of the clasper in the form
of a heavily sclerotized, downturned spike; dorsal branch of the clasper
slender and finger-like, bearing a mesal pair of short thin processes. Tenth
segment with dorsomedian plate triangular, the lateral margins concave
in dorsal view; superior appendages somewhat mitten-like in shape, the
broadest part ventrad; mesal appendages each with an extremely slender
sickle-like process, each frequently crossing the other behind the phallus,
and with a flat clavate process arising separately at the base. Phallus of the
same basic structure as the other species in the genus, but with a pair of
slender sclerites in the endotheca; extensile appendage armed with curious
truncate bristles.

19

|. falcifer

I ZF> ee

es Ss
19a ee
“a “ —
“4
YY f
N

I. comans

Figs. 18-19—Male genitalia in Indomolannodes spp. 18. 1. falcifer sp. n. (holo-
type). 19. I. comans sp. n. (holotype). a, lateral view; 5, ventral
view; c, dorsal view; d, caudal view; e, phallus (from paratypes).

20

Female Genitalia (Fig. 20). Resembling most closely /. decurvatus, but
distinguished by characters given in the key. Posterior margin of the eighth
sternum with a narrow V-shaped median notch. Apical margin of the tenth
segment roughly trilobate in ventral view.

TYPES (all collected by F. Schmid). Holotype, male: INDIA, Assam,
United Jaintia and Khasi Hills: Rumkheng, 4500 ft., 24 March 1960.
Paratypes: INDIA, Assam, Manipur: Chingsao, 3800 ft., 14 June 1960,
39, at light; Hkayam Boum, 8000 ft., 20-23 June 1960, 2°, at light;
Huiahu, 5000 ft., 2 July 1960, 16 12, at light; Khopum, 2500 ft., 27
May 1960, 18 39, at light; Mattiyang, 2800 ft., 17 June 1960, 29, at
light; Sirohi Kashong, 6200 ft., 10 June 1960, 3 ¢, at light; ibid., 7000 ft.,
6-7 June 1960, 13 292, at light; ibid., 7000 ft., 11-13 July 1960, 19, at
light; Tairenpokpi, 4000 ft., 31 May 1960, 1 ¢, at light. INDIA, Assam,
United Jaintia and Khasi Hills: Jarain, 2800 ft., 13 April 1960, 1 ¢ ; Lait-
lyngkot, 5000 ft., 15 March 1960, 1 ¢ 29, at light; Rumkheng, 4500 ft.,
24 March 1960, 36 12, at light (type series); ibid., 20-23 March 1960,
1 (topotype); Serrarim, 5800 ft., 3 Oct. 1960, 1 9; Syntung, 4000 ft.,
11 April 1960, 1 ¢, at light.

The name given to this species means sickle-bearing, in reference to
the curved blades on the mesal appendages of the tenth segment.

DISTRIBUTION. Known from Manipur and the Khasi Hills in Assam.

HABITAT AND BIOLOGY. Records for adults of this species range from eleva-
tions of 2500 to 8000 feet, and from March to October. Collections of
adults were made in the vicinity of turbulent streams in densely wooded
areas (where they were taken along with /. decurvatus at some localities),
and medium-sized rivers.

Indomolannodes comans sp. n.

ADULTS. Length of fore wing: male 6-7 mm.; female 7-8 mm. Medium
brown in colour; antennae with light, narrow, alternating bands; overall
structure typical for the genus, but distinctive in having the femora, and
especially the tibiae and the tarsi of the fore legs of the male densely
clothed with long whitish hairs; the hairs on the warts of the head and of
the pronotum are unusually long and dense as well. Fore wings with
yellowish patches on a medium brown hairy covering, largely in the apical
half, but with two larger patches in the basal portion. Venation as in
I. decurvatus.

This species is most readily distinguished by characters of the male and
female genitalia.

Male Genitalia (Fig. 19). Most closely resembling /. falcifer, but dis-
tinguished most readily by characters given in the key. Ninth segment as in
I. falcifer, very narrow dorsally, the lateral portion broadest above the
mid-line, and tapering ventrad from that point. Posterior margin of the
eighth sternum rounded dorsad into a heavily sclerotized caudal plate, the
lateral corners of which are extended caudad into a pair of bidentate lobes.

