Genera of the Trichoptera of Canada and Adjoining or Adjacent United States

THE INSECTS AND ARACHNIDS OF CANADA PART 7 Genera of the Trichoptera of Canada and Adjoining or Adjacent United States

F. Schmid Eastern Cereal and Oilseed Research Centre Ottawa, Ontario Research Branch Agriculture and Agri-Food Canada

NRC RESEARCH PRESS Ottawa 1998

NRC Monograph Publishing Program Editor: R.H. Haynes,

OC, FRSC

(York University)

Editorial Board: W.G.E. Caldwell, FRSC (University of Western Ontario); P.B. Cavers (University of Western Ontario); G. Herzberg, CC, FRS, FRSC (NRC, Steacie Institute for Molecular Sciences); K.U. Ingold, OC, FRS, FRSC (NRC, Steacie Institute for Molecular Sciences); W. Kaufmann (Editor-in-Chief Emeritus, Annual Reviews Inc., Palo Alto, CA); M. Lecours (Université Laval); W.H. Lewis (Washington University); L.P. Milligan, FRSC (University of Guelph); G.G.E. Scudder, FRSC (University of British Columbia); E.W. Taylor, FRS (University of Chicago); B.P. Dancik, Editor-in-Chief, NRC Research Press (University of Alberta) Inquiries: Monograph Publishing Program, NRC Research Press, National Research Council of Canada, Ottawa, Ontario K1A 0R6, Canada

Correct citation for this publication: Schmid, F. 1998. The Insects and Arachnids of Canada. Part 7. Genera of the Trichoptera of Canada and Adjoining or Adjacent United States. NRC Research Press, Ottawa, Ontario, Canada. 319 p.

The Insects and Arachnids of Canada Part 1. Collecting, Preparing and Preserving Insects, Mites and Spiders, compiled by J.E.H. Martin, Biosystematics Research Institute, Ottawa, 1977. Part 2. The Bark Beetles of Canada and Alaska (Coleoptera: Scolytidae), by D.E. Bright, Jr., Biosystematics Research Institute, Ottawa, 1976. Part 3. The Aradidae of Canada (Hemiptera: Aradidae), by R. Matsuda, Biosystematics Research Institute, Ottawa, 1977. Part 4. The Anthocoridae of Canada and Alaska (Heteroptera: Anthocoridae), by L.A. Kelton, Biosystematics Research Institute, Ottawa, 1978. Part 5. The Crab Spiders of Canada and Alaska (Araneae: Philodromidae and Thomisidae), by C.D. Dondale and J.H. Redner, Biosystematics Research Institute, Ottawa, 1978. Part 6. The Mosquitoes of Canada (Diptera: Culicidae), by D.M. Wood, P.T. Dang and R.A. Ellis, Biosystematics Research Institute, Ottawa, 1979.

© 1998 National Research Council of Canada All rights reserved. No part of this publication may be reproduced in a retrieval system, or transmitted by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the National Research Council of Canada, Ottawa, Canada K1A 0R6. Printed in Canada on acid-free paper ISBN 0-660-16402-7 NRC No. 41649

Canadian Cataloguing in Publication Data Schmid, F. Genera of the Trichoptera of Canada and Adjoining or Adjacent United States (The Insects and Arachnids of Canada; Part 7) Translation of: Généra des trichoptères du Canada et des Itats adjacents. Issued by the National Research Council of Canada. Includes glossary, bibliographical references, and index. ISBN 0-660-16402-7 1. Caddisflies — Canada — Classification. 2. Caddisflies — United States — Classification. I. National Research Council of Canada. II. Title. QL517.1S52 1998

595.7′45.0971

C98-980018-0

Table of Contents Acknowledgments . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . Purpose of Manual . . . . . . . . . . . . . . . . . . Characters of the Order . . . . . . . . . . . . . . . . Classification . . . . . . . . . . . . . . . . . . . . . List of Supraspecific Taxa of Nearctic Trichoptera . Family Rhyacophilidae Stephens . . . . . . . . . . . Family Glossosomatidae Wallengren . . . . . . . . Family Hydroptilidae Stephens . . . . . . . . . . . . Family Philopotamidae Stephens . . . . . . . . . . . Family Arctopsychidae Martynov . . . . . . . . . . Family Hydropsychidae Curtis . . . . . . . . . . . . Family Polycentropodidae Ulmer . . . . . . . . . . Family Hyalopsychidae Lestage . . . . . . . . . . . Family Psychomyiidae Curtis . . . . . . . . . . . . Family Limnephilidae Kolenati . . . . . . . . . . . Family Uenoidae Iwata . . . . . . . . . . . . . . . . Family Goeridae Ulmer . . . . . . . . . . . . . . . . Family Lepidostomatidae Ulmer . . . . . . . . . . . Family Phryganeidae Leach . . . . . . . . . . . . . Family Brachycentridae Ulmer . . . . . . . . . . . . Family Sericostomatidae Stephens . . . . . . . . . . Family Helicopsychidae Ulmer . . . . . . . . . . . Family Beraeidae Wallengren . . . . . . . . . . . . Family Leptoceridae Leach . . . . . . . . . . . . . . Family Odontoceridae Wallengren . . . . . . . . . . Family Calamoceratidae Ulmer . . . . . . . . . . . Family Molannidae Wallengren . . . . . . . . . . . List of Abbreviations Used in the Figures . . . . . . Figures 1 to 754 . . . . . . . . . . . . . . . . . . . . Glossary . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . Index . . . . . . . . . . . . . . . . . . . . . . . . . .

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Acknowledgments I am indebted to Dr. G.B. Wiggins for having supplied additional information on the ecology and biology of a large number of genera discussed in this work, as well as his complete checklist of the Nearctic species. I thank also J.C. Morse for the complete list of the supraspecific taxa given in the following pages. I am very grateful to J.E. O’Hara for improving the quality of the English translation of the text and for his assistance with the publication of this book. My thankfulness also to E.C. Becker for his endless patience in reading the final proofs.

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Introduction This book, translated from French, is an updated English edition of the Généra des Trichoptères du Canada et des États adjacents, originally published in 1980. Much progress in our knowledge has been made since then. The present edition has therefore been revised. With Canada’s extensive system of rivers and lakes, it hardly comes as a surprise that the order Trichoptera is well represented in this country. But, until recently, only very few comprehensive studies were available (Betten et al. 1934; Ross 1944, 1972). Among recent general studies one may cite: Morse (1975), Schmid (1981, 1982, 1983), Nimmo (1986, 1987), Weaver (1988), Armitage and Hamilton (1990), and Armitage (1991). This work does not provide information on the biology, ecology, collecting methods, morphology, preserving, and studying of specimens. These aspects have been discussed comprehensively by Betten et al. (1934), Ross (1944, 1946, 1947, 1950, 1956, 1965, 1967), and Wiggins (1996). The reader will find these works most informative. This book deals with the adult stage only. For the larval stage, one should refer to the remarkable Larvae of the North American Caddisfly Genera, 2nd edition, by Wiggins (1996). The classification of several taxa of Trichoptera is currently in a period of change and instability. In the following pages, I have adopted a traditionalist and conservative attitude and have not incorporated several recently-made changes in the two suborders and the limnephilids and some related families. I do not accept the existence of families based on larval characters only and not also on imaginal ones. These changes have been made for psychological convenience more than objective scientific reasons.

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Purpose of Manual My goal has been to provide a synopsis of all Canadian supraspecific taxa, i.e., suborders, families, subfamilies, genera, and subgenera. The geographic region under consideration extends beyond the boundaries of Canada and includes all the adjoining American States and Alaska. This work has been written for both practical and scientific purposes. It is my hope it will prove interesting as well as useful. It is intended not only for the Trichoptera specialist but also for the amateur. The layman will find clear, concise descriptions that are accompanied by a number of figures illustrating most of the characters mentioned and by complete dichotomous keys. Each genus is represented by one of its most common species, with illustrations of the venation and genitalia in both sexes. Descriptions do not apply to all known taxa but only to the species occurring in our area. The few taxonomic discussions that occur throughout the text have been kept to a minimum and can be easily disregarded. To permit identification down to the species level, wherever possible within a genus, names have been provided for all the species belonging to genera containing fewer than five species. In addition, a few distinguishing characters not requiring illustration have been included. Likewise, revisions are mentioned where appropriate. With the professional in mind, I have provided systematic descriptions. All the main characters of the taxa are given and illustrated as they define these taxa, even if they are not immediately useful for purposes of identification. Thus, occasional neoformations of the legs, palpi, and antennae have been illustrated. I have described and often figured the abdominal hemogill system and the internal gland of sternite V, wherever present. Also, more importantly, I have elucidated the morphology of the genital segments and established the identity and homologies of most of the appendages in all families and genera, and this with a degree of certainty that naturally may not be absolute but that I generally consider satisfactory (except in the case of the Hydroptilidae). For this purpose I relied on Nielsen’s excellent work on the subject (1957), though in exploring new ground I have occasionally found myself at variance with this author. The list of abbreviations used to identify the appendages in the figures is given on pages 203–204 along with explanations.

2

Characters of the Order The most useful taxonomic characters of the Trichoptera reside in the head and its appendages, the thoracic notal warts, and the legs, wings, and genitalia. The head has 2 bare or short-haired, compound eyes. The ocelli are present or absent. The vertex has a variable number of warts. The first or first 2 antennal segments display some degree of modification (Figs. 2, 42, 606, 608, 751, 762, 852). The maxillary palpi are the site of numerous variations and often of sexual dimorphism, and the number of segments can vary from 5 to 1 (Figs. 1, 102, 689, 690, 751, 753). Labial palpi are almost always three segmented. The legs bear a variable number of spines. The tibiae are equipped with spurs whose number is important. The front tibiae have 0 to 3 spurs and the middle or hind tibiae, 1 to 4. The numbers are expressed in formulas, such as 1,1,1 or 3,4,4, which indicate the number of spurs on each pair of legs—front, middle, and hind, respectively. The wings vary considerably in shape and are often the site of sexual dimorphism (Figs. 82, 126, 127, 845, 846). The venation supplies basic characteristics. Primitively, the venation is fairly simple, and any variations that do occur virtually always constitute simplifications. In the complete state (Figs. 41, 333), the veins of the front wings are as follows: a costal vein (C), followed by a subcostal vein (Sc), then the 1st radius (R1). The radial sector (RS) separates from the base of the 1st radius and forks once, thereby forming the discoidal cell (d), which may be open or closed by a crossvein. Each of the branches forks in turn, R2 and R3 forming apical Fork I, and R4 and R5, Fork II. The media (M) divides in the same way: first forming the median cell (m), which may be open or closed by a crossvein. A second division yields M1 and M2, which form Fork III, and M3 and M4, Fork IV. The 1st cubitus (Cu1) always originates from the base of the media, forming the thyridial cell (th). It then divides into Cu1a and Cu1b, thereby creating Fork V. The 2nd cubitus does not fork. There are 3 anal veins (A1, A2, and A3) which coalesce at a more or less basal level to form 2 anal cells. The venation of the hind wings is similar to that of the front wings in primitive families, but in specialized families it is often somewhat simplified. M3 and M4 very constantly merge into M3+4, with the result that 3

Fork IV is always absent. The number of anal veins can vary from 1 to 5, and these veins are usually divergent. Use of the apical wing forks is not very much in favor in the North American literature, whereas it is in the literature from other continents. I have made use of apical forks in this text. It is so convenient that I have found it an indispensable tool. The number of forks can be grouped into formulas like the spurs of the legs, and these formulas, e.g., I, II, III, IV, V or I, V, immediately and with little written explanation, express the degree of specialization of the vein system of a given wing. These formulas have been indicated on all the figures of venation. The abdominal hemogill system has been reported only in the Arctopsychidae, Hydropsychidae, and Thamastes dipneumus Schmid. I examined most of the genera for this system and found it in the following families: Arctopsychidae, Hydropsychidae, Polycentropodidae, Hyalopsychidae, Limnephilidae, Goeridae, Lepidostomatidae, Phryganeidae, Brachycentridae, Leptoceridae, Helicopsychidae, Calamoceratidae, and Molannidae. It is particularly well developed in the Arctopsychidae, Hydropsychidae, Phryganeidae, and some Leptoceridae. The system consists of clusters of simple, rarely forked, tubes on the pleurites of virtually all the abdominal segments. It is usually completely retracted behind a fold of the teguments and, more often than not, is inconspicuous. A simple technique, which I have described elsewhere (Schmid 1968b, p. 13), induces turgescence of the system. These hemogill tubes are probably modifications of the pupal gills. The conditions under which they operate are not known, and thus far they have not been found to be of any taxonomic use (Figs. 125, 186, 212, 699, 782, 826, 859, 862). Hitherto, the internal gland of abdominal sternite V had been identified only in the Rhyacophilidae, Agapetus, and the Arctopsychidae. I have also studied this structure and found it in all of the families, with the exception of the Philopotamidae, Goeridae, Lepidostomatidae, Leptoceridae, Odontoceridae, Calamoceratidae, and Sericostomatidae. It undoubtedly plays a role in pheromone production, but to date has not proved to be of any taxonomic value. It is always more developed in the % than in the & and empties into a slight concavity on the sternite or at the base of a lobe or on a filament of variable length (Figs. 35, 46, 146, 147, 164, 165, 178, 189, 439, 518, 519, 691, 862). The broad structural lines of the genitalia provide excellent characters for classification of the genera, while the details of the shapes provide essential characters for diagnosis of the species. A specimen that has been mutilated and deprived of its genitalia usually does not lend itself to determination, except if it belongs to a species with characteristic coloring, which is rarely the case. 4

All of the figures in this work are based on abdomens previously treated with KOH and studied in beech creosote. There is some slackening of the appendages, which should be taken into account when comparing the prepared genitalia with dry or alcohol-preserved specimens. Similarly, all erectile organs are shown in a state of complete turgescence in order to reveal their actual structure (Figs. 5, 57, 196, 329, 685, 875). In most collection specimens, these organs are retracted or invaginated and cannot be seen, and this should be taken into account in the course of identification (compare Figs. 119 and 120). Variations in the male genitalia are so great that it is extremely difficult to provide an overall description. Generally, segment IX is the 1st to show sexual changes and it is almost always in a single piece, i.e., not divided into tergite and sternite. Ventrally, it bears 1- or 2-segmented inferior appendages. Segment X is more or less prominent and bears the preanal, superior, and intermediate appendages and the anal opening. The armature of segment X differs in the Limnephilidae and is occasionally highly complex. Male Trichoptera do not have any cerci. The phallic apparatus is contained within the phallocrypt that opens under segment X. The apparatus consists of the phallotheca, the endotheca, the aedeagus, and 2 parameres. The first two parts are often developed to the detriment of the last three, which are often lost. In the &, there are such substantial variations in the genitalia, that I shall examine them when describing the suborders. A detailed description of the groundplan of Trichoptera is now available (Schmid 1989, p. 18–21). It has been shown (Schmid 1989, p. 110) that most of the primitive, monticole, rheophilic, cold-stenothermic lineages of Nearctic Trichoptera originated in the Oriental region. The most primitive species of each of these lineages and the largest number of their species are actually localized there. They secondarily spread to the Nearctic region at various periods of the Cenozoic that cannot be precisely determined. In the cases of the more specialized, more widely spread low-altitude lentic and eurythermal lineages, the present state of our knowledge does not permit any hypothesis on their origin. The Nearctic region does not seem to have been the area of origin of any larger lineages of caddisflies. Its fauna seems to be of a “colonial" nature and mainly the result of local differentiation of foreign-originated elements.

5

Classification The order Trichoptera consists of two suborders: the Annulipalpia and the Integripalpia, each composed of superfamilies. Wiggins (1996, p. 9–14) did not use these suborders, but restricted himself to the superfamilies. More recently, Weaver and Morse (1986, p. 153) introduced the concept of infra-order, between the suborders and the superfamilies. I do not adopt this category here, finding it of little use and causing unnecessary taxonomic inflation. Descriptions of the groundplan of each suborder are now available (Schmid 1989, p. 21–22). In the adults, the characters of the suborders reside in the palpi, venation, and genitalia of both sexes. Venation and the male genital system have undergone such extensive differentiation within each suborder that there are few clear-cut and constant distinctive characters. The &, on the other hand, is far more conservative than the % and has two totally different and distinctive types of genitalia. They remain remarkably constant in the Annulipalpia, with the characters becoming only slightly more pronounced within the family series (Schmid 1970, p. 32).

Suborder Annulipalpia Martynov Annulipalpia Martynov, 1924, p. 18 Larvae campodeiform, collectors-gatherers. They are free-living or construct turtle shell-like or silky purse-like cases (Rhyacophiloidea), or, generally, fixed retreats, which are always much larger than the larvae themselves, allowing them to move freely in their retreats (Hydropsychoidea) by means of elongated, movable anal legs. In the adults, maxillary palpi consistently 5-segmented in both sexes. Segment V either entire (Rhyacophiloidea) or annulate and flagellate (Hydropsychoidea). In the front wings, discoidal cell usually short and the median cell closed. Apical fork I rarely sessile, usually petiolate or absent. & median legs sometimes flattened. In the %, segment IX either well developed or very reduced dorsally. Segment X often membranous. Phallic apparatus rarely including the aedeagus and the parameres. The apparatus usually consists of only the phallotheca and endotheca, and the latter may be more or less enlarged and equipped with spines and processes. 6

In the &, the abdomen comprises 11 segments, all of which are clearly visible in some of the Rhyacophilidae and Glossosomatidae (Fig. 16; Schmid 1989, Fig. 32, p. 21; Schmid 1970, plates XLIX, LII). Generally, segment IX absent or virtually so (Hydropsychoidea). This occasionally also applies to segment VIII (Hydroptilidae). Segment XI consisting of 2 contiguous membranous lobes, consistently equipped with 1- or 2-jointed cerci. There is a single anovaginal opening at the apex of segment X, under segment XI. In the Hydropsychoidea (but not in the Rhyacophiloidea), segment X forms a vulvar scale closing the anovaginal cavity underneath. Vaginal apparatus located at the end of a vestibule, which occasionally is eversible (Rhyacophiloidea). In the case of the Hydropsychoidea, the apparatus is formed by the vestibule walls. The Annulipalpia comprise two superfamilies: the Rhyacophiloidea and Hydropsychoidea, which group the most primitive families of the order along with families exhibiting gradual and greater, even occasionally extreme, specialization. Nine families are represented in Canada.

Suborder Integripalpia Martynov Integripalpia Martynov, 1924, p. 18 Larvae eruciform or suberuciform, shredders-detritivores, constantly living in tight-fitting portable cases to which they are attached by short, anchoring, slightly movable anal legs. In the adults, maxillary palpi consistently 5-segmented in the &. In the %, there may be 5 to 1 segments, and the last one is never flagellate. In the front wings, discoidal cell generally long, median cell open, and apical fork I usually sessile. In the %, segment IX generally shortened dorsally. Segment X often roof-shaped and conspicuous. Phallic apparatus generally including the aedeagus. Parameres occasionally present. In the &, there are 10 abdominal segments, all of which are consistently well developed. It is therefore not possible to determine whether the last one is segment X or segment XI, as in the case of the Annulipalpia. I consider it to be segment X. In some families, this segment bears a pair of appendages inserted at the base of segment X. They are clearly not homologous with the cerci in the Annulipalpia. Unlike the Annulipalpia, which have only one anovaginal opening, the Integripalpia have two: an anal opening at the apex of segment X and a vaginal opening generally between segments VIII and IX or on segment IX. There is often a supragenital plate above the vaginal opening, which is also protected underneath by the generally trilobate vulvar scale. The vaginal apparatus displays varying degrees of complexity and is located in the middle of the vaginal chamber. 7

The Integripalpia comprise four superfamilies: the Limnephiloidea, Phryganoidea, Leptoceroidea, and Sericostomatoidea, which include 13 native Canadian families. None of them is truly very primitive, and some are amongst the most specialized of the order.

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List of the Supraspecific Taxa of Nearctic Trichoptera ANNULIPALPIA Rhyacophiloidea Rhyacophilidae

Rhyacophila Pictet Himalopsyche Banks

Glossosomatidae Glossosomatinae

Hydroptilidae

Glossosoma Curtis Anagapetus Ross Anseriglossa Ross Ripaeglossa Ross Synafophora Martynov

Agapetinae

Agapetus Curtis

Protoptilinae

Culoptila Mosely Matrioptila Ross* Protoptila Banks

Ptilocolepinae

Palaeagapetus Ulmer

Hydroptilinae Hydroptilini

Agraylea Curtis Hydroptila Dalman Oxyethira Eaton Paucicalcaria Mathis and Bowles*

Leucotrichiini

Alisotrichia Flint* Anchitrichia Flint* Leucotrichia Mosely Zumatrichia Mosely*

Neotrichiini

Mayatrichia Mosely Neotrichia Morton

Ochrotrichiini

Ochrotrichia Mosely Metrichia Ross*

9

Orthotrichiini

Ithytrichia Eaton Orthotrichia Eaton

Stactobiini

Stactobiella Martynov

Tribe incertae sedis Dibusa Ross* Superfamily incertae sedis Hydrobiosidae Hydropsychoidea Philopotamidae

Atopsyche Banks* Philopotaminae

Dolophilodes Ulmer Fumonta Ross* Sisko Ross* Wormaldia McLachlan Doloclanes Banks*

Chimarrinae

Chimarra Stephens

Arctopsychidae Hydropsychidae

Polycentropodidae

10

Arctopsyche McLachlan Parapsyche Betten Oestropsinae

Centromacronema Ulmer* Leptonema Guérin-Méneville* Macronema Pictet* Macrostemum Kolenati

Hydropsychinae

Cheumatopsyche Wallengren Hydropsyche Pictet Mexipsyche Ross & Unzicker* Plectropsyche Ross* Potamyia Banks Smicridea McLachlan* Rhyacophylax Müller*

Diplectroninae

Diplectrona Westwood Homoplectra Ross Oropsyche Ross* Cernotina Ross Cyrnellus Banks Neureclipsis McLachlan Paranyctiophylax Tsuda Polycentropus Curtis Polyplectropus Ulmer*

Hyalopsychidae

Phylocentropus Banks

Ecnomidae

Austrotinodes Schmid*

Psychomyiidae

Psychomyiinae

Lype McLachlan Psychomyia Latreille Tinodes Curtis

Paduniellinae

Paduniella Ulmer*

INTEGRIPALPIA Limnephiloidea Limnephilidae

Dicosmoecinae

Allocosmoecus Banks Allomyia Banks Amphicosmoecus Schmid Cryptochia Ross Dicosmoecus McLachlan Ecclisocosmoecus Schmid Ecclisomyia Banks Eocosmoecus Wiggins & Richardson Ironoquia Banks Manophylax Wiggins Moselyana Denning* Onocosmoecus Banks Pedomoecus Ross Rossiana Denning

Apataniinae

Apatania Kolenati

Pseudostenophylacinae Pseudostenophylax Martynov Limnephilinae Limnephilini

Anabolia Stephens Arctopora Thomson Asynarchus McLachlan Clistoronia Banks Clistoronia s. str. Banks Clistoroniella Schmid Grammotaulius Kolenati Halesochila Banks Hesperophylax Banks Lenarchus Martynov Paralenarchus Schmid 11

Lenarchus s. str. Martynov Prolenarchus Schmid Leptophylax Banks Limnephilus Leach Nemotaulius Banks Macrotaulius Schmid Philarctus McLachlan Platycentropus Banks Psychoronia Banks* Stenophylacini

Chyranda Ross Clostoeca Banks Hydatophylax Wallengren Philocasca Ross Sphagnophylax Wiggins & Winchester Pycnopsyche Banks

Chilostigmini

Chilostigma McLachlan Chilostigmodes Martynov Desmona Denning Frenesia Betten & Mosely Glyphopsyche Banks Grensia Ross Homophylax Banks Phanocelia Banks Psychoglypha Ross

Uenoidae

Goeridae

Farula Milne Neophylax McLachlan Neothremma Dodds & Hisaw Oligophlebodes Ulmer Sericostriata Wiggins, Weaver & Unzicker Goerinae

Goera Stephens Goeracea Denning Goerita Ross*

Lepaniinae

Goereilla Denning Lepania Ross

Lepidostomatidae Lepidostomatinae

Theliopsychinae 12

Lepidostoma Rambur Lepidostoma s. str. Rambur Mormomyia Banks Neodinarthrum Weaver Nosopus McLachlan Theliopsyche Banks

Brachycentridae

Phryganoidea Phryganeidae

Adicrophleps Flint* Amiocentrus Ross Brachycentrus Curtis Brachycentrus s. str. Curtis Oligoplectrodes Martynov Sphinctogaster Provancher Sychnothrix Flint* Eobrachycentrus Wiggins Micrasema McLachlan Yphriinae

Yphria Milne*

Phryganeinae

Agrypnia Curtis Banksiola Martynov Beothukus Wiggins & Larson Fabria Milne Hagenella Martynov Oligostomis Kolenati Oligotricha Rambur Phryganea Linnaeus Ptilostomis Kolenati Neuronella Banks*

Sericostomatoidea Sericostomatidae

Agarodes Banks Agarodes s. str. Banks Psiloneura Banks Gumaga Tsuda*

Helicopsychidae

Cochliopsyche Müller* Helicopsyche von Siebold

Beraeidae

Beraea Stephens

Leptoceroidea Leptoceridae

Leptocerinae Athripsodini

Ceraclea Stephens Ceraclea s. str. Stephens Athripsodina Kimmins

Leptocerini

Leptocerus Leach

Mystacidini

Mystacides Berthold

Nectopsychini Oecetini Setodini

Nectopsyche Müller Oecetis McLachlan Setodes Rambur

13

Triaenodini

Odontoceridae

Calamoceratidae

Molannidae

Triaenodes McLachlan Ylodes Milne

Triplectidinae

Triplectides Kolenati*

Odontocerinae

Marilia Müller Namamyia Banks* Nerophilus Banks* Parthina Denning* Psilotreta Banks

Pseudogoerinae

Pseudogoera Carpenter* Anisocentropus McLachlan* Heteroplectron McLachlan Phylloicus Müller* Molanna Curtis Molannodes McLachlan

Genera marked with an asterisk (*) are not included in this manual, since they are not represented in our area.

Key to the Families 1a Very small (1–5 mm) and very hairy species with both pairs of wings in shape of very narrow, tapered lamellae with very long fringes, especially in hind wings (Figs. 58, 63). Flat, raised, triangular mesoscutellum. Front tibia never with more than single spur. Lateral ocelli close to eye margin or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydroptilidae, in part, p. 30 1b Species without these characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Ocelli present (Figs. 1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b Ocelli absent (Fig. 573) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 3a M of hind wings simple (Fig. 535), or if 2-branched (Fig. 544) then each branch distinct from base, or discoidal cell open at base, RS double beginning at base of wing (Figs. 537, 544) . . . . . . . . . . . . . . . Uenoidae, p. 134 3b M of hind wings 2-branched (Fig. 601) or 3-branched (Fig. 594) . . . . . . . . . 4 4a A of front wings simple (Fig. 601) or composed of 3 veins reaching edge of wing independently (Fig. 594) . . . . . . . . . . . . . . . Goeridae, in part, p. 142 4b A of front wings with 3 branches, confluent and forming 2 or 3 closed cells (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

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5a Maxillary palpi 3-segmented (Fig. 253). . . . . . . . . . . Limnephilidae %, p. 74 5b Maxillary palpi 4-segmented (Fig. 689). . . . . . . . . . . Phryganeidae %, p. 156 5c Maxillary palpi 5-segmented (Fig. 254). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a Maxillary palpi with 5th segment flexible, about twice length of 4th (Fig. 102) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Philopotamidae, p. 42 6b Maxillary palpi with 5th segment not flexible, barely longer than 4th. . . . . 7 7a Maxillary palpi with first 2 segments very short and equal in length and 5th segment ending in a point (Fig. 1). Discoidal cell in both pairs of wings open (Fig. 3). Spurs: 3,4,4 . . . . . . . . . . . . . . . . . . . . . Rhyacophilidae, p. 18 7b Maxillary palpi with first 2 segments subglobular and equal in length and 5th segment not ending in a point (Fig. 20). Discoidal cell in front wings closed (Fig. 22). Spurs: 2,2,4 or 0,4,4 or 2,4,4 . . . . . . . . . . . . . . . . . . . . . 8 8a Posterior cephalic warts large, arcuate, the two nearly meeting on meson. Pronotum with a mesal pair of warts nearly touching (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydroptilidae, in part, p. 30 8b Posterior cephalic warts oval or round and widely separated on meson. Pronotum with a mesal pair of warts well separated (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glossosomatidae, p. 21 8c Maxillary palpi with 2nd segment clearly longer than 1st (Fig. 254). Discoidal cell on both wings closed (Fig. 333) . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9a Front tibia with single spur (Fig. 332) . . . . . . . . . . . . Limnephilidae &, p. 74 9b Front tibia with 2 or more spurs . . . . . . . . . . . . . . . . . Phryganeidae &, p. 156 10a Fifth segment of maxillary palpi flagellate, long, flexible, ringed, and generally much longer than 4th (Figs. 137, 185) . . . . . . . . . . . . . . . . . . . . . . . 11 10b Fifth segment of maxillary palpi not flagellate and not much longer than 4th (Figs. 771, 861) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 11a Long discoidal cell in front wings. Median cell open or closed and beginning apical to proximal tip of discoidal cell (Fig. 187, 193). Third segment of maxillary palpi inserted before apex of 2nd segment which is spinose (Figs. 185, 199, 205) . . . . . . . . . . . . . . . . . . . . . . . Polycentropodidae, p. 60 11b Short discoidal cell in front wings. Median cell closed and beginning basal to proximal tip of discoidal cell (Figs. 213, 224). Third segment of maxillary palpi inserted at apex of 2nd segment (Fig. 230) . . . . . . . . . . . . . . . . . . . 12 12a Spurs: 3,4,4. Antennae of % longer than front wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalopsychidae, p. 67 12b Spurs: 2,4,4. Antennae of % shorter than front wings . . . . . . . . . . . . . . . . . 13 13a Thyridial cell on front wings very small, located at base of the wing, without contact with median cell. Hind wings narrower than front wings, FI absent, discoidal cell open and anal area reduced (Fig. 220, 224) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psychomyiidae, p. 69

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13b Thyridal cell in front wings large, medianly positioned and in contact with thyridial cell. Hind wings equal in width to front wings or wider; FI almost always present; discoidal cell closed and anal area well developed (Figs. 127, 148) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14a Antennae thick (Fig. 123). Both pairs of wings similar in shape and width. Postcostal cell in front wings short and wide (Fig. 127) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctopsychidae, p. 47 14b Antennae thin (Fig. 163). Hind wings wider than front wings and differently shaped. Postcostal cell in front wings long, usually narrow (Fig. 148). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydropsychidae, p. 51 15a Discoidal and thyridial cells of front wings absent (Figs. 772, 863) . . . . . 16 15b Discoidal and thyridial cells of front wings present (Fig. 793) . . . . . . . . . 17 16a Middle tibia with 4 small spurs and large number of black spines. Mediumsized brown species . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molannidae, p. 200 16b Middle tibia with 2 large spurs and few black spines. Small black species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beraeidae, p. 179 17a Middle tibia without preapical spurs but with black spines . . . . . . . . . . . . 18 17b Middle tibia with preapical spurs and with or without black spines (Fig. 719) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 18a Very thin antennae, longer than front wings. Very long, slender maxillary palpi (Fig. 781) clothed with dense, erect hairs. Pronotum partially concealed under mesonotum . . . . . . . . . . . . . . . . . . . . . . . Leptoceridae, p. 181 18b Antennae and maxillary palpi thicker, antennae shorter than front wings (Fig. 753). Pronotum clearly visible from above . . . . . . . . . . . . . . . . . . 19 19a Hind wings with basal half of costal margin equipped with hooks and forming a broad costal angle (Fig. 765) . . . . . . . . . . . . . . . Helicopsychidae, p. 177 19b Hind wings without hooks or costal angle . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20a Front wings with crossvein between R1 and R2. FI merges with discoidal cell for long distance and Cu2 ends on Cu1b (Fig. 755). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sericostomatidae, p. 174 20b Front wings without crossvein between R1 and R2+3. FI merges with discoidal cell for a short distance and Cu2 and Cu1b joined by crossvein (Figs. 734, 735) . . . . . . . . . . . . . . . . . . . . . Brachycentridae, in part, p. 168 21a Median cell of front wings closed (Fig. 853). Antennae much longer than front wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calamoceratidae, p. 198 21b Median cell of front wings open or absent (Fig. 834). Antennae slightly or much longer than front wings . . . . . . . . . . . . . . . . . . Odontoceridae, p. 194 21c Median cell of front wings open or absent. Antennae shorter than front wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

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22a Thyridial or subthyridial cell of front wings broadened at apex (Figs. 576, 586). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goeridae, in part, p. 142 22b Thyridial or subthyridial cell of front wings not broadened at apex . . . . . . 23 23a Spurs: 2,4,4 and hairy. Middle tibia without black spines (Fig. 609) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidostomatidae, p. 148 23b Spurs: 2,3,3 or 2,4,4 and not hairy. Middle tibia with black spines (Fig. 719) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycentridae, in part, p. 168

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Family Rhyacophilidae Stephens Rhyacophilidae Stephens, 1836, p. 148 Type genus: Rhyacophila Pictet Ocelli present (Fig. 2). Maxillary palpi 5-segmented in both sexes. First 2 segments very short, 2nd subglobular. Last segment of maxillary and labial palpi ending in a small point (Fig. 1). Spurs: 3,4,4 (Fig. 4). Middle legs of & not flattened. Wings smooth ovals. Both pairs in both sexes similarly shaped. Venation complete (Fig. 3), with all forks present, I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. R1 in front wings divided into R1a and R1b at tip. In both pairs of wings, discoidal and median cells open. Thyridial cell in front wings very long. Internal gland of abdominal sternite V small. Abdominal hemogill system absent. % genitalia (Fig. 5): segment IX ring-shaped, occasionally forming an apicodorsal lobe overhanging other genital parts. Segment X variously shaped, with occasionally a highly complex structure. Preanal appendages occasionally present. The intermediate appendages occurring in the other families are referred to here as anal sclerites. They are virtually always small and may be present or absent, paired or unpaired. Inferior appendages very large, subhorizontal and 2-segmented, with 2nd segment virtually always indented in a manner characteristic of the species. Phallic apparatus situated between inferior appendages and joined to base of latter by means of a sclerotized connection. This apparatus primitive and complete in structure. It comprises a phallotheca, endotheca, aedeagus, and generally 2 more or less spiniform parameres. Occasionally there is a ventral lobe below and a dorsal lobe above aedeagus. & genitalia (Figs. 6 and 7): segment VIII in shape of a ring or truncated cone, often with apical notches separating lobes that, in shape, are characteristic of species. Segment IX present or not distinguishable from intersegmental membrane VIII–X. The last segments and their connecting membranes generally stretch considerably into a long, flexible, retractile ovipositor containing 2 pairs of sclerotized apodemal rods serving as sites of insertion for the muscles retracting the ovipositor. Single-jointed cerci. Intersegmental membrane VIII–X often invaginated for a variable distance within segment VIII. Vulvar scale absent and anovaginal opening at tip of segment X. Vaginal apparatus located at the end of a very long vaginal 18

vestibule with such fine membranous walls that they are virtually invisible. The apparatus consequently appears to be free inside abdomen. During copulation, the last 3 segments retract, the vaginal vestibule turns inside out like the finger of a glove and the vaginal apparatus protrudes from the abdomen. The structure of this apparatus provides an excellent source of species-specific characters. Rhyacophilidae contain only two genera, Rhyacophila Pictet and Himalopsyche Banks, the 2nd one being a specialized offshoot of the former. 1a Mesoscutellum without long hairs. Length of anterior wing under 20 mm.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhyacophila, p. 19 1b Mesoscutellum with a tuft of long thin hairs. Length of anterior wing above 20 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Himalopsyche, p. 20

Genus Rhyacophila Pictet

Rhyacophila Pictet, 1834, p. 181 Type species designated by Ross, 1944: Rhyacophila vulgaris Pictet Revisions: Schmid, 1970, 1981; Nimmo, 1971 As defined here, Rhyacophila has all the characters mentioned in the family description. I shall not add any here and simply figure the head, the anterior leg, the venation, and the genitalia of both sexes for 2 species (Figs. 1–7). The genus Rhyacophila is large, flourishing, and Holarctic and Oriental in distribution. Some 500 species have been described to date and more than 50 of these live in Canada and the bordering American states. The genus is composed of four subgenera, all of which are represented in Canada, but the boundaries between these subgenera are so fluid that it is impossible to supply a clear, simple diagnosis, much less provide a dichotomous key. It is therefore preferable not to name these subgenera but to consider them “branches”. The latter divide in turn into “twigs” and a large number of groups of species (Schmid, 1970, Fig. 8). Larvae of Rhyacophila live in all, even temporary, types of running water, have a restricted range, and inhabit all mountainous regions at every altitude. They are not found, however, in the Arctic. This genus originated in the Oriental region, where the greatest number of species and the most primitive species occur (Schmid 1989, p. 110). One of the outstanding features of the Rhyacophila is the beautiful shapes of the % genitalia. The large size and complexity of the genital system facilitate determination. For purposes of identification, it is advisable 19

to remove the left inferior appendages so that the internal parts normally concealed by this appendage can be examined. In the case of the &, the abdomen should be treated with KOH to release the vaginal apparatus. Genus Himalopsyche Banks

Himalopsyche Banks, 1940, p. 197 Type species by original designation: Rhyacophila tibetana Martynov Revision: Schmid, 1966 Vertex very convex. Ocelli large. Mesoscutellum with a tuft of long, thin hairs. Anterior wings strongly spotted. Venation similar to that of Rhyacophila. % genitalia (Figs. 8, 9): segment IX short, regularly annular, and without apicodorsal lobe. Xth segment absent and replaced by the very large, complex preanal appendages. Anal sclerite without cleft and broadly 2-branched. Second article of inferior appendages barely incised. Phallic apparatus small and blunt: aedeagus bifid and parameres platelike. & genitalia (Fig. 10): segment VIII very massive, with a shallow lateral concavity. Last segments barely extended in a thick ovipositor. The genus Himalopsyche is closely related to Rhyacophila, of which it is merely a specialized off-shoot. It is easily characterized by a small number of features, as given above. Only one species, phryganea Ross, is known from the Nearctic region, recorded from California to Washington. The species is striking in its strongly spotted anterior wings and its large size; length of anterior wings: 21–23 mm. This genus originated in the Oriental region. This is where all species occur, with the exception of phryganea (Schmid 1989, p. 110).

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Family Glossosomatidae Wallengren

Glossosomatidae Wallengren, 1891, p. 163 Type genus: Glossosoma Curtis Ocelli present. Maxillary palpi 5-jointed in both sexes. First 2 are very short and 2nd particularly globular. Last segments of maxillary and labial palpi without pointed tips (Fig. 20). Cephalic and prothoracic warts well separated (Fig. 12). Abdominal hemogill system absent. Internal gland of sternite V variously developed. There are such major differences among the three subfamilies with respect to the wings and venation that it is impossible to provide an overall description. The only two characters that are consistently present are of little significance and are virtually absent in the Protoptilinae. These characters are: in front wings, bifurcations R1-RS and M-Cu1 are ogival, symmetrical, and located very close to base of wing, and veins RS, R4+5, and R5 are aligned in a straight line (Figs. 22, 32). % and & genitalia. A description of the genitalia at the family level is also not possible since the degree of specialization at the subfamily, genus, and even subgenus levels is so great. Mention can be made, however, of the fact that the intermediate appendages in the % have disappeared, the inferior appendages are single-jointed, and the phallic apparatus is positioned fairly high in the abdomen. In the &, the genitalia are closely related to those of the Rhyacophilidae. The cerci are always present, the anovaginal opening occurs at the tip of segment X, and the vulvar scale is absent. The Glossosomatidae thus constitute a highly diverse family, but the various taxa can be easily derived from one another phylogenetically. Furthermore, the larval and pupal stages are far more uniform and leave no doubt as to the unity of the family. The family is divided into three subfamilies: the Glossosomatinae, Agapetinae, and Protoptilinae. In the first two subfamilies, the considerable basic differences in the hind wings and genitalia of the % suggest that differentiation occurred at a very early stage. The Protoptilinae are the most highly specialized and the most markedly different of the three subfamilies. There is no justifiable reason for maintaining the tribes designated by Ross (1956) as Glossosomatini and Anagapetini. 21

Larvae of the Glossosomatidae inhabit running water. 1a Spurs: 0,4,4 or 0,3,3. Front and hind wings long and very narrow, venation reduced and faint (Fig. 37) . . . . . . . . . . . . . . . . . . . . . . Protoptilinae, p. 27 1b Spurs: 2,4,4. Front and hind wings fairly evenly elliptical, venation complete or almost complete (Figs. 32, 41). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Hind wings: closed discoidal cell and long R1 ending at wing margin (Figs. 22, 41) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glossosomatinae, p. 22 2b Hind wings: open discoidal cell and R1 short, ending at R2+3 (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agapetinae, p. 26

Subfamily Glossosomatinae Wallengren This subfamily comprises only the type genus. Genus Glossosoma Curtis

Glossosoma Curtis, 1834, p. 216 Monobasic type species: Glossosoma boltoni Curtis Revisions: Ross, 1956; Nimmo, 1974; Schmid, 1982 Spurs: 2,4,4. Middle legs of & usually flattened and ciliate (cf. Fig. 124). Wings regularly oval-shaped, both pairs equal in width. Venation complete (Figs. 22, 41) with all forks present: I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. On both pairs, discoidal cell closed and median cell open. In front wings, R1 divided into R1a and R1b. Symmetrical R1-RS and M-Cu1 forks close to and equidistant from base. RS, R4+5, and R5 follow one another in roughly the same alignment. Small, subtriangular discoidal cell, slanting slightly upward, since FII shares a much longer boundary with the cell than FI. Anal veins of the % often forming a callosity. In hind wings, R1 present and long and parallel with R2. Sternites VI and VII provided with ventral lobes and plates more highly developed in % than in &. % genitalia (Figs. 24–31): highly differentiated in various subgenera, making it difficult to provide a generic description. Segment X reduced to a small membranous lobe, usually completely concealed by large, preanal appendages, which lie against it. Intermediate appendages absent. Inferior appendages large and single-jointed or absent. Well-developed phallic apparatus complete but occasionally simplified by loss of parameres and situated in middle of segment IX, above inferior appendages. & genitalia (Figs. 28, 29): very homogeneous, closely related to those of Rhyacophila. Similarly, they are stretched into a long ovipositor and 22

equipped with 2 pairs of long apodemal rods. Segment VIII developed into a regular tube, which occasionally is complicated by proximal and apical notches. Segment IX generally conspicuous and distinctly sclerotized. Segment X very elongate in relative terms and highly sclerotized dorsally and laterally. Segment XI ovoid with 2 tiny cerci. Vaginal apparatus located at end of a long membranous vaginal vestibule, as in the case of the genus Rhyacophila, and is evaginated during mating. The genus Glossosoma is fairly heterogeneous and has been divided into 10 subgenera (Ross, 1956), only 6 of which are valid. Of this number, three are represented in Canada. To them I add Anagapetus, which Ross isolated in a special tribe, which in fact is merely a subgenus of Glossosoma. These subgenera are based solely on primary and secondary sexual characters, and this makes && unclassifiable other than by association with the respective %%. Only members of the subgenus Anagapetus can be recognized by their black color and nonenlarged middle legs. The genus Glossosoma is medium-sized with a Holarctic and Oriental distribution. It is of Oriental origin, with its most primitive subgenera, Lipoglossa Martynov and Muroglossa Ross, being located there (Schmid 1989, p. 110). A dozen species reside in Canada in cold running water. 1a Inferior appendages of % very large, strongly curved downward, and cleft at the end (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagapetus, p. 23 1b Apicolateral margin of segment IX extending strongly backward, forming a capsule containing virtually all the other genital parts (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ripaeglossa, p. 24 1c Aedeagus small with vertical parameres completely concealed in genital cavity and a tendon connecting phallotheca to preanal appendages (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synafophora, p. 24 1d Tendons joining phallotheca to preanal appendages developed into two large appendages framing the aedeagus. Inferior appendages absent (Figs. 30, 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anseriglossa, p. 25

Subgenus Anagapetus Ross

Anagapetus Ross, 1938, p. 109 Monobasic type species: Anagapetus debilis Ross Anagapetini Ross, 1956, p. 127 Middle legs of & not flattened. Anal cells on front wings of % without callosity (Fig. 41). Sternite VII of % with large ventral lobe. % genitalia (Fig. 24): segment IX ring-shaped. Simple triangular preanal appendages. Inferior appendages very large, strongly curved downward, and with apical cleft separating 2 narrow, strongly spined branches. They are connected to segment X by upper basal lobe. Phallic complex 23

short and stout, greatly reduced, and located very high above inferior appendages. Anagapetus was isolated by Ross into a special tribe on the basis of a suture separating the upper and lower parts of the mesepisternum. This character is not constant and is insignificant. The venation and genitalia of the % indicate that Anagapetus is simply a subgenus of Glossosoma, related to but less specialized than the Asian subgenus Lipoglossa. Anagapetus is a small eastern Palearctic and western Nearctic subgenus. It is represented in Canada by three species only, bernea Ross, hoodi Ross, and debilis Banks. %% are quite distinct by the shape of the preanal and inferior appendages. && can be identified by the vaginal apparatus only. Larvae of all three species live in cooler headwater sections of mountain streams of the western part of the continent. Subgenus Ripaeglossa Ross

Ripaeglossa Ross, 1956, p. 152 Type species by original designation: Glossosoma parvulum Banks Middle legs of & flattened. Anal cells in front wings of % enlarged with a round callosity. Sternite VI of % with large oval plate and sternite VII with blunt lobe. % genitalia (Fig. 25): apicolateral margins of segment IX strongly extending backward into a semitransparent capsule almost completely enclosing the genital parts. Segment X less strongly reduced than in other subgenera and equipped with a median sclerite. Protruding preanal appendages generally small, but occasionally forming a long whiplike structure emerging from genital capsule. Inferior appendages large, horizontal, and always protruding from the capsule. Very long aedeagus, occasionally accompanied by short, thick parameres and occasionally joined to base of inferior appendages by a connecting lobe, depending on groups of species. Ripaeglossa is the only genus among the Trichoptera in which the genital armature is contained within a real capsule. It is an exclusively western Nearctic genus and is represented in our region by 10 species. Larvae generally live in large rapid streams. Subgenus Synafophora Martynov

Synafophora Martynov, 1927, p. 165 Type species by original designation: Synafophora minutum Martynov Mystrophora Klapalek, 1892, p. 459 (preoccupied) Monobasic type species: Mystrophora intermedium Klapalek 24

Eomystra Martynov, 1934, p. 84 Monobasic type species: Eomystra dulkejti Martynov Mystrophorella Kloet and Hincks, 1944, p. 97 Nomen novum for Mystrophora Klapalek, 1892, nec Kayser, 1871 Klapalekia Botosaneanu, 1955, p. 792 Nomen novum for Mystrophora Klapalek, 1892 Middle legs of & flattened. Anal cells in front wings of % with a large callosity (Fig. 22). Inner apical spur on middle legs of % thickened and curved (Fig. 23). Sternite VI of % with ventral plate and sternite VII with large lobe (Fig. 27). % genitalia (Fig. 26): segment IX reduced ventrally. Large preanal appendages forming lobes that are blunt or narrowed into spines. Inferior appendages large and horizontal, simple or complex. Small aedeagus with 2 short, erect, vertical, spine-bearing parameres, completely concealed under preanal appendages. The latter are connected to phallotheca by a tendon. There was considerable confusion in the past over the name of this subgenus. It was designated by no fewer than five names, as shown above. The only valid and correct name is Synafophora Martynov. Synafophora has a Holarctic distribution and includes four Canadian species: intermedium Klapalek (Holarctic and transcontinental), verdona Ross (British Columbia, Alberta, and Alaska), lividum Hagen (Quebec, Ontario, and New Hampshire), and nigrior Banks (from Newfoundland to Ontario). In the former two species the inferior appendages are simple, whereas in the latter two they are deeply trifid. Larvae live in rapid streams of all sizes. Subgenus Anseriglossa Ross

Anseriglossa Ross, 1956, p. 157 Type species by original designation: Glossosoma penitum Banks Middle legs of & flattened. Anal cells in front wings of the % with a large callosity covered with erect scaly hairs. Sternite VI in % with large oval ventral plate. % genitalia (Figs. 30, 31): segment IX reduced laterally and ventrally where it forms 3 asymmetrical lobes. Preanal appendages in simple, long, oval lobes. Phallotheca joined to the base of preanal appendages by 2 strong tendons which serve as site of insertion for 2 very stout appendages. The latter widen before ending in a point. They frame the aedeagus, which is asymmetrical, very long, flattened, then tapered at end. It is equipped 25

with stout apical spines on the side only. There are no parameres and no inferior appendages. Anseriglossa is related to the Oriental and European subgenus Glossosoma s. str., but is quite sufficiently different to be considered as a good distinct subgenus. Anseriglossa includes only a single species, penitum Banks, found in British Columbia and further south.