21

1. excavatus

|. comans

Figs. 20-24—Female genitalia in Indomolannodes spp. 20. I. falcifer sp. n.
21. I. decurvatus sp. n. 22. I. excavatus sp. n. 23. I. comans sp. n.
24. I. incurvatus sp. n. a, lateral view; 6, ventral view.

De

Ventral branch of the clasper in the form of a short, stout, heavily sclero-
tized hook; dorsal branch of the clasper very much larger, in lateral view
comprising a wide basal area and a slender terminal process. Tenth segment
with dorsomedian plate triangular, the lateral margins concave in dorsal
view; lateral appendages somewhat mitten-like in shape as in I. falcifer,
but with the broadest part dorsad; mesal appendages each with a dorsal
process in the form of a clavate strip, and with a ventral elongate, slender
pointed process. Phallus much as in /. falcifer, phallotheca very broad,
endothecal sclerites larger and curved, bristles on the extensile appendage
pointed, ventral piece short and beak-like.

Female Genitalia (Fig. 23). Most closely resembling /. incurvatus, but
distinguished by characters given in the key. Posterior margin of the eighth
sternum slightly excavate in ventral view. Apex of the tenth segment
extended into an elongate median process, terminating in a rounded knob.

TYPES (all collected by F. Schmid). Holotype, male: INDIA, Assam,
United Jaintia and Khasi Hills: Umlangshor, 4100 ft., 18 April 1960, at
light. Paratypes: INDIA, Assam, United Jaintia and Khasi Hills: Jarain,
2800 ft., 13 April 1960, 2 ¢, at light; Mawpran, 3500 ft., 8 April 1960,
1 $; Mawpyut, 4000 ft., 14 April 1960, 1. ° ; Nongjni, 3750 ft., 19 April
1960, 3 6 1 9, at light; Thangrain, 3000 ft., 22 April 1960, 1 @, at light.

The name given to this species refers to the dense coating of long hairs
on the front legs of the male.

DISTRIBUTION. Known only from the Khasi Hills in Assam.

HABITAT AND BIOLOGY. Adults of this species were taken only in April, and
at elevations ranging from 2800 to 4100 feet, somewhat lower than some of
the other species. Adults were collected at lights located close to both large
and small rivers. At several localities this species was taken along with
Molanna paramoesta Wiggins.

Systematic Considerations

The genera Indomolannodes and Molannodes are certainly closely related.
Both show essentially the same type of venation. Basic similarity between
the male and female genitalic structures is also clear, and the males in
both genera possess the pair of curious pronotal pouches, as already
described. The close interrelationships between the two are in contrast to
those shown with Molanna, where the veins of the wings are still further
reduced from the primitive condition in a very specialized way, and the
wings are rolled around the body in a highly characteristic manner. The
genitalic structures are quite different, with the male genitalia in Molanna
showing considerable simplification in structural detail from the condition
in Indomolannodes and Molannodes. The pronotal pouches are absent in
Molanna. The evidence tends to support the interpretation that Jndomolan-
nodes and Molannodes represent a more primitive level of molannid evo-
lution than does Molanna. New evidence will be available when the
immature stages of Indomolannodes become known.

rf

If this interpretation, based entirely on morphological evidence, is
correct, it relates in an interesting way to another observation. The species
of Molanna are for the most part inhabitants of lakes and larger rivers,
but those of Molannodes and Indomolannodes are clearly confined to
cooler, lotic habitats. This invasion of more lentic, downstream-type
environments by groups derived from ancestors inhabiting the cooler,
upstream sections is a pattern of evolution typical for the Trichoptera
(Ross 1956) and for other aquatic insect groups, too (Corbet 1962).

Within the genus Indomolannodes three groups of species are clearly
evident. Comparison of male genitalia leaves little doubt that these groups
comprise /. falcifer and comans, I. decurvatus and incurvatus, and I. exca-
vatus. Analysis of the female genitalic structure lends no particular support
to these groupings, but neither does it militate strongly against them. It is
of interest to see that in J. excavatus, the one species of the genus in which
M, and Mz are separated in the fore wing of the male and thus the same
as Molannodes in this respect, the structure of the phallus also shows a
strong similarity to that of Molannodes.