Subfamily Agapetinae Martynov Agapetinae Martynov, 1913, p. 11 Type genus: Agapetus Curtis This subfamily is represented in Canada by a single genus. Genus Agapetus Curtis

Agapetus Curtis, 1834, p. 217 Type species designated by Westwood, 1840: Agapetus fuscipes Curtis Revision: Schmid, 1982 Spurs: 2,4,4. Second segment of maxillary palpi particularly globular. Middle legs of & flattened and strongly ciliate. In %, internal gland of sternite V very large, spherical in shape but secondarily complicated with strongly sclerotized walls. It opens laterally through a narrow slit (Fig. 35). Sternite VI with long appendage, which is much smaller in & than in %. Front wings regularly oval with complete venation. Hind wings clearly smaller and narrower with slightly reduced venation. Venation (Fig. 32): in front wings, R1 does not divide at tip and crossvein R1–R2+3 absent. Otherwise, venation resembles that of Glossosoma, with triangular, slightly oblique discoidal cell. In hind wings, R1 terminates at R2+3, close to anterior end of latter. Open discoidal cell, anastomosis in middle of wing, and A3 ending at A2 at base of wing. % genitalia (Figs. 33, 34): segment IX very robust although occasionally reduced dorsally. Segment X bulky, forming large concave roof enclosing the phallic apparatus; segment membranous over most of surface but equipped at tip with wide variety of sclerotized reinforcements, spines, and points. Intermediate appendages absent. Preanal appendages small and elongate. Inferior appendages large, single-segmented, horizontal, devoid of spines or provided with teeth. Phallic apparatus small and slender and consisting of an aedeagus and 2 asymmetrical parameres. 26

& genitalia (Fig. 36): extended into an ovipositor of medium length with apodemal rods. Segment VIII entire and laterally flattened. Segment IX completely lost. Segment X membranous but with a sclerotized reinforcement toward upper part of lateral sides. Segment XI ovoid, bearing 2-jointed cerci with 2 long, very slender segments. Short vaginal vestibule. Small, simple, and barely conspicuous vaginal apparatus. In all likelihood, it is slightly eversible during copulation. The genus Agapetus is large and distributed in the Holarctic, Oriental, and Australian regions, and in Africa at high altitudes. About 30 species occur in the Nearctic region, with 8 recorded from the eastern and central parts of Canada, from Nova Scotia to Ontario and further south. Larvae are confined to cold running water, in mountainous areas.

Subfamily Protoptilinae Ross Protoptilinae Ross, 1956, p. 149 Type genus: Protoptila Banks Very small species (length of anterior wings: 2.75–4 mm). Both wings narrow, pointed, with rather long fringes. Protoptilinae superficially resemble Hydroptilidae. They can be distinguished by lateral ocellae far from eye margin, longer antennae, no erect hairs on anterior wings, and anastomosis straight. Coloration of anterior wings very dark, with a transverse, white line on anastomosis. Spurs: 0,4,4 or 0,3,3. Middle legs of & not flattened. Venation (Fig. 37): in front wings, discoidal cell present and closed and median cell open. Crossveins of anastomosis in alignment. Forks I, II, and III or I, II, III, and IV present. FI especially wide. In hind wings, indistinct nervation and forks II, or II and III, present. % genitalia: tergite and sternite VIII enlarged and confluent along much of their length. Segment IX small and entirely invaginated within segment VIII. Segment X membranous and barely visible. Preanal appendages prominent, simple or 2-branched, curving downward. Inferior appendages inconspicuous or lost. Phallic apparatus very large, complex, and sometimes strongly modified. & genitalia not elongated into ovipositor and consequently devoid of apodemal rods. Segment VIII massive. Segment IX completely lost. Segments X and XI apparently fused into large, triangular lobes with singlejointed, well-developed cerci at apex. Vaginal apparatus very large, complex, sometimes strongly modified. Protoptilinae are widespread in the Nearctic and Neotropical regions, with a few species in Oriental and Palearctic Asia. Curiously, its 27

most primitive genera are located at both ends of this very large area, Nepaloptila Kimmins in Central Himalaya and Toluaca Schmid in Chile. In Canada, the subfamily is represented by two genera only. Larvae live exclusively in running, but not too cold water. 1a Phallic apparatus between preanal appendages. % sternite VIII platelike, elongated posteriorly (Figs. 38, 39). Vaginal apparatus with very long rod in ventral position (Fig. 40) . . . . . . . . . . . . . . . . . . . . . . . . . . . Protoptila, p. 28 1b Phallic apparatus below preanal appendages (Fig. 18). % sternite VIII not elongated posteriorly. Vaginal apparatus without long rod (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Culoptila, p. 29

Genus Protoptila Banks

Protoptila Banks, 1904, p. 215 Type species by original designation: Beraea? maculata Hagen Revision: Schmid, 1982 Spurs: 0,4,4. Venation (Fig. 37): anterior wings with FI, II, and III. Posterior wings with FII only. % genitalia (Figs. 38, 39): sternite VIII extending strongly posteriorly into large, generally bifid plate, probably replacing lost inferior appendages. Segment IX well developed, but completely invaginated within segment VIII. Its apicoventral margin strongly elongated backward. Xth segment membranous and inconspicuous. Preanal appendages prominent, simple or 2-branched, curving downward. Inferior appendages inconspicuous or lost. Phallic apparatus between preanal appendages, very large and complex at base. Aedeagus thickened at tip, with 2 large spines, which may be parameres inserted on an erectile base. & genitalia (Fig. 40): ventral part of sternite VIII with two ciliated extensions. Segments X and XI apparently fused into large triangular lobes, with single-jointed, well-developed cerci at apex. Ventral portion of segment X forming a lip serving as site of insertion for 2 similar, cercalshaped lobes. Anovaginal vestibule very wide and gaping, with sclerotized walls. It contains the vaginal apparatus, which is small and equipped with slender sclerotized rod as long as 2 abdominal segments. The genus Protoptila is large and widely distributed throughout the Neotropical and Nearctic regions. Five of the more northerly species live also in Canada. Larvae are found in running, not very cold water and occasionally in large slow-moving rivers. 28

Genus Culoptila Mosely

Culoptila Mosely, 1954, p. 336 Type species by original description: Culoptila aluca Mosely This genus was established for several Mexican species. Subsequently, several Nearctic species were included. Originally, the genus was based on the very large size of the % tegulae. In the only Canadian species, cantha Ross, these tegulae are of normal size, as in Protoptila. However, several characters of the venation and genitalia show that this species is rightfully placed in Culoptila. Venation: apical forks I, II, III, and IV present in anterior wings and probably II and III in posterior ones. Spurs 0,3,3. % genitalia (Fig. 18): tergite and sternite VIII narrowly contiguous, regularly short and not extended ventro-posteriorly. Segment IX entirely contained in VIII, strongly reduced and modified. Its ventral part forming thin, horizontal plate. Segment X invisible or absent. Preanal appendages forming two parts: one thin horizontal segment and a more voluminous and vertical part. Phallic apparatus far below preanal appendages, of very large size and of complex and modified structure. Phallocrypt large and extending backward as ogival roof concave downward. Phallic apparatus with large horizontal spine surrounded with membranous and erectile lobes. & genitalia (Fig. 19): segments X and XI fused into large dorsal part bearing cerci. Vulvar lip broadly concave. Vaginal apparatus simple, large, partially membranous, and without long rod. C. cantha, the only Canadian species of the genus, has a wide Nearctic distribution and has been recorded from Saskatchewan. Nothing is known about the biology of its larval stage.

29

Family Hydroptilidae Stephens Hydroptilidae Stephens, 1836, p. 151 Type genus: Hydroptila Dalman The family Hydroptilidae is composed of two subfamilies. These are so different from each other that it is impossible to give a general familial description of the adult characters. The taxonomic unity of the family appears mainly in the larval stage. 1a Wings broad, obtusely rounded at apex, with venation complete (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptilocolepinae, p. 30 1b Wings forming very narrow, tapered lamellae, with very long fringes, especially in hind wings (Figs. 44, 52) . . . . . . . . . . . . . . . . Hydroptilinae, p. 31

Subfamily Ptilocolepinae Martynov Ptilocolepinae Martynov, 1913, p. 22 Type genus: Ptilocolepus Kolenati This subfamily is of relict nature and represents a link between the Glossosomatidae and the Hydroptilinae. It contains only two very small genera: Ptilocolepus Kolenati, European and Oriental in distribution, and Palaeagapetus Ulmer, Nearctic and East Palearctic in distribution. Adults have the habitus of small Agapetinae. Genus Palaeagapetus Ulmer

Palaeagapetus Ulmer, 1912, p. 35 Monobasic types species: Palaeagapetus rotundatus Ulmer (from Baltic amber) Posterior cephalic warts and pronotal warts close to median line. Ocelli present, the lateral ones close to eye margin (Fig. 11). Spurs 2,4,4. Wings broad, obtusely rounded at apex and with venation complete in both wings (Fig. 17). In anterior wings, FI, II, III, IV, and V present. Discoidal cell closed and median cell open. In posterior wings, FI, II, III, 30

and V present. Discoidal and median cells as in anterior wings. Three anal veins present. % genitalia (Figs. 13–15): segment IX massive and with a median subhorizontal line. Inferior appendages broadly fused to segment IX, more or less prominent at apex. Segment X entirely contained in segment IX and so reduced that its components are no longer identifiable. Phallic apparatus short, blunt, and much reduced in structure. & genitalia: last segments extended into an ovipositor. Segment XI with well-developed uni-articulated cerci. Vaginal apparatus tripartite (Fig. 16). The genus Palaeagapetus contains one fossil species, two Nearctic and four east Palearctic species. In Canada, celsus Ross is recorded from Quebec, New Brunswick, and further south; nearcticus Banks (= guppyi Schmid) occurs in the west, on Vancouver Island and southward to California. Larvae live in small, cold seepage springs. Larval cases composed of pieces of liverwort. This genus has the reputation of being very primitive, due to the opinion of Ross (1956). It is surely a very primitive Hydroptilidae, but the male genitalia are among the most reduced by specialization in the whole order Trichoptera.

Subfamily Hydroptilinae Stephens The Hydroptilinae group are the species that are generally referred to as the Microtrichoptera. These insects share a characteristic appearance: they are minute with narrow, lamellar wings provided with very long fringes that compensate for the narrowness of the wings. The antennae are short and all parts of the body are profusely clothed with hairs. In morphology, these insects are among the most specialized and varied, making it difficult to provide a general description. Head short with prominent, very hairy eyes with relatively large ommatidia. Vertex with large, more or less movable posterior warts. Ocelli absent or present. When present, lateral ocelli are very close to inner margin of eyes. Antennae always shorter than front wings and always shorter in & than in % and often comprising moniliform segments. Maxillary palpi 5-segmented in both sexes. The 2 basal segments are very short. Spurs range from 0,2,3 to 1,3,4. Abdominal hemogill system absent. Internal gland of sternite V small, with an opening on a tiny, seta-bearing knob (Fig. 46). 31

Wings forming narrow strips provided with fringes several times longer than wide. The membrane is densely clothed with appressed hairs, whereas the veins may bear short, erect setae. Venation fairly difficult to see in the Hydroptilidae, but a technique I have described elsewhere (Schmid 1959, p. 4) makes venation readily visible. There is considerable variation and the veins may be slightly or considerably reduced or even secondarily complicated. Interpretation is often a matter of conjecture. In the generic descriptions, I shall therefore only briefly mention its characters. % genitalia also display considerable diversity. They consist of appendages which, in our present state of understanding, usually cannot be identified with any degree of certainty. The following interpretations are therefore approximate. The genitalia are completely concealed by a very dense pilosity. In the figures, they have been shown hairless. Abdominal sternite VII with a point or a ventral appendage. Segment IX well developed but usually deeply invaginated within segment VIII. Segment X occasionally bulky, forming a roof over the phallic apparatus. Preanal appendages absent. Intermediate appendages present or absent and at times fused together to form an unpaired structure. Inferior appendages single-segmented, variable in size, occasionally forming an upper branch which could be taken for the intermediate appendages. Phallic apparatus always very long and slender and comprises a long tubelike phallotheca, which is directly extended by the aedeagus. The latter is also tube-shaped and occasionally spined. Endotheca obliterated and lost. Parameres also seem to be lost, but some genera have a corkscrew-like spine at base of the aedeagus and this could be one of the modified parameres. & genitalia very simple and homogeneous throughout the generic series. It is similar to the genus Rhyacophila but simplified through loss of segments VIII and IX. Abdominal segment VII is the first one to show any sexual changes. It consists of a single part in the shape of a ring or truncated cone, with ventral lobes or seta-bearing tubercles. It is endowed with a pair of anterior apodemal rods. Segments VIII and IX lost or virtually so. Segment X slightly developed and also equipped with apodemal rods. Segment XI forms 2 contiguous, ovoid lobes with single-jointed cerci. Vulvar scale absent. Anovaginal opening between segments VII and X. Vaginal apparatus simple, with little sclerotization. It is located at the end of a membranous vaginal vestibule, but unlike the vaginal systems in the previous two families, it does not seem to evaginate during mating. The Nearctic genera of Hydroptilinae have been classified into 6 tribes. I do not use them here, finding them of little use in the framework of a manual. The Hydroptilidae make up a huge family with a worldwide distribution. Only eight genera have been reported in Canada, and to this list I add 32

Leucotrichia Mosely and Stactobiella Martynov, which have been captured in bordering American states. There are more than 80 species in Canada and their larvae occur in water of all types, from cold springs to marshes and large lakes. They are not found, however, at high altitudes or very far north. In the case of some genera, the following key applies only to the %%. Since the Hydroptilidae are present in large numbers locally, there will be no trouble in identifying the && by association of the sexes. 1a Ocelli absent (Fig. 57) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b Ocelli present (Fig. 42) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a Spurs: 0,3,4. % genitalia asymmetrical (Figs. 65, 66). . . . Orthotrichia, p. 36 2b Spurs: 0,2,4. % genitalia symmetrical (Figs. 59, 60). . . . . . Hydroptila, p. 35 3a Spurs: 1,3,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucotrichia, p. 38 3b Spurs: 0,2,3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotrichia, p. 39 3c Spurs: 0,2,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mayatrichia, p. 40 3d Spurs: 0,3,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a Wings very narrow and lanceolate with only 2 veins at tips (Fig. 75) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxyethira, p. 37 4b Species with wider wings and more complex venation . . . . . . . . . . . . . . . . . 5 5a Inferior appendages in % very large, slightly asymmetrical, the length far exceeding height (Figs. 93–95) . . . . . . . . . . . . . . . . . . . Ochrotrichia, p. 39 5b Inferior appendages in % not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a Front wings relatively wide, with more than 7 veins at tips (Fig. 44) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agraylea, p. 33 6b Front wings not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7a Segment IX in % with 2 long anterior apodemal rods (Fig. 70) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stactobiella, p. 36 7b Segment IX in % without these rods (Figs. 52, 54) . . . . . . . Ithytrichia, p. 34

Genus Agraylea Curtis

Agraylea Curtis, 1834, p. 217 Type species, subsequently designated by Westwood, 1840: Agraylea sexmaculata Curtis Ocelli present. Vertex with 3 pairs of barely prominent warts (Fig. 42). Antennae in % slightly over half length of front wings. Spurs: 0,3,4. Wings not very narrow in comparative terms, with only moderately long fringes. Venation fairly complete (Fig. 44). In front wings, R1 long, 33

terminating at wing margin. Apical forks I, II, and III present. In hind wings, forks I, II, III, and V present. % genitalia (Figs. 45–47): sternite VII with a large, simple or bifid median ventral lobe. Segment IX massive, slightly invaginated within the preceding segment, with lateral median margins extending far backward in tonguelike structure over inferior appendages. Segment X forming fold over phallic apparatus and with downward-curving spur-like part supporting complex from below. Inferior appendages faintly prominent, massive, consisting of 2 or 3 distinct lobes. Phallic apparatus stout but not very long and provided with a coiled paramere. & genitalia (Figs. 49, 50): segment VII consists of a tergite and a sternite. The latter has simple relief on ventral side with an apicoventral row of small seta-bearing warts. A rudimentary segment VIII may be present, but it is difficult to be sure. Vaginal apparatus: Fig. 51. The genus Agraylea is small and of Holarctic distribution. It comprises three Nearctic species, all reported from Canada, i.e., multipunctata Curtis, a Holarctic species that is transcontinental in Canada, costello Ross, which occurs in Ontario and Quebec, and saltesa Ross, recorded from British Columbia and southward. In the first species, ventral lobe of sternite VII bifid and the inferior appendages trilobate, whereas in costello Ross, ventral lobe of sternite VII simple and inferior appendages bilobate; saltesa Ross is characterized by the conical process of sternite VII and the shape of the inferior appendages. Larvae of these species occur in lakes, ponds, and in tranquil parts of large rivers. Genus Ithytrichia Eaton

Ithytrichia Eaton, 1873, p. 139 Type species by original designation: Ithytrichia lamellaris Eaton Ocelli present. Posterior cephalic warts slightly detached from their substrate. Antennae nonmoniliform and over half length of front wings. Spurs: 0,3,4. Wings of medium width with moderately reduced venation (Fig. 52). In front wings, R1 moderately long, ending on Sc. RS 4-branched and M 3-branched. In hind wings, RS 4-branched and M simple. % genitalia (Figs. 53, 54): sternite VII with a fine ventral point. Segment IX massive, completely open dorsally, membranous ventrally, ending in 2 small lateral knobs. Segment X developed into simple membranous plate occupying the space left by dorsal opening of segment IX and overhanging phallic apparatus. Intermediate appendages apparently present, but tiny, inconspicuous, and within apical angles of segment IX. Inferior appendages forming long, simple, horizontal lobes inserted 34

virtually at base of segment IX and partially embedded under it. Phallic apparatus medium-sized with large coiled paramere. & genitalia (Fig. 55): segment VII with row of apical seta-bearing tubercles and smooth relief on underside. Vaginal apparatus: Fig. 56. The genus Ithytrichia is very small and of Holarctic distribution. It comprises only two Nearctic species, only one of which, clavata Morton, has been reported in Canada. It has a transcontinental distribution. Its larvae develop in mosses submerged in running water. Genus Hydroptila Dalman

Hydroptila Dalman, 1819, p. 125 Monobasic type species: Hydroptila tineoides Dalman Ocelli absent. In %, posterior cephalic warts movable and can be raised like valves (Fig. 57). They conceal an erectile organ bearing brushlike or scaly pheromonic androconia. These organs vary considerably from species to species but are not convenient for species identification since they are difficult to observe in the state of turgescence. Cephalic warts of & normal. Antennae of % moniliform, less than half the length of front wings. Spurs: 0,2,4. Hind femora with very long hairs. Wings of medium width and moderately reduced venation (Fig. 58). In front wings, R1 moderately long, ending on Sc. RS 4-branched and M 3-branched. In hind wings, RS 3-branched and M 2-branched. % genitalia (Figs. 59–61) displaying considerable variation, making it difficult to provide a generic description. Abdominal sternite VII with stout ventral appendage. Segment IX very long laterally and deeply invaginated within segment VIII. Apicolateral angles often elongated into triangular lobes extending alongside other genital parts. Segment X variable in size and complexity, forming a roof over the phallic apparatus. Inferior appendages always well developed, generally horizontal and simple in shape, but in some groups, such as the spatulata group, they have an upper branch which lies alongside segment X. Very large phallic apparatus. Coiled paramere present and aedeagus often forming apical elbow. & genitalia (Fig. 62): segment VII lobate or with apical crenulations bearing long setae. Vaginal apparatus simple and slender. The genus Hydroptila is huge, highly varied and cosmopolitan. It is divided into a large number of very distinct groups of species, plus a number of isolated species. There are nearly 80 Nearctic species, about 40 of which have been recorded from our area. The species have an extensive ecological tolerance. Larvae are found in lakes and running water of every type. 35

Genus Orthotrichia Eaton

Orthotrichia Eaton, 1873, p. 141 Type species by original designation: Hydroptila angustella McLachlan Clymene Chambers, 1873, p. 114 (preoccupied) Monobasic type species: Clymene aegerfasciella Chambers Revision: Kingsolver and Ross, 1961 Ocelli absent. Vertex with large posterior warts, partially detached from their substrate. Antennae relatively long, two-thirds length of front wings; nonmoniliform. Spurs: 0,3,4. Front wings uniformly dark in middle with silvery anterior and posterior margins. Venation (Fig. 63) only moderately reduced although wings quite tapered at tips. In front wings, R1 long, ending on wing margin. RS 4-branched and M 3-branched. In hind wings, RS 3-branched and M 2-branched. % genitalia (Figs. 64–67): sternite VII with large appendage with club-shaped spines. Segment IX well developed and massive. Segment X barely distinct from preceding segment, forming a highly asymmetrical roof over phallic apparatus and often equipped with a slender spine on the left side. Inferior appendages small, simple, lunulate, with unpaired basal plate between. Phallic apparatus very large, almost as long as abdomen, with coiled paramere and very slender aedeagus. & genitalia (Fig. 68): sternite VII forming a very short tube with small ventral tonguelike protuberance. Vaginal apparatus membranous and very small. The genus Orthotrichia has a Holarctic, African, and Oriental distribution. There are six Nearctic species, three of which have been recorded from Canada: cristata Morton (Quebec, Ontario, Manitoba, and British Columbia), curta Kingsolver and Ross (Quebec), and aegerfasciella Chambers (Quebec and Ontario), which are distinguished primarily by the shape of segment X and of the inferior appendages. Larvae of these species live primarily on submerged aquatic plants found in lakes and gentleflowing watercourses. Genus Stactobiella Martynov

Stactobiella Martynov, 1924, p. 37 Monobasic type species: Stactobia ulmeri Siltala Tascobia Ross, 1944, p. 124 Type species by original designation: Stactobia palmata Ross 36

Ocelli present. Posterior cephalic warts very prominent and partially detached from their substrate. Antennae in % very short. Spurs: 0,3,4. Wings fairly strongly tapered. Venation quite considerably simplified (Fig. 69). R1 in front wings very short and ending on Sc. RS forms only very short fork and M 3-branched. In hind wings, RS simple and M 4branched. % genitalia (Figs. 70–72): sternite VII without ventral appendage. Segment IX with reduced and open ventral portion. This segments extends anteriorly into pair of long apodemal rods. Segment X completely membranous, framing phallic apparatus dorsally and laterally. Intermediate appendages fused into slender unpaired lobe. Inferior appendages long and slender, symmetrical or asymmetrical, simple or complex. Phallic apparatus evenly slender with fairly large tip; no coiled paramere. & genitalia (Fig. 73): sternite VII a simple tube with regular row of small seta-bearing tubercles. Vaginal apparatus: Fig. 74. The genus Stactobiella is small, Holarctic, and comprises six Nearctic species. Two of them have been reported in Canada and southward: palmata Ross (Alberta, Manitoba) has denticulate, asymmetrical inferior appendages, whereas in delira Ross (British Columbia) these appendages form simple, symmetrical lobes. These species occur close to fast-flowing and often very small streams. Genus Oxyethira Eaton

Oxyethira Eaton, 1873, p. 143 Type species by original designation: Hydroptila costalis Curtis sensu Eaton Ocelli present. Posterior cephalic warts small and undetached from their substrate. Antennae of % nonmoniliform and longer than half the length of front wings. Spurs: 0,3,4. Wings quite strongly narrowed with especially long fringes. In dry specimens, wings curve strongly upward at the apex. Venation (Fig. 75) greatly simplified but in proportion to the narrowness of wings. No generic significance should therefore be attached to these variations. In front wings, R1 moderately long, ending at wing margin. RS 3-branched and M 3- or 2-branched. In hind wings, RS simple and M 2-branched. % genitalia simplified and not very prominent (Figs. 76–78). Sternite VII with a fine ventral point. Segment VIII stout with apicolateral angles occasionally equipped with points or spines. Segment IX extremely elongated and completely invaginated within segment VIII. Segment X small, more or less complex, forming a membranous area over 2 small, highly 37

sclerotized intermediate appendages. Inferior appendages barely prominent. Phallic apparatus very long. Coiled paramere present, occasionally highly developed. Tip of aedeagus displaying varying degrees of complexity. & genitalia (Fig. 79): sternite VII highly developed, forming large ventral plate protecting last segments on underside. Segment X complex in shape, with dorsal protuberance. Segment XI fairly long, with relatively large cerci. Vaginal apparatus membranous and simple. The genus Oxyethira is large and Holarctic, African, and Oriental in distribution. It comprises more than 35 Nearctic species, about 18 of which are native to our area and southward. Larvae inhabit submerged aquatic plants found in lakes and gentle-flowing or quiet watercourses.

Genus Leucotrichia Mosely

Leucotrichia Mosely, 1934, p. 157 Type species by original designation: Leucotrichia melleopicta Mosely Only 2 ocelli present. Pronounced sexual dimorphism in anterior portion of body. Antennae of %: first 6 segments, short, thick, and clothed with very dense hairs (Fig. 82). Two pairs of very large, fairly strongly erectile warts on top of head, densely covered with scaly hairs. Tegulae highly developed, also erectile and densely clothed with modified pilosity (Fig. 81). Spurs 1,3,4. Wings fairly wide, but distinctly acuminate. Venation both simplified and secondarily complicated (Fig. 80). In front wings, R1 long, ending on wing margin. RS secondarily divided into 6 branches. M 2-branched. In hind wings, an additional short vein, parallel to base of R1 and equipped with row of hooked setae probably acting as frenulum. RS 4-branched and M simple, joining RS for some distance midlength. % genitalia (Figs. 83–85): sternite VII without ventral appendage. Sternite VIII deeply cleft at apex. Segment IX completely open ventrally, but without anterior apodemal rods. Segment X simple, forming a series of lateral plates quite strongly sclerotized. Intermediate appendages fused into single, spiniform part, located very low, between the inferior appendages. The latter small, slender, and contiguous. They are concave on the upper surface, surround intermediate appendage, and bear short lateral spine on upper surface. Phallic apparatus without paramere but with robust, highly complicated aedeagus consisting of membranous lobes and spines. & genitalia (Fig. 86): segment VII blunt with row of small apicoventral tubercles bearing long setae. Long, slender vaginal apparatus. 38

The genus Leucotrichia occurs only in the Americas and is particularly well represented in Mexico and Central America. It comprises only three Nearctic species, none of which has been reported in Canada. The species pictipes Banks, however, is widespread transcontinentally. Genus Neotrichia Morton

Neotrichia Morton, 1905, p. 72 Monobasic type species: Neotrichia collata Morton Cyllene Chambers, 1873, p. 124 (preoccupied) Monobasic type species: Cyllene minutisimella Chambers Ocelli present. Antennae two-thirds length of front wings; nonmoniliform. Posterior cephalic warts slightly detached from their substrate. Spurs: 0,2,3. Wings extremely narrow and tapered with particularly long, thick fringes. Venation considerably simplified, consequently most veins thick (Fig. 87). In front wings, R1 very short, ending on Sc. RS 2-branched and M simple. In hind wings, RS simple and M bifid. % genitalia (Figs. 88–90): segment VII without ventral appendage. Segment IX not very elongated nor deeply invaginated within preceding segment. Segment X forming large membranous dorsal lobe above phallic apparatus and bearing underneath several small, variously complex, more markedly sclerotized protuberances. Inferior appendages bipartite, forming leaflike upper lobe laterally and 2 stout spurlike parts ventrally. Phallic apparatus with very long and thick phallotheca and correspondingly smaller aedeagus. Coiled paramere present. & genitalia (Fig. 91): last segments generally slightly more elongated than in other genera. Sternite VII with simple specifically characteristic ventral relief and small seta-bearing apical tubercles. Slender vaginal apparatus. The genus Neotrichia contains the smallest known Nearctic Trichoptera. The genus is solely Neotropical and Nearctic in distribution and comprises 14 species, 6 of which have been recorded from our region. Larvae inhabit the fast-flowing portions of large rivers. Genus Ochrotrichia Mosely

Ochrotrichia Mosely, 1934, p. 162 Type species by original designation: Ochrotrichia insularis Mosely Polytrichia Sibley, 1926, p. 102 (preoccupied) Monobasic type species: Ithytrichia confusa Morton 39

Revision: Denning and Blickle, 1972 Ocelli present. Posterior cephalic warts slightly detached from their substrate. Antennae virtually as long as front wings, nonmoniliform and very hairy. Spurs: 0,3,4. Wings not strongly acuminate, with some parts of venation reduced and others secondarily divided (Fig. 92). In front wings, R1 long, ending on wing margin. M 2-branched, whereas RS with a succession of divisions and no fewer than 6 branches. In hind wings, RS 5-branched and M simple. % genitalia very prominent and asymmetrical (Figs. 93–95). Segment VII with fine ventral point. Segment IX fairly strongly invaginated within segment VIII and consisting of 2 distinct parts: a horizontal dorsal part, concave on underside, and a ventrolateral part with typical family shape and very prominent apicolateral angles. Segment X very large, horizontal, surrounding phallic apparatus and consisting of large compact mass of highly asymmetrical and complex plates, lobes, and spines. Inferior appendages very large, oval, very elongate, horizontal and strongly concave on inner surface. The 2 inner margins with asymmetrical tooth-bearing and dentiform projections. Phallic apparatus forming long, very slender, simple tube. Coiled paramere absent. & genitalia (Fig. 96): last segments drawn out into relatively long ovipositor. Segment VII massive, truncated, ending in blunt lobes, and equipped with pair of apodemal rods inserted on posterior margin. At base, intersegmental membrane VII–X invaginated within segment VII for variable distance and displaying granulated reticulation on inner ventral side. Vaginal apparatus Fig. 97. The genus Ochrotrichia comprises about 70 species and is found only in the Nearctic and Neotropical regions. Three species have been recorded from Canada: tarsalis Hagen has a wide distribution and is found from Mexico to New York, Ontario, and Manitoba, stylata Ross is known from British Columbia, and potomus Denning from Manitoba. Six other species have also been recorded from neighboring states. Larvae occur in many types of running water and occasionally even in temporary streams.

Genus Mayatrichia Mosely

Mayatrichia Mosely, 1937, p. 182 Type species by original designation: Mayatrichia ayama Mosely Ocelli present. Posterior cephalic warts large, triangular, partially detached from their substrate. Antennae half length of front wings, nonmoniliform, and comprising at least 20 segments. Spurs: 0,2,4. 40

Wings moderately tapered with moderately reduced venation. Venation (Fig. 98): in front wings, R1 short. RS 3-branched and M 2-branched. In hind wings RS and M 2-branched. % genitalia (Figs. 99, 100): sternite VII equipped with long spiniform point. Segment IX robust, looking triangular from side but not strongly invaginated within segment VIII. Lower margins tucked under inferior appendages. Segment X consists of a membranous lobe over phallic apparatus, a sclerotized dorsolateral plate, and a median lobe located under phallic apparatus and ending in a downward-curving hook. Inferior appendages 2-branched. Upper branch forms a very prominent lobe on either side of lower part of segment X. Lower branch is large quadrangular piece with a deep indentation at extremity, and base inserted into segment IX. Phallic apparatus moderately long, slender, simple, and devoid of coiled paramere. & genitalia (Fig. 101): segment VII forms a simple tube. Vaginal apparatus fairly large and only slightly complex. The genus Mayatrichia is small and found only in the Americas. It comprises four Nearctic species, one of which, ayama Mosely, has a very wide distribution extending from Mexico to Maine and Montana and including Quebec, Ontario, Saskatchewan, Manitoba, and Alberta. Larvae of this species occur in the fast-flowing sections of large rivers.

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Family Philopotamidae Stephens

Philopotamidae Stephens, 1829, p. 316 Type genus: Philopotamus Stephens Occipital portion of head well developed and extending in wide curve far behind the eyes (Figs. 102, 103). Ocelli present. Maxillary and labial palpi of considerable length, with very long 5th segment forming a flexible flagellum. Maxillary palpi with 5 segments; 2nd twice the length of 1st, and 4th half the length of 5th (Fig. 102). Spurs: 2,4,4 or 1,4,4. Middle legs in & not flattened. Abdominal hemogill system and internal gland of abdominal sternite V absent. Regularly oval wings with both pairs similarly shaped in both sexes. Venation usually complete with all forks present: I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. Discoidal cell present and closed on both wings and median cell closed in front wings. Crossveins C-Sc and R1–R2+3 present in front wings. In hind wings, 2 or 3 anal veins. Veins in front wings slightly crowded toward anterior margin, with the result that the cells in posterior half are broader than the others. % genitalia considerably simplified: segment IX ring-shaped but greatly reduced dorsally. Preanal appendages present, in shape of elongated lobes or knobs. Segment X roof-shaped, only slightly sclerotized or membranous, simple or bilobed. Intermediate appendages present or absent. One- or 2-segmented inferior appendages. Phallic apparatus with aedeagus and parameres consistently absent. The apparatus consists of phallotheca and endotheca only, which are variously modified. & genitalia: last segments may or may not be drawn out into an ovipositor. Apodemal rods generally present. Segment IX lost or virtually so. Segment XI particularly hairy. One- or 2-segmented cerci. Anovaginal vestibule short with membranous walls. Vulvar scale absent. Anovaginal opening at the apex of segment X. Vaginal apparatus very simple and not eversible during mating. The Philopotamidae are divided naturally into two subfamilies: the Philopotaminae, a primitive subfamily, and the Chimarrinae, with clearly more specialized venation and genitalia. The family is ubiquitous but rather poorly represented in Canada. There are only a dozen species in this 42

country, grouped into three genera. All of them occur in lotic environments. 1a Spurs: 2,4,4. Discoidal cell in front wings of long ogival shape; FIV present (Figs. 104, 111) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Philopotaminae, p. 43 1b Spurs: 1,4,4. Discoidal cell in front wings short and wide with anterior tip thickened; FIV absent (Fig. 116) . . . . . . . . . . . . . . . . . . Chimarrinae, p. 45

Subfamily Philopotaminae Stephens Philopotaminae Ulmer, 1903, p. 116 Type genus: Philopotamus Stephens Spurs: 2,4,4. Venation in front wings thin and regular. Discoidal cell in front wings ogival. Thyridial cell large. FIV present and particularly wide. Anal veins in hind wings do not form loop. % genitalia: segments VII, VIII, and IX same size as preceding segments with the result that the genital parts are prominent and large compared to abdomen. Segment IX evenly shaped. Segment X roof-shaped and fairly thin, entire or bilobed with free, elongated preanal appendages and no intermediate appendages. Inferior appendages very large and horizontal, consisting of 2 simple, well-defined segments equipped on inner surfaces with various tubercles or spines. Phallic apparatus highly variable in size and equipped with small endothecal spines. & genitalia always somewhat extended into ovipositor, which is never very long. Segment VIII comprises a tergite and a sternite. Apodemal rods proportional in length to degree of elongation of the last segments. Cerci always bi-articulated. The Philopotaminae are represented in Canada by only two genera which occur in running water. 1a FI in both wings with a petiole of variable length. Hind wings with 3 free anal veins (Fig. 104) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dolophilodes, p. 43 1b FI in both wings sessile or absent. Hind wings with 2 free anal veins (Fig. 111) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wormaldia, p. 44 1c Wings reduced to tiny scales (overwintering &&) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dolophilodes distinctus Walker

Genus Dolophilodes Ulmer

Dolophilodes Ulmer, 1909, p. 125 Monobasic type species: Dolophilodes ornatus Ulmer 43

Trentonius Betten and Mosely, 1940, p. 11 Type species by original designation: Trentonius distinctus Walker Revisions: Schmid, 1982; Armitage, 1991 Spurs: 2,4,4. In both wings, FI with petiole of variable length (Fig. 104). In hind wings, 3 anal veins extending freely to wing margin. Sternites VII and VIII in % without ventral plates or lobes. % genitalia (Figs. 105–107): preanal appendages variable in size, always ear-shaped or short-lobed. Segment X either slightly or completely cleft into 2 lobes. Inferior appendages with 2nd segment either shorter than 1st or subequal. Phallic apparatus deeply modified into a very unusual structure. The phallotheca looks like a membranous internal sac, variable in size and occasionally very large, ending posteriorly in a point which is slightly evaginated under segment X. Lower left portion concave and equipped with one membranous, erectile cylinder ending in a long eversible spine. Endotheca completely contained within phallotheca and equipped with a large, conspicuous phallotremal sclerite that is eversible during mating. & genitalia (Figs. 108, 109) not very long. Segment VIII consists of a tergite and sternite, the latter with a barely perceptible apodemal rod. The corresponding rod in segment X is small but distinctly longer. Segment XI very stout. The genus Dolophilodes is large and Asian, Australian, South African, Neotropical, and Nearctic in distribution. It is divided into several subgenera but only the type subgenus is represented in Canada; it contains five species found in the cold running waters of the mountains of eastern and western Canada. The genus originated in the Oriental region, where the greatest number and the most primitive species occur (Schmid 1989, p. 110). Genus Wormaldia McLachlan

Wormaldia McLachlan, 1865, p. 140 Type species designated by Ross, 1949: Hydropsyche occipitalis Pictet Dolophilus McLachlan, 1868, p. 301 Monobasic type species: Dolophilus copiosus McLachlan Paragapetus Banks, 1914, p. 202 Monobasic type species: Paragapetus moestus Banks Dolophiliella Banks, 1930, p. 230 Type species by original designation: Dolophiliella gabriella Banks Revisions: Schmid, 1982; Armitage, 1991 44

Spurs: 2,4,4. Hind femora with long silky hairs. Venation (Figs. 110, 111): in both wings, FI absent or present, then sessile. Three anal veins in hind wings but A1 joins A2 at base and does not reach wing margin. Abdominal sternites VII, VIII, and IX occasionally with large ventral plates or lobes. % genitalia (Figs. 112, 113): preanal appendages always developed into elongated finger-shaped lobes. Segment X in form of ogival roof, uncleft at tip. Inferior appendages with 2nd segment equal to or longer than 1st. Phallic apparatus fairly small, consisting of a broad-based tubelike phallotheca containing an invaginated spine-bearing endotheca. & genitalia (Figs. 114, 115) extended into fairly long ovipositor and equipped with apodemal rods of proportionate length. Segment VIII comprising both a tergite and sternite. Segment X fairly large. Segment XI small with 2-segmented cerci. Vaginal apparatus very small, simple, and annular. The genus Wormaldia is medium-sized and of Holarctic, Oriental, African, and Neotropical distribution. Of the 12 Nearctic species, 6 occur in our area. One of these species, gabriella Banks, belongs to the relicta group that has an Oriental origin. It is in the Oriental region that the largest number of species occur (Schmid, 1991a, p. 93).