Some generalization concerning the distribution of the species of Indo-
molannodes seems appropriate, especially in view of the extensive collecting
which was carried out in these areas by Dr. Schmid. From present evidence,
the only species living in the Himalayas is J. incurvatus, which extends
from Garhwal (west of Nepal) to the Kameng Frontier Division (northern
Assam, east of Bhutan). Its close relative 7. decurvatus is known from
Burma and Manipur (eastern Assam). Of the second group, J. falcifer is
known from Manipur and the Khasi Hills, and J. comans from the Khasi
Hills alone. The remaining species I. excavatus is known only from Mani-
pur. Four of the five known species of Indomolannodes, representing all
three species groups, are, then, evidently confined to Assam and western
Burma, strongly suggesting that the radiation of these species did occur
there.

Of the species of Molanna, evidence of the enormously high ecological
versatility of M. paramoesta is seen from the fact that its distribution
extends over much of the Indian subcontinent, where it occurs in a wide
variety of habitats at elevations ranging from 200 to 7400 feet. The
Siberian and Japanese counterpart of this species, M. moesta, is also very
widespread, and it must be concluded that these two species represent an
extremely viable and adaptable segment of the Molannidae. By contrast,
the two Indian species here recognized for the first time, Molanna saetigera
and crinita, are evidently so highly specialized for their montane environ-
ment that they have been unable to exploit the lower elevations as M. para-
moesta has done. And these two Indian species of Molanna show no close
relationship with others of the genus.

24

Key to the Holarctic Genera of the Molannidae

1. Dorsum of head with a single pair of lateral, elongate warts

(GE oraal) ues, Prete ie ome ts «tat. Beene eMac ciias whee 2
Dorsum of head with two pairs of small lateral warts (Fig.
1D) TIRE ee I rk eh ee se fen Mie Rites Us Ora sh desea ot ooo Molanna

2. Clasper of male comprised of a single piece of regular out-

line, somewhat triangular in lateral view; fore wing of
male with M, and Mg separate, as in the females... Molannodes

Clasper of male highly complex, comprised of a dorsal and

a ventral branch, the dorsal branch usually slender and

finger-like, the ventral branch usually hook-like (Figs.

13-19); fore wing of male with M, and Ms united
(usually), and thus distinct from the females (Fig. 16).... Indomolannodes

Acknowledgments

My appreciation is extended to Dr. Schmid for the opportunity of
studying this extremely interesting collection. Dr. H. E. Evans, Museum
of Comparative Zoology, Harvard University, kindly made it possible for
me to study the holotype of Molanna moesta Banks. Additional Asian
specimens of Molanna of assistance in clarifying the status of Molanna
falcata Ulmer were loaned by Dr. O. S. Flint of the U.S. National Museum.
All illustrations were prepared by Mr. Anker Odum, artist in the Depart-
ment of Entomology and Invertebrate Zoology, Royal Ontario Museum.

Literature Cited

BANKS, N.
1906 New Trichoptera from Japan. Proc. Ent. Soc. Wash., vol. 7,
nos. 2—3, pp. 106-113.
CORBET, P. S.
1962 A biology of dragonflies. H. F. & G. Witherby, London. 247 pp.
FISCHER, F.C. J.

1964 Trichopterorum catalogus. Vol. V. Phryganeidae; Limnocentro-
podidae; Molannidae. Nederlandsche Entomologische Vereeniging,
Amsterdam. 214 pp.

MARTYNOVY, A. V.

1910 Les Trichoptéres de la Sibérie et des régions adjacentes. II-e partie.
Ezheg. Zool. Muz., vol. 15, pp. 351-429.

1957 Trichoptera of the Amur region. Part I. Trudy Zool. Inst., Leningr.,
t: TI, 1933-35, pp: 205=395.

NIELSEN, A.

1957 A comparative study of the genital segments and their appendages

in male Trichoptera. Biol. Skr., vol. 8, no. 5, 159 pp.

25

ROSS, H. H.
1952

1956

TSUDA, M.
1942

ULMER, G.
1908

26

The caddisfly genus Molannodes in North America. Ent. News,
vol. 63, no. 4, pp. 85-87.

Evolution and classification of the mountain caddisflies. Univ.
Illinois Press, Urbana. 213 pp.

Zur Kenntnis der koreanischen Trichopteren. Mem. Coll. Sci. Kyoto
Univ., ser. B, vol. 17, no. 1, art. 5, pp. 227-237.

Japanische Trichopteren. Dt. Ent. Z., vol. 3, pp. 339-355.

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