Subfamily Chimarrinae Rambur Chimarrrhides Rambur, 1842, p. 498 Type genus: Chimarrha (recte Chimarra) Stephens This subfamily is represented in Canada by a single genus. Genus Chimarra Stephens

Chimarra Stephens, 1829, p. 318 Monobasic type species: Phryganea marginata Linnaeus Revisions: Schmid, 1982; Armitage, 1991 Spurs %,&: 1,4,4. Venation at base of wing and anterior portion of front wings thickened and generally arranged in a somewhat irregular pattern, thereby modifying the shape of some cells (Fig. 116). Discoidal cell consequently short and very wide, with anterior tip thickened and RS slightly sinuate. Median and thyridial cells very small. FIV absent and FV not especially wide. In hind wings, discoidal cell also small and Sc markedly thickened. Three anal veins, but A1 and A2 join to form loop and extend together to wing margin. 45

% genitalia (Figs. 117–120): segments VII, VIII, and IX, progressively smaller, with the result that the genitalia are small compared to the abdomen and not very prominent. Tergite and sternite VIII short and contiguous laterally. Sternite VIII with ventral point. Segment IX complex in shape, deeply invaginated within preceding segment, and with stout ventral lobe. Segment X small, membranous, with no specific shape. Preanal appendages form small, hairy knobs. Intermediate appendages well developed and of a specifically characteristic shape. Inferior appendages singlesegmented, rather small, with fairly complex inner relief. Phallic apparatus consists of a tubelike phallotheca arising from a bulbous base; its lower apical edge extended and curved into a sclerotized hook. Endotheca membranous and completely contained within phallotheca. When evaginated, endotheca displays lobes and oddly shaped spines in some species (Figs. 119, 120). & genitalia (Figs. 121, 122) large and blunt. Tergite and sternite VIII completely fused into short cylinder open dorsally, with apical rim bearing few tubercles surmounted by long setae. Segment X short, complex in shape, and equipped with short apodemal rod. Segment XI large and bulbous, with 2-segmented cerci. Vaginal apparatus simple and partially membranous. The genus Chimarra is huge, virtually cosmopolitan in distribution but with strong tropical affinities. There are 20 Nearctic species, only 4 of which occur in central and eastern Canada, from Manitoba to Newfoundland: socia Hagen, obscura Walker, feria Ross, and aterrima Hagen. Members of the genus can be easily recognized by their all black coloring. They inhabit gently flowing water, are often abundant locally, and are strongly attracted to light.

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Family Arctopsychidae Martynov Arctopsychidae Martynov, 1924, p. 25 Type genus: Arctopsyche McLachlan Female clearly larger and heavier than %. Ocelli absent. Five-segmented maxillary palpi in both sexes, with first 2 segments very short and subequal, 4th slightly shorter than 3rd, and 5th forming a long flagellum. Antennae slightly thickened, particularly in %, with very short, globular scape. Antennae slightly crenulated along basal half where each segment bears a dark cross groove (Fig. 123). Spurs: 2,4,4 and very large. Abdominal pleurites II to VII equipped with hemogills that are generally invaginated behind a fold of skin. In a turgescent state (Fig. 125), they appear to consist of 1 or more basal bulbs per segment, each bearing large number of slender and unbranched gill tubes. Internal gland of sternite V long and narrow with an opening on a slight protuberance. Roughly oval wings. Both pairs in both sexes almost same in shape, but hind wings have regularly rounded lower margin. Venation very similar in the two genera comprising the family, with all forks present: I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. In both wings, discoidal cell small and triangular. In front wings, discoidal, median, and thyridial cells closed. Thyridial cell especially long proximally. Crossveins C-Sc, Sc-R1, and R1–R2+3 present. Cu2 terminates at A before wing margin. Postcostal cell very wide. On hind wings, crossveins Sc-R1 present. Four free anal veins. % genitalia very different in the two genera. Segment IX fairly well developed dorsally. Preanal appendages present, free or fused with segment X. Intermediate appendages also present, free or fused together. Inferior appendages large or small and always 2-segmented. Phallic apparatus large, located very high in abdomen, and comprising a tubelike phallotheca containing small, invaginated, membranous, erectile endotheca. Aedeagus and parameres absent. & genitalia with last 2 segments short and therefore without apodemal rods. Tergite VIII very large, extending quite far down on the sides. Corresponding sternite proportionately reduced, ending in 2 large apicoventral lobes covering base of segment X. Segment IX absent. Segment X short, with smooth relief, and devoid of concavities to accommodate inferior appendages of male during mating. Dorsal margin bearing 2 small warts 47

equipped with slender brush of very long setae. Ventrally and posteriorly, segment X forms a large membranous vulvar scale. Anovaginal opening located at apex of segment X. Vaginal vestibule membranous and short. Vaginal apparatus complex and noneversible during mating. Cerci present, mono- or bi-articulated. The Arctopsychidae constitute a small family very closely related to the Hydropsychidae. It can be considered either a distinct family or a subfamily of the latter. It has an Oriental and Nearctic distribution with one Holarctic species: ladogensis Kolenati. The Arctopsychidae are confined to running water and contain only two genera, both of which are represented in Canada. Both genera originated independently in the Oriental region, where the greatest number and the most primitive species occur (Schmid 1989, p. 110). 1a Eyes bare. % genitalia prominent and not invaginated within preceding segment (Fig. 128). Middle legs of & flattened (Fig. 124) Arctopsyche, p. 48 1b Eyes hairy. % genitalia only slightly prominent and invaginated within preceding segment (Fig. 133). Middle legs of & not flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapsyche, p. 49

Genus Arctopsyche McLachlan

Arctopsyche McLachlan, 1868, p. 300 Monobasic type species: Aphelocheira ladogensis Kolenati Revision: Schmid, 1968b; Nimmo, 1987 Eyes bare. Third segment of maxillary palpi twice as long as thick and not much longer than 4th segment. Middle legs of & very flattened and ciliated (Fig. 124). Abdominal hemogill system highly developed, consisting of 2 to 4 bulbs per segment, each equipped with 8 to 40 tubes, 1.5 times the length of a single segment. Venation (Figs. 126, 127): discoidal cell on both wings and median cell on front wings small. % genitalia (Figs. 128, 130): segment IX almost as high as segment VIII and not invaginated within latter. Preanal appendages free and long, slender, and oval. Intermediate appendages form very stout, long, single or double, smooth or crenulated spines. Segment X completely membranous and either very short or in the shape of a long slender tube, depending on the species group. Inferior appendages 2-segmented, but reduced in size and complex in shape: 1st segment massive, with a point or dorsal lobe and 2 or 3 ventral points and 2nd segment small and inserted between these points. Phallic apparatus very large and stout, consisting of a phallotheca in shape of long tube containing the endotheca. The latter membranous and erectile and ends in an S-shaped phallotremal sclerite. 48

& genitalia (Figs. 131, 132): apicolateral margins of segment X form slightly prominent winglets, homologous with apicolateral angles of the Hydropsychinae. Segment XI strongly extended on underside. Cerci biarticulated. Vaginal apparatus very large and complex. The genus Arctopsyche is small, with a Palaearctic, Oriental, and Nearctic distribution. It includes five North American species, three of which are native to Canada: grandis Banks and inermis Banks, which are closely related but distinct and occur in the western part of the country in lotic biotopes, and ladogensis Kolenati, holarctic, which is found across the country in lentic environments. Genus Parapsyche Betten

Parapsyche Betten et al., 1934, p. 181 Monobasic type species: Arctopsyche apicalis Banks Revision: Schmid, 1968b; Nimmo, 1987 This genus closely resembles Arctopsyche by characters of body, wings, and genitalia of &, but differs considerably in genitalia of %, which are more highly specialized. Eyes with fairly long, dense pilosity. Palpi slightly longer than in Arctopsyche, with 3rd segment of maxillary palpi at least 3 times longer than wide. This segment strongly convex on inner surface. Middle legs in & nonflattened and ciliated. Hind femora very long and corresponding tarsi shortened, especially in &. Abdominal hemogill system consists of a single bulb per segment, each bearing 8 to 10 thick and moderately long terminal tubes. Venation identical to that of Arctopsyche, but with definitely larger discoidal and median cells in front wings. % genitalia (Figs. 133–135): segment IX fairly small and partially enclosed within segment VIII to which it is connected by long membranes. It is short, and the dorsal portion protrudes vertically as a result of the low position of intermediate appendages. Preanal appendages completely and firmly fused with base of intermediate appendages where they persist only as scars. Intermediate appendages large, horizontal, in shape of slightly sclerotized lamellae, fused together only at base or along virtually their entire length, and equipped with sensory microsetae. They are joined to apex of phallocrypt by conspicuous internal tendon. Inferior appendages fairly large, slanting slightly upward, and 2-segmented. First segment simple, large and 2nd segment inserted at apex or in middle of 1st and partially fused with latter. Phallic apparatus consisting of a tube-shaped phallotheca and an erectile, membranous endotheca equipped with paired external phallotremal sclerites in shape of downward-curving hooks. 49

Phallotheca accompanied by unpaired, oval, dorsal lobe arising from upper apical margin of phallocrypt and protruding from abdominal cavity. & genitalia (Fig. 136) very similar to those in Arctopsyche. Apicolateral edge of segment X not raised in a winglet and segment XI does not extend so far down. Cerci mono-articulated. Vaginal apparatus smaller and almost completely membranous. The genus Parapsyche is of Oriental and Nearctic distribution and includes seven Nearctic species. Three of these are Canadian: almota Ross and elsis Milne inhabit the lotic regions of British Columbia, Alberta, and southward, and apicalis Banks is found from Newfoundland to Ontario and southward. The species are characteristic of cold, swift streams.

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Family Hydropsychidae Curtis Hydropsychidae Curtis, 1835, p. 544 Type genus: Hydropsyche Pictet Ocelli absent. Eyes occasionally very large in %. Maxillary palpi 5segmented in both sexes. Basal segments vary in length but 5th segment is always very long and flagellate. Middle legs of & generally flattened. Spurs usually 2,4,4. Abdominal hemogills always present. Sternite V often with lobe containing an opening for the internal gland. Front and hind wings differently shaped: front wings more or less in evenly shaped bands that are truncated under apex, whereas hind wings broader and rounded. Venation complete with usually all forks present: I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. Discoidal and median cells always closed in front wings. Very long thyridial cell. In hind wings, Sc ends at R1. Discoidal and median cells open or closed. There are 4 free anal veins extending to wing margin. % genitalia very similar in all genera. Segment IX generally short but well developed dorsally. Segment X developed into large roof-shaped mass above phallic apparatus and forming various lobes in shapes characteristic of species. Preanal appendages always present but completely integrated with segment X and reduced to simple, poorly differentiated tubercles bearing a cluster of short setae. Intermediate appendages always absent. Inferior appendages strongly slanting upward, consisting of 2 simple, slender segments curving inward and forming thin 2-segmented caliper-like structure. Large phallic apparatus, bulky at base, and consisting exclusively or almost exclusively of phallotheca. Endotheca absent or tiny, simple or complex. Aedeagus and parameres absent. & genitalia also very homogeneous. Sternite VIII more or less completely separated into 2 large lobes by apicoventral cleft. Segment IX absent. Segment X large and appearing triangular in side view. It extends far down on sides, producing sclerotized strip along margins of its ventral side, which forms membranous vulvar scale. Dorsal end of segment X with brushes of stout setae. Segment XI basically in shape of long oblique rectangle with short setae and 2 small, rounded, papillary lobes framing cerci that have generally kept vestigial 2nd segment. Vulvar scale large, elongated, mostly membranous, hinged at base to lower angles of segment X. It can generally fold down, uncovering a gaping anovaginal opening 51

under the mass of segment X. Anovaginal vestibule short, very high, with a ceiling forming complex folds (the “median plate" described by some American authors). Small, simple vaginal apparatus located at end of vaginal chamber. The Hydropsychidae constitute a large cosmopolitan family. They are one of the main families of trichopterous insects in our country with respect to the number of species and the profusion of individuals found locally. They occur in all types of low-altitude lotic environments. They are grouped into three subfamilies, all represented in Canada. 1a Discoidal and median cells in front wings small. Discoidal cell in hind wings open (Fig. 139). Front wings brown with strong yellow stripes. Antennae very long and thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oestropsinae, p. 52 1b Abdominal sternite V with filament (Figs. 164, 165). Hind wings rather broadly rounded (Fig. 166). Antennae slightly crenulated (Fig. 163) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplectroninae, p. 57 1c Abdominal sternite V with at most one lobe or one blunt protuberance (Figs. 146, 147). Hind wings narrower at apex (Figs. 148, 154). Antennae thin and cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . Hydropsychinae, p. 53

Subfamily Oestropsinae Brauer Oestropsidae Brauer, 1868, p. 264 (as a family) Type genus: Oestropsis Brauer This subfamily is represented in Canada by a single genus. Genus Macrostemum Kolenati

Macrostemum Kolenati, 1859, p. 168 Type species designated by Ulmer 1957: Hydropsyche hyalinata Pictet Revision: Nimmo, 1987a Fairly distinct secondary sexual dimorphism. Globular head with bulging face and large malar space. Very fine antennae, 1.5 times as long as front wings in % and 1.3 times as long in &. Anterior cephalic warts very large in % (Fig. 138) and slightly smaller in &. Maxillary palpi with first 2 segments short (Fig. 137). Flagellum 1.3 times longer in % than in &. Middle legs of & markedly flattened. Hind tibiae of % with very long hairs. Sternite V with a finger-shaped lobe in % and a tiny protuberance in &. Abdominal hemogill system consisting of few simple, thick tubes. 52

Wings faintly pubescent. The 2 pairs of wings fairly different in shape. Front wings forming long, slender ellipsoid, and hind wings a triangle. Hind wings 1.5 times wider than front wings in %; in &, they are distinctly narrower than in males and have a costal angle. Frenulum consisting of a row of very regular hooks. Venation (Fig. 139) somewhat irregular and slightly incomplete in hind wings. In front wings, Sc joins R1 slightly before it ends. Forks I, II, III, IV, and V present, the 1st fork petiolate. Discoidal and median cells small, thyridial cell very long. Postcostal cell rather wide. A1 very long. In hind wings, only forks II, III, and V present. R1 and R2+3 merge with Sc. Open discoidal cell. M separates from base of RS. Intercubital cell containing a fold extending Cu1b to wing base. % genitalia (Figs. 140, 141): segment IX very elongate laterally, but short dorsally and ventrally. Segment X forming 2 winglike lobes broadened posteriorly, with inner basal portion membranous. Inferior appendages long, evenly slender, and consisting of 2 subequal segments. Phallic apparatus fairly thick at base, swelling into pyriform structure at tip, which is simple. & genitalia (Figs. 142, 143): sternite VIII split at apicoventral edge forming 1 median and 2 lateral lobes. Segment X appears triangular in side view. It is extended on underside by 2 strong ribs that keep vulvar scale closed. Segment XI not very developed. Two papillary lobes slender and far apart, with space between filled with small membranous tubercles. Small, 2-jointed cerci. The vaginal apparatus almost completely membranous. The genus Macrostemum is large, almost cosmopolitan, and with strong tropical affinities. There are three Nearctic species of which only one, zebratum Hagen, lives in Canada where it is widespread in the eastern part of the country and further south. It inhabits all watercourses of some size but is found in greatest profusion in the larger rivers. Adults are brightly colored and easy to identify; they fly in large numbers at dawn and dusk, but are occasionally seen in broad daylight.

Subfamily Hydropsychinae Ulmer Fine antennae slightly longer than front wings, especially in %. Maxillary palpi with 2nd segment clearly longer than 1st; 3rd and 4th shorter (Fig. 144). Top of head with more than 2 pairs of warts. Middle legs of & flattened. Abdominal hemogills consisting of elongated bulbs bearing simple, apicolateral tubes. Front wings in shape of fairly even strips with truncated apex. Hind wings slightly wider and regularly rounded at the posterior margin, but 53

narrow at tips. Venation complete with forks I, II, III, IV, and V present in front wings and I, II, III, and V or II, III, and V in hind wings. In front wings, discoidal and median cells fairly small. FI and FIII petiolate. A converges with Cu2 at tip. In hind wings, discoidal cell closed and median cell open or closed (Figs. 148, 154). % genitalia: segment IX fairly short, forming more or less protuberant apicolateral angle. Segment X occasionally a single mass, at times bilobed or forming points. Inferior appendages slender and 2-jointed. Phallic apparatus simple or complex at apex. & genitalia: segment X forms only narrow, barely sclerotized lower strip reinforcing vulvar scale. Ceiling of phallocrypt with complex sclerotized folds. The Hydropsychinae contain most of the species of the family. The subfamily is represented in Canada by only three genera. 1a In hind wings, trunks of M and Cu1 parallel and very close together. FI present (Figs. 148, 154) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b In hind wings, trunks of M and Cu1 not parallel and not so close together; FI absent (Fig. 157) . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheumatopsyche, p. 56 2a In front wings, crossveins M3+4-Cu1 and Cu1-Cu2 close together. Median cell in hind wings open (Fig. 154) . . . . . . . . . . . . . . . . . . Potamyia, p. 55 2b In front wings, crossveins M3+4-Cu1 and Cu1-Cu2 far apart. Median cell in hind wings closed (Fig. 148). . . . . . . . . . . . . . . . . . . . . Hydropsyche, p. 54

Genus Hydropsyche Pictet

Hydropsyche Pictet, 1834, p. 199 Type species designated by Ross, 1944: Hydropsyche cinerea Pictet = instabilis Curtis Ceratopsyche Ross and Unzicker, 1977, p. 208 Type species by original designation: Hydropsyche bronta Ross Revision: Nimmo, 1987a In %, claws on 3 legs equal in size but asymmetrically twisted and concealed in tuft of thick black bristles that are more developed on inner side (Fig. 145). Spurs: 2,4,4. Sternite V in % with more or less developed lobe, depending on species. It contains an opening for large internal gland with double caecum (Fig. 146). In &, this gland is tiny and spherical with opening on small protuberance (Fig. 147). Venation (Fig. 148): in front wings, crossveins M3+4-Cu1 and Cu1Cu2 far apart. In hind wings, FI present and petiolate, median cell closed, and trunks of M and Cu1 parallel and very close together. 54

% genitalia (Figs. 149–151): segment IX forming prominent apicolateral angle provided with long setae. Segment X simple or complex, massive or forming lobes or points characteristic of species. Inferior appendages with 2nd segment stout and cone-shaped. Phallic apparatus slightly curved at base, with tip always bilobed, simple or complex. It comprises a phallotheca with a short cleft at tip in simulans species group. In instabilis species group, the phallotheca not cleft but with short erectile endotheca forming more or less complex lobes and equipped with spiny framework which is also variously complex. These characters are of no generic significance but apply simply to groups of species. & genitalia (Figs. 152, 153): tergite VIII notched in middle of apex, occasionally with slightly curved apicolateral angles. Sternite VIII divided into 2 large lobes over only half its length. Segment X forming blunt, seta-bearing lateral angle. A cavity or depression, characteristic in shape of species, located in middle of the lateral sides for insertion of tip of inferior appendages of % during mating. The genus Hydropsyche is by far the most homogenous of the order Trichoptera. Dividing it would be pure taxonomic inflation. Its components are mere species groups. The genus Hydropsyche is found worldwide except in Neotropical America. Some 80 Nearctic species are known, half of which exist in our area. Larvae inhabit all types of running water and also large lakes. Some species, such as recurvata Banks, have a clear preference for the major watercourses where they may occur in such profusion that they become a serious nuisance, specially at night around lights.

Genus Potamyia Banks

Potamyia Banks, 1900, p. 259 Type species by original designation: Macronema flavum Hagen Revision: Nimmo, 1987a This genus is fairly unusual since it presents a blend of characters found in the subfamily Macronematinae and the genera Hydropsyche and Cheumatopsyche. Like the Macronematinae, it has a strong sexual dimorphism, a globular head with a bulging face and wide malar space. Antennae very long and fine, especially in %, with globular 1st segment. Front wings with very few hairs and hind wings broad, especially in %, and fairly pointed at tips. Sc and R1 in front wings strong, and discoidal and median cells convex. Cu2 and A converge before reaching wing margin, and R1 in hind wings merges with Sc before the wing edge (Fig. 154). 55

Like Hydropsyche, % claws of 3 pairs of legs asymmetrical and concealed with thick, black hairs. Hemogill system large and arborescent, FI present in hind wings, and trunks of M and Cu1 parallel and very close together. Also, sternite VIII in & divided over only half its length. Like Cheumatopsyche, Potamyia displays reduction in size of gland of sternite V and protuberance on this sternite. Crossveins M3+4-Cu1 and Cu1-Cu2 very close together and genitalia in both sexes similar in type, except for two characters. Potamyia has only a few exclusive characters. They are: fairly strongly spined front tarsi in %, absence of crossveins Sc-R1 and R1–R2+3 in front wings, spurs: 0,4,4 in the % and 1,4,4 in &, segment X of % thin, bilobed at the extremity, without a median tonguelike structure (Figs. 155, 156). The genus Potamyia is very small and Siberian and Nearctic in distribution, with only one species in North America: flava Hagen. This species is very widely distributed and found in southern Ontario and Manitoba. It shows a definite preference for major watercourses and occasionally occurs in large enough numbers to be a nuisance. Genus Cheumatopsyche Wallengren

Cheumatopsyche Wallengren, 1891, p. 142 Monobasic type species: Hydropsyche lepida Pictet Revisions: Gordon, 1974; Nimmo, 1987a This genus is very similar to Hydropsyche and differs only in a small number of characters. The insects are smaller and slenderer in build. Claws in % variously deformed in some species, but not in others. Internal gland of sternite V and protuberance similar in both sexes and identical to those of female Hydropsyche. Venation (Fig. 157): in front wings, crossveins M3+4-Cu1 and Cu1Cu2 close together. In hind wings, FI absent, median cell open, and M and Cu1 stems neither parallel nor close together. % genitalia (Figs. 158–160) very similar to those of Hydropsyche, but less robust. Apicolateral angle of segment IX less prominent and positioned lower. Segment X not so massive, ending in 2 or 4 seta-bearing lateral lobes, characteristic in shape of species, and separated by barely protuberant median tonguelike structure. Inferior appendages slender, with 2nd article often hook-shaped. Phallic apparatus very thick at base, always simple and ending in 2 endothecal papillae. & genitalia (Fig. 161, 162): tergite VIII not notched at the tip. Sternite VIII divided over its entire length into 2 hairy separate valves. Segment X 56

similar to that of Hydropsyche, but apicolateral angle indistinct and lateral concavity less consistently present. The genus Cheumatopsyche is also virtually cosmopolitan. It has strong African affinities and is absent from Neotropical America. There are 40 Nearctic species, half of which are found in our area. The species inhabit all types of running water, even some polluted rivers, and at times exist in such profusion in large rivers that some, like speciosa Banks and campyla Ross, can constitute a serious nuisance.

Subfamily Diplectroninae Ulmer Diplectroninae Ulmer, 1951, p. 303 Type genus: Diplectrona Westwood This subfamily is represented in our area by two genera. 1a Front wings with crossveins M3+4-Cu1 and Cu1-Cu2 close together. Hind wings broadly rounded, with Sc and R1 markedly curved upward at their ends (Fig. 166). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplectrona, p. 57 1b Front wings with those crossveins not close together. Hind wings less broadly rounded, with Sc and R1 barely curved upward at their ends (Fig. 171). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homoplectra, p. 58

Genus Diplectrona Westwood

Diplectrona Westwood, 1840, p. 49 Type species by original designation: Aphelocheira flavomaculata Stephens nec Pictet = Diplectrona felix McLachlan Revision: Nimmo, 1987a Antennae as long as front wings, fairly fine, but slightly crenulated. Segments with length far exceeding thickness and bulging in middle. Basal segments with a dark transverse groove (Fig. 163). Maxillary palpi with 2nd segment much longer than 1st and 3rd and 4th segments progressively shorter. Vertex with 4 large warts. Middle legs of & not flattened. Abdominal sternite V with bulb prolonged, forming very long slender tube slightly longer in % than in &. Within the segment, there is a gland, large and oval in % and smaller and spherical in &. It discharges through opening at tip of basal bulb (Figs. 164, 165). Abdominal hemogill system consisting of small basal bulbs with tubes that fork repeatedly, producing an arborescent complex. Front wings fairly broad at level of anastomosis. Hind wings wide, rounded and blunt at tip. Venation (Fig. 166): in front wings, FI and FIII 57

petiolate. Discoidal cell small, median and thyridial cells large. Cu2 and A strongly curved, which considerably widens postcostal cell. Crossveins M3+4-Cu1 and Cu1-Cu2 close together. In hind wings, Sc very thick, R1 very thin, and both sinuate with the result that subcostal cell is very narrow at base and wider at tip. Discoidal cell narrow, FI and FIII petiolate. % genitalia (Figs. 167, 168): segment IX without any well-defined apicolateral angle. Segment X barely distinct from segment IX and forming 2 outer and 2 inner lobes. Inferior appendages with very long basal segment. Phallic apparatus fairly evenly thick, ending in 2 endothecal papillae. & genitalia (Figs. 169, 170): sternite VIII completely cleft ventrally, forming 2 separate valves. Segment X without any lateral cavity, extending far down on sides and forming stout sclerotized rib that slants upward over vulvar scale. Ceiling of anovaginal cavity and vaginal tract simple. The genus Diplectrona is widespread in the Old World, except in Africa, and displays a clear Oriental affinity. It has only three Nearctic species, only one of which has been reported in Canada. D. modesta Banks has a wide distribution across the eastern part of the continent, from Florida to Newfoundland and Ontario. It is found only in springs and small, clear, fast-flowing rivers and streams. Adults emerge in the spring but never in large numbers in any one place. Genus Homoplectra Ross

Homoplectra Ross, 1938, p. 119 Type species by original designation: Homoplectra alseae Ross Aphropsyche Ross, 1941, p. 78 Type species by original designation: Aphropsyche aprilis Ross = Diplectrona doringa Milne Revision: Nimmo, 1987a Eyes small, situated rather anteriorly. Antennae moderately stout, with 2nd segment only half length of scape. Vertex convex, with posterior warts large. Filament and inner gland of sternite V not as developed as in Diplectrona. Both wings of similar and rather oval shape. Venation (Fig. 171): in front wings, crossveins M3+4-Cu1 and Cu1-Cu2 not close together. In hind wings, Sc and R1 barely curved at their ends. Stems of M and Cu not parallel and not so close together. % genitalia (Figs. 172–175): segment IX not very developed, reduced ventrally, and with a ventral apical extension. Segment X very large, not distinct from segment IX at its base, and forming an apical upward-directed 58

hook. Inferior appendages very long and thin; 2nd article entirely fused with 1st. Phallic apparatus secondarily much complicated by very strong development of several sclerotized branches, whose identity cannot be assigned. & genitalia (Figs. 176, 177): sternite VIII almost completely cleft mesally. Segment X without lateral clasper receptacle. Segment XI with welldeveloped bi-articulated cerci. The genus Homoplectra is easily recognized by the complexity of the phallic apparatus. It contains 10 Nearctic species, mostly located in Oregon and California. None has been recorded from our country, but doringa Milne has a large eastern distribution and has been found in New Hampshire. Its biology is unknown.

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Family Polycentropodidae Ulmer Polycentropodidae Ulmer, 1906, p. 83 Type genus: Polycentropus Curtis Small, robust, heavy-bodied, very hairy insects with wings generally densely mottled with gold. Ocelli absent. Top of head with large warts and clothed with particularly abundant, erect, silky hairs. Antennae thick and short-segmented. Maxillary palpi 5-segmented in both sexes, with the 2 basal segments usually short and subequal and the 5th forming a long flagellum. Third segment always inserted slightly before apex of 2nd, which is spiny (Figs. 185, 199, 205). Spurs: 3,4,4 or 2,4,4, very hairy. Middle legs in & flattened. Abdominal hemogill system comprising 2 simple erectile tubes or 1 lobe and 1 bulb (Fig. 186). I observed the system in all the genera, with the exception of Paranyctiophylax, where it probably escaped my notice. Gland of abdominal sternite V fairly large and opening at the base of a filiform, generally well-developed lobe (Figs. 178, 189). Wings with a densely hairy membrane, fairly wide, with both pairs fairly similar in shape; elliptical tip. Venation complete with all forks present: I, II, III, IV, and V on the front wings and I, II, III, and V on the hind wings. A slight tendency toward simplification. Thus, FI and FIII may be missing from both wings and FI is always petiolate. In front wings, crossveins Sc-R1 and R1–R2 are generally present. Discoidal, median, and thyridial cells long and closed. Thyridial cell always with point of contact with median cell. Discoidal cell constantly begins basal to median cell. In hind wings, R1 may be variably reduced and converges with Sc. Discoidal cell open or closed. Four or 2 free anal veins. % genitalia robust, massive, and not very homogeneous within the series of genera. Segment IX well developed ventrally but absent or virtually so dorsally. Segment X forms membranous lobe and is rarely partially sclerotized. Preanal appendages present, ear-shaped, more or less elongated, but very wide at base. Intermediate appendages always present, generally in shape of spines or spurs, occasionally vestigial but always fused at extreme base with preanal appendages. Lower inner angles of resulting structure extend medially into complete or incomplete bridge under phallic apparatus. Inferior appendages single-segmented, more or less complex in shape, subhorizontal, and never fused to each other. Phallic apparatus large, located very high in abdomen, between preanal appendages. 60

It comprises a phallotheca, endotheca, and aedeagus, but is always deeply, although occasionally not obviously, modified in some way through reduction of one of its parts. Parameres always absent. & genitalia, on the other hand, are very homogeneous within the family. Sternite VIII completely divided into 2 lobes in shape of free, widely spaced valves. Segment IX absent. Segment X large with ventral side forming large vulvar scale closing anovaginal cavity underneath. Segment XI with apical angles stretched into smooth papillae framing cerci, which they resemble. Anovaginal opening under segment X with ceiling never complex. Vaginal apparatus more or less complicated and may be source of useful specific characters. Unfortunately, they have not yet been studied. In Europe, the family Polycentropodidae has been considered distinct from the Psychomyiidae since its creation in 1906. In America, Ross (1944) reduced the two families to a single one on artificial grounds, and other authors unfortunately followed suit. Later Ross (1967) fortunately restored the Polycentropodidae to family status. This is the approach I have adopted here as both families can immediately be distinguished from each other by the maxillary palpi, the venation and genital systems of both sexes, irrespective of the genus under consideration. The family Polycentropodidae is medium-sized, cosmopolitan in distribution, and found in all types of lentic and lotic environments, provided the water is not too turbulent. It is divided into two subfamilies: the Polycentropodinae and the Pseudoneureclipsinae. Only the former is represented in Canada. 1a Spurs: 2,4,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cernotina, p. 65 1b Spurs: 3,4,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a FIII present in hind wings (Fig. 179) . . . . . . . . . . . . . . . . . Neureclipsis, p. 61 2b FIII absent from hind wings (Fig. 187) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a FI present in front wings and at times in hind wings (Fig. 187) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polycentropus, p. 62 3b FI absent from both pairs of wings (Fig. 200). . . . . . . . . . . . . . . . . . . . . . . . . 4 4a Maxillary palpi with 2nd segment 3 times shorter than 3rd (Fig. 185) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paranyctiophylax, p. 64 4b Maxillary palpi with 2nd segment barely shorter than 3rd (Fig. 199) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyrnellus, p. 65

Genus Neureclipsis McLachlan

Neureclipsis McLachlan, 1864, p. 30 Monobasic type species: Phryganea bimaculata Linnaeus 61

Revisions: Nimmo, 1986; Armitage and Hamilton, 1990 Maxillary palpi with first 2 segments very short. Spurs: 3,4,4. Venation (Fig. 179) complete and with little modification; all forks present: I, II, III, IV, and V in front wings and I, II, III, and V in hind wings. In front wings, crossvein C-Sc absent. In hind wings, discoidal cell closed and R1 somewhat reduced, clearly separate from Sc and joined to latter by distinct crossvein. % genitalia (Figs. 180–182): segment IX strongly elongated laterally. Segment X stout, often very long and semimembranous. Preanal appendages free or fused with base of intermediate appendages or absent. Intermediate appendages in long spines or reduced to knob. Lower inner angles of these appendages form continuous bridge under phallic apparatus. Inferior appendages very large, subhorizontal, and calliper-like with curved parts. Phallic apparatus very large, comprising long phallotheca with wide opening underneath at base and membranous endotheca at tip. Endotheca fairly complex and obscurely structured. Aedeagus very small or absent. & genitalia (Figs. 183, 184) with segment X small and lobes of sternite VIII large, well developed, and unconnected. Vulvar scale semimembranous and occasionally carinate. Vaginal apparatus simple and mainly membranous. The genus Neureclipsis is small with nine species. It is Holarctic in distribution. It includes five Nearctic species, only three of which are represented in Canada: bimaculata Linnaeus (Holarctic) which has small preanal appendages fused with the base of long intermediate appendages, valida Walker (Manitoba, Ontario, and Quebec) in which the preanal appendages have disappeared and the intermediate appendages are spiniform and inserted very anteriorly, and crepuscularis Walker (from Newfoundland to Manitoba), in which the preanal appendages are free and the intermediate appendages reduced to small knobs. The genus Neureclipsis is very widely distributed in Nearctic America. Species occur in several biotopes but have an affinity for large rivers and lentic habitats and fly during almost the entire warm season.

Genus Polycentropus Curtis

Polycentropus Curtis, 1835, pl. 544 Type species by original designation: Polycentropus irroratus Curtis = Hydropsyche flavomaculata Pictet Revisions: Nimmo, 1986; Armitage and Hamilton, 1990 62

Maxillary palpi with first 2 segments very short (Fig. 185). Spurs: 3,4,4. Venation (Figs. 187, 188) slightly simplified and somewhat modified, with forks I, II, III, IV, and V in front wings and I, II, and V or II and V in hind wings. In front wings, crossveins C-Sc, Sc-R1, and R1–R2 present. Hind wings with slight sexual dimorphism: wings slightly wider in % than in & and apical veins are more divergent. Subcostal cell wide, Sc thickened, R1 and M very thin. Sc and R1 join then sharply diverge. F1 present or absent, and discoidal cell open or closed. There is occasionally a crossvein between Cu2 and A1. % genitalia (Fig. 190) vary considerably between species groups. Segment IX always stout laterally and ventrally and at times forming apicoventral plate (aureolus group). Segment X small, membranous, shapeless, occasionally virtually unsclerotized (remotus group) and, rarely, slightly sclerotized (confusus group). Preanal appendages generally small and free, but at times fused with base of the intermediate appendages (confusus group). Intermediate appendages always present, occasionally small (cinereus group), at times curved broadly downward (interruptus group), and at times divided into 4 long spines (variegatus group). Inner apical angles of intermediate appendages drawn inward, forming an open or closed bridge under phallic apparatus and occasionally bearing a downward-curving spur (remotus group). Inferior appendages medium-sized and always in some way bilobed. Phallic apparatus also highly variable in structure. In interruptus group, it comprises a short phallotheca, an almost vestigial endotheca, and a long, curved aedeagus (Fig. 190). In confusus group, aedeagus lost and endotheca well-developed and spine-bearing. In other groups, tip of phallic apparatus complex and obscurely structured. & genitalia (Figs. 191, 192): segment X small. Ventral lobes of sternite VIII far apart and unconnected. Vulvar scale particularly large and at times carinate. Ceiling of anovaginal opening occasionally with sclerite. Vaginal apparatus large, simple or complex. The genus Polycentropus is fairly large and distributed almost worldwide. It is especially well represented in the Holarctic region. It is fairly heterogeneous. In Europe it is considered distinct from two other genera, i.e., Plectrocnemia Stephens in which the discoidal cell on the hind wings is closed and Holocentropus McLachlan in which FI is absent from the hind wings. Probably not without reason, American authors have included these genera in Polycentropus, but subsequent studies will show whether they constitute separate lineages and should be given generic status. The genus comprises 45 Nearctic species, 33 of which live in our area, in all provinces. They are particularly numerous in the eastern portion of the country and quite a few species are transcontinental. They 63

inhabit all lotic biotopes, and are found, although less often, in still water and occasionally temporary ponds. Genus Paranyctiophylax Tsuda

Paranyctiophylax Tsuda, 1942, p. 265 Type species by original designation: Paranyctiophylax kisoensis Tsuda Revisions: Nimmo, 1986; Armitage and Hamilton, 1990; Neboiss, 1993 Maxillary palpi with 2 basal segments very short. Spurs: 3,4,4. Venation (Fig. 193) slightly simplified through absence of FI from both wings and of FIII from hind wings. In front wings, particularly long discoidal cell. Three anal veins forming 2 markedly convex loops without any crossvein between them and ending at virtually the same point. In hind wings, Sc slightly thickened, R1 very thin and joining Sc for short distance. Particularly short and triangular discoidal cell. % genitalia very homogeneous (Figs. 194–196): segment IX robust, but only half as high as the abdomen. Segment X small, membranous, but always more or less partially sclerotized. Preanal appendages very wide, but protruding little, massively shaped, and slightly notched. Intermediate appendages forming downward-curving spurs arising from lower inner angles of preanal appendages that meet in bridge under phallic apparatus. Inferior appendages not very prominent, flattened when viewed caudally, forming basal heel and 2 apical lobes. Phallic apparatus quite unusual in its development. Phallotheca reduced and faintly sclerotized. Endotheca long, cylindrical, and provided at base with 2 long spines that are not parameres; these disappeared at early stage in Hydropsychoidea. Tiny, membranous aedeagus containing large phallotremal sclerite. Lower apical angle of endotheca forms paired or unpaired lobe endowed with prodigious erectile power. It has number of folds and is equipped with spines on lateral sides and underneath. Aedeagus seems to arise from point halfway along endotheca. & genitalia (Figs. 197, 198): ventral lobes of sternite VIII small, far apart, but joined together by body of sternite from which they have separated. Segment X simple, very high, with median margins on underside close together, providing only narrow anovaginal opening that is closed from below by small vulvar scale. Segment XI short and high. The genus Paranyctiophylax is small, very homogeneous, and found worldwide, except in Europe. It has tropical affinities. It comprises nine Nearctic species, five of which have been reported in Canada. They live primarily in the eastern portion of the country and inhabit virtually all still water and calm sections of running water. Species are very similar in 64

appearance and can be distinguished primarily by the development of the phallic apparatus, which should be examined when turgescent (Fig. 196). Our species were placed in Nyctiophylax Brauer until recently. Genus Cyrnellus Banks

Cyrnellus Banks, 1913, p. 88 Type species by original designation: Cyrnellus minimus Banks Revisions: Nimmo, 1986; Armitage and Hamilton, 1990 Small, very heavy-bodied insects with a very hairy body and wings. Maxillary palpi with 2nd segment almost as long as 3rd. Latter arises before tip of 2nd segment, as in all genera of family (Fig. 199). Spurs: 3,4,4. Hind wings not wider than front wings. Venation (Fig. 200) slightly simplified and unmodified. In front wings, FI absent and median cell open. In hind wings, veins in middle of wing thinned. R1 short, ending early at Sc; FI and FIII absent and discoidal cell open. % genitalia (Figs. 201, 202): ventrally, segment IX extends far forward. Segment X large, broadly flared posteriorly, pilose, and moderately sclerotized. Preanal appendages in long ovals. Intermediate appendages small, faintly sclerotized, and projected downward. Basal portion forming 2 paired triangular lobes under phallic apparatus. Inferior appendages large, simple, robust, and equipped with strong cone-shaped protuberance on inner side. Phallic apparatus simple, short, thick, comprising phallotheca and simple, spine-free endotheca with large phallotremal sclerite. & genitalia fairly elongate (Figs. 203, 204): sternite VIII visible between its large lateral lobes. Segment X long with base concealed behind sternite lobes. Segment XI vertical. Anovaginal opening, a long narrow slit, half-closed by long, bifid vulvar scale. Vaginal apparatus very simple. The genus Cyrnellus is small and very homogeneous, with a wide distribution in Neotropical America. A single species, fraternus Banks, is widely distributed in the Amazon River basin, throughout Central America, and the United States. It has not been captured in Canada, but has been, reported in Minnesota. It has a preference for large rivers but is occasionally encountered near small rivers, lakes, and reservoirs. Genus Cernotina Ross

Cernotina Ross, 1938, p. 136 Type species by original designation: Cernotina calcea Ross Revisions: Nimmo, 1986; Armitage and Hamilton, 1990. 65

Maxillary palpi with first 2 segments very short. Spurs: 2,4,4. Wings narrow and elongate with hind wings slightly narrower than front wings, but blunt-ended. Venation quite considerably simplified (Fig. 206). In front wings, crossvein C-Sc absent, Sc short, FI and FIII absent. Discoidal cell short and median cell open. In hind wings, Sc and Cu1 remarkably thickened and R1 joins Sc at very early stage. FI and FIII absent, discoidal cell open. Only 2 anal veins. % genitalia (Fig. 207) with considerable variation in development of various appendages. Segment IX triangular and low. Segment X arises from upper angles of segment IX and is moderately sclerotized, bifid, and hairy. Preanal appendages very elongate, ending in long, calliper-shaped ribs, usually equipped with inner teeth and spines. Intermediate appendages probably lost or integrated into inner portion of preanal appendages. Base of latter structures in shape of 2 faintly sclerotized, hairy lobes that extend under phallic apparatus but without forming continuous bridge. Inferior appendages large and complex with stout upper branch. Phallic apparatus simple and slender and consists of phallotheca, that is reinforced along its upper basal portion, and endotheca containing oval phallotremal sclerite. & genitalia fairly elongate (Figs. 208, 209): sternite VIII reduced, with 2 slender lateral lobes and still visible between the two. Segment X elongate, notched dorsally, and with concealed anterobasal angles. Segment XI large, slanting downward and forward. Anovaginal opening generally forming slit with closed labia. Vulvar scale lost. Vaginal apparatus long and clearly sclerotized. The genus Cernotina is large and particularly well represented in Neotropical America. Twelve species occur in the United States but only two of these, pallida Banks and spicata Ross, have been captured in southern Ontario.

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Family Hyalopsychidae Lestage

Hyalopsychinae Lestage, 1925, p. 97 Type genus: Hyalopsyche Ulmer In the first edition of this handbook (Schmid, 1980), I elevated the long forgotten subfamily Hyalopsychinae to family level, for the inclusion of the two closely related genera Hyalopsyche Ulmer (African, Oriental, and Australian regions) and Phylocentropus Banks (Baltic amber, Oriental, and Nearctic regions). In 1973, Ross and Gibbs recognized some interesting larval synapomorphies between Phylocentropus and Dipseudopsis Walker and suggested the inclusion of Phylocentropus in the Dipseudopsinae, then a subfamily of the Polycentropodidae. More recently, Wells and Cartwright (1993), discovered interesting new larval and female synapomorphies between Hyalopsyche and Phylocentropus. They showed that these two genera are closely related, what was already known, and suggested also that the Hyalopsychidae should merge with the Dipseudopsidae. The discoveries of these authors are interesting, but their suggestion is not acceptable (see also Weaver and Malicky, 1994). The estimation of the degree of relationship of two different lineages should not rest only on their common characters, but also take into consideration the weight and amplitude of their differences. Careful evaluation should be made of this evidence. The adult characters of the Hyalopsychidae and the Dipseudopsidae are so different, especially their strikingly dissimilar habitus and size, that each of these two lineages surely deserves to retain its family status. Hyalopsychidae and Dipseudopsidae certainly are sister families, having a common origin, but sorority is not identity. These two families are not related to Polycentropodidae, as usually thought, but to Ecnomidae, Psychomyiidae, and Xiphocentronidae. The Hyalopsychidae are represented in Canada by only a single genus.

Genus Phylocentropus Banks

Phylocentropus Banks, 1907, p. 130 Type species by original designation: Holocentropus placidus Banks 67

Acrocentropus Betten et al., 1934, p. 213 Monobasic type species: Polycentropus lucidus Hagen Revisions: Schmid, 1983; Schuster and Hamilton, 1984 Eyes markedly globular and protuberant, especially in % (Fig. 210). Antennae longer than front wings, clearly thickened, especially in %, and consisting of very short segments, each with cross furrow. Maxillary palpi reduced in size, with first 2 segments short and the 5th not very long. Labial palpi considerably reduced (Fig. 211). Spurs: 3,4,4. Abdominal hemogill system present and poorly developed (Fig. 212). Both pairs of wings elliptical and slightly narrower in & than in %. Venation complete (Fig. 213), with all forks present on both pairs of wings and all sessile except FIII. Thyridial cell long, in contact at one point with median cell. Crossvein C-Sc present. Four free anal veins on hind wings. % genitalia (Figs. 214–217): segment IX consists of tergite and sternite, both well developed but narrow laterally where they hinge together as in Psychomyiidae. Segment X fairly large and membranous. Preanal appendages resembling oval ears. Intermediate appendages greatly reduced, free or fused with segment X or each other. Two lobes at base of appendages form bridge under phallic apparatus. Inferior appendages small, single-segmented, simple or complex, with numerous spines on inner surface and not fused together. Phallic apparatus small and consisting of phallotheca and endotheca. Tip of endotheca complex and obscurely structured. Aedeagus probably absent. & genitalia (Figs. 218, 219): sternite VIII without lateral lobes but developed into concave tonguelike structure protecting segment X from below. Segment IX absent. Segment X large and massive, forming sharply pointed vulvar scale that is loosely joined to but articulated with segment. Segment XI small, with lower apical angles curving upward and 2 papillae framing well-developed cerci. Ceiling of anovaginal cavity and vaginal apparatus simple. The genus Phylocentropus is small, Oriental and Nearctic in distribution, with five North American species, three of which are represented in Canada: lucidus Hagen in which the intermediate appendages are fused together (Fig. 217), placidus Banks in which the intermediate appendages are fused to segment X (Fig. 216), and carolinus Carpenter in which the preanal appendages are free and thin. The first two species have a large distribution from Nova Scotia to Ontario and Manitoba; the third is recorded from Quebec and Ontario. The larvae burrow in the silt of sandy rivers and lakes. The genus likely originated in the Oriental region (Schmid 1989, p. 110). 68

Family Psychomyiidae Curtis

Psychomidae Curtis, 1835, pl. 561 Type genus: Psychomyia Latreille Fairly slender, medium-sized or small insects, not very pubescent, with front wings uniformly brown. Ocelli absent. Maxillary palpi 5-segmented in both sexes with first 2 segments variable in length and 3rd never inserted before apex of 2nd which is spine-free. Fifth segment flagellate. Slender antennae with fairly long segments. Spurs: 2,4,4. Middle legs in & flattened, except in Tinodes. Abdominal hemogill system absent. Sternite V with or without tiny internal gland that does not open onto filiform lobe. Wings long and narrow with hind wings slightly reduced and lanceolate. Venation slightly simplified in front wings and greatly reduced in hind wings. In front wings, FI absent, discoidal cell short, and thyridial cell very small, situated at extreme base of wing and without any contact with median cell. The latter long and always begins ahead of discoidal cell. In hind wings, Sc or R1 shortened, FI absent, and discoidal cell open; FIII occasionally absent. One or 2 free anal veins. % genitalia generally slender, simple or complex, but with certain constant basic characters that allow them to be unequivocally differentiated from those of Polycentropodidae. Segment IX secondarily divided into a tergite and sternite. Tergite well developed dorsally, narrowing laterally toward bottom. Symmetrically, sternite well developed ventrally, narrowing laterally toward top. Tergite hinged to sternite, and preanal and intermediate appendages primitively inserted on resulting angle. Segment X small and membranous, except in Lype. Preanal appendages always considerably extended, especially in Tinodes. Intermediate appendages well developed in Tinodes and lost in Lype and Psychomyia. They form a complete bridge under phallic complex in Tinodes. In Psychomyia and Lype in which intermediate appendages are lost, this bridge absent in former genus, but partially reformed in latter by lower margins of segment X. Inferior appendages fairly stout, subhorizontal, and 2-segmented. First segments stout and partially fused together at their base. Second segments more or less simple and somewhat reduced. Phallic apparatus located very high, under segment X, and comprising long phallotheca, obliterated or vestigial endotheca, and tiny, upwardly curved aedeagus. 69

& genitalia extended to variable degree into ovipositor as in Rhyacophilidae, but is differently structured. In Psychomyiidae, last 2 segments are distinctly more specialized, rigid, and nonretractile. Intersegmental membranes poorly developed and apodemal rods absent. Elongation is produced by segments X and XI, which gradually become attenuate. Sternite VIII without ventrolateral lobes. Since these lobes have become increasingly separated in Arctopsychidae–Hydropsychidae–P olycentropodidae series, their absence can only be interpreted as a loss, i.e., a specialization. Segment IX absent. Segment X strongly elongated. Segment XI ovoid, with cerci always present, but without papillae to frame them. Anovaginal opening remarkably long and narrow, forming thin slit along entire length of segment X. Vaginal apparatus complex and largely membranous. The Psychomyiidae constitute a fairly small family that is cosmopolitan in distribution. It is found in all lotic and lentic habitats. It is divided into two subfamilies: the Psychomyiinae and the Paduniellinae, both of which are represented in North America. Only the former, however, occurs in Canada. The family most likely originated in the Oriental region, in which the most primitive genus occurs (Schmid 1989, p. 110). 1a In hind wings, Sc very short and R1 long, extending to wing margin (Fig. 220) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lype, p. 70 1b In hind wings, Sc long or absent and R1 extending to Sc or wing margin (Figs. 224, 231) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Third segment of maxillary palpi longer than 2nd. Hind wings not acuminate. FIII present (Fig. 224) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinodes, p. 71 2b Third segment of maxillary palpi shorter than 2nd (Fig. 230). Hind wings acuminate. FIII absent (Fig. 231) . . . . . . . . . . . . . . . . . . Psychomyia, p. 72

Genus Lype McLachlan

Lype McLachlan, 1879, p. 422 Type species designated by Ross, 1944: Anticyra phaeopha Stephens Maxillary palpi with 3rd segment shorter than 2nd. Middle legs of & flattened. Front wings with fairly long apical area. Hind wings distinctly narrower than front wings. Venation (Fig. 220): in front wings, FIII sessile or with short petiole. In hind wings, Sc very short, R1 long, extending to wing margin. FIII present. Two anal veins and a crossvein between M3+4 and Cu1. 70

% genitalia very simple (Figs. 221, 222): sternite IX very elongate. Tergite IX slender, broadly and firmly fused with segment X, but free at tip. Segment X massive, moderately sclerotized, surrounding phallic apparatus, with lower lateral sides of segment tucked under apparatus. Preanal appendages forming long ovals. Intermediate appendages lost. Inferior appendages with first 2 segments short and partially fused together. Second segment simple, long, and caliper-like with subapical prominence on inner surface. Phallic apparatus long, narrow, comprising tubular phallotheca, vestigial endotheca and aedeagus movable and concave toward top. & genitalia (Fig. 223): tergite and sternite VIII fused into single part. Segments X and XI considerably extended and gradually tapered. Ceiling of anovaginal opening simple. Tiny vulvar scale and moderately complex vaginal apparatus. The genus Lype is very small and found throughout the Holarctic region, Africa, and India. It groups species that are amazingly similar considering how isolated they are geographically. There is only one Nearctic species, diversa Banks, which is widely distributed in the eastern part of the continent and has been reported from New Brunswick to Ontario with a single record from Alberta. Larvae live in cool, running water. Genus Tinodes Curtis

Tinodes Curtis, 1834, p. 216 Monobasic type species: Tinodes lurida Curtis = Phryganea waeneri Linnaeus Revisions: Schmid, 1983; Armitage and Hamilton, 1990 Maxillary palpi with 3rd segment longer than the 2nd. Middle legs of & not flattened. Wings not greatly narrowed. Hind wings forming definite but rounded angle posteriorly. In front wings, apical area short. Venation (Fig. 224): in front wings, relatively large discoidal cell. FIII and FIV petiolate. In hind wings, Sc long. R1 ends at Sc at an early stage and is joined to R2+3 by a crossvein. FIII present. Two free anal veins and a crossvein between M3+4 and Cu1. % genitalia (Figs. 225–227): sternite IX low and fairly stout. Tergite IX very slender laterally and hinged to sternite. Segment X membranous and vestigial. Preanal appendages elongated into rods and sharing with tergite same point of insertion on sternite IX. Intermediate appendages remarkably developed; inserted on upper portion of sternite IX and initially vertical and separate; they subsequently fuse together, curve horizontally backward, then once again separate. Horizontal portion equipped 71

with many stout spines. Inferior appendages stout. First segment complicated in shape, whereas 2nd simple and arising before apex of first. The first 2 segments fused together at base and are extended by long anterior apodemal tonguelike structure serving as site for muscle attachment; inner portion of these segments developed into large sagittal appendage, which acts as guide for phallic apparatus; it is highly sclerotized and curves backward and downward. Phallic apparatus fairly long and situated very high above preanal appendages. It comprises phallotheca and endotheca that are not clearly distinguishable from each other and tiny, immovable, upwardly curved aedeagus. & genitalia (Figs. 228, 229): segment VIII comprises separate tergite and sternite. Segments X and XI considerably extended, with gradually tapering ends. Ceiling of anovaginal opening and vaginal apparatus complex. Tiny vulvar scale. The genus Tinodes is medium-sized and cosmopolitan in distribution although it does not occur in the Neotropical region. It is especially well represented in the mountain system extending from the Mediterranean to China. In Nearctic America, there are 12 species, none of which has been captured in Canada. One species, provo Ross and Merkley, is widely distributed in the western United States and has been reported in Idaho. It occurs in fairly warm running water. Genus Psychomyia Latreille

Psychomyia Latreille, 1829, p. 263 Type species designated by Ross, 1944: Psychomyia annulicornis Pictet = Psychomyia pusilla Fabricius Quissa Milne, 1936, p. 89 Monobasic type species: Psychomyia flavida Hagen Revisions: Schmid, 1983; Armitage and Hamilton, 1990 Maxillary palpi with 3rd segment shorter than 2nd (Fig. 230). Middle legs of & flattened. Wings long and narrow, especially hind wings which are acuminate. Venation (Fig. 231): in front wings, median cell relatively short and anal cells very unequal, with A2 ending at A1 and not at A3. In hind wings, Sc absent, R1 long, and R2+3 very short, ending at R1. FIII absent. Only 1 anal vein, and no crossvein between M3+4 and Cu1. % genitalia (Fig. 232): sternite IX small and compact. Tergite IX very slender, with long cleft at extremity that is clearly sclerotized. Preanal appendages large and very elongate. Intermediate appendages lost. Segment X vestigial. Inferior appendages with first 2 segments considerably 72

shortened, fused together, and embedded in sternite IX. Second segment long and well developed, simple or bifid. There is large genital cavity for phallic apparatus that is unusually shaped: phallotheca has thick, horizontal basal portion giving rise to cylindrical part that extends upward, then curves backward and upward, ending in hook that is probably the only remaining vestige of aedeagus. & genitalia (Fig. 233) do not form long ovipositor. Segment X relatively short; anovaginal opening fairly wide and vulvar scale large and prominent. The genus Psychomyia is medium-sized, with a wide Holarctic and Oriental distribution. There are only three Nearctic species, two of which, flavida Hagen and nomada Ross, live in Canada. P. flavida is widely distributed. It inhabits all lotic biotopes and is occasionally found in large numbers in one place. The %% are very rare locally, and the species is considered an example of optional parthenogenesis (Corbet et al., 1966). P. nomada is rare and has been reported in Quebec; lumina Ross has been recorded from Washington. Compared to the other species, flavida displays such extensive modification of the % genitalia that it could not be used as a model for the generic description provided above. The description was therefore based on the two other Nearctic species that, moreover, are very similar to many other non-Nearctic forms. The main modifications encountered in flavida are: (Fig. 232) considerable enlargement of tergite IX, with inner side profusely equipped with spines. Tergite fused with preanal appendages that are reduced and joined to sternite VIII, passing over sternite IX. First segment of inferior appendages greatly reduced, very complex, and bearing 2 long spines. Second segment widely bifid, with upper branches parallel to phallic apparatus.

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Family Limnephilidae Kolenati Limnephilidae Kolenati, 1848, p. 30 Type genus: Limnephilus Leach Revision: Schmid, 1955 The Limnephilidae are medium-sized or large caddisflies that vary considerably in structure, color, and appearance. Their overall appearance can therefore also prove useful for determination purposes. Head usually short and wide, but occasionally with occipital part more or less developed. Ocelli present. First antennal segment not longer than head. Maxillary palpi 3-segmented in %, with 1st segment very short and other 2 long and subequal. In &, palpi 5-jointed (Figs. 253, 254). Legs bearing generally well-developed spines. Spurs ranging from 1,3,4 to 1,1,1 but usually 1,3,4. Abdominal hemogill system present only in Dicosmoecinae and Apataniinae and consisting of small number of simple tubes. Internal gland of sternite V almost always small, with opening in slight depression or reticulated area (Fig. 439). The 2 pairs of wings fairly differently shaped. Front wings narrow at base, distinctly wider at level of anastomosis and elliptical at apex. Hind wings wider, with very ample anal area. Venation very constant and almost complete, with forks I, II, III, and V present in both pairs of wings. Sexual dimorphism rare and always weak. In front wings, discoidal and thyridial cells closed and very long. Median cell open. Anastomosis bipartite, consisting of a more or less broken line with variable slant. In hind wings, discoidal cell generally long and occasionally open. Four or 5 anal veins. Variations in venation occur only in Dicosmoecinae and Apataniinae and always represent simplifications. % genitalia are so fundamentally varied that six subfamilies had to be established. I have divided them into two conveniently named groups: the Polyphorae in which the genitalia primitively comprise 6 pairs of appendages, but which often lose some of them through specialization, and the Oligophorae in which the genitalia consistently comprise 3 pairs of appendages. & genitalia less varied than % genitalia. Segment VIII unmodified, although sternite is occasionally concave. Segment IX generally consists of tergite and sternite. Segment X is either simple or else more or less 74

indented tube. Appendages present or absent. Subanal plate occasionally present. Supragenital plate generally present. Vaginal opening on segment IX or between segments VIII and IX. Vulvar scale often trilobate. Vaginal apparatus complex and rather strongly sclerotized, with or without vestibule. Segment IX primitively short and annulate. Segment X not very large or virtually absent, with complex and highly variable armature (Schmid, 1955, Fig. 3). Preanal appendages rarely present and generally lost. Each of intermediate appendages is divided into 2 horizontal U-shaped branches. In Polyphorae, these branches are referred to as outer and inner branches because of their shape. In Oligophorae, the branches are differently shaped and named: inner branches have retained the name of intermediate appendages, whereas outer branches are referred to as superior appendages. The latter are thus present only in the Limnephilidae and should not be confused with the preanal appendages occurring in the other families although there is some resemblance. The superior appendages vicariously assume the role of the preanal appendages. The preanal and superior appendages are thus analogues and not homologues. In Polyphorae, there are also occasionally lower branches and a subanal plate. Inferior appendages 1- or 2-segmented and fused or articulated with segment IX. Phallic apparatus situated very low between inferior appendages. It retains its entire primitive conformation and is rarely simplified. Phallotheca merges with phallocrypt and takes over its functions. Endotheca membranous and erectile, as is generally the case, but inserted at base and not at extremity of phallotheca (Schmid, 1970, Fig. 3 is incorrect in this regard). Aedeagus long and slender and parameres rod-shaped or spiniform. As indicated in the Introduction, I do not accept families based on larval characters only. I consider these taxa as artificial and created for reasons of psychological convenience. The Limnephilidae are by far the largest trichopteran family in high latitudes of the Northern Hemisphere. The family is represented in all cold and temperate regions in the world, except in South Africa. In Canada there are currently almost 200 species inhabiting all lotic and lentic environments. They are found at very high altitudes and latitudes. The family was divided into six subfamilies (Schmid 1955, p. 17), five of which are found in Canada: Dicosmoecinae, Apataniinae, Neophylacinae, Pseudostenophylacinae, and Limnephilinae. Later on, the Neophylacinae were elevated to familial rank under the name Uenoidae. 1a Discoidal cell in hind wings open at apex. Pterostigma of front wings convex, anteriorly truncated, and strongly marked. Sc ending on crossvein C-R1 (Fig. 321) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apataniinae, p. 91 1b Discoidal cell in hind wings ogival and closed (Fig. 234). Crossvein C-R1 usually absent (Fig. 324) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

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2a Huge phallic apparatus with small aedeagus and parameres forming very large membranous lobes, strongly erectile and equipped with spines (Fig. 329). Sternite VIII in & with 2 seta-bearing concavities (Figs. 330, 331) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudostenophylacinae, p. 93 2b Species without these characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a FI in front wings joins discal cell for at least one-fourth length of latter (Fig. 301) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicosmoecinae, in part, p. 76 3b F1 in front wings joins discal cell for a much shorter distance (Fig. 234) . . 4 4a Segment X in % with more than 2 pairs of appendages (Figs. 236, 255) or, if this is not the case, 1- or 2-segmented inferior appendages hinged to segment IX (Figs. 261, 273). Vaginal opening of & on segment IX, between ventral lobes of latter (Figs. 245, 293). . . . . Dicosmoecinae, in part, p. 76 4b Segment X in % with 2 pairs of appendages. Inferior appendages in % singlesegmented and rigidly fused with segment IX (Fig. 334). Vaginal opening of & between segments VIII and IX, under ventral lobes of segment IX (Fig. 338) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnephilinae, p. 94

Subfamily Dicosmoecinae Schmid Dicosmoecinae Schmid, 1955, p. 29 Type genus: Dicosmoecus McLachlan Very wide, short head with protuberant eyes. Antennae almost always crenulated underneath. Maxillary palpi in % slender and variable in length. Spurs generally 1,3,4. Pronotum short. Abdominal hemogill system present, comprising small number of simple tubes on each segment. Wings large or medium-sized. Front wings elliptical at tips, and hind wings are as wide as or wider than front wings. Venation less constant than in other subfamilies and generally complete, with forks I, II, III, and V present in both pairs of wings. Some forks, such as FV in front wings and FI in hind wings, occasionally petiolate. FV in hind wings absent in Allomyia. Discoidal cell in both pairs of wings always closed. % genitalia very variable in size and complexity. Tergite VIII always undifferentiated, except in Ironoquia. Segment IX annulate, generally well developed dorsally and often indented for insertion of base of inferior appendages. Segment X slightly or not at all prominent with armature of often complex and variable branches. Complete armature comprising preanal appendages, lower, inner, and outer branches, and subanal plate, but it never occurs in its entirety in any given genus. Thus, preanal appendages and outer branches are never present at same time. Any one of branches or appendages may be missing in a given genus, except for inner branches that are always present. Thus, preanal appendages are absent in Dicosmoecus and Onocosmoecus, outer branches do not occur in Allomyia and Cryptochia, and lower branches and subanal plate are missing in 76

Ecclisomyia and Ecclisocosmoecus. Inferior appendages variable in size and generally articulated but not fused with segment IX. They may be in shape of large subcircular and 2-segmented callipers (Dicosmoecus, Eocosmoecus, Onocosmoecus), or 2nd segment may be more or less fused with 1st (Amphicosmoecus, Pedomoecus) or else may have completely disappeared (Allocosmoecus). Phallocrypt generally very long and reinforced with sclerotized bands joining phallotheca to inner margins of inferior appendages. Endotheca always present. Aedeagus generally large, but occasionally membranous and reduced (Ecclisomyia, Amphicosmoecus) or even absent (Rossiana). Parameres generally well developed and spine-bearing, occasionally partially fused with aedeagus (Dicosmoecus, Onocosmoecus) or even absent (Rossiana). & genitalia: segment IX entire or divided into tergite and sternite. Tergite IX short and blunt. Segment X, generally forming incised tube and more or less distinct from segment IX at base. Appendages almost always absent. Subanal plate present or absent. Sternite IX divided into 2 lobes framing vulvar scale, very variable in development and often concave on inner side, next to vaginal opening. Supragenital plate always present. Vaginal opening along entire length of segment IX. Vulvar scale formed by segment IX only and generally trilobate. There is occasionally a long vaginal vestibule, which is simple and cylindrical in Dicosmoecus and Onocosmoecus or complex in Ecclisomyia. Vaginal apparatus well developed and sclerotized. The Dicosmoecinae constitute a medium-sized subfamily with a Holarctic and southern Neotropical distribution. It is especially well represented on the northern Pacific rim. None of the genera is flourishing and almost all are relicts. A few of them are monobasic. Virtually all are highly differentiated and represent their own, and very unequal, stages of evolution, which makes the subfamily extremely heterogeneous and consequently difficult to describe. I include in the Dicosmoecinae the genera Allomyia, Manophylax, Moselyana, Pedomoecus, and Rossiana, since they fit into this subfamily by their adult characters. But, by their immature characters they do not and instead are considered as incertae sedis awaiting further investigation. 1a Large species with front wings over 20 mm long and thoracic pleura bearing tufts of woolly hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b Species without these characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a Inferior appendages of % in shape of large, circular, 2-segmented callipers (Fig. 236). Vaginal system of & preceded by long cylindrical vestibule (Fig. 239) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicosmoecus, p. 78 2b Inferior appendages of % small and single-jointed (Figs. 255, 257). Vaginal system of & without long vestibule (Figs. 259, 260) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Allocosmoecus, p. 81

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3a Discoidal cell in front wings very long, but with R2+3 and R4+5 parallel and very close together on basal half of cell (Fig. 308) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecclisocosmoecus %, in part, p. 89 3b Discoidal cell in front wings small and very narrow, joined by FI for more than half of cell length (Fig. 295). Small black species . . . Rossiana, p. 87 3c Discoidal cell in front wings very long, average in width, joined by FI for one-fourth or more of cell length (Fig. 301). Medium-sized, rust-colored species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3d Species without these characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a Inner side of inferior appendages with a stout spur or several long spines (Fig. 302). Vulvar scale prominent and single-lobed (Fig. 306) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecclisomyia, p. 88 4b Species without these characters and of eastern distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ironoquia, p. 90 4c Species without these characters and of western distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecclisocosmoecus, in part, p. 89 5a FI in hind wings absent (Fig. 280) . . . . . . . . . . . . . . . . . . . Manophylax, p. 85 5b FI in hind wings petiolate (Fig. 272) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5c FI in hind wings sessile (Fig. 234) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6a FV in hind wings absent. FI in hind wings with short petiole (Fig. 272). Small brown species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allomyia, p. 84 6b FV in hind wings present. FI in hind wings with long petiole (Fig. 266). Small rust-colored species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pedomoecus, p. 83 6c FV in hind wings present. FI in hind wings with short petiole (Fig. 287). Small black species with rust-colored pronotum . . . . . . Cryptochia, p. 86 7a Inferior appendages distinctly 2-segmented (Figs. 241, 246) . . . . . . . . . . . . . 8 7b Inferior appendages indistinctly 2-segmented. Inner branches of segment X small and lower branches slender and broadly curved downward (Fig. 261). Segment X of & ending in 2 small, slender, widely spaced points (Figs. 264, 265) . . . . . . . . . . . . . . . . . . . . . . Amphicosmoecus, p. 82 8a Anterior wings orange, with large imprecise brown areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onocosmoecus, p. 80 8b Anterior wings uniformly dark brown. . . . . . . . . . . . . . . . Eocosmoecus, p. 81

Genus Dicosmoecus McLachlan

Dicosmoecus McLachlan, 1875, p. 122 Type species designated by Ross, 1944: Stenophylax palatus McLachlan Revision: Wiggins and Richardson, 1982 78

Quite large species. Eyes very prominent and ocelli very large (Fig. 235). Top of head and thorax densely clothed with short macrochaetae. Thoracic pleura covered with fairly dense pilosity of woolly hairs. Spurs: 1,3,4. Wings brown with veins strongly marked in front wings. Front wings form fairly narrow ellipses with very long apical areas. In hind wings, anal area is highly developed. Sparse hairs in front wings and also in hind wings, except in anal area where there is dense cover. Venation (Fig. 234) unmodified, with all cells narrow and all forks present and sessile. In front wings, discoidal cell 3 or 4 times the length of its petiole. Anastomosis in broken line and slanting slightly bodyward anteriorly. In hind wings, very long discoidal cell and first 3 forks narrow at base. % genitalia (Figs. 236–238): segment IX evenly short all around except ventrally where it forms a blunt tonguelike extension. Laterally deeply indented for insertion of inferior appendages. Segment X large and fairly prominent. Preanal appendages absent or integrated with outer branches of segment X. These branches form medium-sized, elongated, and occasionally bifid lobes. Inner branches forming elongated, contiguous ovals. Lower branches as long as inner branches and located laterally. Subanal plate fairly large. Inferior appendages in shape of stout callipers. First segment particularly short, subglobular with length barely greater than width. Inner side strongly concave, forming distinct dentate carina. Second segment slightly shorter than 1st and abruptly narrowing shortly after base. Phallocrypt extremely long with lateral strips joining phallotheca to upper basal angle of 1st segment of inferior appendages. Endotheca long. Aedeagus completely sclerotized, simple, and slender. Parameres long pointed rods that are fused with aedeagus along their basal half. Free portion contiguous with aedeagus and bears row of more or less slender spines in arrangement characteristic of species. & genitalia (Figs. 239, 240): tergite and sternite IX almost completely separate. Segment X in shape of elongated tube with concave underside. Latter displaying fairly complex relief. Subanal plate tiny. Sternite IX prominent and divided into 2 large bipartite folds framing vaginal opening. Supragenital plate poorly developed. Vulvar scale small and cordiform. Vaginal vestibule forms long, fairly evenly sclerotized tube. Vaginal apparatus flattened. The genus Dicosmoecus has a Palearctic and Nearctic distribution. Eight closely related species have been recorded from North America. Wiggins and Richardson (1982) revised them and reduced their number to four, three of which are represented in our area: atripes Hagen and gilvipes Hagen are indeed very closely related and very widespread in the mountains of the western part of the continent; obscuripennis Banks is more isolated and more septentrional in distribution: Alaska, Yukon 79

Territory, and Northwest Territories. Larvae occur in a wide range of cool rivers and streams, but downstream from headwater sites. Genus Onocosmoecus Banks

Onocosmoecus Banks, 1943, p. 357 Type species by original designation: Dicosmoecus tristis Banks Revision: Wiggins and Richardson, 1986 Fairly large insects with wide, orange-tinted wings. Ocelli mediumsized. Spurs: 1,3,4. Top of body sparsely covered with macrochaetae. Thoracic pleura without woolly covering. Wings with veins not strongly marked. Front wings broadly elliptical and uniformly brown or orange-tinted with few indistinct cloudy spots and clothed with fine erect hairs. Hind wings colorless and virtually smooth, anal area included. Venation similar to that of Dicosmoecus but with systematically wider cells. In front wings, discoidal cell 2.5 to 3 times longer than its petiole, and anastomosis slanting strongly bodyward posteriorly. In hind wings, discoidal cell also roughly 3 times longer than its peduncle and anatomosis fairly irregular. % genitalia (Figs. 241–243): segment IX not so short and without ventromesal extension. Outer and inner branches of segment X fairly long, robust, straight, and subequal in length. Lower branches absent or vestigial and subanal plate very wide and bifid. Inferior appendages long, with 1st segment at least twice as long as high. Inner surface slightly concave with carina inconspicuous or absent. Second segment distinctly shorter than 1st and gradually tapered. Phallocrypt short. Aedeagus partially membranous and erectile, with pair of stout spines and with strongly and uniformly sclerotized spermatic duct. Parameres slender, fused with aedeagus for some distance, and equipped with few large spines. & genitalia (Figs. 244, 245): segment IX massive and not separated into tergite and sternite. Segment X blunt and more or less bifid. Subanal plate absent. Ventral portion of segment IX forming very large lobes framing vaginal opening. Supragenital plate large and triangular. Vulvar scale trilobate. Vaginal vestibule forming short cylindrical tube that is uniformly sclerotized. Fairly simple vaginal apparatus situated in large membranous pouch. The genus Onocosmoecus has an East Palearctic and Nearctic distribution. Six very closely related species have been described from North America. Wiggins and Richardson (1986) revised them and found that they all represent mere intraspecific variations of a single species: unicolor Banks. This species is transcontinental in distribution. Larvae occur 80

in slow waters and pool areas of cool rivers and streams and also in the littoral zone of cool lakes. Genus Eocosmoecus Wiggins and Richardson

Eocosmoecus Wiggins and Richardson, 1989, p. 356 Type species by original designation: Drusinus frontalis Banks Rather large insects, with front wings uniformly dark brown. Pleura and femora yellow to orange. Thorax with long, black, sparse setae. Dorsal warts of thorax pale against surrounding area. Thoracic pleura without woolly pilosity. Spurs: 1,3,4. Wings with veins not strongly marked. Anterior wings broadly elliptic, uniformly brown, and with sparse, erect hairs. Hind wings colorless and smooth, including anal area. Venation similar to that of Onocosmoecus, with all cells broad. % genitalia (Figs. 246–250): segment IX evenly short all around, except for ventral, obtuse, tonguelike extension. Segment X with outer and inner branches short, blunt, subequal in length, both much shorter than subanal plate. Lower branches absent. Subanal plate divided into pair of pointed, divergent lobes. Inferior appendages with first segment twice as long as high. Inner face slightly concave and without carina. Second segment shorter than 1st and gradually tapering. Phallocrypt short. Aedeagus completely membranous and erectile and with well-visible spermatic duct. Parameres entirely distinct from aedeagus, rodlike, strongly sclerotized, and bearing apical or subapical spines. & genitalia (Figs. 251, 252): segment IX divided into tergite and sternite. Tergite IX very massive. Sternite IX well developed as pair of large lobes. Segment X short, broad, and cleft into 2 widely divergent lobes. Supragenital plate large and not fused with segment X. Vulvar scale thick, ogival, and undivided. Vaginal vestibule and apparatus long. The genus Eocosmoecus contains two species previously included in Onocosmoecus. Wiggins and Richardson (1989) later isolated them in a separate generic entity; frontalis Banks (British Columbia, Washington, Oregon) shows angular outer branches of segment X and parameres bispinose apically; schmidi Wiggins (British Columbia, Idaho, Montana) has rounded outer branches and parameres with a row of spines on the apical half. Both species live in small spring streams. Genus Allocosmoecus Banks

Allocosmoecus Banks, 1943, p. 365 Monobasic type species: Allocosmoecus partitus Banks 81

Adults belonging to this genus show a remarkable resemblance to Dicosmoecus. They are very large, veins in front wings are lined with brown, and hind wings have wide anal area and dense pilosity. Top of body profusely covered with short macrochaetae and thoracic pleura have thick woolly covering. Ocelli smaller. Spurs: 1,2,2. Discoidal cell in both pairs of wings shorter. Colors showing great contrast. Body is dark, brown-black, with light rust-colored thoracic warts. Face, palpi, and legs also light rust. Wings darker and contrasted in % but uniformly brownblack in &. % genitalia (Figs. 255–258), on other hand, differ markedly from those of Dicosmoecus but derived from same type. Segment IX short dorsally and especially laterally, but elongated ventrolaterally and indented for insertion of inferior appendages. Segment X inconspicuous. Preanal appendages absent. Outer branches consist of large, wide, ear-shaped appendages located fairly low. Inner branches located very high and form 2 elongated, slightly divergent lobes. Lower branches situated very low, above inferior appendages, and create a vast concave space extending to inner branches. Lower branches consist of 2 stout, heavily sclerotized, horizontal, sharp spurs. Inferior appendages greatly reduced in size, single-jointed, and shaped into 2 horizontal lobes. Phallocrypt not very long, but high and each side equipped with a lateral carina. Endotheca rather short. Aedeagus short, flaring laterally at extremity, and equipped with 2 erectile lateral lobes. Parameres free, forming slender, seta-bearing rods. & genitalia (Figs. 259, 260): segment IX well developed dorsally but very short laterally and ventrally where it forms 2 very small lobes framing vulvar scale. Segment X clearly distinct from segment IX and forming 2 large, horizontal, ear-shaped lobes above and 2 small points below anus. Subanal plate absent. Supragenital plate large, membranous, and not prominent. Vulvar scale simple, blunt oval. Vaginal vestibule short. Vaginal apparatus large, flattened, and accompanied by 2 irregularly shaped, highly sclerotized lateral folds. The genus Allocosmoecus contains only a single species, partitus Banks, which is found in the headwaters of small cool streams in the mountains of the western part of the continent, from California to British Columbia. The genitalia of the % are an unusual feature in this genus, which is closely related to the Neotropical genus Monocosmoecus Ulmer. Genus Amphicosmoecus Schmid

Amphicosmoecus Schmid, 1955, p. 49 Type species by original designation: Dicosmoecus canax Ross Medium-sized rust-colored species with yellow-brown front wings. Spurs: 1,2,4. 82

Wings fairly wide with anal area of hind wings very wide. Front wings are slightly granular and bear short, erect hairs. Venation complete, with all forks present and sessile. Discoidal cells in both pairs of wings 2.5 times longer than their petioles. % genitalia (Figs. 261–263): segment IX short dorsally and very long laterally and ventrally where it is indented for insertion of inferior appendages. Segment X virtually absent. Inner branches forming short subvertical lobes completely fused with dorsal part of segment IX. Outer branches absent. Preanal appendages present, forming fairly long ovals. Lower branches very long, located between preanal appendages, strongly sclerotized, and curve broadly downward. Subanal plate absent. Inferior appendages large, slanting upward, calliper-shaped, and comprising 2 conspicuous segments that are almost completely fused together. Phallic apparatus situated fairly high. Phallocrypt very long and reinforced with sclerotized strip. Endotheca well developed. Aedeagus small, membranous, and capable of telescoping within itself. Tip of aedeagus and spermatic duct strongly sclerotized. Parameres reduced, slender, fused at base with aedeagus, and bearing only a few setae. & genitalia (Figs. 264, 265): segment IX consisting of tergite in broad contact with sternite. Segment X small, forming very short tube, ending in 2 slender, widely spaced points, and with 2 large oval appendages. Subanal plate absent. Large supragenital plate. Sternite IX forming 2 large lobes that are thick and concave on upper and inner sides. Vulvar scale with 3 barely marked lobes. Vaginal vestibule shallow and vaginal apparatus simple. The genus Amphicosmoecus comprises only a single species, canax Ross, found from British Columbia to Saskatchewan and further south. It has been captured near rivers of different sizes and degrees of turbulence, but also near lakes. Genus Pedomoecus Ross

Pedomoecus Ross, 1947, p. 150 Type species by original designation: Pedomoecus sierra Ross Fairly small, uniformly orange-yellow species. Spurs: 1,2,2. Wings large but not very wide. Front wings regularly elliptical at tips and hind wings barely wider than front wings. Very fine and fairly dense hairs on membrane of both pairs. Venation complete, with all forks present, although not all sessile (Fig. 266). In front wings, discoidal cell narrow and not longer than its petiole. FIII pointed. Anastomosis parallel to body and fairly strongly fragmented. In hind wings, R1 very thin, discoidal cell short, FI as long as its petiole and FIII with short peduncle or pointed. 83

% genitalia (Figs. 267–269): segment IX considerably reduced, slightly elongated dorsally, and represented ventrally only by a narrow band barely distinct from inferior appendages. Segment X reduced, only slightly sclerotized, with all branches lost or indistinct, forming notched roof above anal cavity. Subanal plate small. Inferior appendages large, barely protruding, 2-segmented but not calliper-shaped, slanting sharply upward and immovable. Basal segments large, bulky, thick at base, appearing flattened when viewed transversely, forming vast circular, concave surface, directed backward. Upper apical margin stretched into more or less sharp spur. Second segment comprising 2 branches forming blunt lobes and clearly distinct from 1st segment, except dorsally where it fuses with underside of apical spur of 1st segment. Phallic apparatus large and short; aedeagus simple with large phallotremal sclerite, an unpaired, stout, dorsal point and 2 trilobate, spine-bearing parameres. & genitalia (Figs. 270, 271): segment IX comprises tergite and sternite, distinct from each other. Tergite massive but sternite considerably reduced, forming 2 slender lobes. Segment X clearly distinct from segment IX, forming 2 large triangular parts. Subanal plate absent. Supragenital plate very large and as wide as segment IX. Vulvar scale embedded in sternite VIII, very wide, not prominent, with 3 lobes: a strongly protruding, membranous median lobe and 2 lateral, barely indicated triangles. Vaginal opening very wide and gaping. Vaginal apparatus large, complex, very wide, and without vestibule. The genus Pedomoecus includes only a single species, sierra Ross. It is rare and local and found from California to British Columbia and Alberta in cold, fast-flowing streams. Genus Allomyia Banks

Allomyia Banks, 1916, p. 120 Monobasic type species: Apatania tripunctata Banks Imania Martynov, 1935, p. 298 Type species by original designation: Imania sichotalinensis Martynov Revision: Ross, 1950 Small, slender, brown or dark beige insects. Antennae fine and not crenulated. Spurs: 1,3,4. Hind wings slightly wider than front wings and uniformly covered with fine hairs. Venation complete in front wings and slightly reduced in hind wings (Fig. 272). In front wings, R1 occasionally slightly curved at the level of pterostigma and at times joined to Sc by crossvein. Discoidal cell twice as long as its petiole. All forks sessile and anastomosis slightly 84

concave in direction of wing base. In hind wings, discoidal cell small, FI with a short petiole and anastomosis faintly broken and strongly slanting. Cu1a, and therefore FV, absent. % genitalia (Figs. 273–276): segment IX short laterally and fitting around base of inferior appendages for some distance. Armature of segment X simple but difficult to interpret. There are probably 2 inner branches, which are reduced and firmly fused together into short median lobe, and 2 ovoid outer branches resembling preanal appendages. The latter absent. In addition, there are 1 or 2 pairs of lower parts, which probably are single or double lower branches. Inferior appendages 2-segmented, very large, and slanting sharply upward. First segment elongate, with a long line of contact with segment IX. Two spiniform, lobed or plated appendages arise from inner basal angles. Second segment crescent-shaped. Inner surface of the 2 crescent horns bearing stout coneshaped tubercles. Phallocrypt narrow, moderately deep, and reinforced with lateral sclerotized bands connected to inner margin of inferior appendages. Endotheca short. Aedeagus membranous, with stout spermatic duct. Spine-shaped parameres that seem to be divided into 2 parts in bifosa Ross, thomasi Nimmo, and tripunctata Banks. In these species, only the basal pair constitutes the parameres, and the apical pair is actually the basal portion of the aedeagus. This double, sclerotized base is detached from the membranous apical portion and is movable in relation to the basal pair. & genitalia (Figs. 277, 278): segment IX consists of 2 generally clearly distinct parts. Tergite IX little developed and not distinct from segment X. The latter comprises 2 horizontal plates that are strongly sclerotized underneath. Sternite IX consisting of 2 well-developed lobes framing vaginal opening. Subanal plate absent. Supragenital plate well developed. Underside forms ceiling of genital cavity. Vulvar scale simple, membranous, and probably erectile. Vaginal opening gaping and vaginal vestibule short and very wide. The genus Allomyia is a fairly diverse East Palaearctic and Nearctic genus with a dozen species, most of which are North American. Seven have been reported in the mountains of western Canada and southward where the larvae inhabit springs and cold watercourses at every altitude. A. bifosa is especially common and flies in June, even before the snow has completely melted. The erectile appendage of segment V is an unusual feature (Schmid, 1968a).

Genus Manophylax Wiggins

Manophylax Wiggins, 1973b, p. 10 Type species by original designation: Manophylax annulatus Wiggins 85

Madeophylax Huryn and Wallace, 1984, p. 286 Type species by original designation: Madeophylax altus Huryn and Wallace Small, rather dark brown insects. Vertex with posterior warts ovoid. Spurs: 1,2,4. Wings rather broadly elliptical at apex. Hind wings slightly broader than front wings. These are uniformly brown with dense coating of hairs. Venation (Figs. 279, 280) complete in front wings and somewhat reduced in hind wings and with slight sexual dimorphism in anterior wings. In both sexes, in front wings, crossvein Sc-R1 absent, discoidal cell narrow and 3 times longer than its petiole. Forks I, II, III, and V present. In %, fork III sessile and Cu2 curiously ends midway at A1+2+3. In &, FIII petiolated and Cu2 ends at wing margin. In posterior wings of both sexes, forks II, III, and V present. % genitalia (Figs. 281–284): segment IX very short all around, but forming a mediolateral posterior angle. Segment X with both inner branches forming a long oval plate, connected mesally by membranes and with setose lateral margins. Outer branches absent. Preanal appendages ovoid and strongly sclerotized. Lower branches long, very thin, and with little sclerotization. Inferior appendages 2-segmented. First segment thin and 3 times longer than thick. Second segment ovoid and with strong apical spine. Phallocrypt shallow. Phallotheca well sclerotized and extended lateroventrally into rodlike strongly sclerotized parts that could be parameres fused at their base with the phallotheca. Endotheca absent. Aedeagus long, spatulate, and somewhat troughlike. & genitalia (Figs. 285, 286): segment IX consists of 2 very distinct parts separated by a membranous area. Tergite IX simple and not extending laterally downward. Sternite IX forming quadrate parts laterally to vulvar scale and with their inner basal angles extended anteriorly. Segment X in 2 obtusely triangular lobes. Subanal plate absent. Supragenital plate more than twice as wide as long, terminating in 2 rounded lobes and a shallow median notch. Vaginal vestibule short and wide. Vaginal apparatus simple. The genus Manophylax contains two species: altus Huryn and Wallace from North Carolina and annulatus Wiggins, recorded from Idaho. Larvae were found in a small stream cascading over boulders down a steep slope; they apparently live under a thin film of flowing water on flat rocks. Genus Cryptochia Ross

Cryptochia Ross, 1950, p. 425 Type species by original designation: Parachiona pilosa Banks 86

Small, entirely black insects with a rust-colored pronotum. Eyes small. Vertex strongly bulging. Spurs: 1,3,4. Wings evenly elliptical, with hind wings barely wider than front wings and uniformly clothed with dense hairs. Venation complete with all forks present (Fig. 287). In front wings, discoidal cell narrow and 1.5 times longer than its petiole. FIII pointed. In hind wings, discoidal cell as long as its petiole and FI with short petiole. Anastomosis in slightly broken line and slanting sharply toward body, rearward. % genitalia (Figs. 288–291): segment IX strongly developed and completely surrounding inferior appendages. Segment X reduced, forming 2 sclerotized plates in crescent arrangement around anus. Its armature unusually developed. Preanal appendages consisting of small, ovoid lobes. Outer branches absent. Inner branches very large, forming very elongated horizontal roof. Underside of segment X with 2 paired lobes that undoubtedly constitute divided lower branches. Subanal plate absent. Inferior appendages 2-segmented but reduced and not prominent. Each with first segment developed into kidney-shaped plate completely embedded in segment IX. Second segment situated to upper end of 1st segment, as a very long, slender, subvertical rod. Phallocrypt bulky and joined to base of inferior appendages. Endotheca short. Aedeagus very large, sclerotized, and bent downward. Parameres in very stout spines. & genitalia (Figs. 292, 293): sternite VIII broadened and concave on underside. Segment IX completely embedded in preceding segment, markedly reduced, and composed of 2 parts. Vestigial tergite and sternite concave on side of vaginal opening. Segment X comprising 2 large triangular plates, concave underneath. Subanal plate absent. Supragenital plate very wide, trapezoid. Vulvar scale very large and consisting of 3 subequal lobes. Vaginal cavity vast and gaping. Vaginal vestibule absent. Vaginal apparatus very large and complex. The genus Cryptochia is exclusively Nearctic and includes seven very closely related species, two of which, pilosa Banks and furcata Denning, have been reported in British Columbia. They are found near small, cold, mountain streams. Genus Rossiana Denning

Rossiana Denning, 1953, p. 165 Type species by original designation: Rossiana montana Denning Small, entirely black species. Eyes small and covered with fairly long hairs. First antennal segment clearly longer than head. Maxillary palpi of % greatly reduced and modified and do not reach base of antennae. Last 2 segments are very small and inserted before apex of 1st segment (Fig. 294). 87

Both pairs of wings long, narrow, and smooth ellipses, with membrane densely clothed with appressed hairs. Venation (Fig. 295) similar on both pairs, complete, with all forks present and sessile, but remarkable on account of modifications it has undergone. In front wings, crossveins C-Sc and Sc-R1 present. On both pairs of wings, discoidal cell strongly reduced in length and width, joined by FI for over half of cell length. FII, although not petiolate, beginning well after end of discoidal cell. In hind wings, veins in middle of wing extremely fine and unstable, depending on specimens. % genitalia (Figs. 296–298): segment IX not very elongate, fairly irregular in shape, forming a ventral tonguelike structure bordering inferior appendages. Segment X not large. Outer branches absent. Preanal appendages consisting of downward-slanting lobes constricted at midlength. Inner branches distinctly lower, straight, horizontal, and slightly enlarged at tips. Inferior appendages comprising 2 well-developed segments. First segment massive, high, and very short. Inner surface exhibiting sharp, complex relief and forms 2 carinae. Second segment slightly longer than 1st, and slender with concave underside. Phallic apparatus comprising subcylindrical phallotheca joined to base of inferior appendages and short, erectile endotheca bearing 2 tufts of spines separated by long phallotremal sclerite. Aedeagus and parameres lost. & genitalia (Figs. 299, 300): segments IX and X reduced, not readily distinguishable from each other and forming very blunt mass. Subanal plate absent. Supragenital plate short, very wide, and bluntly rounded and concave underneath. Vulvar scale forming simple, very broadly curved bulge with median lobe corresponding to stout internal carina. It is flanked by 2 semimembranous lobes that probably arise from the upper apical angles of sternite VIII. Large, fairly complex vaginal apparatus without any vestibule. The genus Rossiana contains only a single species, montana Denning, which is found in Washington state, Montana, and British Columbia. It occurs very locally and inhabits cold mountain streams. Genus Ecclisomyia Banks

Ecclisomyia Banks, 1907, p. 123 Type species by original designation: Ecclisomyia conspersa Banks Medium-sized reddish-brown species with front wings speckled with light color. Spurs: 1,2,4 or 1,3,4. Wings of smooth elliptical shape. Hind wings barely wider than front wings. The latter slightly grainy with sparse, erect hairs. Anal area on hind wings also grainy and covered with erect setae. Venation complete with all forks present and sessile (Fig. 301). In front wings, discoidal cell 3 times 88

length of its petiole and joined by FI for over almost half of cell length. Anastomosis in strongly broken line and slanting slightly toward body, rearward. In hind wings, discoidal cell barely longer than its petiole. % genitalia (Figs. 302–305): segment IX short dorsally and ventrally but long laterally, always completely embedded in segment VIII. Lateral sides unsclerotized and reinforced with anterior rib. Segment X virtually absent, with simplified armature, forming lateral wings. Preanal appendages absent. Outer branches oval, simple, and slightly concave. Inner branches completely fused together into slender horizontal lobe, longer than outer branches. Lower branches absent. Inferior appendages apparently single-segmented and embedded in segment IX without indenting the latter’s margin. They consist of fairly irregular lobes with large spur or tuft of long spines on inner side. Extreme base of appendages with spur or spines probably constitutes 1st segment, and apical lobe 2nd segment. Phallocrypt variable in length, fairly strongly sclerotized, without lateral ribs but with dorsal reinforcement. Endotheca completely sclerotized. Aedeagus and parameres fused into a compact, rigid, and strongly sclerotized unit. & genitalia (Figs. 306, 307): segment IX forming single, large, solid, simple part. Segment X cone-shaped, with an elongated anal opening directed downward. Subanal plate absent. Large supragenital plate, strongly concave on underside, sheathing vulvar scale, which is long, simple, membranous, and movable. Vaginal vestibule long, irregular, and reinforced with curved sclerotized bands. Vaginal apparatus flattened. The genus Ecclisomyia includes half a dozen species of East Palaearctic and Nearctic distribution. Three species inhabit this continent, two of which are common in the western Nearctic region, i.e., conspersa Banks (in which the inferior appendages bear a stout inner spur and the phallocrypt is very long) and maculosa Banks (in which the inferior appendages bear a tuft of spines and the phallocrypt is short). Larvae of these species occur in all types of running water. E. conspersa is especially common and begins to emerge in May, near the snow. Genus Ecclisocosmoecus Schmid

Ecclisocosmoecus Schmid, 1964, p. 830 Type species by original designation: Ecclisocosmoecus spinosus Schmid Medium-sized rust-colored species with front wings densely speckled with light spots. Spurs: 1,2,4. Wings elliptical and fairly wide. Hind wings clearly wider than front wings. In front wings, membrane slightly grainy and clothed with bristles. In hind wings, anal area not particularly hairy. Venation complete with all 89

forks present and sessile (Fig. 308). In front wings, discoidal cell 4 to 5 times as long as its petiole, but with basal two-thirds strongly tapered in %, with the result that R2+3 and R4+5 lie very close together. In resulting furrow, there are tiny scales covered with row of erect, convergent setae. In &, discoidal cell is narrow but normally shaped. In hind wings in both sexes, discoidal cell 3 to 4 times longer than its petiole. % genitalia oddly, but only superficially, resemble those of Limnephilinae (Figs. 309–311). Segment IX small and much lower than segment VIII, short dorsally and ventrally, but very long laterally where apical margin folds broadly inside genital cavity; dorsally forming very large cavity within tergite VIII. Segment X virtually absent but with sclerotized winglike structures extending laterally. Preanal appendages absent. Outer branches in shape of oval lobes. Inner branches paired, consisting of 2 long spurs, concave along median line. Lower branches absent. Inferior appendages single-segmented, blunt, not very prominent, concave on inner side, and heavily clothed with pilosity. Phallocrypt thick, shallow, equipped with 2 sclerotized apicodorsal ribs. Phallic apparatus very small. Well-developed endotheca. Aedeagus reduced, membranous, and erectile, with spermatic duct thick and sclerotized. Parameres form stout spine-bearing spurs. & genitalia (Figs. 312, 313): segment IX consisting of single part, with ventral angles folding into 2 large, not very prominent lobes on either side of vaginal opening. Segment X virtually indistinct from segment IX, ending in 2 winglike structures overhanging anus. Subanal plate absent. Supragenital plate bottle-shaped and concave at lower end where it forms ceiling of vaginal cavity. Vulvar scale embedded in sternite VIII and comprising 3 slightly protuberant lobes, lateral ones largely inconspicuous. Vaginal apparatus flattened, preceded by very short vertical vestibule and reinforced with 2 sclerotized bands. The genus Ecclisocosmoecus contains only two species, one of which is found in the Eastern Palearctic region and the other, scylla Milne, in the Nearctic region. It is found in the western portion of the continent, from Oregon to British Columbia. Larvae occur in cold, swift-moving mountain streams.

Genus Ironoquia Banks

Ironoquia Banks, 1916, p. 121 Type species by original designation: Chaetopterygopsis parvula Banks Allophylax Banks, 1907, p. 119 (preoccupied) Type species by original designation: Halesus punctatissimus Walker 90

Caborius Navas, 1918, p. 362 Nomen novum for Allophylax Banks, 1907 Revision: Schmid, 1951 Medium-sized or small insects with wide, blunt, rust-colored, and finely speckled front wings. Two sexes with slightly different wing shapes. In %, front wings wide and obtusely rounded at tips. Hind wings barely wider than front wings and slightly notched at lower margin. In &, wings are slightly smaller and narrower. Long, fine, erect hairs on front wings. Anal area of hind wings in % slightly grainy. Venation complete and very similar to that of Ecclisomyia (Fig. 301) with all forks sessile and FI merging for a long distance with discoidal cell on front wings. % genitalia (Figs. 314–317): tergite VIII developed into apicodorsal plate. Segment IX short dorsally but stout laterally and ventrally and fitting around inferior appendages. Segment X barely protruding. It has a complex armature but all branches short. Outer branches consist of slightly concave lobes. Inner branches form upward-slanting spurs. Preanal appendages absent. Lower branches fairly prominent. A large, trilobate subanal plate occasionally present. Inferior appendages single-segmented, extensively merging with segment IX, cone-shaped or upward-slanting. Phallocrypt large and reinforced with lower sclerotized band joined to upper inner angle of inferior appendages. Phallic apparatus large. Aedeagus robust, strongly erectile at tip where it may bear coneshaped spurs. Parameres spine-bearing or pectinate. & genitalia (Figs. 318–320): segment IX consisting of 2 parts. Tergite short and massive. Segment X forming blunt, cone-shaped tube truncated obliquely downward. Subanal plate absent. Sternite IX forming 2 blunt ventral lobes concave on inner surface. Supragenital plate small. Vaginal opening wide, short, without vestibule. Vulvar scale trilobate, protruding only slightly. Vaginal apparatus blunt. The genus Ironoquia is small, Holarctic, and contains four Nearctic species, two of which have been reported in the eastern and central parts of Canada: punctatissima Walker is fairly large and robust and parvula Banks is much smaller with a reduced anal area on the hind wings. Larvae of these species inhabit essentially ponds and temporary streams.

Subfamily Apataniinae Wallengren Apataniidae Wallengren, 1886, p. 73 Type genus: Apatania Kolenati This subfamily is represented in Canada by only a single genus. 91

Genus Apatania Kolenati

Apatania Kolenati, 1848, p. 75 Type species designated by Fischer, 1967: Phryganea vestita Kolenati nec Zetterstedt = Apatania wallengreni McLachlan Radema sensu Ross, 1944, p. 181 Revision: Schmid, 1953, 1954a Head relatively narrow with sides bulging. Eyes small. Maxillary palpi only slightly developed. Spurs: 1,2,2 or 1,2,4. Abdominal hemogill system well developed. Wings medium-sized and constant in shape among species; similar in both sexes. Front wings elongated and obliquely elliptical. Hind wings barely wider than front wings, with posterior margin convex but slightly indented. Fine, very dense pilosity, uniformly distributed on both wings. Venation complete, with all forks present (Fig. 321). In front wings, R1 joined to C by crossvein which also cuts off Sc. Pterostigma slightly leathery, more so in % than in &. Anastomosis in single, irregularly broken line. Discoidal cell rather short and curved slightly upward. FI and FIII narrow or pointed. In hind wings, discoidal cell open, and FI very short. M forks virtually at the level of Cu1. % genitalia of very variable complexity, as in Dicosmoecinae (Figs. 322–324). Segment IX ring-shaped but slightly shortened dorsally, forming 2 slender, apicodorsal lobes parallel or integrated with branches of segment X. The latter more or less prominent. Preanal appendages present, free or integrated with outer branches, which are large and occasionally bifid. Inner branches distinct or fused together. Lower branches and subanal plate absent. Inferior appendages large, robust, always 2-segmented, with 2nd segment simple or complex. Phallic apparatus variable in length. Phallocrypt more or less deep and reinforced with 2 lateral sclerotized bands forming sclerotized connection with extension of upper inner angle of inferior appendages. Endotheca well developed. Aedeagus variable in length and occasionally equipped with apical spines. Parameres spiniform. & genitalia (Figs. 325, 326): segment IX blunt, consisting of single part in primitive species, but laterally narrower in other species, forming 2 ventral lobes around vaginal opening. Segment X short, appendage-free, in often-complex tube shape, with anal opening toward top or bottom. Subanal plate absent. Supragenital plate membranous or rigid, occasionally fused with segment X and extending far into vaginal cavity. Vulvar scale consisting of single lobe arising from sternite VIII, membranous and probably erectile. Vaginal chamber large, without any vestibule, extending along entire length of segment IX. Vaginal apparatus complex. 92

The genus Apatania is large and of Holarctic distribution. The genitalia of the % display considerable variation. Twelve species have been reported in our area. They have been classified into four groups that vary considerably in level of specialization. Since the wing and body characters remain extremely constant, there is no reason to consider them subgenera. Apatania species are essentially stenotherms, dwelling in cold springs in the mountains of eastern and western North America. The most common species of the genus, zonella Zetterstedt, however, is lentic and inhabits lakes of the entire circumboreal region, extending as far north as conditions for invertebrate life allow. It is generally parthenogenetic.

Subfamily Pseudostenophylacinae Schmid Pseudostenophylacinae Schmid, 1955, p. 102 Type genus: Pseudostenophylax Martynov This subfamily is represented in Nearctic America by a single genus. Genus Pseudostenophylax Martynov

Pseudostenophylax Martynov, 1909, p. 281 Type species designated by Mosely, 1936, as: Pseudostenophylax fumosus Martynov Drusinus Betten et al., 1934, p. 359 Type species by original designation: Drusinus uniformis Betten Revision: Schmid, 1991 Head very wide with eyes and ocelli large and very prominent. Antennae thick and strongly crenulated on inner surface. Maxillary palpi of % long and stout. Spurs: 1,3,3 or 1,3,4. Abdominal hemogill system absent. Wings fairly broadly rounded with hind wings not much wider than front wings. Membrane of front wings grainy and clothed with erect setae. Venation complete with forks I, II, III, and V present on both pairs of wings and all of them sessile. In front wings, discoidal and thyridial cells very long. % genitalia simple with 3 pairs of appendages (Figs. 327–329). Posterior portion of tergite VIII developed into a rounded, sclerotized area, densely covered with short, thick spinules. Segment IX short dorsally but very convex and strongly developed laterally. Superior appendages forming small lobes bearing long hairs; base of these lobes broadly lining genital 93

cavity. Intermediate appendages large or small. Inferior appendages singlejointed, ventrally located, forming horizontal, shovel-like plates. Phallocrypt shallow but very wide. Phallic apparatus short, thick, and of very large size. Phallotheca and endotheca very short and reduced. Aedeagus short and thick with apical portion far more slender than basal part. Parameres very large, membranous, and endowed with prodigious erectile powers; apical or subbasal part thick, highly sclerotized, and equipped with long spines. & genitalia forming a very wide, massive unit with complex relief (Figs. 330, 331). Sternite VIII with wide concavities clothed with erect setae. Segment X small, developed into 2 blunt lobes, and without appendages. Supragenital plate absent or membranous. Vulvar scale unusually structured, consisting of a tiny median lobe surrounded by 2 large, very high, thick, lateral lobes. Inner surfaces of these lobes forming a hollow serving as vestibule for vaginal apparatus that is simple in structure and positioned very posteriorly. The genus Pseudostenophylax is large, with an Oriental distribution centered around the Himalayas and central China. Only two species are Nearctic: edwardsi Banks, in which the intermediate appendages consist of large masses forming a grainy horizontal surface, occurs from California to British Columbia; sparsus Banks, whose intermediate appendages are slender and vertical, is widespread in the eastern and central part of the continent. It consists of two subspecies: sparsus sparsus whose anterior wings are brown with many light spots and sparsus uniformis Betten, in which the anterior wings are uniformly brown. The genitalic subspecific characters are very few, involving very slight details only. Both species inhabit small streams which generally arise from springs and are occasionally intermittent. This subfamily originated in the Oriental region, where all species occur except for the two Nearctic ones (Schmid 1989, p. 110).

Subfamily Limnephilinae Kolenati Head shape, eye size, and antennal and palpal thickness variable. Eyes often hairy. In %, 1st segment of front tarsi occasionally shorter than 2nd. Spurs vary considerably in number. The usual pattern is 1,3,4 but can be reduced to 1,1,1. Abdominal hemogill system absent. Wings also vary considerably in size and shape. Front wings fairly narrow, truncated or rounded at tips. Hind wings constantly wider, with anal area highly developed. Pilosity of front wings varies considerably. Color of front wings varied and often bright, with contrasting hues, whereas hind wings consistently colorless. Venation (Fig. 333) of primitive, complete type with forks I, II, III, and V present on both pairs of wings. Variations are very minor and always same in both sexes. In front 94

wings, discoidal cell 1 to 3 times longer than its petiole. Anastomosis variously broken, at times concave in direction of the body and at others, slanting forward or rearward. Forks rarely petiolate. In hind wings, anastomosis more or less broken and generally slanting rearward. % genitalia consistently simple, with 3 pairs of appendages. Reductions are rare and the major structures vary little. On this stable base occur a large number of secondary variations involving primarily the shape of the appendages, thus creating an extremely varied lineage. Tergite VIII often with spinule-covered apical protuberance. Segment IX more or less long laterally and ventrally but considerably shortened dorsally where it is reduced to a narrow strip. Segment X virtually absent and represented only by superior and intermediate appendages. Superior appendages ear-shaped and rounded and concave, inerm or dentate. Intermediate appendages developed into plates or spurs; always very sclerotized, slanting upward, and usually arising from triangular lateral sclerites. Anal opening between intermediate appendages. Inferior appendages consistently single-segmented, rigidly fused with segment IX, only slightly prominent and slanting upward. Phallic apparatus always complete, including phallotheca, endotheca, aedeagus, and parameres, with large variations in shape of aedeagus and parameres. & genitalia: segment IX almost always consisting of distinct tergite and sternite. Tergite IX shaped like large cone and extended directly beyond segment X. Primitively, segment X, with or without appendages, in shape of simple tube, but may be variously and occasionally so deeply indented that its different parts are reduced and separated into lobes or scales. Sternite IX developed into 2 large, very blunt and generally barely prominent lobes. Supragenital plate present or absent. Vaginal opening between sternites VIII and IX. Vulvar scale usually trilobate, thick and fleshy, with sharp relief on inner side, partially or entirely forming vaginal vestibule. Median lobe produced by an elongation of sternite VIII and lateral lobes by 2 extensions of vaginal apparatus. These extensions have become external. Vaginal apparatus similar to that of other subfamilies. The Limnephilinae are the largest, most flourishing, and most widely distributed of the various subfamilies. Unlike the Dicosmoecinae, which comprise small genera differing considerably from one another, the Limnephilinae have achieved a stable and unique status onto which are superimposed numerous, but less pronounced, generic variations. The species are found throughout the Northern Hemisphere and inhabit all types of lotic and lentic environments. The subfamily is divided into four natural tribes: the Limnephilini, Stenophylacini, Chilostigmini, and Chaetopterygini. Only the first three are represented in our area. The characters defining them are fairly subtle and not very stable and as a result, a key cannot be used to distinguish these tribes. The genera will therefore keyed as one group. 95

1a Stout insects, with short, rounded, broad wings; front wings with thick veins and bearing long, erect bristles (Fig. 451) . . . . . . . Sphagnophylax, p. 119 1b Insects without these characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Front wings slightly indented at apex (Figs. III, X) . . . . . . . . . . . . . . . . . . . . 3 2b Front wings not indented at apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3a Head and pronotum especially elongate and grainy (Figs. II, III). Front wings leathery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemotaulius, p. 101 3b Head and pronotum not elongate and front wings not leathery (Fig. IX) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphopsyche, p. 125 4a R5 of hind wings strongly lined with brown (Fig. I). Grammotaulius, p. 100 4b R5 of hind wings unlined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a In front wings, only 2 anal cells present, since A2 lacks distal end (Fig. 420) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platycentropus, p. 115 5b In front wings, 3 anal cells present since A2 is complete (Fig. 333). . . . . . . 6 6a Spur of front tibia short, wide, and triangular (Fig. 403) . Philarctus, p. 112 6b Spur of front tibia normal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7a Front wings strongly striped with black (Fig. IV) . . . . . . Halesochila, p. 105 7b Front wings differently colored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a Anastomosis of front wings in one part (Fig. 520) . . . . Homophylax, p. 131 8b Anastomosis of front wings in 2 parts (Fig. 333) . . . . . . . . . . . . . . . . . . . . . . 9 9a Head and pronotum especially elongate and grainy in texture. Hind wings deeply indented below apex (Fig. 353) . . . . . . . . . . . . Leptophylax, p. 102 9b Species without these characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a In hind wings, Sc and R1 joined by a crossvein slightly before their extremities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydatophylax, p. 117 10b In hind wings, Sc and R1 not joined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a FIII of both pairs of wings pedunculate (Fig. 528) . . . . . . Phanocelia, p. 132 11b FIII of both pairs of wings sessile (Fig. 333) . . . . . . . . . . . . . . . . . . . . . . . . 12 12a Pterostigma of front wings slightly leathery, with R1 and R2 strongly curved and parallel at that point (Figs. 507–509) . . . . . . . . . . . . . . . . . . . . . . . . 13 12b Pterostigma of front wings without these characters . . . . . . . . . . . . . . . . . . 18 13a Front wings wide and rounded, with strongly divergent apical veins (Fig. 507) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frenesia, p. 124 13b Front wings narrower, with subparallel apical veins (Figs. 508, 509, X) . 14

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14a Head and pronotum clothed with many long, erect setae (Fig. X). R1 of front wings ending at wing margin close to R2 (Fig. 508) . . . . Grensia, p. 127 14b Species without these characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15a Spurs: 1,1,1. Front wings strongly speckled with grey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chilostigmodes, p. 126 15b Spurs more numerous, and different coloring of front wings . . . . . . . . . . . 16 16a Front wings with large brown areas . . . . . . . . . . . . . . . . . Chilostigma, p. 128 16b Front wings rust-colored, with a longitudinal white stripe (Fig. XII) . . . . . 17 17a % intermediate appendages large, longer than preanal appendages (Fig. 501). & segments IX and X with paired mesal and lateral lobes (Fig. 505) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psychoglypha, p. 129 17b % intermediate appendages small and shorter than preanal appendages (Fig. 510). & tergite IX fused with segment X into sclerotized cap extended apically in 2 pointed lobes (Fig. 516) . . . . . . . . . . Desmona, p. 130 18a Front wings with longitudinal silver stripe produced by dense, flattened hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hesperophylax, p. 121 18b Front wings without this stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19a Maxillary palpi extremely large. In %, apex of 2nd segment extends beyond 1st antennal segment (Figs. 460, 461). . . . . . . . . . . . . . . Chyranda, p. 122 19b Maxillary palpi of usual length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20a Large, rust-colored, heavy-bodied insects. Inferior appendages in % vertical (Figs. 433, 434). Vulvar scale in & not trilobate (Fig. 438) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pycnopsyche, p. 118 20b Species without these characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21a Dorsal part of segment IX of % strongly developed and very conspicuous (Figs. 379, 386). & genitalia as in Figs. 383, 394 . . . . . . . . . . . . . . . . . . 22 21b Dorsal part of segment IX in % forming only a narrow band embedded in segment VIII (Figs. 433–435). & genitalia different . . . . . . . . . . . . . . . . 23 22a Small, pale species with sparsely speckled front wings . . . Arctopora, p. 107 22b Large and medium-sized species with strongly speckled front wings (Figs. VI–VIII). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lenarchus, p. 108 23a Superior appendages of % large and indented, forming lunules (Figs. 409, 414). Front wings strongly speckled or striped . . . . . . Clistoronia, p. 114 23b Species without these characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 24a Anastomosis of hind wings in regular zigzag pattern parallel to body (Fig. 333) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnephilus, in part, p. 98 24b Anastomosis of hind wings essentially unbroken and slanting toward body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

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25a Intermediate appendages of % much smaller than superior appendages and knob-shaped or lamelliform (Fig. 358). Ventral lobes of sternite IX in & very large (Figs. 361, 362) . . . . . . . . . . . . . . . . . . . . . . Asynarchus, p. 103 25b Intermediate and superior appendages of % differently proportioned . . . . 26 26a Front wings rust-colored and regularly speckled with many, tiny, dark spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anabolia, in part, p. 104 26b Front wings dark brown and regularly speckled with many, tiny light spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anabolia, in part, p. 104 26c Front wings greyish, pale, with large clear, indistinct areas. Sc and R1 of hind wings converge before reaching margin (see Fig. 321) (British Columbia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clostoeca, p. 123 26d Front wings with other coloring. Sc and R1 of hind wings parallel before reaching margin (transcontinental) . . . . . . . . . . Limnephilus, in part, p. 98

Genus Limnephilus Leach

Limnephilus Leach, 1815, p. 136 Monobasic type species: Phryganea rhombica Linnaeus Limnophilus Burmeister, 1839, p. 929 Monobasic type species: Phryganea rhombica Linnaeus Algonquina Banks, 1916, p. 121 Type species by original designation: Stenophylax? parvula Banks Anabolina Banks, 1903, p. 244 Type species by original designation: Anabolina diversa Banks Apolopsyche Banks, 1916, p. 121 Monobasic type species: Stenophylax minusculus Banks = Limnephilus dispar McLachlan Rheophylax Sibley, 1926, p. 107 Type species by original designation: Limnephilus submonilifer Walker Zaporota Banks, 1920, p. 342 Type species by original designation: Zaporota pallens Banks Revision: Ruiter, 1995 Limnephilus is a vast and very important genus containing two-thirds of the species included in the tribe. As a result, its characters are those of the tribe once the genera that are relatively easy to characterize have been taken away. Limnephilus is made up of species groups that are not sufficiently unlike one another to be raised to the genus level. It is therefore a heterogeneous genus and cannot be defined in terms of constant, clear-cut characters. Some understanding of the Limnephilidae, however, should make 98

it intuitively possible to distinguish a member of the genus Limnephilus from representatives of the other genera. The description given here is therefore simplified and applies only to the more typical species. Head relatively long and narrow with eyes protruding only slightly. Antennae stout, thick, and slightly shorter than front wings. Maxillary palpi long and slender. Pronotum proportional in length to length of head. Front legs with 1st tarsal segment generally longer than 2nd in both sexes. In some groups, sexual dimorphism is present, and 1st segment of % is shorter than 2nd (Fig. 332). Often a black brush of hairs at base of femur and apex of tibia. Wings medium-sized or fairly small. Front wings generally strip-like and barely wider at level of anastomosis than at base; obliquely truncated under apex. Hind wings markedly wider than front wings. Venation (Fig. 333): in front wings, discoidal cell narrow and 1.5 to twice as long as wide. Anastomosis often in regular zigzag pattern and parallel to body. In hind wings, discoidal cell fairly variable in length, anastomosis in zigzag pattern, as on front wings. Very abrupt forking of median vein with unusual arrangement of forks due to pronounced curve of all branches. % genitalia (Figs. 334–336): tergite VIII often developed into apicodorsal protuberance covered with spinules. Segment IX well developed and forming massive, rigid unit with the appendages. Superior appendages highly variable in shape and size, concave or not concave, spine-free or equipped with fine crenulations or large teeth. Intermediate appendages forming triangular, upward-slanting plates or spurs. Inferior appendages with part fused with segment IX very thin and free portion poorly developed, spine-free or dentate. Aedeagus often membranous at base and with slightly extensible extremity. Parameres usually present, shaped into long, slender rods ending in 1 or 2 branches equipped with fine hairs or stout spines. Occasionally one of the branches erectile. & genitalia (Figs. 337, 338): segment IX well developed, with both parts more or less briefly touching. Tergite IX never highly developed. Segment X in strongly sclerotized tube with more or less indented end. Appendages large and free or more or less fused with segment X or even completely lost. Ventral lobes of sternite IX usually very large and more or less extensively touching. Supragenital plate ogival, large, and free. Vulvar scale always trilobate with angular, subequal lobes. The genus Limnephilus is large with a Holarctic distribution. It contains some 200 species, half of which are Nearctic and 70 Canadian, some of them Holarctic. The genus has a broad ecological tolerance and a strong affinity for lentic habitats, lakes, permanent or temporary ponds, and marshes, but some species also inhabit rivers and springs. It is particularly well represented at low altitudes, with many subarctic species. Some forms, however, are found high in the mountains. 99

Genus Grammotaulius Kolenati

Grammotaulius Kolenati, 1848, p. 30 Type species designated by Milne, 1935: Phryganea interrogationis Zetterstedt Revision: Schmid, 1950b Species large, robust, and fairly lightly colored. Head relatively elongate with well-developed occipital part. Eyes relatively small. Antennae stout. Maxillary palpi long and slender. Pronotum relatively elongate. Macrochaetae on top of body well developed. Spurs: 1,3,4. Front wings forming regular, lightly speckled strips. Hind wings with very large anal area. R5 of both pairs of wings strongly lined with dark color (Fig. I). Venation: in front wings, discoidal cell 2.5 to 3 times longer than its petiole. Anastomosis strongly slanting forward. In hind wings, anastomosis strongly broken and parallel to body. Discoidal cell very long, and median forks abrupt. % genitalia (Figs. 339–342): a very massive, rigid unit. Tergite VIII without a spinule-covered area. Segment IX always elongate and very robust laterally and ventrally, with midportion strongly convex. Superior appendages stout, at times large and thin, and at others, short, massive, and thick, frequently notched and always equipped with stout crenulations. Intermediate appendages in subquadrangular lamellae and as long as superior appendages or else short and triangular. Lateral sclerites of segment X large. Inferior appendages weakly developed, slender, and barely protruding; fused portion forming only narrow fold at segment IX and free portion short, simple, and spine-free. Phallic apparatus very stout and similar to that of many Limnephilus. Parameres not erectile, simple or bifid.

Fig. I. Grammotaulius betteni Hill-Griffin 100

& genitalia (Figs. 343–345): tergite and sternite IX extensively touching. Tergite IX narrow and fairly elongate. Segment X consistently with 2 free, blunt, and prominent appendages, as very large, short, and highly sclerotized cylinder with fairly small indentation. Ventral lobes of sternite IX extremely large and blunt, contiguous for some distance, very prominent and usually as long as segment X. Supragenital plate small. Vulvar scale large, consisting of 3 usually wide, blunt, subequal, and fairly strong prominent lobes. Vaginal vestibule very wide, funnel-shaped, and horizontal, with inner and outer margins upturned. The genus Grammotaulius is small and of Holarctic distribution. It contains six Nearctic species, five of which are represented in our area. Larvae of these species inhabit primarily ponds, small lakes, and still, gently flowing watercourses. Genus Nemotaulius Banks

Nemotaulius Banks, 1906, p. 107 Type species by original designation: Grammotaulius brevilinea McLachlan Revision: Schmid, 1952a Head very elongate in relative terms, with vertex flat and occipital margin forming distinct ridge. No cephalic warts but a granulated surface covered with sparse, thick, very short macrochaetae. Pronotum two-thirds length of head, with same granulated surface and same macrochaetae, but in more pronounced form. Mesonotum with very sharp relief (Figs. II, III). Antennae thick, shorter than front wings, with 1st segment especially long. Maxillary palpi long and slender. Spurs: 1,3,4. Wings large. Front wings clearly indented and scalloped at extremity, with leathery membrane and covered with very short hairs; rust-brown and

Figs. II and III. Nemotaulius hostilis Hagen 101

slightly speckled. Hind wings very broad and only slightly indented under apex (Fig. III). Venation: in front wings, discoidal cell fairly wide and 3 times as long as its petiole. Anastomosis slightly oblique and faintly broken. In hind wings, discoidal cell and anastomosis similarly arranged. M very abruptly forks at level of beginning of discoidal cell. % genitalia (Figs. 346–349): tergite VIII with a small area of coarse, scattered spinules. Segment IX massive and well developed laterally. Superior appendages medium-sized, massively shaped, thick, concave, and spine-free. Intermediate appendages rather small, shaped like thick spurs, very divergent, and roughly as long as the superior appendages. Lateral sclerites of segment X large and transversely positioned. Inferior appendages fairly low, with prominent extremities; free portion massive, with upper surface developed into ridges separated by concavities. Phallic apparatus large and stout. Parameres not erectile, upward-curving, in shape of simple strips equipped with spines and setae. & genitalia (Figs. 350–352): tergite IX fairly narrow, triangular, and moderately developed. Appendages absent. Segment X extends beyond segment IX without any sharp discontinuity; it is in a short, massive, thick-walled, and hairy tube with fairly small indentation. Ventral lobes of sternite IX form 2 very large, only slightly prominent and extensively touching masses. Supragenital plate small. Vulvar scale very large, wide, lamelliform, and completely embedded in sternite VIII but its outline can be distinguished from latter. Median lobe small and wedged between subquadrangular lateral lobes. Vaginal vestibule narrow, deep, and horizontal. The genus Nemotaulius is small, Holarctic in distribution, and contains seven species, only one of which, hostilis Hagen, is widely distributed in North America. This species has been classified in the subgenus Macrotaulius Schmid. It is one of the largest Canadian caddisflies and is immediately recognizable by its leathery front wings notched at the extremity. It is common and distributed transcontinentally, from Newfoundland to British Columbia and Alaska, but is not found very far north. Its larvae occur in still water, especially in small ponds and marshes with substantial vegetation.

Genus Leptophylax Banks

Leptophylax Banks, 1900, p. 252 Monobasic type species: Leptophylax gracilis Banks Insects with small, narrow wings with a very pronounced shape. Eyes small with diameter barely half length of head. Occipital portion strongly developed, extending far behind eyes. Cephalic warts large, but indistinct since entire top of head is very grainy and bears numerous macrochaetae. 102

Antennae short and thick. Maxillary palpi thin and moderately long. Pronotum two-thirds as long as head, very angular and seta-bearing. Legs short and thick. Spurs: 1,3,4. Wings small, with front wings forming even, narrow, and very elongated ellipses. Hind wings very sharply indented under apex. Front wing membrane slightly leathery, faintly speckled, with very short, scattered hairs. Venation characteristic (Fig. 353): in front wings, discoidal cell narrow and 3 to 4 times longer than its petiole. Apical area short. Anastomosis very strongly and irregularly broken and slanting slightly forward. FIII occasionally with short petiole. In hind wings, discoidal cell also long. Anastomosis not strongly broken and slanting forward, in direction of body. % genitalia (Figs. 354, 355): tergite VIII without spinules. Segment IX moderately long laterally and ventrally; dorsally, it bears blunt, rounded lobe connected by a membrane to dorsal margin of intermediate appendages. Superior appendages large, notched, quadrangular, thin, concave, and spine-free. Intermediate appendages long and slender. Inferior appendages large and tapering, with free portion long, slender, and ending in fine, sclerotized point. Phallic apparatus large and well developed. Aedeagus strongly pleated at base and large at extremity, which is concave at top. Parameres long, thin, and seta-bearing. & genitalia (Figs. 356, 357): tergite IX short and small. Appendages large and free. Segment X in very blunt cone with thin and highly sclerotized walls and ending in 2 acute points. Ventral lobes of sternite IX very large, prominent, but not very thick, slightly concave on outer surface and far apart. Supragenital plate small. Vulvar scale embedded in sternite VIII, with median lobe long and thin, and lateral lobes narrow and concave on inner surface. Vaginal apparatus small. The genus Leptophylax is monobasic. Its only species, gracilis Banks, has not yet been captured in Canada. It is, however, widespread in the northeastern part of the United States, from Minnesota to New York. Genus Asynarchus McLachlan

Asynarchus McLachlan, 1880, p. 26 Type species by original designation: Asynarchus McLachlan = Phryganea lapponica Zetterstedt

fusorius

Revision: Schmid, 1954b Medium-sized species with dark wings finely speckled with a light color. Head not very wide and fairly elongate. Eyes not very protruding. Short and very long macrochaetae interspersed. Maxillary palpi long and slender. Pronotum short. Spurs: 1,3,4. 103

Wings of medium size but occasionally reduced in &. Front wings resembling those of Limnephilus in shape but wider at anastomosis and more rounded at tip. Hind wings very wide and not indented under apex. Front wings with a brown background; generally, a large number of tiny light-colored, irregularly shaped specks and larger spots on thyridium, on anastomosis, and at extremity of M4+5. Venation unremarkable. % genitalia (Figs. 358–360): tergite VIII spinule-free. Segment IX highly developed laterally where it is strongly convex. Superior appendages large or medium-sized, with fairly variable conformation; they are occasionally thin, fairly strongly concave, with sclerotized apical margin; at other times, apical margin curves inward, producing posterior surface with stout toothlike structure. Intermediate appendages always small, located under superior appendages, and in subtriangular lamellae or subspherical knobs. Lateral sclerites of segment X small. Inferior appendages with relatively prominent part fused with segment IX; only small portion of inferior appendages is free, and it consistently ends in 2 thick points. Phallic apparatus large. Aedeagus occasionally erectile at base. Parameres slender and usually bifid. Subapical branch short, flattened, and equipped with rows of setae. Apical branch slender, bearing small tubercles or very fine hairs. & genitalia (Figs. 361, 362): tergite IX short, barely prominent, and in contact with the sternite. Appendages often bulky, usually flattened laterally and fused together at base. Segment X under appendages, thinwalled and with only slight indentation. Ventral lobes of sternite IX always very large, high, massive, and generally extensively touching each other. Lateral portions of these lobes protrude, in some instances even extending beyond apex of segment X. Supragenital plate large. Vulvar scale half embedded in sternite VIII, with lateral lobes usually in shape of long, slanting bands. Vaginal vestibule large, triangular with inner margin not turned upward. The genus Asynarchus is Holarctic and Boreo-Alpine in distribution. It includes 15 species, 9 of which have been reported in our area. Several, such as batchawana Denning, montanus Banks, and mutatus Hagen, are transcontinental, whereas others are confined to the mountains of western Canada. Larvae inhabit very diverse environments: temporary or permanent marshy ponds, small lakes, more or less turbulent watercourses, or high-altitude glacial lakes.

Genus Anabolia Stephens

Anabolia Stephens, 1837, p. 229 Type species designated by Westwood, 1840: Limnephilus nervosus Curtis 104

Revision: Schmid, 1950a Medium-sized, heavy-bodied, and rust-colored insects. The && occasionally slightly brachypterous. Slightly variable head shape. Maxillary palpi fairly long. Pronotum relatively wide. Spurs: 1,3,4. Front wings wide, slightly truncated, rounded or parabolic at apex, with faintly leathery or grainy membrane. Front wings rust-colored and speckled with fine dark spots or else brown and evenly speckled with small light-colored spots but without any discolored areas. Venation fairly variable. In front wings, discoidal cell 1.3 to 3 times as long as its petiole. Anastomosis rectilinear and slanting forward or slightly broken and concave in direction of body. In hind wings, anastomosis always broken and slanting rearward. % genitalia (Figs. 363–366): tergite VIII spinule-free. Segment IX well developed laterally, rigid, and very sclerotized. End of anal cavity completely sclerotized and without discontinuity extending to inner surface of superior appendages. Latter are large, robust, highly sclerotized, massive, and almost consistently equipped with teeth and carinae. Intermediate appendages forming stout, more or less triangular, downward-directed lamellae, equal in length to superior appendages or longer. Lateral sclerites of segment X very stout, often prominent and horizontal. Inferior appendages with lower portion fused as narrow ridge and free portion forming long, slender, sharp, and strongly sclerotized, horizontal lobe. Phallic apparatus large. Aedeagus short, thick, and membranous at base. Parameres slender, virtually always bifid and spine-bearing. & genitalia very blunt and only slightly prominent (Figs. 367–369). Relative development of tergite and sternite IX fairly variable. Tergite IX often small. Appendages sometimes free and sometimes fused with base of segment X. The latter always small, cone-shaped and cleft dorsally and ventrally; base more or less wide, depending on whether it is fused or not with appendages. Ventral lobes of segment IX always very large and massive, occasionally fairly prominent and far apart, but at times blunt and contiguous. Supragenital plate small. Vulvar scale medium-sized, with fairly variably shaped lobes. The genus Anabolia is of Holarctic distribution and contains 15 species, 4 of which are Canadian: consocia Walker, ozburni Milne, bimaculata Walker, and sordida Hagen have a wide distribution. They inhabit marshes, temporary or permanent ponds, and still watercourses. Genus Halesochila Banks

Halesochila Banks, 1907, p. 119 Type species by original designation: Halesus taylori Banks 105

A beautiful, medium-sized species with black-striped front wings (Schmid 1950d) (Fig. IV). Head short and wide with very large eyes. Cephalic warts small. Maxillary palpi large and stout. Spurs: 1,3,3. Wings large with front wings regularly rounded at apex and hind wings broad and without indentation. Venation: in front wings, pterostigma is slightly leathery, and discoidal cell fairly wide and 1.5 to 2 times as long as its petiole. Anastomosis almost rectilinear and slanting slightly toward body, forward. In hind wings, anastomosis virtually unbroken and slanting slightly toward body, rearward. Discoidal cell beginning well ahead of median forks, which diverge little. % genitalia (Figs. 370–375): tergite VIII spinule-free. Segment IX short ventrally, well developed laterally, and fairly elongate dorsally. Superior appendages large, close together, located dorsally, fairly sclerotized, spine-free, strongly concave underneath, and contiguous. They form a vault over other genital parts. Intermediate appendages shaped into stout, upward-curving spurs. Lateral sclerites of segment X forming very thick, horizontal plates dividing the apical cavity in two. Inferior appendages forming thin, fairly narrow bands bordering segment IX and quite strongly concave on inner surface. Upper portion of inferior segments bearing a long, curved, slender, and barely sclerotized branch on inner surface. Phallic apparatus large and slender. Aedeagus with folds at base and small cupule at apex. Parameres long, narrow, with row of apical spines.

Fig. IV. Halesochila taylori Banks & genitalia (Figs. 376–378): segment IX very large, massive, and consisting of a single part. Fairly well developed dorsally and bearing 2 large lateral concavities. Ventrally, lower angles are strongly concave on inner surface but merge mesally. Segment X small, barely prominent, consisting of 2 short and wide horizontal lobes, 2 small, triangular appendages, and 1 very short, only slightly sclerotized tube. Supragenital plate large. Vulvar scale fairly small and composed of subequal lobes. Vaginal vestibule horizontal. 106

The genus Halesochila is monobasic. Its only species, taylori Banks, occurs in British Columbia, Alaska, and southward in small lakes and temporary or permanent ponds. It takes flight in September and October. Genus Arctopora Thomson

Arctopora Thomson, 1891, p. 1591 Monobasic type species: Phryganea trimaculata Zetterstedt Lenarchulus Schmid, 1952b, p. 164 Type species by original designation: Phryganea trimaculata Zetterstedt Revision: Schmid, 1952b Small insects with front wings widely spotted with grey (Fig. V). Head fairly elongate. Eyes and cephalic warts small, and vertex strongly bulging. Palpi thick and fairly short. Spurs: 1,3,4. Pronotum short. Wings fairly wide, with front wings fairly even in width, but clearly truncated at the apex. Hind wings not very wide and indented below apex. Venation: in front wings, discoidal cell wide and slightly shorter than its petiole. Anastomosis moderately broken, concave, and slanting slightly toward body rearward. FV often petiolate.

Fig. V. Arctopora trimaculata Zetterstedt % genitalia (Figs. 379–382): tergite VIII with tuft of long, stout setae apicodorsally. Segment IX elongate laterally; dorsally as long as laterally and forming paired or unpaired protuberances covering base of appendages. Superior appendages small, highly sclerotized, thick, and convex on all sides. Intermediate appendages small, blunt, and massive. Lateral sclerites of segment X fairly large and strongly prominent. Inferior appendages small, without any free portions; they are in thin, fairly wide lamellae and do not extend as far as middle angle of segment IX. Phallic apparatus 107

medium-sized. Aedeagus slender and nonerectile at base. Parameres also very narrow except at apex where they widen, bearing rows of hairs and divided into several points by semicircular indentations. & genitalia (Figs. 383–385): tergite IX well developed, prominent, and almost completely overhanging segment X, which is small and not protruding. This segment annuliform, barely sclerotized, and complex in outline. Appendages absent. Ventral lobes of sternite IX large, suboval, massive, and contiguous for some distance. Supragenital plate large but only slightly prominent. Vulvar scale fairly large and half embedded in sternite VIII; median lobe long and narrow, lateral lobes subquadrangular, divergent, and oblique. Vaginal vestibule V-shaped. The genus Arctopora is small and of Holarctic distribution. It contains three species, all of which are Nearctic. A. trimaculata Zetterstedt is Holarctic and occurs in Alaska; it has 2 small sclerotized points on the dorsal margin of segment IX. A. pulchella Banks is transcontinental and has been captured from Newfoundland to British Columbia; it is distinguished by 2 large, blunt, contiguous lobes extending the dorsal margin of segment IX. A. salmon Smith has been reported in Idaho only and has blunt, distantly spaced lobes on the dorsal margin of segment IX. These species live in marshes, temporary or permanent ponds, and still streams. Genus Lenarchus Martynov

Lenarchus Martynov, 1914, p. 222 Type species designated by Schmid, 1952b: Asynarchus productus Morton Revision: Schmid, 1952b Medium-sized or large insects with finely and extensively speckled wings (Figs. VI–VIII). Head fairly consistently short and wide, with prominent eyes. Maxillary palpi long and stout. Legs often with dark-colored bands and numerous long erect spines. Spurs: 1,3,4. Wings large or medium-sized and somewhat variable in shape since they are truncated or more or less rounded at tip. Hind wings wider than front wings and not indented. Venation: in front wings, discoidal cell of variable width, 1 to 2.5 times as long as its petiole. Anastomosis broken to variable degree, more or less concave, and slanting fairly sharply toward body, forward. In hind wings, anastomosis strongly broken, concave, parallel to body or slanting slightly rearward. % genitalia: tergite VIII spinule-free. Genital appendages large, massive, rigid, and arising from a robust segment IX. Latter fairly elongate laterally, generally short ventrally, and always strong dorsally. When free, superior appendages large, thick, very sclerotized, massive, and equipped 108

with points. Intermediate appendages developed into elongated plates. Inferior appendages protruding little and so firmly fused with segment IX that suture is barely visible; these appendages usually consist of 2 thin plates extending apical margin of segment IX and included as part of segment since they compensate for a certain shortening of latter. Phallic apparatus long and stout. Aedeagus with folds at base. Parameres slender, ending in 2 flattened, converging branches equipped with rows of setae. & genitalia fairly variable: segment IX generally composed of 2 distinct parts. Appendages usually present. Segment X developed into more or less sclerotized tube with more or less uneven edge and thickened, often hairy walls. Ventral lobes of sternite IX occasionally very large and blunt, but usually long and slender like appendages and far apart. Supragenital plate short and wide. Vulvar scale relatively large. Lateral lobes more or less quadrangular but always oblique and divergent. The genus Lenarchus is small and of Holarctic distribution. It is divided into three subgenera on basis of appendage shape and nature of % dorsal plate. The species occur in lentic biotopes: small lakes, temporary or permanent ponds and marshes. Some are found at fairly high altitude. 1a Dorsal portion of segment IX of % bulging, not stretched out and equipped with 2 divergent spurs (Figs. 386–388). . . . . . . . . . . Prolenarchus, p. 109 1b Dorsal portion of segment IX of % not bulging but stretched out into bifid plate and overhanging other genital parts (Figs. 390–392) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lenarchus, p. 108 1c Dorsal portion of segment IX of % neither bulging nor stretched out, but serving as base for large dorsal plate formed by fusion of superior appendages (Figs. 397–399) . . . . . . . . . . . . . . . . . . . . . . . . . Paralenarchus, p. 111

Subgenus Prolenarchus Schmid

Prolenarchus Schmid, 1952b, p. 170 Type species by original designation: McLachlan

Asynarchus

bicornis

Front wings finely and evenly speckled with grey (Fig. VI). % genitalia (Figs. 386–388): dorsal portion of segment IX forming a large, very obtuse bulge overhanging only base of appendages. Two small, very sclerotized, divergent, spurlike horns lie beneath this rim, between superior appendages. Latter of fairly unusual shape and size, uniformly sclerotized, and concave posteriorly. Intermediate appendages large, prominent, and curved upward. Lateral sclerites of segment X mediumsized, massively shaped, and positioned for some distance against intermediate appendages. Inferior appendages without any free portion or any prominent angle and included in width of segment IX. Aedeagus thin and 109

nonerectile. Parameres slender and enlarged at apex into concave, setabearing plate. & unknown. The subgenus Prolenarchus contains only two species, one of which, keratus Ross, is found in the Nearctic region. It is known from only a few isolated specimens captured in Quebec, Ontario, Michigan, Alberta, and Minnesota.

Fig. VI. Lenarchus (Prolenarchus) keratus Ross Subgenus Lenarchus s. str. Martynov

Lenarchus Martynov, 1914, p. 222 Type species designated by Schmid, 1952b: Asynarchus productus Morton Front wings usually strongly speckled with brown, with light-colored areas in middle of wing, on external side of anastomosis, and at tip of M3+4 (Figs. VII, VIII). % genitalia (Figs. 390–393): segment IX very elongate all around, forming dorsal plate variously developed and shaped, but consistently stout. In most species, this structure has reversed direction of appendages, which slant downward. Superior appendages fused with underside of dorsal plate, not very large, blunt, convex, fairly sclerotized, and laterally positioned. Intermediate appendages developed into points or plates. Lateral sclerites of segment X weakly developed. Inferior appendages very short, included in width of segment IX, sometimes with very long, slender, and more or less sclerotized free portion. Aedeagus with entire basal portion erectile. Parameres narrow and bifid at tip where they are surmounted by rows of setae. & genitalia (Figs. 394–396) with varied characters. Appendages present and slender or absent. Segment X forming variously developed tube with more or less indented edge. Ventral lobes of sternite IX small, 110

Fig. VII. Lenarchus (Lenarchus) productus Morton

Fig. VIII. Lenarchus (Lenarchus) rho Milne strongly protuberant, and far apart or else, large and contiguous for long distance. Lateral lobes of vulvar scale large but not very distinct from sternite VIII. Median lobe long and deeply wedged between the lateral lobes. The subgenus Lenarchus is of Holarctic distribution and contains three North American species. L. crassus Banks is distributed across the country from Newfoundland to British Columbia and has a wide dorsal plate ending in 2 triangular points. L. expansus Martynov (Siberia, Alaska and Yukon) has a longer dorsal plate ending in 2 widely spaced quadrangular lobes. L. rho Milne (British Columbia and southward) has a narrow, drawn-out dorsal plate ending in 2 small, divergent lobes.

Subgenus Paralenarchus Schmid

Paralenarchus Schmid, 1952b, p. 191 Type species by original designation: Limnephilus vastus Hagen 111

Wings of very variable size and coloration. When of usual size, anterior wings densely spotted with brown. % genitalia (Figs. 397–399): segment IX very elongate laterally and moderately developed dorsally. Dorsal plate not formed by segment IX but by superior appendages that are strongly developed and more or less fused together. Plate systematically large, very sclerotized, and forming vault over other genital parts. Intermediate appendages in shape of stout spurs projecting horizontally or occasionally reduced and small. Lateral sclerites of segment X always very large. In species with long intermediate appendages, sclerites form 2 wide, funnel-shaped concavities but in species with reduced appendages, each sclerite consists of 2 long points that are longer than appendages themselves and that can eventually replace them. Inferior appendages generally included in width of segment IX and are occasionally equipped with long, free, slender branch. Phallic apparatus similar to that of Lenarchus. & genitalia (Figs. 400–402): tergite IX short and wide. Appendages large and free. Segment X developed into stout tube with more or less indented edge. Ventral lobes of sternite IX long, slender, and far apart, although internally extending to median line. Supragenital plate small. The subgenus Paralenarchus is small and found only in the mountains of the western Nearctic region. All five species have been captured in our area and southward. Genus Philarctus McLachlan

Philarctus McLachlan, 1880, p. 80 Monobasic type species: Philarctus bergrothi McLachlan Small, thick, heavy-bodied insects with rather small wings. Head fairly elongate, with eyes and cephalic warts small. Antennae thick and shorter than front wings. Maxillary palpi poorly developed. Pronotum relatively elongate, with dense pilosity. Legs stout but not very long. In %, front femur and tibia strongly thickened. Latter slightly flattened, without black spines and ending in a carina. Tibia and femur bear thick, coarse, black brush over their entire length. Short, flat triangular apical spur. Tarsal segments very short (Fig. 403). Spurs: 1,3,4. Wings similar to those of Limnephilus but small, parabolic at apex, and dull yellow-brown. Venation: in front wings, discoidal cell 1.5 to 2 times as long as its petiole. Anastomosis unbroken and slightly oblique. In posterior wings, discoidal cell equally long, but anastomosis more strongly broken and parallel to body. FIII pointed. % genitalia (Figs. 404–406): tergite VIII spinule-free. Segment IX strongly developed, very long laterally, and dorsally forming a blunt protuberance densely covered with long setae and extensively tucked around 112

superior appendages. These very large, massive, regularly and fairly strongly sclerotized, contiguous in back of anal cavity, and sealing off the latter like a pair of window shutters. Their inner surface therefore not visible, and lower inner angles protrude and curve downward. Intermediate appendages (or lateral sclerites?) located under superior appendages and shaped like horizontal disks with slightly concave upper side. Inferior appendages with barely prominent fused part and long, slender free part inerm. Aedeagus similar to that of many Limnephilus and ending in truncate tip. Parameres slender, but strongly widened at apex and bearing spines on margin. & genitalia (Figs. 407, 408): tergite IX short, blunt, protuberant, and extending downward on sides. Appendages large, free, and prominent. Segment X large, blunt, thick, slightly sclerotized, very hairy, and in shape of fairly strongly indented tube with broad cleft on lower surface. Ventral lobes of sternite IX fairly small, massive, and widely spaced. Supravaginal plate short and wide. Median lobe of vulvar scale long and narrow. Lateral lobes short, thick, subtriangular, horizontally positioned, and wider than long. The genus Philarctus is small, and widespread in the Boreo-Alpine regions of Palaearctic Asia and North America. Six species have been described, but this number should be reduced to probably three. A single species, quaeris Milne, lives in our area, but it is probably synonymous with the Central Asian species przewalskii McLachlan. It is widespread from the Northwest Territories to Manitoba and southward to Colorado and occurs in marshes, ponds, small lakes, and slow-moving watercourses. Genus Clistoronia Banks

Clistoronia Banks, 1916, p. 119 Monobasic type species: Halesus magnificus Banks Beautiful, large insects with brightly colored front wings. % genitalia: tergite VIII spinule-free. Segment IX very short all around but extending very high. Superior appendages large, stout, crescent-shaped, and very sclerotized. In lateral view, triangular with strongly indented apical margin. Intermediate appendages in horizontal spurs that are either free or fused. Subanal plate large. Lateral sclerites of segment X forming 2 hairy, sclerotized, horizontal lobes positioned against segment IX and dividing apical cavity. Inferior appendages weakly developed, with narrow, fused portion and long, cone-shaped, inerm portion projecting inward. Phallic apparatus large. Aedeagus with folds at base. Parameres spine-bearing, simple or bifid. & genitalia: tergite IX more or less reduced. Variably sized appendages, strongly flattened dorsoventrally and completely fused either with 113

segment IX or segment X. The latter varies in shape and is blunt or long and slender. Ventral lobes of sternite IX differ greatly in shape, depending on subgenus. Subgenital plate present or absent. Lateral lobes of vulvar scale angular, subtriangular, and rounded. Clistoronia is divided into two subgenera which are fairly different with respect to the characters of the body, wings, and & genitalia but are very similar with respect to the characters of the male genitalia. 1a Medium-sized species with front wings bearing longitudinal brown and lightcolored stripes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clistoroniella, p. 115 1b Large species with front wings strongly speckled with dark brown and lightcolored spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clistoronia, p. 114

Subgenus Clistoronia s. str. Banks

Beautiful, large insects. Front wings strongly speckled, more than 17 mm in length. Spurs: 1,3,3. Macrochaetae of top of head short, fine, and sparse, whereas those of pronotum are very stout and profuse. Wings fairly large. Front wings widen slightly at level of anastomosis and are truncated at apex. Hind wings fairly narrow. Venation: in front wings, discoidal cell 1.5 to 2 times as long as its peduncle. Anastomosis slightly broken, almost rectilinear, and slanting forward. In hind wings, anastomosis strongly broken and fairly distinctly concave in direction of body. FIII slants sharply at base. % genitalia (Figs. 409–411): segment IX with distinctly convex middle portion that is indented above the inferior appendages. Superior appendages large and strongly sclerotized, with apical margin strongly indented. In front view, upper and lower margins blunt, highly sclerotized, and continuous with stout inner carina that is also highly sclerotized and forms a concave posterior surface. Intermediate appendages fused into single, cylindrical part. Inferior appendages embedded in segment IX. Parameres ending in 2 stout, fairly long, profusely spined branches. & genitalia (Figs. 412, 413): tergite IX vestigial. Appendages very large, flattened dorsoventrally, and covering segment X. The latter comprises long, slender, bifid dorsal part and highly sclerotized, deeply indented ventral part. Lobes of sternite IX large, concave toward back, and strongly splayed horizontally. Supragenital plate present and triangular. Median lobe of vulvar scale pointed and of equal width and length. Lateral lobes subtriangular. The subgenus Clistoronia contains three species located in the western Nearctic region. Only one occurs in our area, magnifica Banks, which is found in Alaska, Alberta, British Columbia, and southward, where it inhabits ponds and small lakes; formosa Banks has been recorded from Idaho. The former has anterior wings intensely and regularly spotted with 114

dark brown; the latter has reddish-brown anterior wings irregularly dotted with colorless spots. Subgenus Clistoroniella Schmid

Clistoroniella Schmid, 1955, p. 155 Type species by original designation: Asynarchus flavicollis Banks Medium-sized insects, with front wings less than 15 mm in length and bearing a series of longitudinal brown stripes. Spurs: 1,3,4. Macrochaetae on top of body long, fine, and sparse. Wings distinctly larger than those of Clistoronia s. str. Front wings widen considerably at level of anastomosis and bluntly rounded at apex. Hind wings very wide and blunt. Venation fairly different from that of true Clistoronia. In front wings, discoidal cell fairly narrow and slightly longer than its petiole. Anastomosis regularly and strongly broken and parallel to body. In hind wings, anastomosis also strongly broken, but slanting toward body, rearward. Discoidal cell fairly long, beginning slightly ahead of median fork, which is not strongly divergent. % genitalia (Figs. 414–416): segment IX with middle portion faintly convex and without indentation above inferior appendages. Superior appendages not very robust. Small inner carina is only part of appendages that is highly sclerotized, and is not sufficiently prominent to create a posterior surface. Intermediate appendages completely distinct. Inferior appendages weakly embedded in segment IX. Parameres simple and strongly spine-bearing. & genitalia (Figs. 417, 418): tergite IX narrow and elongate. Appendages small, rectangular, contiguous, and covering segment X. The latter comprises dorsal part in 2 triangular, wing-shaped structures and ventral part forming large fold. Ventral lobes of sternite IX large and virtually contiguous. Supragenital plate absent. Median lobe of vulvar scale angular and of greater length than width. Lateral lobes forming rounded ovals. The subgenus Clistoroniella is monobasic, the only representative of which, flavicollis Banks, is found in British Columbia and Alaska.

Genus Platycentropus Ulmer

Platycentropus Ulmer, 1905b, p. 13 Type species by original designation: Halesus maculipennis Kolenati = Phryganea radiata Say Hylepsyche Banks, 1916, p. 121 Monobasic type species: Limnephilus indistinctus Walker 115

Fairly large insects with front wings extensively spotted with brown or yellow. Head short and wide, with large, protuberant eyes and very small cephalic warts. Maxillary palpi very long and stout. In %, 1st segment half length of 2nd. Spurs: 1,3,3. In both sexes inner apical spur of hind legs lanceolate with 3 carinae formed by dense alignments of very short hairs (Fig. 419). Wings not very large. Front wings narrow at base, widening considerably at level of anastomosis and weakly truncated under apex. Hind wings blunt and slightly wider than front wings. Venation (Fig. 420): in front wings, discoidal cell wide and distinctly longer than its petiole. Anastomosis fairly strongly broken, concave in direction of body and subparallel to latter. A2 absent on its posterior half. In hind wings, anastomosis also strongly broken, concave in direction of body and slants slightly rearward. Discoidal cell long. % genitalia (Figs. 421–423): tergite VIII spinule-free. Genital parts generally barely prominent, forming rigid, massive unit as result of considerable development of segment IX. Dorsally, latter comprises 1 plate or 2 highly sclerotized points overhanging base of genital parts. Segment IX strongly constricted in middle portion, increasing in size dorsally and ventrally. Superior appendages fairly large, convex, very blunt, and strongly sclerotized. Intermediate appendages usually in shape of 2 thick, highly sclerotized, horizontal spurs. Lateral sclerites of segment X large, horizontally positioned, and dividing apical cavity. Subanal plate large. Inferior appendages small, barely prominent, in shape of fairly short, faintly protruding lamellae extending segment IX, but without any free portion. Phallic apparatus very large. Aedeagus fairly narrow, occasionally pleated at base, and equipped with apical points. Parameres fairly variable: at times, in short spines and at other times, very long and setabearing. & genitalia (Figs. 424, 425): tergite IX long, stout, and extended without discontinuity by segment X, which has blunt cone shape and is fairly strongly indented, with thick walls and sclerotized inner surfaces. In amicus Hagen, there are free appendages. Ventral lobes of sternite IX developed into short, wide, transverse plates with fairly protruding external angle. Supragenital plate small and short. Vulvar scale comprising long, slender median lobe and straight, subquadrangular lateral lobes. The genus Platycentropus is small and exclusively Nearctic in distribution. It contains three species that inhabit the central and eastern parts of the continent. In radiatus Say (from Nova Scotia to Manitoba), the dorsal part of segment IX forms a simple bulge. In indistinctus Walker (from Newfoundland to Ontario), this part is bilobed and curved downward between the superior appendages. P. amicus Hagen is distributed from Quebec to Alberta. The dorsal part of segment IX forms 2 large horizontal points. These species are strongly eurythermal and inhabit a 116

large number of biotopes ranging from cold streams to marshes, lakes, and warm ponds. They have a preference for dense aquatic vegetation. Genus Hydatophylax Wallengren

Hydatophylax Wallengren, 1891, p. 73 Monobasic type species: Stenophylax infumatus McLachlan Astenophylax Ulmer, 1907, p. 32 Type species designated by Milne 1935: Phryganea arga Harris Revision: Schmid, 1950c Large or medium-sized, variously colored, and often beautiful species (Fig. IX). Head very short and wide with large ocelli and strongly protruding eyes. Antennae thick, crenulated on lower face, with 1st segment as long as head. Maxillary palpi long and stout. Spurs: 1,3,4. Wings with varied coloring, plain or speckled, and variously shaped but usually fairly elongate. In front wings, R1 often darkened at base, and on hind wings, joining Sc before reaching margin. % genitalia (Figs. 426–429): tergite VIII with very fine spinules. Segment IX short and strongly elongate vertically. Anal cavity deep, generally noneversible, lined by superior appendages, which are wide and form a sort of horizontal platform extending backward and mesally. Intermediate appendages small, cylindrical, and fused with mesal edges of upper appendages. Lateral sclerites of segment X bulky, triangular, and serving as supports for superior appendages. Inferior appendages long, vertical, and extensively fused with segment IX with which they form a unit; their free portion long and slender. Phallic apparatus very small. Aedeagus spinefree, thin, and curved upward. Parameres located above aedeagus and reduced to thin filament arising from membranous bulb.

Fig. IX. Hydatophylax argus Harris 117

& genitalia (Figs. 430–432): segment IX very elongate vertically. Tergite IX and segment X firmly fused into short, only slightly indented tube with no prominent angles. Sternite IX forms soft, nonprotruding membranous area, very elongate vertically and rarely divided into 2 lobes. Supragenital plate small. Vulvar scale fairly prominent, in one piece, single-lobed, thick, fleshy, highly sclerotized, and with complex relief on inner surface. Vaginal opening wide but situated so low that it is no more than narrow, transverse slit. The genus Hydatophylax is small and of Holarctic distribution. It contains 12 species, only 4 of which are native to this country. They are easily recognizable by their appearance. H. argus Harris (from Newfoundland to Manitoba) is characterized by its large size and beautiful coloring (Fig. IX). H. victor Banks (Nova Scotia and Quebec) is a small orange species. H. hesperus Banks is a large yellow-brown form, resembling a Pycnopsyche and found in British Columbia and southward. H. variabilis Martynov is found in the Arctic regions of Asia, but has also been captured in Alaska; it is recognizable by its dark brown wings and occasionally reduced size. Larvae of these species inhabit small and medium-sized watercourses. Genus Pycnopsyche Banks

Pycnopsyche Banks, 1905, p. 9 Type species by original designation: Limnephila scabripennis Rambur Allegophylax Banks, 1916, p. 118 Type species designated by Fischer, 1969: Phryganea subfasciata Say Eustenace Banks, 1916, p. 118 Monobasic type species: Stenophylax limbatus McLachlan Revision: Betten, 1950 Fairly large, heavy, strong-bodied, and consistently rust or orange colored insects. Antennae thick and shorter than front wings. First segment of maxillary palpi in % half length of 2nd segment. Variable number of spurs: 1,2,2, 1,3,3 or 1,3,4. Wings not very large and variously shaped. Venation: in both pairs of wings, discoidal cell short and wide. Anastomosis a strongly concave curve. % genitalia (Figs. 433–436): tergite VIII virtually always with neoformations consisting of 2 dorsal appendages or 2 dorsolateral plates or 2 lateral appendages flanking superior appendages on external side. These formations in shape characteristic of species. Segment IX short and fairly 118

elongated vertically. Often shallow, noneversible anal cavity. Superior appendages of variable size, lining anal cavity and forming small horizontal platform. Intermediate appendages cone- or spine-shaped, often small, occasionally vestigial and fused with inner margins of superior appendages. Lateral sclerites of segment X forming unpaired transverse plate. Inferior appendages vertical, not prominent, with long, slightly bulging fused portion and well-developed, often dentate, free portion, which at times vicariously replaces reduced intermediate appendages. Phallic apparatus small. Aedeagus fairly large and membranous. Parameres reduced to membranous bulb bearing tuft of setae or equipped with spines or ending in single filament. & genitalia (Figs. 437, 438): segment IX a single piece. Segment X very short and so thoroughly integrated with segment IX that boundary no longer visible. Appendages weakly prominent and occasionally indistinct. Supragenital plate fairly large, subtriangular, and separated from ventral side of segment IX by large membranous area. Vulvar scale as in Hydatophylax, single-lobed, thick, distinctly sclerotized, and with sharp inner relief. The genus Pycnopsyche is medium-sized and exclusively Nearctic in distribution. It contains 15 heavy, strong-bodied, orange-colored species, 11 of which have been captured in our area. Most of them are located in eastern Canada, but two, subfasciata Say and guttifer Walker, extend westward to the foot of the Rocky Mountains. They inhabit primarily cold, forest watercourses and streams flowing through deciduous woods. They take flight at the end of the summer and in the fall.

Genus Sphagnophylax Wiggins and Winchester

Sphagnophylax Wiggins and Winchester, 1984, p. 1853 Type species by original designation: Sphagnophylax meiops Wiggins and Winchester Very distinctive insects, with short broad wings, front wings with prominent erect bristles, stout-bodied, with short, thickened appendages. Head comparatively very long, with occipital region very prominent. Eyes small and with large ommatidia. First segment of antennae shorter than head. Antennae and palpi thick. % foreleg with dense strips of short, black setae along ventral edge of femora and facing edge of tibia. Spurs 1,2,2. Wings short and broad, little longer than abdomen, with thick veins and densely covered with erect, strong bristles on veins and membrane. Venation (Fig. 451): individually quite instable and irregular. In front wings, discoidal and thyridial cells large and equal in size. Apical area 119

short. In hind wings, discoidal and thyridial cells of inequal length. Four anal veins. % genitalia (Figs. 445–450): segment IX short dorsally and ventrally, longer laterally, and fairly extended vertically. Anal cavity shallow. Superior appendages appearing long laterally; they form sort of horizontal platform extending mesally. Intermediate appendages small, briefly cylindrical, and fused with mesal edge of superior appendages. Lateral sclerites of segment X prominent, strongly clavate, coarsely granulated, and asymmetrically crossed over each other. Inferior appendages vertical, not prominent, with long, slightly bulging fused part and well-developed, coarsely granulated, and dorsoventrally depressed free portion. Aedeagus flattened dorsoventrally and with spatulated apical part. Parameres spiniform, spiniferous, and arising from base of aedeagus. & genitalia (Figs. 452–454): tergite and sternite IX broadly fused with each other; ventral lobes not prominent and widely separated. Segment X not prominent and forming 2 dorsal and 2 lateral blunt lobes. Anal cavity widely gaping and open ventrally. Supragenital plate membranous and broadly triangular. Vulvar scale with median lobe long and shortly bilobed; lateral lobes narrow and in very oblique position. Vaginal apparatus broad. Sphagnophylax has been subjected to elaborate but inconclusive taxonomic speculation (Wiggins and Winchester, 1984). These authors considered it as a Limnephilinae tribe incertae sedis. I tentatively consider this genus as related to Hydatophylax, Pycnopsyche, and Philocasca, since the % genitalia show striking similarities in many characters. However, the & genitalia do not confirm this relationship. The genus Sphagnophylax comprises a single species, meiops Wiggins et al., known from Yukon Territory and Northwest Territories. Adults were taken near a small isolated pond, in the Arctic maritime tundra. Genus Philocasca Ross

Philocasca Ross, 1941, p. 111 Type species by original designation: Philocasca demita Ross Revision: Wiggins and Anderson, 1968 Medium-sized species with yellow-brown front wings. Spurs: 1,2,2, 1,2,4, or 1,3,4. Wings large and rounded, with anal area of hind wings particularly well developed. Venation with no distinguishing features. % genitalia (Figs. 440–442): tergite VIII without spine-bearing areas. Segment IX very elongate laterally and ventrally, but very short dorsally. Segment X forming deep, horizontal anal cavity, extensively lined by 120

superior appendages that are oval and slightly indented at extremity. Intermediate appendages reduced and forming simple spurs, and continuous with inner apical angles of superior appendages. Lateral sclerites of segment X developed into small lobes that are faintly protruding in Canadian species, but very prominent and longer than intermediate appendages in other, more southerly forms. Inferior appendages small, low, forming 2 short knobs positioned against segment IX. Phallocrypt short and wide. Phallic apparatus short and thick. Aedeagus completely membranous with clearly visible spermatic duct. Parameres forming simple, highly sclerotized spines. & genitalia (Figs. 443, 444): segment IX composed of 2 well-defined parts. Tergite IX not very distinct from segment X which forms 2 slender and slightly divergent lobes, with 2 small points on inner surface framing anus laterally. Ventral lobes of sternite IX barely prominent and fused laterally with vulvar scale. Subanal plate absent, and supragenital plate large and ogival in shape. Vulvar scale without median lobe, lateral lobes very wide and slightly indented on upper margin. The genus Philocasca is exclusively western Nearctic in distribution and comprises seven species, all of which are rare and occur very locally. One of them, demita Ross, has terrestrial larvae. Only two species, thor Nimmo and alba Nimmo, have been captured in Alberta. P. banksi Denning and antennata Banks have been found in Idaho, Washington, and Montana. Genus Hesperophylax Banks

Hesperophylax Banks 1916, p. 118 Monobasic type species: Platyphylax occidentalis Banks Revision: Parker and Wiggins, 1985, p. 2443 Large or medium-sized insects with front wings striped with brown and silver, producing disruptive mimetic coloring. Head short and very wide. Eyes large. Dorsum of body completely covered with very dense, fine, silky, hairs. Maxillary palpi very slender. Spurs: 1,2,2. Wings usually large and similar in shape to those of Limnephilus, but hind wings relatively narrow and without any subapical indentation. Front wings with very dense pilosity of fine hairs, which in itself creates fairly vivid designs. In subradial cell, a silver stripe formed by row of silky hairs perpendicular to veins. The stripe divided in two in apical area. These stripes underlined with grey, creating a certain resemblance with genus Psychoglypha. Venation: in front wings, discoidal cell narrow and much longer than its petiole. Anastomosis slightly broken but concave and slanting in direction of body, anteriorly. In hind wings, discoidal cell and anastomosis similar to those in front wings. 121

% genitalia (Figs. 455–457): tergite VIII with large spinule-covered area. Segment IX short all around. Superior appendages very large, blunt, thin, and only slightly sclerotized, with lateral portion large, vertical, and concave. Inner side of superior appendages completely lining end and floor of vast, noneversible apical cavity. Intermediate appendages completely fused into unpaired appendage, situated at entrance to apical cavity and inserted on lateral sclerites of segment X. These form bulky apical margin. Inferior appendages fairly large and subvertical, ending in long, fairly slender, free portion. Phallic apparatus fairly small, short, and upward-curving. Aedeagus simple, slender, and nonerectile. Parameres much shorter, with basal part surmounted by 2 tufts of thin, sclerotized, spineshaped, and variously twisted and indented strips. & genitalia (Figs. 458, 459): sternite VIII forming large fold, clothed with long setae, on apical margin. Tergite IX very short and weakly developed. Appendages free, long, and thin or shaped into wide horizontal disks, fused with dorsal side of segment X, which is short, with fairly little sclerotization and covered with fine hairs. Ventral lobes of sternite IX small, complex, barely prominent, and fused with sides of large, thick, prominent supragenital plate. Vulvar scale with median lobe long and narrow and lateral lobes developed into oblique, slightly protruding transverse strips. The genus Hesperophylax is of Nearctic distribution and contains five closely related species, four of which occur in our area: occidentalis Banks, alaskensis Banks, consimilis Banks, and designatus Walker. The 1st two occur in the mountains of western Canada, whereas designatus Walker is transcontinental; consimilis is recorded from Idaho. These species are fairly eurythermal and inhabit all types of watercourses, particularly small and occasionally temporary rivers. They also occur in still water in the north and at high altitudes.

Genus Chyranda Ross

Chyranda Ross, 1944, p. 283 Type species by original designation: Asynarchus centralis Banks Fairly small, slender insects, uniformly rust-colored. Antennae fine and very long, particularly in % where they clearly extend beyond apex of front wings. Maxillary palpi considerably developed in %. First segment tiny, and apex of 2nd segment on level with 4th antennal segment (Figs. 460, 461). Spurs: 1,3,3. Wings large; in %, narrow and elongate, with apex forming narrow parabola. In &, shorter and wider. Venation: FIII pointed on front wings, otherwise no distinctive features. 122

% genitalia (Figs. 462–464): genital parts slightly prominent and not invaginated. Tergite VIII spinule-free. Segment IX short. Superior and intermediate appendages medium-sized, shaped into vertical plates forming succession of concavities, and with inner surfaces uniformly sclerotized. Segment X developed into large plate raised in middle into tubelike structure containing anal opening. Subanal plate absent. Inferior appendages very wide, completely fused with segment IX and without any free portion. Aedeagus very small and spine-free. Parameres spiniform, variable in size, and generally asymmetrical, with right paramere longer than left. & genitalia (Figs. 465, 466): tergite IX short. Segment X small, forming 2 large upper lobes and 1 smaller lower lobe. Ventral portion of sternite IX not prominent. Supragenital plate small. Vulvar scale with 2 large lateral lobes but no median lobe. Vaginal apparatus wide and complex. The genus Chyranda is monobasic. The only species, centralis Banks, has a very large but discontinuous range. It is widely distributed in the western provinces where it is common, and has been reported in Quebec but is uncommon there. Larvae are found primarily in streams and seem to have a preference for submerged clusters of dead leaves. Genus Clostoeca Banks

Clostoeca Banks, 1943, p. 352 Type species by original designation: Clostoeca sperryae Banks = Anisogamus disjunctus Banks Medium-sized species with front wings indistinctly and extensively speckled with grey. Antennae fine and as long as front wings. Maxillary palpi of medium length. Legs long and thin. Spurs: 1,3,4. Wings large with apex in shape of long parabola in % and clearly smaller and truncated in &. Venation: in front wings, discoidal cell not longer than its petiole. Anastomosis faintly broken, slightly concave, and virtually parallel to body. In hind wings, Sc and R1 converge or have a point of contact before they reach margin. Anastomosis strongly broken and slanting sharply toward body, rearward. % genitalia (Figs. 467–469): tergite VIII spinule-free. Segment IX short all around, completely membranous dorsally, strongly invaginated under tergite VIII, and slanting downward. Superior appendages situated dorsolaterally and forming small sclerotized cones. Anal opening between superior appendages. Intermediate appendages completely fused into small, slender, highly sclerotized point inserted in large, broadly rounded plate lying against middle margins of segment IX. This plate forms several concavities and 2 sharp toothlike structures at base of median appendage. Inferior appendages in shape of elongated cones, projecting upward. 123

Phallic apparatus small, short, and thick. Aedeagus simple. Parameres developed into regularly shaped rods equipped with a few apical spines. & genitalia (Figs. 470, 471): tergite IX developed into narrow, regularly shaped tube ending in 2 blunt lobes almost completely enclosing segment X. The latter also tubular, but very short and open on dorsal side. Ventral lobes of sternite IX only slightly prominent and widely separated by membranous space. Supragenital plate membranous. Vulvar scale simple, without median lobe, forming thick, fleshy fold with broad indentation in middle and wide depression on inner surface. The genus Clostoeca is monobasic. Its single species, disjuncta Banks, is quite widespread in the mountains of the western Nearctic region, including British Columbia. It inhabits turbulent streams. Genus Frenesia Betten and Mosely

Frenesia Betten and Mosely, 1940, p. 165 Type species by original designation: Limnephilus difficilis Walker Revision: Schmid, 1952c Medium-sized species with wide rust-colored and rounded wings. Maxillary palpi in % small, fairly thick, and slightly flattened. Spurs: 1,2,2. Wings short and wide, with front wings evenly rounded at apex and hind wings blunt and without any subapical indentation. Membrane of front wings grainy and covered with short setae. Venation (Fig. 507): widening of front wings results in strong apical expansion of discoidal and subradial cells, roughly straight anastomosic alignment and distinct divergence of apical veins. FIII pointed or with short petiole. Discoidal cell very large. In hind wings, discoidal cell also large and anastomosis strongly oblique and slightly broken. % genitalia (Figs. 472–475): tergite VIII with spinule-covered area. Segment IX developed dorsally into very wide vertical plate lining bottom of anal cavity. Latter completely eversible. Superior appendages small and elongate. Intermediate appendages fairly large and spine-shaped. Lateral sclerites of segment X well developed, bearing small sclerotized lobes. Inferior appendages developed into faintly protruding oblique cones, concave in inner surface, and joined to small, concave plate lining apical cavity. Phallic apparatus very large and thick. Parameres thick and much shorter than aedeagus. & genitalia (Figs. 476–478): tergite IX large and well developed on sides that are tucked around segment X. Latter in shape of very short cylinder, truncated very obliquely upward and flanked by 2 lobes. Ventral 124

lobes of sternite IX not protruding, but membranous and barely distinct from each other. Supragenital plate obtuse. Vulvar scale small but protruding and comprising subequal lobes. The genus Frenesia is found exclusively in the eastern Nearctic region and comprises two species: missa Milne, in which the inferior appendages are very blunt, and difficilis Walker, in which the extremity of the inferior appendages is strongly extended. Both species occur in waterlogged land, springs, and cold-water streams. The larvae are occasionally semi-aquatic. The adults emerge in November and overwinter in that stage. These species are widespread from Nova Scotia to Minnesota.

Genus Glyphopsyche Banks

Glyphopsyche Banks, 1904, p. 141 Type species by original designation: Glyphopsyche bryanti Banks = Phryganea irrorata Fabricius Revision: Schmid, 1952c Fairly large species with front wings strongly colored with brown (Fig. X). Slight, very slender body. Wings much larger in % than in &. Body of % very small. Front wings slightly indented subapically, and membrane sparsely covered with short hairs. Hind wings very broad. Venation: in front wings, discoidal cell 3 times as long as its petiole. FI narrow at base. FII very wide and FIII pointed or equipped with short petiole. Discoidal and subradial cells wide at apex. Anastomosis virtually straight and parallel to body or slanting slightly forward. % genitalia (Figs. 479–481): tergite VIII with 3-lobed area densely clothed with spinules. Superior appendages small, bilobed, and extensively fused with segment X. Intermediate appendages completely fused

Fig. X. Glyphopsyche irrorata Fabricius 125

together and upward-curving. Subanal plate large, horizontal, with reinforcements, particularly on underside. Inferior appendages quadrangular and very prominent. Phallic apparatus of medium size, but wide and strongly sclerotized. Parameres very small and thick. & genitalia (Figs. 482–484): tergite IX very short. Segment X in shape of short tube that is as wide as tergite IX and that it extends without discontinuity. Ventral lobes of sternite IX barely distinct from each other and completely fused with both tergite IX and segment X from which they can barely be distinguished. Supragenital plate small. Vulvar scale also small but fairly thick, with massive, wide lateral lobes and slender median lobe. The genus Glyphopsyche is solely Nearctic in distribution and probably contains only a single species, irrorata Fabricius. A second species, missouri Ross, may be only an ecological form. G. irrorata is found transcontinentally, from Newfoundland to California and Alaska. It occurs in marshes, ponds, small lakes, and generally in essentially still water. Adults emerge in September, overwinter in that stage, and fly until May. Genus Chilostigmodes Martynov

Chilostigmodes Martynov, 1914, p. 260 Monobasic type species: Chilostigmodes forcipatus Martynov Revision: Schmid, 1950c Small, slender insects with front wings strongly mottled with grey. Maxillary palpi of % moderately developed. First segment over half length of 2nd segment which is subequal to 3rd. Legs long and thin. Spurs: 1,1,1. Wings relatively very large compared to body. Front wings fairly wide at apex, with few small setae on membrane. Hind wings strongly indented subapically. Venation (Fig. 509): in front wings, discoidal cell slightly longer than its petiole. Anastomosis located relatively close to apex, parallel to body, and evenly and very sharply broken. Apical cells short and FIII sessile. In hind wings, anastomosis as strongly broken as in front wings and closer to apex. Discoidal cell longer than in front wings. % genitalia (Figs. 485–488): tergite VIII spinule-free. Segment IX very elongate laterally and not produced ventrally into tonguelike structure. Superior appendages large, very prominent, inwardly curved, and resembling stout callipers. Base of superior appendages firmly fused with sclerites of segment X that are developed into 2 wide plates. Latter completely fused with bottom of apical cavity. Intermediate appendages very strongly reduced, as 2 small lobes fused with inner side of superior appendages. Inferior appendages cone-shaped, slender, and consisting of single part. Phallic apparatus long and thin. Aedeagus without basal tubercles and parameres very slender. & not described. 126

The genus Chilostigmodes comprises a Siberian species and a very closely related Canadian species, areolatus Walker. It occurs very locally, yet is found throughout the country from Newfoundland to Alberta and Alaska. It inhabits ponds and small marshy lakes, emerges in October, and overwinters in the adult stage. Genus Grensia Ross

Grensia Ross, 1944, p. 201 Type species by original designation: Limnephilus praeteritus Walker Revision: Schmid, 1950c Hairy, heavy-bodied insects with wings strongly mottled with brown (Fig. XI). Head thick and convex. Eyes small and anteriorly positioned. Palpi thick. Head and pronotum profusely clothed with fine, long hairs. Spurs: 1,2,2. Medium-sized wings with front wings obtusely rounded at extremity and hind wings slightly indented subapically. Setae of membrane short and profuse. Venation (Fig. 508): in front wings, R2 ending at wing margin, close to R1. Anastomosis fairly close to apex, not strongly broken, and slanting fairly strongly toward body, forward. % genitalia (Figs. 489–492): tergite VIII with wide and very deep cleft in middle of apicodorsal side; cleft half closed by membranes, and fine spinules clothe apical angles of tergite. Segment IX robust, forming a wide dorsal-vertical strip embedded in tergite VIII. Superior appendages large, strongly sclerotized and bifid. Intermediate appendages comprising 2 small concave plates, almost completely fused together and only slightly prominent. Lateral sclerites of segment X large, forming 2 widely divergent points. Inferior appendages comprising an external part concave anteriorly and as large as 2 inner concavities facing it and directed backward.

Fig. XI. Grensia praeterita Walker 127

Medium-sized phallic apparatus. Aedeagus large and membranous at apex and without basal tubercles. Parameres thin. & genitalia (Figs. 493–495): tergite IX narrow but fairly elongate. Segment X with 2 triangular, prominent dorsal parts and small, barely protruding ventral scale. Ventral lobes of sternite IX fairly large, faintly prominent, and poorly differentiated. Supragenital plate absent. Vulvar scale large, triangular, strongly concave at top, with lateral lobes triangular; median lobe lost. Vaginal vestibule large and vast. The genus Grensia is monobasic and its only species, praeterita Walker, is of Arctic and circumboreal distribution. It is found primarily above the treeline and inhabits mainly lakes but also occasionally rock pools.

Genus Chilostigma McLachlan

Chilostigma McLachlan, 1876, p. 187 Monobasic type species: Chilostigma sieboldi McLachlan Revision: Schmid, 1950c Medium-sized species with front wings extensively speckled with brown. Head fairly narrow, relatively long, and strongly bulging. Palpi of % moderately long. Spurs: 1,2,2. Medium-sized wings shaped like those of Grensia, but with hind wings distinctly wider. Veins of front wings thickened and, like the membrane, bearing large numbers of erect setae. Venation: in front wings, anastomosis strongly broken, relatively close to apex, and slanting fairly strongly toward body, forward. FIII wide and discoidal cell short. In hind wings, discoidal cell short and triangular. % genitalia (Figs. 496–498): tergite VIII with area clothed with spinules. Segment IX forming prominent, ventral, tonguelike structure. Superior appendages bifid, very small, and extensively fused with segment X. Intermediate appendages poorly developed, as 2 slightly sclerotized, virtually flat lobes fused together. Inferior appendages large, prominent, and comprising 2 parts, both strongly developed. On inner side, appendages form a large concave plate completely fused with segment X. Phallic apparatus very long and thin, with aedeagus almost completely membranous. & genitalia (Figs. 499, 500): tergite IX small and very short, with apicoventral angles extending into 2 lamellae, which undoubtedly are ventral lobes, but poorly differentiated. Segment X forming very short and wide tube, located low. Supragenital plate wide and prominent. Vulvar 128

scale very large, strongly concave upward, and consisting of very large lateral lobes and tiny median lobe. Vaginal vestibule deep and wide. The genus Chilostigma comprises two very closely related species, one of which is European and the other, itascae Wiggins, has been reported in Minnesota, where it was captured on the surface of snow in February, in a wet meadow not far from a gently-flowing stream. Genus Psychoglypha Ross

Psychoglypha Ross, 1944, p. 201 Type species by original designation: Psychoglypha avigo Ross Revisions: Schmid, 1952c; Denning, 1970 Beautiful species with large, light, virtually smooth, gold-colored wings with a longitudinal silver stripe (Fig. XII). Cephalic and prothoracic warts densely clothed with setae. Maxillary palpi long and thin. Spurs: 1,2,2 or 1,3,3. Wings relatively large, with front wings narrow and truncated at apex, and hind wings very wide and more or less indented. Front wing membrane fine and not very pilose. Characteristic coloring of front wings: rustyellow with longitudinal white stripe extending to subradial and 4th apical cells. Venation: in front wings, pterostigma long and narrow. Discoidal and subradial cells wide at apex. Anastomosis slanting toward body, forward, and slightly broken. Discoidal cell slightly longer than its petiole. In hind wings, it is very long. % genitalia (Figs. 501–504): tergite VIII with 1 or 2 spinule-clothed areas. Segment IX elongate laterally forming wide and distinct concavity, which extends onto base of inferior appendages. Ventrally, segment IX produces a tonguelike plate. Anal cavity wide, noneversible, shallow, and

Fig. XII. Psychoglypha bella Banks 129

divided into 2 by membranous fold. Superior appendages of medium size, occasionally simple but usually consisting of 2 widely spaced lobes. Intermediate appendages of variable size, concave on upper side, always fused together or joined by membrane; they form a part with flared lateral sides, lining back of anal cavity and extending to margin of segment IX, creating 2 fairly deep concavities. Inferior appendages prominent and developed into plates that are concave on upper side; they are so firmly fused with segment IX that suture has disappeared; they almost always consist of 2 parts: outer part protrudes sharply whereas inner part very small. Phallic apparatus very long and thin. Aedeagus slightly widened at base where it bears small tubercles. Parameres very thin and spiniform. & genitalia (Figs. 505, 506): segment IX generally large. Tergite comprises 2 wide, very prominent lobes, which are concave on inner surface and almost completely cover segment X. Latter consists of 2 dorsal points and 1 membranous, blunt, subanal plate. Ventral lobes of sternite IX small and fused with tergite from which they cannot always be distinguished. They form 2 lateral plates. Supragenital plate large. Vulvar scale large, thick, quadrangular and comprising 2 wide, lateral lobes and 1 very small median lobe. Vaginal apparatus extremely complex. The genus Psychoglypha is restricted to the Nearctic region and contains 14 species, 8 of which have been reported in our area. They occur in the mountains of the west, except for subborealis Banks, which is transcontinental and extends as far east as Newfoundland. These species inhabit a wide range of cold-water biotopes, from springs to medium-sized rivers and their side pools. The adults emerge in the fall and overwinter in that stage. Genus Desmona Denning

Desmona Denning, 1954, p. 62 Type species by original designation: Desmona bethula Denning Monophylax Nimmo, 1987b, p. 53 Type species by original designation: Psychoglypha (Monophylax) mono Denning Revision: Wiggins and Wisseman, 1990 Yellowish brown species. Anterior wings with unpigmented, silvery longitudinal strip, partially bordered with dark, posteriorly. An additional setal cluster posteromesad of ocelli. Spurs 1,2,2. Front wings sparsely clothed with short setae. Characteristic color of front wings: mainly light tan, cell A3 and postcostal cell dark brown up to arculus, M through M2 dark brown; subdiscoidal cell and FIII unpigmented. Venation: on front wings, R1 and R2 close together and sharply 130

bent. Anastomosis slightly broken and subparallel to body. Discoidal cell longer than its petiole. On hind wings, R2 ending at apex of R1; discoidal cell narrow. % genitalia (Figs. 510–514): tergite VIII without spinule-clothed area. Segment IX long laterally, short dorsally and ventrally; its ventroapical edge with scoop-like oval extension between inferior appendages, surface transversely striated. Superior appendages small in lateral aspect; broad, horizontal, and concave in dorsal aspect. Intermediate appendages lightly sclerotized, not prominent. Inferior appendages spatulate at apex, base expanded in lateral aspect. Phallotheca long. Endotheca short. Aedeagus partially membranous. Parameres thinly spiniform. & genitalia (Figs. 515, 516): segment VIII extended ventrally into lip at vaginal entrance, terminating in pair of quadrate lobes separated by elongate median notch. Segment IX lacking sternite, tergite fused with segment X into sclerotized cap extended apically in 2 pointed processes. Supragenital plate not prominent. Vulvar scale lacking median lobe. The genus Desmona contains only two west Nearctic species: bethula Denning and mono Denning. The latter has been recorded from California to Washington. Larvae are confined to small springs runs and seepage areas and to littoral zone of small alpine lakes. Genus Homophylax Banks

Homophylax Banks, 1900, p. 255 Monobasic type species: Homophylax flavipennis Banks Revision: Denning, 1963 Large light yellow insects with extremely hairy wings. Antennae thin, roughly as long as front wings. Maxillary palpi of % long and slender. Spurs: 1,3,4. Internal gland of abdominal sternite V very large and complex in % but much smaller in & (Figs. 518, 519). Wings very large, with front wings considerably enlarged at the level of anastomosis and forming blunt parabola at apex. Hind wings fairly blunt and distinctly wider than front wings and profusely covered with fine hair. Venation with fairly unusual characters and displaying strong sexual dimorphism on hind wings (Figs. 517, 520–522). In front wings of %, C occasionally thickened at base and M+Cu1 forming a blade-like structure folded forward (Fig. 520). Anastomosis almost always in middle of wing and in one part, with line slanting slightly rearward and minimally broken. Crossveins long, and forks narrow at base. Discoidal cell twice as long as its petiole, and thyridial cell with long peduncle. Only 1 anal cell, since A2 is lost. In hind wings of %, discoidal cell small, narrow, and located basal to middle of wing. FI petiolate, FII pointed, and FIII with 131

petiole. Veins of anastomosis scattered, and M forks well ahead of proximal end of discoidal cell. In some species, a longitudinal fold covered with scaly bristles in median area, which is reduced. In &, venation of front wings similar to that of %. In hind wings, discoidal cell wider and triangular and located in middle of wing. Anastomosis less irregular, and M forks apical to proximal end of discoidal cell. % genitalia very complex (Figs. 523–525): tergite VIII with smooth, sclerotized protuberance. Segment IX strongly developed laterally and ventrally. Vast, noneversible apical cavity. Superior appendages large and complex in shape, extensively lining apical cavity. Inner and outer margins of superior appendages turned upward and prominent. Outer margin developed into oval, faintly sclerotized, hairy lobe. Inner margin located high, extremely sclerotized, and developed into 2 black points: simple or double upper point and lower point situated above intermediate appendages. Latter in relatively very low position and shaped into very sclerotized, upward-slanting, highly prominent, curved plates. At base, fused with lateral sclerites of segment X, which form protruding lobes lying against both lateral side of segment IX and base of inferior appendages. Inferior appendages developed into fairly small, horizontal plates. Aedeagus very small and stout, in form of small, sclerotized, bifid appendage, inserted into a large, membranous endotheca. Parameres lost. & genitalia as simple as % genitalia are complex (Figs. 526, 527). Tergite IX small. Segment X produced into long, fairly narrow tube, exhibiting relatively little sclerotization and strongly cleft. Ventral lobes of sternite IX forming vertical, prominent plates, set fairly far apart. Supragenital plate concave and bilobed. Vulvar scale blunt and trilobed, with each lobe indented at extremity. Vaginal cavity gaping. The genus Homophylax is medium-sized and confined to the mountains of western Nearctic region. It contains 10 species, 5 of which occur in Alberta, British Columbia, and southward: flavipennis Banks, acutus Denning, andax Ross, baldur Nimo, and crotchi Banks. These insects are always rare and local in distribution and inhabit mountain watercourses and lakes. Genus Phanocelia Banks

Phanocelia Banks, 1943, p. 354 Type species by original designation: Apatania canadensis Banks Revision: Schmid, 1950c Small, pale insects. Maxillary palpi weakly developed. Front tibia of % slightly longer than femur. Spurs: 1,2,2. 132

Wings large, narrow, and elongate. Front wings rounded at apex and hind wings fairly wide and indented below apex. Venation (Fig. 528): in front wings, pterostigma narrow, R1 slightly curved and joined to C by a crossvein. Discoidal cell wide and shorter than petiole. Third apical cell wide at base and wedged between discoidal and subradial cells. Anastomosis in pronounced zigzag pattern. FIII petiolate. In hind wings, discoidal cell long, anastomosis not as broken as on front wings, and FIII shorter than petiole. % genitalia (Figs. 529–532): tergite VIII bilobed at apical margin, clothed with relatively large spinules. Segment IX well developed. Superior appendages forming wide, horizontally positioned cupules delimited by fairly high edge, except toward front. Intermediate appendages long, projecting vertically, and strongly fused at base with superior appendages. Inferior appendages large, stout, concave on inner surface, and consisting of single part. Phallic apparatus large. Aedeagus thin and inerm. Parameres spiniform and very slender. & genitalia (Figs. 533, 534): tergite IX short, very wide, and subquadrangular in side view. Segment X developed into 2 very small, very faintly protruding lobes turned toward each other. Ventral lobes of sternite IX not very prominent singly but together forming a strongly protruding transverse part. Supragenital plate ogival in shape and very short, with width 3 times height. Vulvar scale very large and strongly concave on upper side, with very large, triangular lateral lobes, and tiny, ogival median lobe. Vaginal apparatus with 2 large, sclerotized parts forming fairly long vestibule. The genus Phanocelia is monobasic. Its single species, canadensis Banks, occurs in Alberta, Manitoba, New Brunswick, Nova Scotia, New Hampshire, Quebec, and the Northwest Territories. It inhabits marshes and small lakes and flies in the fall.

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Family Uenoidae Iwata Uenoinae Iwata, 1927, p. 214 Type genus: Uenoa Iwata Uenoidae Botosaneanu, 1976, p. 199 Neophylacinae Schmid, 1955, p. 88 Type genus: Neophylax McLachlan Revision: Wiggins, Weaver and Unzicker, 1985, p. 764 Eyes large and very prominent. Ocelli small, very anteriorly positioned. Occipital portion rather evenly rounded. Antennae very fine, crenulated on underside, and often slightly longer than front wings. Maxillary palpi of % occasionally with neoformations. Spurs ranging from 1,2,2 to 2,4,4. Inner apical spur of hind tibiae sometimes modified in %. Hemogill system absent. Wings fairly variously shaped. Front wings fairly narrow but occasionally widened considerably at level of anastomosis, elliptical or truncated or scalloped at apex. Hind wings not wider than front wings. Latter clothed with thick, appressed hairs. Venation almost always complete in front wings with forks I, II, III, and V present. In hind wings, venation simplified, often displaying sexual dimorphism, depending on genus. % genitalia comprising short parts fitted into compact unit with an unusual appearance, primarily due to considerable development of segment IX. Latter large, solid, and fitting broadly around all genital parts, sometimes including inferior appendages; ventrally strongly convex and developed into plate or point fused with inferior appendages; laterally, occasionally displaying sharp relief or producing appendage. Segment X virtually absent with only its armature visible. This consists of 3 pairs of appendages of difficult interpretation. Inner branches located dorsally and occasionally rounded and concave, but usually elongated, forming roof above phallic apparatus. Anal opening located between 2 inner branches. Occasionally, 2 lower branches protecting phallic apparatus laterally. Preanal appendages small and ovoid or absent. An unusual and more or less pronounced feature in middle portion of segment X: 2 lateral mesal plates, more or less concave and occasionally forming true cavity. Inferior appendages small and not very prominent, often indistinctly 2-segmented and with basal segment reduced in favor of 2nd segment. On inner side, 2 134

basal segments fused to each other and forming unpaired, highly sclerotized mass often rough-surfaced and situated under phallic apparatus. Second segment simple or bifid. Phallic apparatus often emerging in middle of or toward top of segment IX, consistently small, slender, and only slightly movable. Parameres and endotheca often absent. & genitalia: segments IX and X firmly fused together. Segment IX always comprises clearly distinct tergite and sternite. Tergite short or even vestigial. Segment X appendage-free, relatively large, prominent, and consisting of 2 large lateral parts separated by narrow slit on dorsal side and wide opening on ventral side. Anal opening situated between 2 points. On underside, segment X forms vast cavity with membranous bottom, serving as antechamber for vaginal opening. Supragenital plate present or absent. Gaping vaginal opening on segment IX. Vulvar scale simple or bifid, arising from sternite IX. Latter consists of 2 lobes more or less fused laterally with vulvar scale. Vaginal cavity deep, without any vestibule. Vaginal apparatus composed of central part and ventral, unpaired, accessory part. For a long time, the Uenoidae have been classified in the Limnephilidae under the name of Neophylacinae. They now comprise seven genera, five of which are found in the Nearctic region. Neophylax has also an Oriental and Palearctic distribution, whereas the other four are strictly Nearctic. The family has been divided into two subfamilies, the Thremmatinae and the Uenoinae. The former is so weakly defined that it is only mentioned here. 1a Front wings roughly 3 times longer than wide (Figs. 549, 555) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uenoinae, 2 1b Front wings broader (Fig. 535) . . . . . . . . . . . . . . . . . . . . . . . . Thremmatinae, 3 2a M3+4 of front wings absent. Discoidal cell of hind wings twice as long as wide, located before middle of wing (Figs. 555, 556) . . . . . Farula, p. 139 2b M3+4 of front wings present. Discoidal cell of hind wings barely longer than wide, located in middle of wing (Figs. 549, 550) . . . Neothremma, p. 138 2c M3+4 of front wings present. In both wings, discoidal cell joined by FI for almost half of its length (Fig. 562) . . . . . . . . . . . . . . . Sericostriata, p. 140 3a Inferior appendages of % more or less prominent and not embedded in segment IX. Armature of segment X composed of 2 or 3 pairs of appendages (Figs. 540, 541). Ventral lobes of sternite IX in & fused with lateral margins of vulvar scale (Figs. 542, 543) . . . . . . . . . . . . . . . . Neophylax, p. 136 3b Inferior appendages of % not prominent and deeply embedded in segment IX. Armature of segment X composed of 2 membranous lobes and 2 earshaped appendages (Figs. 545, 546). Ventral lobes of sternite IX in & fused together ventral to vulvar scale, which is only slightly sclerotized (Fig. 548) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligophlebodes, p. 137

135

Genus Neophylax McLachlan

Neophylax McLachlan, 1871, p. 111 Monobasic type species: Neophylax concinnus McLachlan Acronopsyche Banks, 1930, p. 227 Type species by original designation: Acronopsyche pilosa Banks Maxillary palpi of % very long and slender. Spurs: 1,2,2, 1,2,3 or 1,3,4. In %, inner apical spur of posterior tibiae modified. In primitive species, simple, very long, and tapered (Fig. 538), but in specialized species, thick at base and surrounded by sort of rigid horn-shaped structure consisting of row of partially fused hairs (Fig. 539). Wings variously shaped. Front wings wide, angular, and occasionally scalloped at apex in small species, but much narrower and elongated in large forms. Venation (Figs. 535–537) complete and constant in front wings, with all forks present: I, II, III, and V. Discoidal cell 2 to 3 times longer than its petiole. Anastomosis strongly and regularly broken. In hind wings, venation simplified and subject to strong sexual dimorphism. Venation varies greatly in %, but little in &. As radial sector occupies a subcentral position, the cells it delimits usually wide. In %, radial sector quadruple, forming consistently sessile forks I and II. In most species, it displays normal configuration (Fig. 535), whereas in others, the 2 branches arise from R1 at extreme base of wing, but at different points (Fig. 537). Discoidal cell consequently open next to body. In all species, M reduced to single branch free only along apical half; basal half coalesces with Cu1 or R4+5, depending on author. Cu1 simple, hence FV absent. Four anal veins. In & (Fig. 536), forks I, II, and V present and RS normal. M free over entire length and double. Cu1a and Cu1b present. Five anal veins. % genitalia (Figs. 540, 541): segment IX massive, forming sclerotized ventral plate. Inner branches produced into stout, contiguous, and often very long points. Occasionally, also 2 lower branches, which then replace inner branches, and latter may be reduced. Middle plates of segment IX small and faintly concave in primitive forms but large and very strongly curved in most specialized species. Extremity of these parts may be very large and protect phallic complex laterally. Preanal appendages small, ovoid, and free in primitive forms, whereas in other forms, fused with middle plate of segment X. Inferior appendages not prominent and very indistinctly 2-segmented. Basal segment more strongly reduced than in other genera and, in addition, consistently forms an unpaired, strongly sclerotized part located under phallic apparatus. Apical segment often bifid, complex in shape and provided with points or teeth. Aedeagus long, stylet-shaped, concealed between inner branches in primitive species; in specialized forms, tiny, straight, and emerging between upper points of inferior appendages. Parameres absent. 136

& genitalia differ from those of Oligophlebodes essentially in structure of vulvar scale. Very small and faintly sclerotized in primitive forms and, on lateral sides of base, fused with ventral lobes of segment IX that are shaped into long appendages. In more specialized forms, vulvar scale far more sclerotized, much longer, and generally bifid. On lateral sides, it is almost entirely fused with ventral lobes of segment IX that, except for free portion, has also become very sclerotized (Figs. 542, 543). The genus Neophylax is medium-sized and found in the Palearctic, Oriental, and Nearctic regions. It includes 22 North American species, 17 of which have been captured in our area. The genus is fairly heterogeneous and includes species that are fairly diversely specialized and that require classification into several groups on the basis of venational and genitalic characters. The species are occasionally quite attractive in color and are especially well represented in the eastern provinces as well as in the mountains of western Canada. All species inhabit lotic environments, and some are characteristic of some types or sections of rivers and streams. Adults mostly fly in the fall.

Genus Oligophlebodes Ulmer

Oligophlebodes Ulmer, 1905, p. 66 Monobasic type species: Oligophlebodes coloradensis Ulmer = Halesus minutus Banks Maxillary palpi of % short, slender, comprising 3 segments of subequal length, raised against face and clothed with long bristles which also occur in large numbers on face and inner side of 1st antennal segment. Spurs: 1,3,3. Inner spur of posterior tibiae of % thickened at base and with few bristles on inner side. Wings of medium size with front wings obliquely elliptical at apex. Venation (Fig. 544) constant, similar to that of Neophylax and always displaying sexual dimorphism in hind wings. In front wings, R1 joined to Sc by a crossvein. Discoidal cell slightly longer than its petiole. Anastomosis slightly broken. In hind wings of %, RS double, situated on anterior part of wing and delimiting narrow cells. R5 absent. FII alone present. As in some Neophylax, R2+3 and R4+5 arise independently from R1 at extreme base of wing, and discoidal cell open against body. Media with 2 simple and completely distinct branches. Venation of & very similar to that of && of concinnus group, with forks I, II, and V present. % genitalia fairly similar to those of Neophylax but simpler and more specialized (Figs. 545, 546). Segment IX very short dorsally with stout lateral protuberance supporting middle plate of segment X and fitted broadly around inferior appendages. Segment X barely prominent, forming bulky dorsal mass with anus in middle. Inner branches in shape of 2 137

rounded, concave appendages separated by large space from which phallic apparatus emerges. Lateral concavities, similar to those of Neophylax, but larger and more complex. Inferior appendages distinctly 2-segmented, small, and completely fitted into an incision of segment IX. Basal segments clearly defined and massive, fused on inner side, forming sclerotized, median, and occasionally bifid protuberance. Terminal segments pointed or hook-shaped and more or less sclerotized. Phallic apparatus comprises phallotheca, large endotheca, aedeagus, and 2 reduced parameres. Aedeagus and parameres in slender rods. Phallic apparatus situated higher than in Neophylax. & genitalia (Figs. 547, 548) very similar to those of Neophylax but vulvar scale is slightly differently structured. Ventral lobes of segment IX consist of massive, barely prominent, barely sclerotized, transverse plates fused on ventral side of vulvar scale and not with lateral margins of latter. The plates also fused with sternite VIII, together forming slightly concave unit lining small and barely sclerotized vulvar scale. The genus Oligophlebodes is small and exclusively western Nearctic in distribution. It contains seven species, six of which have been reported in our area and southward. The species are characteristic of the most rapid sections of mountain streams. Genus Neothremma Dodds and Hisaw

Neothremma Dodds and Hisaw, 1925, p. 127 Monobasic type species: Neothremma alicia Dodds and Hisaw Head not very wide with strongly bulging vertex. Eyes small. Lateral ocelli in very anterior position, behind anterior warts which are large. First antennal segments 1.5 times as long as head, covered on underside and inner surface with long, thick, extremely dense, silky hairs that also occur on face. The latter bears 2 extended, longitudinal warts. Maxillary palpi of % short, slightly thickened, raised against face, and very densely clothed with erect hairs. Spurs: 1,3,4, unmodified. Wings medium-sized, evenly elongate, and roughly 3 times as long as wide. Venation complete on front wings (Figs. 549, 550), with forks I, II, III, and V present, and greatly reduced on hind wings where there is clearcut sexual dimorphism. In front wings, discoidal cell shorter than or equal in length to its petiole, and thyridial cell long and narrow. A2 absent, only one anal cell. In hind wings, only forks I and II present in % and I, II, and V in &. Discoidal cell small, in middle of wing, FI with long petiole and FII wide at base. Media simple. In %, Cu1 simple and fused with Cu2 at base. In &, these 2 veins completely distinct and Cu1 forked, forming FV. Four anal veins. In both sexes, narrow cell between M and Cu1 leathery over virtually its entire length. 138

% genitalia (Figs. 551, 552): segment IX well developed, but not fitted around any appendage; laterally, forming very long, simple or bifid, inwardly projected sclerotized appendage. Segment X small, its armature reduced to 1 or 2 pairs of branches. Preanal appendages present or absent. Inner branches developed into 2 very long, simple spines curving broadly downward, joined at base by membranous mass containing also anal opening; these branches inserted in 2 sclerotized masses which form the ceiling of phallocrypt. Inferior appendages located on very lowest part of segment IX. As in two preceding genera, these appendages were primitively 2-segmented but secondarily acquired a very unusual conformation, with well-developed basal segments fused together, forming thick plate supporting the phallic apparatus. Second segment in upper basal position in relation to 1st and forming a large mass fused with margin of segment X and covered with small tubercles. Phallotheca large, short, membranous, ending in sclerotized part surmounted by few spines. Aedeagus styletshaped. Parameres lost. & genitalia (Figs. 553, 554): segment VIII partially unsclerotized and concave ventrally. Segment IX consisting of 2 distinct parts, with sternite forming 2 large concave parts framing vulvar scale. Segment X forming 2 parts concave on inner surface and framing supragenital plate. Latter very large and not extending to ceiling of vaginal cavity. Vulvar scale very large and broadly bifid, with 2 parts distantly spaced. Vaginal cavity gaping. Vaginal apparatus fairly small, somewhat protruding from vaginal opening and partially membranous. The genus Neothremma is small and found in the western Nearctic region. It contains seven species, two of which have been captured in British Columbia, Alberta, and southward: alicia Dodds and Hisaw and didactyla Ross. They inhabit turbulent mountain rivers and streams and are always rare. Genus Farula Milne

Farula Milne, 1936, p. 116 Type species by original designation: Farula rainieri Milne Head not very wide, with vertex strongly bulging. Eyes small and clothed with short hairs. First antennal segment very long but not densely covered with hairs. Maxillary palpi of % fairly long, slender, consisting of segments subequal in length and, like face, covered with simple, fairly dense hairs. Spurs: 2,4,4, unmodified. Wings medium-sized, evenly elongate and roughly 3 times longer than wide. Venation (Figs. 555, 556) very similar to Neothremma, simplified in both pairs of wings, and exhibiting slight sexual dimorphism on hind wings. In front wings, forks I, II, III, and V present. Discoidal cell 139

fairly short, in middle of wing, and merged for long distance with FI. Thyridial cell short, slightly longer than discoidal cell. M3+4 absent and FV sessile. Three anal veins. In hind wings, only forks I and II present in both sexes. FI with long petiole and FII sessile and wide at base. In %, discoidal cell long, beginning at base of wing. In &, shorter with longer petiole. In both sexes, M and Cu1 simple. In %, Cu1 and Cu2 fused together at base. Cell between M and Cu1 leathery. Four anal veins. % genitalia (Figs. 557–559) with the most complicated feature to be found in the family. Segment IX very elongated all around but interrupted on dorsal side, with or without membranous apicodorsal lobe. Segment X prominent. Its armature consists of 4 pairs of appendages. Preanal appendages distinct. Outer branches developed into 2 very long, thin, wavy spines. Inner branches shorter and thicker with tuft of stout setae. Lower branches form very stout hooks. Inferior appendages represented by 2 long, twisted and completely separate, sclerotized parts. Phallotheca small, barely sclerotized, simple in shape, and slightly movable. Aedeagus stylet-shaped. Parameres lost. & genitalia (Figs. 560, 561) very similar to those of Neothremma, but with following distinguishing characters: lateral lobes of sternite IX smaller. Supragenital plate also smaller, extending far within genital cavity of which it forms the ceiling. Vulvar scale larger, more strongly concave, with wider indentation. Vaginal apparatus larger and completely sclerotized. The genus Farula is small and occurs exclusively in western Nearctic region. It contains 10 species that inhabit mountainous areas. To date, none has been captured in Canada but rainieri Milne has been caught in Washington state. The species inhabit small, torrent-like streams and occasionally are abundant locally. At times they fly in the spring, near snow. Genus Sericostriata Wiggins, Weaver and Unzicker

Sericostriata Wiggins, Weaver and Unzicker, 1985, p. 778 Type genus by original designation: Sericostriata surdickae Wiggins et al. Head with 3 pairs of warts; anterior warts bulky, slightly oblique to median line. First antennal segment as long as or longer than head (Fig. 563). Maxillary palpi 5-segmented in &, 3-segmented in %, held upright, with each segment membranous and inflated into flexible lobe, all lobes contiguous and bearing long, yellowish setae (Fig. 564). Spurs 1,3,4, unmodified. Wings medium-sized, more than 3 times longer than wide. Venation (Fig. 562) complete in front wings, with forks I, II, III, and V present; reduced in hind wings, with forks I and V present and slight sexual 140

dimorphism. In front wings, membrane thickened between bases of Sc and R1. Discoidal cell narrow, much longer than its petiole, and joined by FI for almost half its length. Three anal cells. In hind wings, discoidal cell in basal position and joined by FI for more than half its length. Media simple. In %, Cu1a+b and Cu2 joined basally. In &, these 2 veins distinct from base. Four anal veins. % abdominal segment VI with a short ventromedian process. % genitalia (Figs. 565–569): segment IX longer ventrally than dorsally and not fitted around inferior appendages. Segment X with 3 pairs of branches. Outer branches thickly ovoid. Inner branches in irregular cylinders. Inferior branches longer than preceding ones, in very extended, irregular cones. Preanal appendages and lateral mesal plates absent. Inferior appendages probably 2-segmented and with the 2 segments completely fused to each other, contiguous mesally and very divergent; obtusely conical from side; inner face irregular and with strong medio-superior process bearing a few denticles and supporting phallic apparatus. Latter small and with all components indistinct. & genitalia (Figs. 570–572): tergite VIII unmodified and separated from sternite by prominent membranous area. Tergite IX short, fused with segment X to form rounded roof over anal opening. Sternite IX forming prominent fingerlike lobes at each side of vulvar scale. Supragenital plate fused with segment X. Vulvar scale bears pointed blade-like sclerotized lobe at each side, arising from its inner dorsal edge; posteromedian edge unmodified. The genus Sericostriata contains a single species, surdickae Wiggins et al., recorded from Montana only. Larvae have been found on the upper surface of rocks in rapid to turbulent streams, frequently in the hydropetric zone.

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Family Goeridae Ulmer

Goerinae Ulmer, 1903, p. 81 Type genus: Goera Stephens Goeridae sensu Ross, 1944, p. 256 Head wide, with 3 or 4 pairs of warts on vertex. Ocelli present or absent. First antennal segment large, simple, and slightly thicker in % than in &. Maxillary palpi exhibiting strong sexual dimorphism, very short and 2- or 3-segmented in % and long, 5-segmented, with first 2 segments short, in &. Legs densely clothed with flattened hairs and bearing small spines. Spurs: 2,4,4 or 2,3,4 or 1,2,2. Internal gland of abdominal sternite V absent. Abdominal hemogill system present or absent. Wings densely hairy, fairly evenly elliptical, with both pairs equal in width. Faint sexual dimorphism. Venation similar in both sexes and barely simplified, with forks I, II, III, and V or I, II, and V present on both pairs. In front wings, discoidal cell closed. Thyridial cell of variable length. Thyridial or subthyridial cells sometimes widened at extremity. FV occasionally petiolate. One or 3 anal veins. In hind wings, discoidal cell open or closed. Four anal veins. % genitalia: segment IX variously developed, with or without dorsal median lobe. Segment X reduced or membranous. Preanal appendages present. Inner and outer branches both present but not together in Canadian genera. Inferior appendages large, 2-segmented, horizontal, with 2nd segment simple or bifid and inserted at apex of 1st. Phallic apparatus large, variable in composition, but always without parameres, situated between inferior appendages and joined by sclerotized connection to upper basal angle of these appendages. & genitalia: segment IX comprising tergite and sternite either joined or separate. Segment X indistinct from segment IX at base and without appendage. Vaginal opening on segment IX, with or without vulvar scale. Vaginal apparatus simple. The Goeridae are of medium size and cosmopolitan distribution, although they do not occur in Neotropical America. They are particularly well represented in the Oriental region. 142

In Canada, the family comprises four genera so different from one another that it is difficult to give a precise familial description of the adults. The unity of the family appears mainly in the larval characters. There are two subfamilies. 1a Ocelli absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goerinae, p. 143 1b Ocelli present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepaniinae, p. 145

Subfamily Goerinae Ulmer There are two Canadian genera in this subfamily: Goera Stephens and Goeracea Denning. They are very different and show only a few common characters. Ocelli absent. Venation: in front wings, crossveins SC-R1 present. Median cell open. Thyridial or subthyridial cells widened at their extremity. In hind wings, median cell open. %& genitalia without any common characters relevant to subfamilial level. The only ones are of familial status. 1a % abdominal sternite VI with 1 or more sclerotized points (Fig. 577). Rust or brown coloring. Spurs: 2,4,4. . . . . . . . . . . . . . . . . . . . . . . . . Goera, p. 143 1b % abdominal sternite VI without sclerotized points. Black coloring. Spurs: 2,3,4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goeracea, p. 144

Genus Goera Stephens

Goera Stephens, 1829, p. 28 Type species designated by Westwood, 1840: Phryganea pilosa Fabricius Revision: Schmid, 1983 Head very short, with very prominent eyes. Posterior cephalic warts particularly large and round (Fig. 573). Maxillary palpi of % very small and comprising 2 segments. A large, membranous, strongly erectile lobe equipped with flattened scales inserted behind 2nd segment (Fig. 574). Spurs: 2,4,4. Abdominal sternite VI with ventral row of 8 or 10 long spines in comb-like arrangement in % (Fig. 577) and 1 or 2 shorter points, surrounded by a few tiny teeth in &. Abdominal hemogill system present and consisting of tufts of simple tubular branches, the length of a segment. Wings rust-colored or brownish and distinctly narrower in & than in %. Venation (Fig. 576): in front wings, discoidal cell short and united for long distance with FI and FII. FIII petiolate. Thyridial cell very long. 143

Subthyridial cell constricted before extremity, then dilated and bare at apex. In hind wings, R1 evanescent at extremity, discoidal cell open, and FIII with long petiole. M1+2 separating at very early stage from M3+4. Cu2 and A1 fused at base. % genitalia (Figs. 578–581): segment IX very elongate dorsolaterally, invaginated within tergite VIII and, on ventral side, considerably shortened anteriorly. Dorso-median lobe of segment IX very long, simple or bifid. Segment X greatly reduced and embedded in segment IX. Preanal appendages long, slender, and club-shaped. Outer branches absent. Inner branches forming very long, slender, and pointed spines. Inferior appendages with 1st segment short and very thick and 2nd segment bilobed and not always entirely distinct from 1st segment at base. Upper lobe coneshaped and very hairy and lower lobe strongly sclerotized and almost bare. Phallic apparatus very long, consisting of tiny phallotheca, virtually obliterated endotheca, and large, partially membranous aedeagus (Fig. 581). Phallotremal sclerite in deep concavity, well ahead of apex of aedeagus. & genitalia (Figs. 582, 583) fairly long and slender. Tergite and sternite IX broadly touching. Segment X long and slender, bifid, and forming large ogival supragenital plate. Vulvar scale not lobed. Vaginal apparatus simple, elongate, preceded by short, partially sclerotized vaginal vestibule. The genus Goera is large, with a Holarctic, African, Australian, and especially Oriental distribution. It varies considerably in the structure of the erectile lobe of the maxillary palpi of the %. In the Nearctic region, it is represented by six species, four of which are found in Canada where they occur in the eastern and central parts of the country. In calcarata Banks (Quebec and Nova Scotia), the genitalia are short, and the dorsal median lobe of segment IX is bifid. In tungusensis Martynov (= radissonica Harper and Méthot) (Quebec), the genitalia are also short, but the dorsal median lobe of segment IX is simple; in stylata Ross (Ontario) and fuscula Banks (Quebec), the genitalia are greatly elongate. Larvae of these species inhabit running swift and cold water and occasionally lakes.

Genus Goeracea Denning

Goeracea Denning, 1968, p. 24 Type species by original designation: Goerita genota Ross Revision: Wiggins, 1973 Eyes small and barely prominent. Vertex fairly strongly bulging, with posterior warts not very large but elongate. Maxillary palpi of % short, with 3 segments in shape of simple ovals (Figs. 584, 585). Spurs: 2,3,4. Abdominal hemogill system absent. 144

Wings black, with the hind wings blunt-ended. In %, there are coarse scales along Sc, R2, and R3 and in anal area. Venation (Fig. 586): in front wings, discoidal cell very long and narrow. FIII sessile. Thyridial cell not so long and widened at its extremity. The subthyridial cell also wider at its end. In hind wings, discoidal cell closed and moderately long, first 3 forks pointed or with very short petioles. M1+2 and M3+4 separate at late stage. Cu2 free, but first 2 anal veins fused at base. % genitalia (Figs. 587–589): segment IX much lower than segment VIII, stout, very elongate laterally and ventrally, but very short dorsally and without dorso-median lobe. Segment X developed into large, membranous, bifid plate. Inner branches absent. Outer branches short and thick. Preanal appendages present, but fused to external base of outer branches. Inferior appendages with 1st segment large and forming thickened hook at lower apical angle. Second segment simple, long, and slender. Phallic apparatus short, thick, and consisting of tubular phallotheca and endotheca, which is mostly membranous, but which also has tubular base and equipped with few short spines. Aedeagus absent. & genitalia (Figs. 590, 591): very blunt. Tergite IX short and provided with 2 thickened dorsal triangular structures. Sternite IX separate from tergite IX and forming 2 large angular lobes. Supragenital plate absent. Vulvar scale very large, strongly sclerotized, and shaped into bluntangled triangle. Vaginal apparatus simple, small, and located at end of short vestibule that is wider than apparatus itself. The genus Goeracea contains only two species inhabiting the mountains in the western part of the continent. G. genota Ross has been captured in British Columbia and further south where it occurs in cold and rocky streams in mountainous areas; oregona Denning is a more meridional species.

Subfamily Lepaniinae Wiggins Lepaniini Wiggins 1973, p. 29 Type genus: Lepania Ross Lepaniinae sensu Schmid 1981b, p. 186 This subfamily also comprises two very different genera. General coloration black or dark grey. Ocelli present. Male maxillary palpi 3-segmented, small, with little modification and smaller than labial palpi. Vertex with posterior warts small. Venation: in front wings, crossvein Sc-R1 absent. One or 3 anal veins. In hind wings, M1+2 and M3+4 separate rather apically. Four anal veins. 145

%& genitalia without common characters of subfamilial level. Segment IX without dorsomedian lobe. This subfamily is localized in the mountainous area of the west Nearctic region. 1a Front wings with both discoidal and thyridial cells subequal, short, and triangular (Fig. 601). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepania, p. 147 1b Front wings with discoidal and thyridial cells unequal, long, and narrow (Fig. 594) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goereilla, p. 146

Genus Goereilla Denning

Goereilla Denning, 1971, p. 207 Type species by original designation: Goereilla baumanni Denning Eyes rather large. Vertex with 4 pairs of warts, posterior forming long oval. Maxillary palpi of % much smaller than labial ones; 1st article very short, 2nd longer, and 3rd shorter and thinner than 2nd (Figs. 592, 593). Spurs:1,2,2. Wings elliptical, both pairs of similar shape, with anal area of hind wings reduced. Venation (Fig. 594): in front wings, discoidal cell closed, long, and narrow. FII very narrow at base. Thyridial cell long and not widened apically. Three anal veins, not converging and not forming a cell. In hind wings, subradial cell broad and discoidal cell small. FI and FIII very narrow at base, but sessile. FII with a long pedicel. % genitalia (Figs. 595–598): segment IX short, with its ventral apical edge prominent. Segment X comprising only 2 large, blunt parts, of complex shape and ending in 2 sharp points. Preanal appendages very large, free, concave internally, and excised apically. Inferior appendages far above ventral side of segment IX and of conical, blunt shape. Second article ovoid. Phallic apparatus comprising rather complex phallotheca and simple endotheca equipped with a dozen small spines. Aedeagus absent. & genitalia (Figs. 599, 600): tergite IX entirely membranous and obliterated. Sternite IX small, partially fused with sternite VIII and forming 2 lateral blunt lobes, each side of vulvar scale. Segment X in 2 large, triangular, prominent lobes. Supragenital plate as large, transverse rectangle. Vulvar scale as very blunt, simple lobe. The genus Goereilla comprises only a single species, baumanni Denning, recorded from Montana. Larvae live in the watery organic ooze and muck of spring seepage areas. 146

Genus Lepania Ross

Lepania Ross, 1941, p. 102 Type species by original designation: Lepania cascada Ross Head short and broad, with 3 pairs of warts, posterior ones broadly oval. % maxillary palpi smaller than labial ones and with 3 articles of progressive length. Spurs: 1,2,3. Wings elliptic and of same width in 2 pairs; posterior ones incised posteriorly in middle of their length. Wing coupling well developed. Venation (Fig. 601): in front wings, discoidal cell short. Thyridial cell triangular and very short. FI and FII only present, very narrow at base. M 2-branched. Only one anal vein. In posterior wings, R1 incomplete. Discoidal cell short. FI, FII, and FV only present. % genitalia (Figs. 602, 603): segment IX well developed, but ventrally membranous. Preanal appendages very large. Segment X composed of inner branches, outer branches, and medio-dorsal, paired, accessory lobe. Inferior appendages with 1st article very short. Second segment very long, thin, and curved. Phallic apparatus with endotheca bearing 2 pairs of stout, sclerotized processes. & genitalia (Figs. 604, 605): sternum VIII with median longitudinal incision; on each side a large sclerite. Tergite IX reduced. Sternite IX forming 2 lobes laterally framing vulvar scale. Segment X forming 2 large lobes. Supragenital plate broadly ogival. The genus Lepania contains a single species, cascada Ross, recorded from Washington and Oregon. Larvae live in the muck of spring seepage areas. This genus has developed extremely peculiar imaginal characters, most of which are typically limnephilid. The genus was first considered as a limnephilid (Ross, 1941). Later, I considered it as a dicosmoecine (Schmid 1955), then displaced it in the Apataniinae (Schmid, 1968a). Larval characters indicate that the right place for Lepania is in a special subfamily of the Goeridae (Wiggins, 1973).

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Family Lepidostomatidae Ulmer Lepidostomatinae Ulmer, 1903, p. 89 Type genus: Lepidostoma Rambur Lepidostomatidae Ross, 1944, p. 258 Head short, wide, with bare eyes. Cephalic appendages and wings exhibiting strong sexual dimorphism. && show little variation, whereas %% have developed significant secondary sexual characters. Ocelli absent. First or 2nd antennal segment of % elongate and thickened, simple or forked, and densely clothed with scales. In &, 1st segment long, simple, slender, barely thickened, and scale-free (Figs. 606–608). Maxillary palpi of % occasionally shorter than labial palpi and 1-, 2-, or 3-jointed, bulging or partially erectile or densely clothed with scales or bearing tufts of modified hairs. Maxillary palpi of & simple and 5-jointed. Legs with weakly developed spines and densely covered with flattened hairs which also cover spurs (Fig. 609). Spurs: 2,4,4. Abdominal hemogill system well developed and consisting of branches forming simple tubes equal to a segment in length. Gland of sternite V absent. Wings developed into smooth ellipses, fairly narrow, with 2 pairs of subequal width. Wing shape virtually same in both sexes. Strong sexual dimorphism in pilosity and venation. In %, both pairs have either large scaly areas or rows of spines, folds filled with flattened setae and spiny thickenings. Costal area of front wings occasionally enlarged and reflexed backward. In &, both pairs are consistently covered with thick, simple, and regular pilosity. Venation: almost complete in &, with forks I, II, III, and V present in front wings and I, II, and V in hind wings. In front wings, discoidal and thyridial cells long and median cell open. In hind wings, discoidal cell short, open or closed. In %, venation distinctly simpler, FIII of front wings and FII of hind wings usually absent, but some veins are secondarily divided. Arrangement of veins highly variable from species to species and often modified in relation to specialized scaly areas (Figs. 610, 611, 619, 620, 628, 629, 637, 638, 649–652). % genitalia: segment IX usually fairly regularly short. Segment X roof-like, often complex in shape and occasionally asymmetrical. Among its various lobes, it is no longer possible to distinguish the preanal and intermediate appendages. These appendages are obviously absent in some groups, but probably present in modified form in others. Inferior appendages 148

more or less elongate, horizontal or upward-slanting, complex in shape, and usually 2-segmented. Second segment often indistinguishable from lobes formed by 1st segment. When it can be distinguished, it arises from the inner surface of 1st segment. Phallic apparatus virtually complete, comprising phallotheca, endotheca, and aedeagus. Occasionally parameres also present, but in this case, endotheca often obliterated. Phallotheca joined by means of sclerotized connection to base of inferior appendages and apicolateral margins of segment IX. & genitalia: tergite VIII greatly developed and sternite VIII somewhat reduced, forming concave surface designed to hold the egg mass temporarily. Segment IX simple roof-shaped and sometimes equipped with 2 apodemal anterior horns. Segment X small, greatly reduced, and so thoroughly integrated with segment IX that it can hardly be distinguished from latter. Appendages and anal opening apparently absent. Vulvar scale wide and simple. Vaginal opening gaping and located at apex of segment VIII. Vaginal apparatus simple. The Lepidostomatidae constitute a large, ubiquitous family, although it does not occur in most of Australian and Neotropical regions. It is particularly well represented in the Oriental region. It is one of the most remarkable families in the order from the standpoint of range, inconsistency, and fanciful nature of the secondary characters of the %, and they have inspired McLachlan to label this family a curiosity shop. Some Oriental genera, such as Eodinarthrum Martynov, Dinarthrum McLachlan, and Dinarthrodes Ulmer, are among the most amazing trichopterans that exist. In North America, the family comprises roughly 80 species, which largely occur in the mountains of the eastern and western parts of the continent. The classification of the Lepidostomatidae has been rather unstable in the past, but a recent excellent revision by Weaver (1988) has now clarified the situation. Nearctic Lepidostomatidae comprise two genera, Lepidostoma Rambur and Theliopsyche Banks, with the former genus comprising four subgenera. 1a Posterior cephalic warts bluntly oval (Fig. 606). Sternite VIII of % without a ventromedian process. In &, vaginal apparatus barely longer than its width and vulvar scale simple (Fig. 617) . . . . . . . . . . . . . . . Lepidostoma, p. 150 1b Cephalic warts long and narrow (Fig. 648). Sternite VIII of % with ventromedian process (Fig. 653). In &, vaginal apparatus longer than its width and vulvar scale with 4 points (Fig. 657). . . . . . . . . . . . . . Theliopsyche, p. 154

Subfamily Lepidostomatinae Ulmer Only one North American genus is included in this subfamily. 149

Genus Lepidostoma Rambur

Lepidostoma Rambur, 1842, p. 493 Type species designated by Ross, 1944: Lepidostoma squamulosum Rambur = Phryganea hirta Fabricius Revision: Weaver, 1988 As defined here, Lepidostoma has all the characters mentioned in the family description. I shall not add any more here. The genus is now composed of four subgenera. 1a Male and female segment IX with dorsal paired setose warts (Figs. 630, 631, 635). Eastern, usually montane . . . . . . . . . . . . . . . . . . . Mormomyia, p. 152 1b Male and female segment IX without dorsal setose warts . . . . . . . . . . . . . . . 2 2a Parameres absent. Phallocrypt with lateral straps (Figs. 612–615). Female spermatheca with many conspicuous microtrichia, segregated anteriorly in small clusters, sometimes forming a reticulate pattern (Fig. 620a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidostoma, p. 150 2b Parameres present, but sometimes reduced to small lobes (Fig. 624). Phallocrypt without lateral straps (Fig. 612). Female spermatheca with inconspicuous microtrichia, not segregated into reticulate clusters . . . . . . . . . . 3 3a Male front wings with reflexed anterior margin and unique fold in anal region which obscures Cu2 and A1 adjacent to arculus (Fig. 637). Female sternite VIII heavily sclerotized with chalice-shaped area (Fig. 645). Western . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neodinarthrum, p. 153 3b Male front wings usually without reflexed anterior margin and without such fold near arculus, Cu2 atrophied (Fig. 619) except in reosum. Anal groove or median fold sometimes present. Female sternite VIII without sclerotized chalice-shaped area (Fig. 626) . . . . . . . . . . . . . . . . . . . . . . Nosopus, p. 151

Subgenus s. str. Lepidostoma Rambur

Mormonia Curtis, 1834, p. 215 Type species designated by Fischer, 1970: Mormonia gracilicornis Curtis = Phryganea hirta Fabricius Pristosilo Banks, 1899, p. 212 Monobasic type species: Pristosilo canadensis = Mormomyia togata Hagen Notiopsyche Banks, 1905, p. 216 Type species by original designation: Notiopsyche latipennis Banks Oligopsyche Carpenter, 1933, p. 36 Type species by original designation: Notiopsyche carolina Banks 150

Neuropsyche Carpenter, 1933, p. 38 (preoccupied) Type species by original designation: Neuropsyche tibialis Carpenter Flura Milne, 1936, p. 118 (new name for Neuropsyche Carpenter) Type species by replacement: Neuropsyche tibialis Carpenter Male scape longer than head, shorter than in female. Male maxillary palpi 2-segmented, with segmentation obscured; apical segment with scalelike setae. Venation specificially quite variable and slightly dimorphic (Figs. 610, 611). % genitalia (Figs. 612–615): Segment IX short. Segment X with lateral horn-shaped, asymmetrical, sometimes reduced processes that possibly are intermediate appendages. Dorsomesal processes reduced to pair of small lobes. First article of inferior appendages long, narrow, and acuminate or fingerlike; basodorsal process vertical and capitate. Second article in mesal, subapical position. Phallocrypt with lateral straps. Phallotheca and endotheca short. Parameres absent. Aedeagus slender, strongly curved ventrally. & genitalia as in Figs. 616, 617, 620a. This subgenus is quite homogeneous and contains five Nearctic species, two of which are found in our area; L. togatum Hagen has the intermediate appendages hornlike; it is transcontinental, from Alabama and Georgia to Newfoundland and Northwest Territories; L. latipenne Banks has the intermediate appendages in blunt lobes and is eastern in distribution, from Georgia to Nova Scotia. The species seem eurythermal, inhabiting environments such as third- to fifth-order streams or littoral regions of lakes. Subgenus Nosopus McLachlan

Nosopus McLachlan, 1871, p. 114 Monobasic type species: Nosopus podager McLachlan Olemira Banks, 1897, p. 29 Monobasic type species: Olemira americana Banks Eremopsyche Banks 1901, p. 367 Type species by original designation: Eremopsyche frontalis Banks Atomyia Banks, 1905, p. 217 Type species by original designation: Atomyia modesta Banks Alepomyia Banks, 1908, p. 64 Type species by original designation: Alepomyia bryanti Banks 151

Atomyiodes Ulmer, 1911, p. 23 Monobasic type species: Atomyiodes bispinosus Ulmer = Olemira mexicana Banks Alepomyiodes Sibley, 1926, p. 106 Type species by original designation: Lepidostoma wisconsinensis Vorhies = Alepomyia bryanti Banks Arcadopsyche Banks, 1930, p. 129 Monobasic type species: Arcadopsyche prominens Banks Jenortha Milne 1936, p. 119 Type species by original designation: Jenortha canadensis Milne Male scape often simple and cylindrical, but sometimes modified (Fig. 608). Male maxillary palpi often modified, with segmentation usually obscured; basal segment cylindrical; apical segment generally spatulate, mesal surface concave, bearing several modified petiolate scales. Venation strongly dimorphic, specifically quite variable, and sometimes highly modified (Figs. 619, 620, 650). Male front wings with Cu2 usually atrophied. Arculus generally modified. Discoidal cell usually as long as FI; thyridial cell 1.5 times as long as FI. Male hind wings generally not highly modified. FI only always present. Female venation as in Fig. 620. % genitalia (Figs. 621–625): segment IX uniformly short. Segment X about as long as high in lateral view, bearing 1 or 2 apicolateral points. First article of inferior appendages usually triangular and acuminate in lateral view, sometimes regularly slender; basodorsal process variable, usually clavate, directed dorsally or posteriorly; middorsal process horizontal and parallel to 1st article. Phallocrypt without lateral straps. Phallotheca and endotheca short. Parameres present, but variable in shape and size. Aedeagus rather thick, not very long, and slightly curved ventrally. & genitalia (Figs. 626, 627): sternite VIII rectangular, with posterior margin curved outward and with several longitudinal striations. This subgenus is the largest and the most variable. It contains 45 Nearctic species, 19 of which are recorded in our area. Ten of them are restricted to the east, seven to the west, and two are transcontinental. Most species are endemic to mountain streams and springs, but some also occur in lentic environments. Subgenus Mormomyia Banks

Mormomyia Banks, 1907, p. 127 Type species by original designation: Mormomyia vernalis Banks Phanopsyche Banks, 1911, p. 357 Monobasic type species: Phanopsyche grisea Banks 152

Male scape shorter than head, not highly modified, and cylindrical. Male maxillary palpi apparently single-segmented, fingerlike, and slender. Venation strongly dimorphic and specifically quite variable (Figs. 628, 629). Male front wings with thyridal cell petiolate. F5 and each adjacent cell subequal in length. Male hind wings with M and Cu1 unbranched and A1 and A2 fused. Female venation as in Fig. 629. % genitalia (Figs. 630–634): segment IX regularly long, with paired, sometimes huge dorsal setose warts. Segment X large, somewhat circular, membranous in its center, and usually with bi-pointed ventrolateral processes. First article of inferior appendages acuminate and triangular in lateral view, with vertical basodorsal and horizontal middorsal processes. Second article in subapical mesal position. Phallic apparatus usually asymmetrical. Phallocrypt without lateral straps. Phallotheca bulbous or triangular. Endotheca usually small. Aedeagus rather thick and moderately curved ventrally. Parameres present, thick and heavily sclerotized. & genitalia (Figs. 635, 636): segment IX with paired, dorsal setose warts. This subgenus is rather homogenous and contains 15 Nearctic species, 5 of which are recorded from our area. They are eastern in distribution, from Nova Scotia to Quebec. Subgenus Neodinarthrum Weaver

Neodinarthrum Weaver, 1988, p. 75 Type species by original designation: Olemira pluviale Milne Male scape longer than head, slightly shorter than in female. Male maxillary palpi of length subequal to scape, distinctly 2-segmented, each segment cylindrical. Wings narrower than in other subgenera. Front wings with anterior margin reflexed and bearing dark scalelike setae. Venation specifically quite variable and rather dimorphic (Figs. 637, 638, 650): front wings with Cu1 unbranched. Cu2 and A1 interrupted by small fold adjacent to A2 before arculus. A3 about two-thirds as long as A2 and terminating at hind margin. Hind wings with Cu1 unbranched, A1 and A2 apparently fused, and only FI present. Female venation as in Fig. 638. % genitalia (Figs. 639–643): segment IX long dorsally. Segment X with dorsomesal lobes bearing lateral processes that possibly are intermediate appendages and reduced to pair of lateral points. Inferior appendages with 1st article long and narrow. Second article small, acuminate, inserted subapically; subapical process directed posteriorly or reduced; basodorsal process slender, directed dorso-posteriorly. Phallocrypt without lateral straps. Phallotheca short, with rather long tendon. Endotheca short. Parameres present, spiniform, slender, almost as long as aedeagus. Aedeagus slender, strongly curved ventrally. 153

& genitalia (Figs. 644–647): sternite VIII heavily sclerotized and with chalice-shaped area. This subgenus is rather homogeneous and contains eight species distributed in western mountain regions, from Baja California to British Columbia. Three species have been recorded in our area. They are closely related and best recognized by the bi-pointed shape of segment X; aporna Denning, from Montana; pluviale Milne from Alberta and British Columbia, and rayneri Ross from British Columbia. Larvae live in mountain and coastal streams.

Subfamily Theliopsychinae Weaver Only one North American genus is included in this subfamily. Genus Theliopsyche Banks

Theliopsyche Banks, 1911, p. 356 Monobasic type species: Theliopsyche parva Banks Quisilo Milne, 1936, p. 118 Type species by original designation: Silo griseus Hagen Aopsyche Ross, 1938, p. 174 Type species by original designation: Aopsyche corona Ross Revision: Weaver, 1988 Size smaller than in majority of Lepidostoma (length 5–6 mm) and coloration dark brown. Posterior cephalic warts long and narrow. Male scape longer than head and simply cylindrical (Fig. 648). Male maxillary palpi 2-segmented, very long, with each segment distinct and cylindrical. Spurs: 2,4,4. Venation (Figs. 651, 652) slightly dimorphic, not very variable, and more complete than in Lepidostoma. Male front wings with forks I, II, III, and V. Hind wings with forks I and V. Arculus with normal distal configuration. Female front wings (Fig. 652) and hind wings with forks I, II, and V. Hind wings with discoidal cell open in both sexes. % genitalia (Figs. 654–656): segment VIII with a large rounded ventral plate. Segment IX triangular in lateral view. Segment X longer than high; lateral processes usually prominent and spiniform. Preanal and intermediate appendages absent. First article of inferior appendages usually longer than high, appearing bifid or trifid in lateral view. Second article apparently absent. Phallic apparatus thick and straight. Phallotheca elongate and triangular. Endotheca obscured. Parameres absent. Aedeagus with large phallotremal sclerite. 154

& genitalia (Fig. 657): vulvar scale not membranous and forming 4 more or less sharp points. The genus Theliopsyche is found in the Nearctic region only and contains six species, all of which are uncommon. They occur near spring-fed gravelly streams. Only one species, parva Banks, has recently been reported in Quebec. T. grisea Hagen occurs in New Hampshire.

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Family Phryganeidae Leach Phryganidae Leach, 1815, p. 136 Type genus: Phryganea Linnaeus Medium-sized or large, strongly colored, robust, and heavy-bodied insects. & generally larger and heavier than %. Ocelli present and very large. Head with large, oblique occipital warts and well-developed occipital area behind eyes (Fig. 687). Antennae rather short, thickened, with 1st segment short and globular. Subsequent segments also short, with wavy transverse furrow (Fig. 688). Mouthparts fairly strongly extended, with labrum twice as long as its width. Maxillary palpi 4-segmented in % and 5-segmented in & (Figs. 689, 690), with 1st segment very short and 2nd 2 to 3 times longer. Spurs: 2,4,4. Abdominal hemogill system strongly developed, comprising 3 to 5 simple, unforked gill tubes per segment (Fig. 699). Sternite V identical in both sexes, with triangular cleft at anterior margin and unraised pore that serves as opening for internal gland (Fig. 691). Wings not very large by comparison with body, often strongly mottled, and occasionally clothed with dense, very short hairs. Venation very homogeneous in series of genera, virtually complete, with almost all forks present, although their number differs in both sexes (Figs. 662, 692, 693). In front wings, basic number of forks is I, II, III, and V in % and I, II, III, IV, and V in &. In hind wings, I, II, and V in % and I, II, III, and V in &. There are occasionally generic or specific irregularities in &: FIV may be missing from front wings or FIII from hind wings. In front wings, crossvein C-Sc present. Discoidal and thyridial cells long and narrow. Median cell open. FI fused with discoidal cell for long distance and begins well ahead of FII. First anal cell over twice as long as 2nd. Crossvein Cu1bCu2 present, and A1+2+3 ends at the tip of Cu2. In hind wings, discoidal cell very small and triangular. Four free anal veins. % genitalia covered with many very stout setae. Segment IX annulate and often well developed dorsally. Occasionally, on ventroposterior side a concavity equipped with carinae or teeth. Segment X roof-like, simple or complex. Preanal appendages present. Intermediate appendages always absent. Inferior appendages almost always 2-segmented, but modified in shape and movable on segment IX. First articles often complex, joined at base by inner basal plate. Occasionally, they are contiguous or fused together and form posterior concavity equipped with carinae or teeth. This concavity replaces the one occurring on ventral portion of segment IX, 156

when latter inerm. Second segment present, except in Phryganea, and always small, modified, and often inserted before apex of 1st segment. Phallic apparatus strangely resembling that of Hydropsychoidea in structure and location since situated very high above inferior appendages and just below segment X. It is tubular and consists of phallotheca, enclosing membranous endotheca equipped with spines or sclerotized plates, or inerm. Phallotremal sclerite consisting of short tube ending in semicircular capsule. Aedeagus and parameres lost. Phallotheca generally joined by means of sclerotized connection to inner portion of inferior appendages. & genitalia: tergites IX and X firmly fused together to form blunt, flat roof overhanging anal opening. Appendages of segment X absent. Sternite IX enclosed on 3 sides by segment VIII, which is deeply indented to accommodate it, and forming a large 1-, 2-, 3- or 4-lobed vulvar scale at extremity. Supragenital plate reduced to single, inconspicuous membranous fold, except in Hagenella and Banksiola, in which it is prominent and somewhat sclerotized. Ceiling of vaginal cavity membranous and occasionally with sclerotized folds. Vaginal apparatus large, strongly sclerotized, and very complex in structure. The Phryganeidae constitute a small, homogeneous Holarctic family containing roughly 80 species, 26 of which, representing nine genera, occur in our area. Many of the species are distributed across the continent, but they are also found at higher altitudes in the mountain ranges of the east and west and extend fairly far south. Some species are Holarctic, whereas others are confined to the eastern portion of the country. The Phryganeidae have a wide ecological tolerance and inhabit several different types of water habitats. While they have a preference for low-altitude lentic biotopes, some species occur far north in the subarctic region. The striking appearance and coloring of these insects were formerly used as a basis of classification. These characters, however, have proved to be artificial since they do not correspond to genital characters. As a result, the classification of the family was completely updated some years ago (Martynov, 1924, and Milne, 1934). The species are now divided into two subfamilies, the Phryganeinae and Yphriinae, both of which are Nearctic. Only the Phryganeinae live in Canada. 1a Maxillary palpi 4-segmented (Fig. 689) %% . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b Maxillary palpi 5-segmented (Fig. 690) && . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2a Ventroposterior part of segment IX forming a concavity, equipped with carinae or teeth, under or between the base of inferior appendages. Inferior appendages located somewhat above ventral side of abdomen (Figs. 658, 663, 668) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b Ventroposterior part of segment IX simple without concavity. Inferior appendages located at level with ventral side of abdomen (Fig. 673), except in some Agrypnia (Fig. 683) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

157

3a Ventroposterior part of segment IX forming a large concavity delimited by 2 rows of strong teeth (Figs. 668, 670). Inferior appendages 2-segmented. Large rust-brown species. . . . . . . . . . . . . . . . . . . . . . . . . Ptilostomis, p. 161 3b Ventroposterior part of segment IX forming slight concavity delimited on lower side by 2 simple, low carinae (Figs. 658, 659). Inferior appendages apparently single-segmented. Medium-sized orange species, strongly mottled with brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligostomis, p. 159 3c Ventroposterior part of segment IX forming a strong concavity delimited on upper side by high rectangular carina (Figs. 663, 665). Inferior appendages visibly 2-segmented. Medium-sized rust-brown species with brown cross stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hagenella, p. 160 4a Inferior appendages contiguous at base (Figs. 673, 674, 676, 677) . . . . . . . . 5 4b Inferior appendages not contiguous (Fig. 680). . . . . . . . . . . . . . . . . . . . . . . . . 6 5a Inferior appendages low, short, massive, simple in lateral view (Fig. 694). Large, heavy-bodied species with front wings with dark speckles and longitudinal brown stripes . . . . . . . . . . . . . . . . . . . . . . . . . . Phryganea, p. 165 5b Inferior appendages low, short, massive, and complex in lateral view (Fig. 701). Medium-sized species with front wings with dark veins and anterior margin reticulated . . . . . . . . . . . . . . . . . . . . . . . Beothukus, p. 166 5c Inferior appendages prominent, upward-slanting, and ending in 1 or 2 inwardcurved points (Figs. 676, 677). Medium-sized, light-colored species coarsely spotted with brown . . . . . . . . . . . . . . . . . . . . . . . Banksiola, p. 162 5d Inferior appendages vertical, not prominent, completely contiguous to segment IX, forming a vast semicircle, open posteriorly and ending in 2 long points (Fig. 673, 674). Medium-sized brown species striped with brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligotricha, p. 162 6a Dorsal part of segment IX forming an eaves-like structure equipped with a row of stout spines and overhanging segment X. Inferior appendages slender and complex (Figs. 683, 684). . . . . . . . . . . . . . . . . . . Agrypnia, p. 164 6b Dorsal part of segment IX continuous with segment X and without a row of spines. Inferior appendages stouter and simple (Figs. 679, 680). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fabria, p. 163 7a Vulvar scale forming 3 or 4 closely spaced lobes (Figs. 661, 698, 707). . . . 8 7b Vulvar scale forming a more variable number of widely spaced lobes (Figs. 672, 686) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8a Vulvar scale forming 2 short, blunt lateral lobes and 3 fingerlike median lobes (Fig. 661) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligostomis, p. 159 8b Vulvar scale broader than wide (Fig. 707) . . . . . . . . . . . . . Beothukus, p. 166 8c Vulvar scale longer than wide (Fig. 698) . . . . . . . . . . . . . . . Phryganea p. 165 9a Vulvar scale forming prominent, 2-lobed bulb with narrow neck (Fig. 686) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrypnia, p. 164 9b Vulvar scale not forming narrow-necked bulb . . . . . . . . . . . . . . . . . . . . . . . 10

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10a Vulvar scale forming simple, very wide, and semicircular plate (Fig. 682). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fabria, p. 163 10b Vulvar scale forming several points . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a Vulvar scale forming 2 broad-based triangles anterior of which are 2 small, slender lobes (Fig. 667) . . . . . . . . . . . . . . . . . . . . . . . . . . Hagenella, p. 160 11b Vulvar scale forming 4 parallel points situated on same plane (Fig. 675) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligotricha, p. 162 11c Vulvar scale forming 3 weakly prominent points or lobes. . . . . . . . . . . . . . 12 12a R1 in front wings curving before it meets margin (see Fig. 693). Mediumsized species with light-colored wings coarsely spotted with brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Banksiola, p. 162 12b R1 in front wings not curving before it meets margin. Large rust-brown species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptilostomis, p. 161

Genus Oligostomis Kolenati

Oligostomis Kolenati, 1848, p. 80 Type species designated by Martynov, 1924: Phryganea reticulata Linnaeus Body black. Palpi thick. Legs with few short spines. Wings covered with very fine, recumbent hairs. Front wings with more or less fine network of orange markings against brown background. Hind wings with large orange, lunule-shaped spot. Venation: R1 in front wings may be somewhat sinuate before it meets margin. Discoidal cell slightly shorter than its petiole. FIII in hind wings of & as long as its petiole or longer. % genitalia (Figs. 658, 660): segment IX elongate all around with small concavity equipped with carinae or few teeth on ventroposterior side, under inferior appendages. Segment X projected into simple, triangular roof. Preanal appendages of variable size but free, hinged to segment X and bearing apical spines. Inferior appendages positioned fairly high above ventral side of abdomen, developed into simple callipers. In side view, more or less elongate triangular. Second article present but so completely fused with apex of 1st that there is no longer any trace of suture. Phallotheca highly sclerotized, elongate, truncated very obliquely at extremity, with lower apical angle stretched into 2 downward-curving spurs. Endotheca not any longer than phallotheca and spine-free. & genitalia (Fig. 661): ceiling of vaginal cavity without any sclerotized folds. Vulvar scale developed into triangular structure with 3 apical lobes. Its apicolateral angles extend upward and inward to create 2 large lobes forming sclerotized fold above vaginal opening. Vaginal apparatus small, fairly simple, and flattened. 159

The genus Oligostomis is Holarctic in distribution and contains two fairly different Canadian species: ocelligera Walker (Ontario, Quebec, Nova Scotia, and Newfoundland) and pardalis Walker (same distribution but less widespread in the east). In ocelligera, the body is 15–17 mm long and the front wings are indistinctly mottled with orange and brown-grey. In pardalis, the body is over 20 mm long, and the wings have very distinct bright orange stripes. Larvae of both species live in slow-moving stretches of cold streams. Genus Hagenella Martynov

Hagenella Martynov, 1924, p. 79 Type species by original designation: Anabolia clathrata Kolenati Alleodes Banks, 1951, p. 21 Type species by original designation: Neuronia canadensis Banks Body black. Legs with fairly numerous erect, black spines. Wings forming wide ellipses, and both pairs similarly shaped. Membrane covered with very fine, sparse hairs. Front wings brown and very regularly speckled with round, light-colored spots. Venation (Fig. 662): R1 in front wings fairly strongly sinuate before margin. Discoidal cell fairly wide and as long as its petiole. In & hind wings, FIII distinctly shorter than its peduncle. % genitalia (Figs. 663–666): segment IX fairly irregularly elongate all around with large concavity delimited on upper side by high rectangular carina, on ventroposterior side between inferior appendages; walls of concavity very grainy. Segment X forming flat roof unsclerotized at base, except for 2 longitudinal ribs, and ending in several blunt lobes. Preanal appendages developed on lateral side of segment X into 2 large, blunt lobes bearing 1 or 2 long spines. Inferior appendages in simple callipers with massive, angular 1st article and small 2nd article, articulated with but also partially fused to 1st. Phallic apparatus comprising blunt phallotheca equipped with 2 upper basal horns that hinge phallotheca to lateral sides of segment IX. Phallotheca ending in 2 large, blunt lobes. Endotheca short, forming complex unit of lobes covered with tiny warts. & genitalia (Fig. 667): supragenital plate large, very wide, and complex. Vulvar scale very large, ending in 2 triangular lobes framing 2 smaller lobes arising from internal base. Two slender lobes preapically. Vaginal apparatus long and narrow, ending posteriorly in long slender point. The genus Hagenella has a Holarctic distribution, but contains only a single Nearctic species, canadensis Banks, which has been reported in Ontario and Quebec and lives in ponds and marshes.

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Genus Ptilostomis Kolenati

Ptilostomis Kolenati, 1859, p. 198 Type species designated by Fischer, 1964: Ptilostomis kovalevskii var. ß = Phryganea semifasciata Say Neuronella Banks, 1951, p. 22 Type species by original designation: Neuronia angustipennis Hagen Body rust-brown. Wings fairly wide. Front wings orange-yellow, indistinctly mottled with light brown and covered with fairly long, erect hairs. Palpi fairly long. Legs with numerous, long, erect, black spines. Venation: in front wings, R1 barely sinuate before reaching margin. Discoidal cell long, narrow, and slightly longer than its petiole. FI begins at point corresponding to half discoidal length or ahead of this level. In hind wings of &, FIII slightly longer than its petiole. % genitalia (Figs. 668–671): segment IX quite irregularly elongate, deeply indented laterally to accommodate inferior appendages. Tucked under latter ventrally and forming large mass, concave posteriorly, and provided with 2 rows of teeth. Segment X small, very simple, and virtually indistinct from preanal appendages, which are far more developed. Latter either simple (angustipennis) or developed into 2 long, slender, and parallel branches equipped with stout spines. Inferior appendages situated fairly high above ventral side of abdomen and shaped into 2-segmented callipers; 1st articles are massive, concave on inner surface, and slightly divergent; 2nd articles in long, simple, convergent spurs. Phallic apparatus with slightly sclerotized, short phallotheca continuous with endotheca that is long, bulbous, and equipped with single pair of flattened spines. & genitalia (Fig. 672): vulvar scale very wide, ending in 3 small lobes: lateral lobes slender and median lobe forms faintly prominent triangle. Supragenital plate membranous, and ceiling of vaginal cavity without sclerotized folds. Vaginal apparatus very bulky, sclerotized, and complex. The genus Ptilostomis is restricted to the Nearctic region and comprises only four species: ocellifera Walker and semifasciata Say, which are very common and found across the continent, and postica Walker and angustipennis Hagen, which are less common and confined to eastern and central parts of the continent. The last species has been isolated in the subgenus Neuronella Banks. The %% are distinguished particularly by the development of the preanal appendages. Larvae occur mainly in lakes and slow-flowing watercourses.

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Genus Oligotricha Rambur

Oligotricha Rambur, 1842, p. 472 Type species designated by Milne, 1934: Oligotricha chloroneura Rambur = Phryganea striata Linnaeus Body black. Wings wide. Front wings virtually smooth, light-colored, and extensively cross-striped with brown. Palpi large. Legs with fairly numerous erect black spines. Venation: in front wings, R1 fairly markedly sinuate before reaching margin. Discoidal cell wide and slightly shorter than its petiole. FIII of hind wings in & sessile or with very short petiole. % genitalia (Figs. 673, 674): segment IX stout and regularly elongate all around. Segment X forming flat, trapezoidal roof with small indentation at extremity. Preanal appendages reduced, resembling simple warts fused with base of segment X. Inferior appendages reaching ventral side of abdomen and developed into large, 2-segmented callipers, deeply modified in shape. First articles contiguous and vertical, together forming vast semicircular unit open posteriorly, with inner and lower margins forming dentate carinae; external apical angle drawn into long, stout, inwardprojecting spur. Second article inserted above this spur and similarly shaped and subparallel. Phallic apparatus very stout, consisting of arched, strongly sclerotized phallotheca forming blunt lower apical angle and short, thick endotheca equipped with tuberculate sclerotized plates. & genitalia (Fig. 675): ceiling of vaginal cavity with sclerotized folds. Vulvar scale very large, ending in 4 long, slender lobes; 2 inner lobes may form 2 small inner angles. Vaginal apparatus large, with lower median portion equipped with sort of denticulate and strongly sclerotized ridge. The genus Oligotricha is small, Palearctic in distribution and comprises half a dozen species. To date, none of them has been reported in Canada, but the Palaearctic species, lapponica Hagen, has been captured in Alaska. Genus Banksiola Martynov

Banksiola Martynov, 1924, p. 216 Type species by original designation: Phryganea dossuaria Say Revision: Wiggins, 1956 Body brown. Wings not very wide, with front wings covered with very short, dense hairs. Membrane light-colored and strongly speckled with brown (Banksiola dossuaria wings are reminiscent of those of a scorpion fly). Palpi stout. Legs with many erect black spines. Venation: R1 fairly strongly sinuate before meeting margin. Discoidal cell slightly 162

shorter than its petiole. FI starts ahead of point corresponding to half discoidal cell length. Forks I, II, III, V and I, II, V present in % and I, II, III, V and I, II, IV, V in &. % genitalia (Figs. 676, 677): segment IX crescent-shaped in side view. Segment X developed into simple roof, with width exceeding length. Preanal appendages forming small, simple warts at base of segment X. Inferior appendages reaching ventral side of abdomen and consistently 2-segmented. First articles massive and contiguous, occasionally bearing teeth and tubercles on inner surface and forming 2 or 3 apical and subapical branches. Second article reduced to tiny, nonprominent, sclerotized plate located at base of apical branch and constantly present. Phallotheca fairly slender, strongly sclerotized, very obliquely truncated at extremity, and ending in more or less sharp lower apical angle. Endotheca long, developed into 1 or 2 strongly erectile tubes, occasionally provided with lateral bulbs and bearing stout, spiny framework. & genitalia (Fig. 678): supragenital plate membranous, but prominent, forming sclerotized folds at base. Vulvar scale highly sclerotized and ending in three lobes; median lobe widest and occasionally secondarily bifid. Vaginal apparatus large and strongly sclerotized. The genus Banksiola is exclusively Nearctic in distribution with five species, all of them reported in our area. They are eastern except crotchi Banks, which occurs across the country. The species have a wide ecological tolerance and inhabit a fairly broad range of biotopes: lakes, ponds, marshes, and essentially slow-moving watercourses. Genus Fabria Milne

Fabria Milne, 1934, p. 9 Type species by original designation: Neuronia inornata Banks Body brown. Eyes very prominent. Ocelli very large. Palpi strongly thickened. Front tarsi shortened. Middle tarsi and tibiae slightly flattened in both sexes. Wings fairly wide, with front wings almost smooth and very finely and indistinctly mottled with light brown. Venation: in front wings, R1 sinuate before reaching margin. Discoidal cell narrow and longer than its petiole. FI starts well ahead of point corresponding to half length of discoidal cell. In hind wings of &, FIII much longer than its peduncle. % genitalia (Figs. 679–681): segment IX irregularly elongate. Segment X large, very massive, with rectangular shape in side view and extended lower apical angle. Preanal appendages developed into large, weakly protruding warts. Inferior appendages situated very slightly above ventral side of abdomen and in shape of 2-segmented callipers. First articles completely separate from each other, elongated, and end in upper apical point. Second article inserted behind this point and forming triangle 163

with very elongated apex. Phallotheca consists of long, not very sclerotized cylinder, with lower apical angles forming 2 upward-curving points. No sclerotized connection between phallotheca and segment X or inferior appendages. Endotheca as long as phallotheca and developed into simple, spine-free cylindrical tube. & genitalia very wide (Fig. 682): sternite VIII with sharp, complex apical angles. Vulvar scale not produced into any points but forming simple, strongly thickened, very wide lip, describing an arc at apical margin that is slightly indented in middle. Supragenital plate membranous, not prominent, forming few sclerotized folds at base. Vaginal apparatus not very large and situated at end of wide-mouthed funnel-shaped vestibule. The genus Fabria contains only one species, inornata Banks, which is rare and has been reported in Alberta, Manitoba, Ontario, Quebec, and southward. Larvae of this species live in dense beds of submerged aquatic plants in standing or slow-moving waters. Genus Agrypnia Curtis

Agrypnia Curtis, 1835, pl. 540 Monobasic type species: Agrypnia pagetana Curtis Dasystegia Wallengren, 1880, p. 66 Type species designated by Milne, 1934: Phryganea obsoleta Hagen Phryganomyia Banks, 1907, p. 122 Type species by original designation: Asynarchus alascensis Banks = Agrypnia glacialis Hagen Prophryganea Martynov, 1924, p. 78 Type species by original designation: Prophryganea principalis Martynov Jyrvia Milne, 1934, p. 3 Type species by original designation: Neuronia vestita Walker Species in this genus exhibit considerable diversity and variability with respect to size, coloring, wing venation, hair system, and spiny framework of the legs. In some northern or higher altitude populations of some species, the cold seems to influence phenotype development and causes some deterioration, such as reduced eyes, elongated head, thickened antennae, palpi, and legs, shortened front tarsi, compact body, and wings that are discolored and more or less reduced in size, shrunken and, to some degree, poorly developed. Venation, in addition, is somewhat aberrant. As indicated above, Agrypnia has been divided into five genera or subgenera, but the genital characters of both sexes show that they actually constitute a single phylogenetically uniform genus. 164

Front wings shaped into fairly even strips with obliquely truncated extremities. Hind wings in wide triangle shape with rounded posterior margin. Venation: in front wings, crossvein C-Sc occasionally absent. Discoidal cell of variable length. FI starts behind point corresponding to half length of discoidal cell. In hind wings, crossvein M-Cu1a forms strong curve open exteriorly. FIII often absent in &. % genitalia (Figs. 683–685): segment IX fairly uniformly long but with ventral side often upward-slanting and dorsal side forming eaves-like structure bearing long spiniform setae and overhanging segment X. Latter generally positioned fairly low and forming simple roof occasionally equipped with stout spines. Preanal appendages developed into small free lobes. Inferior appendages 2-segmented and may or may not reach ventral side of abdomen. First articles spaced far apart, slender, upward-slanting, with apical and posterior margins bearing 1, 2, or 3 processes in shape of teeth, branches or heels. Second article always present, bulb- or clubshaped, often thickly covered with hairs and always inserted behind apical point of 1st segment. Phallic apparatus long and slender. Phallotheca long tube with long apodemal part, without sclerotized connection either with segment X or inferior appendages; lower apical angle blunt. Endotheca simple, long, and slender with framework of long spines. & genitalia (Fig. 686): segments IX and X firmly fused into wide roof-like structure with somewhat prominent lateral angles. Supragenital plate very wide and membranous, except at base. Vulvar scale complex, forming large, rounded, upper carina, below which lies bulbous, bilobed, carinate, and very hairy piece as long as segment X. Vaginal apparatus large, complex, and strongly sclerotized. The genus Agrypnia is the largest in the family. It is Holarctic in distribution and contains 20 species, 9 of which are widely distributed across Canada and in the mountain ranges in the western part of the continent. They inhabit mainly lentic biotopes and are found in lakes, ponds, marshes, and occasionally gently flowing water. Genus Phryganea Linnaeus

Phryganea Linnaeus, 1758, p. 547 Type species designated by Westwood, 1840: Phryganea grandis Linnaeus Body brown. Wings covered with very fine, appressed hairs. Front wings with stripes and spots in various shades of brown, a case of mimicry. Palpi stout. Legs with many erect black spines. Abdominal hemogill system particularly well developed (Fig. 699). Venation (Figs. 692, 693): in front wings, R1 sinuate before extremity. Discoidal cell very narrow and 2.5 times longer than its petiole. FI starting behind point corresponding to 165

half discoidal cell length. In &, M4 on front wings frequently converges with Cu1a, and FIII on hind wings much longer than its petiole. % genitalia (Figs. 694–696): segment IX of very uneven length all around. Segment X obscurely structured, consisting of very short median part, which actually is probably segment X itself, and 2 bifid lateral parts that, not impossibly, could be the preanal appendages. Lower branches bearing large warts. Inferior appendages single-segmented in Canadian species, reaching ventral side of abdomen and contiguous; triangular in side view, concave on upper inner surface, and displaying complex relief. Sclerotized connection between phallotheca and inferior appendages. Phallotheca short with 2 stout, denticulate, and paired, lower apical angles. Endotheca fairly long, slender, equipped with 2 stout spines on upper basal side and 2 small apical thorns. & genitalia (Figs. 697, 698): supragenital plate membranous, forming complex sclerotized folds at base. Vulvar scale not very wide and developed into 3 slender apical lobes. Median lobe with a short cleft. Vaginal system very large and stout. The genus Phryganea consists of large, heavy-bodied species with mimetic coloring. Eight species are known to date. Only two of these are found in the Nearctic region and have been classified in the subgenus Neophryganea Martynov. Retaining this subgenus is not of concern here. P. cinerea Walker is transcontinental in distribution and can be found from Alaska to Newfoundland and along the mountains in the western part of the continent. P. sayi Milne has a smaller range and has been reported in Ontario, Quebec, and the east central United States.

Genus Beothukus Wiggins and Larson

Beothukus Wiggins and Larson, 1989, p. 1551 Type species by original designation: Ecclisomyia complicata Banks Body medium brown. Vertex with only posterior warts present. Antennae dark brown, with lighter annulation. Legs with well-developed black spines. Front wings with light covering of hairs. Veins outlined with dark brown and reticulation present along anterior margin. Hind wings uniformly colored. Venation similar to Phryganea (Fig. 700). % genitalia (Figs. 701–704): segment IX well developed, especially laterally. Segment X large and complex, consisting of median part narrowly cleft apically, extending laterally as curved flange bearing stout setae; near base, 2 pairs of slender processes of very unequal length. Preanal appendages present, small, and knoblike. Inferior appendages 2segmented. First article scoop-shaped in lateral view; in caudal view, concave mesally, with upper lateral margin forming 3 lobes. Second article 166

small. Phallotheca tubelike and sharply angled downward at its extremity. Endotheca very small and equipped with 2 tiny spines. & genitalia (Figs. 705–707): supragenital plate terminating in 3 hairy lobes. Vulvar scale 3-lobed, the lateral ones simple, fingerlike, the median one shorter and bifid. The genus Beothukus contains only one species, complicatus Banks, conspicuous in the family by its very complex % genitalia. It is very rare and local and recorded from Newfoundland, Quebec, Ontario, and Alberta. The species lives in lakes and bogs.

167

Family Brachycentridae Ulmer Brachycentrinae Ulmer, 1903, p. 85 Type genus: Brachycentrus Curtis Brachycentridae Ross, 1944, p. 260 Head particularly short and wide. Eyes small and hairy (Fig. 715). Vertex with 3 pairs of warts. Ocelli absent. First antennal segment short and slightly thickened. Maxillary palpi 3-segmented in % and 5-segmented in & (Figs. 716–718). Spurs: 2,2,2 or 2,3,3 or 2,4,4. Gland of abdominal sternite V in shape of small sphere with opening on anterior margin of segment. Abdominal hemogill system consisting of 3 to 4 long, simple tubes per segment. Wings evenly elliptical at extremity. Venation slightly simplified and exhibiting faint sexual dimorphism (Figs. 720, 721). In front wings, forks I, II, III, and V present in % and I, II, III, IV, and V or I, II, III, and V in &. In hind wings, forks I and V or I, II, and V present in % and I, II, III, and V or I and V in &. In front wings, discoidal cell small and closed, thyridial cell very long and closed, and median cell open. Cu2 extends to wing margin and joined to Cu1b by crossvein. Anal veins occasionally simplified. In hind wings, discoidal cell open or closed. Three or 4 anal veins. % genitalia: segment IX well developed. Segment X developed into horizontal, bifid plate located fairly low and provided with short spines. The conformation of this plate in Eobrachycentrus suggests that it is not actually segment X, but that the latter is virtually absent and has been replaced with contiguous, more or less fused intermediate appendages. Preanal appendages generally large and situated dorsally. Inferior appendages simple, upward-slanting, and usually single-segmented but occasionally equipped with 2nd segment. Two basal articles joined together at base by V-shaped ventral plate. Phallic apparatus situated midway along segment IX and without any sclerotized connections with neighboring parts, consisting of tubular phallotheca and fairly small, spine-free endotheca. & genitalia: tergite IX small, with an anterior apodemal point. Sternite IX absent and replaced with sternite VIII, which is strongly developed and concave to provide temporary holding place for egg mass and with lateral reinforcements. Segment X small, appendage-free, and concave on 168

inner surface and with anal opening under its extremity. Vaginal opening between sternite VIII and segment X. Vulvar scale absent. Vaginal apparatus simple, slender, and at end of membranous vestibule. The Brachycentridae constitute a fairly small family of Holarctic distribution. Of the five genera represented in North America, four are found in Canada: Brachycentrus Curtis, Micrasema McLachlan, Eobrachycentrus Wiggins, and Amiocentrus Ross. They inhabit all types of running water and occur far north in the subarctic region. 1a Spurs: 2,4,4. Front wings with R1 sinuate at level with pterostigma. FII sessile (Figs. 708, 709) . . . . . . . . . . . . . . . . . . . . . . . Eobrachycentrus, p. 169 1b Spurs: 2,2,3 or 2,3,3. Front wings with R1 sinuate at level with pterostigma. FII sessile (Figs. 720, 271) . . . . . . . . . . . . . . . . . . . . Brachycentrus, p. 170 1c Spurs: 2,2,2. Front wings with R1 not sinuate at level with pterostigma. FII petiolate (Figs. 734, 735) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Male labial palpi as long as maxillary palpi (Figs. 730, 731). Female hind wings with SR 3-branched (Fig. 735) . . . . . . . . . . . . . . Micrasema, p. 172 2b Male labial palpi twice as long as maxillary palpi (Figs. 743, 744). Female hind wings with SR 4-branched (Fig. 745). . . . . . . . . Amiocentrus, p. 173

Genus Eobrachycentrus Wiggins

Eobrachycentrus Wiggins, 1965, p. 1090 Type species by original designation: Eobrachycentrus gelidae Wiggins Head particularly short and wide. Antennae crenulated on inferior face. Maxillary palpi of % very short, 3-segmented, with last segment reduced and pointed (Fig. 710). Maxillary palpi of & well developed. Spurs: 2,4,4. Legs with few spines only. Wings covered with dense, appressed hairs. Front and hind wings subequal in width, but anal area not very large in hind wings. Venation (Figs. 708, 709): in front wings, discoidal cell small but fairly elongate. R1 only slightly sinuate at the level of pterostigma. Forks I, II, III, and V present in both sexes. In %, only 2 short anal veins enclosing small cell, and postcostal area very wide. In &, A1 and A2 very long, forming very long cell. A3 vestigial, and postcostal area narrow. In hind wings, forks I, II, and V present in % and I, II, III, and V in &. % genitalia (Figs. 711, 712): segment IX appearing triangular in lateral view. Preanal appendages small, globular, and pebbled. Segment X forming membranous dorsal lobe and 2 horizontal, strongly sclerotized, very elongate and subparallel parts that are probably the intermediate appendages. Inferior appendages massive and complex, with 1st article 169

crescent-shaped and strongly concave on inner surface. Second segment large, developed into stout horizontal rod inserted far within 1st segment. Phallic apparatus very large, with endotheca short and equipped with an elongated phallotremal sclerite. & genitalia (Figs. 713, 714): tergite IX massive. Segment X small, bifid, and concave on underside. Vaginal apparatus simple and located at end of long, slightly sclerotized vestibule. The genus Eobrachycentrus contains only a single species, gelidae Wiggins, which was discovered in Oregon at high altitudes where it inhabits very cold springs. Since then, it has also been found in the larval stage in the state of Washington and in British Columbia. Genus Brachycentrus Curtis

Brachycentrus Curtis, 1834, p. 216 Monobasic type species: Brachycentrus subnubilus Curtis Sphinctogaster Provancher, 1877, p. 262 Monobasic type species: Sphinctogaster lutescens Provancher Oligoplectrodes Martynov, 1909, p. 294 Type species (first included): Oligoplectrodes potanini Martynov = Brachycentrus americanus Banks Revision: Flint, 1984 Female substantially larger and heavier than %. Antennae faintly crenulated on inferior face and distinctly thicker in % than in &. Maxillary palpi of % short, thick, densely covered with erect hairs raised against face, and comprising 3 subequal segments. Labial palpi of % also thick and distinctly longer than maxillary palpi. In &, maxillary palpi slender and weakly developed, but with thickened 1st segment (Figs. 716–718). Legs covered with short hairs and number of spines (Fig. 719). Spurs: 2,2,2 or 2,3,3. Subapical middle spurs very apically positioned. Sternite VII of % with short ventral plate. Wings covered with short, sparse hairs. Hind wings as wide as front wings, subrectangular, with well-developed anal area. Venation (Figs. 720, 721): in front wings, R1 strongly sinuate at level of pterostigma. Forks I, II, III, and V present in % and I, II, III, IV, and V in &. FII sessile. There are 3 anal veins forming 3 cells. In hind wings, forks I and V present in % and I, II, III, and V in &. M single-branched in % and 3-branched in &. Discoidal cell open. % genitalia (Figs. 722–725): segment IX evenly short all around. Preanal appendages large, strongly sclerotized, dorsally positioned, parallel, and occasionally more or less fused to each other at base. Segment X 170

forming more or less bifid plate equipped with variable number of spines. Inferior appendages single-articulated, subvertical, contiguous to segment IX for nearly half their length; apical half free and simple; lower apical angle well defined, and apical edge generally concave. Phallotheca consists of slender tube prolonged by endotheca that contains large phallotremal sclerite. & genitalia (Figs. 726, 727): all abdominal segments can be easily telescoped, whereas sternites VII and VIII are enlarged and concave, forming a large area for holding the egg mass. Segment X concave on entire undersurface. Vaginal opening gaping under segment X. Vaginal apparatus fairly simple. The genus Brachycentrus is Holarctic and Oriental in distribution and is represented in the Nearctic region by a dozen species, nine of which have been recorded in our area. They have been classified in four subgenera (Flint 1984), which are very closely related and can be distinguished by a small number of % genitalic characters only. In fact, they are not subgenera, but merely species groups. I do not adopt them here, but mention three of them that are represented in our area: Sphinctogaster Provancher has the preanal appendages entirely distinct from each other (Fig. 723). Segment X elongated, more or less bifid. Apex of the inferior appendages slender and simple (type species: Sphinctogaster lateralis Say). Brachycentrus Curtis has the preanal appendages fused to each other at their base. Segment X not elongated and very blunt. Apex of the inferior appendages slender, simple. Anteroventral part of segment IX extended anteriorly (type species: Brachycentrus nigrisoma Banks). Oligoplectrodes Martynov has the preanal appendages entirely fused together into a single dorsal plate. Segment X rather elongated, bifid, and with a pair of rodlike processes in superior position. Mesoventral part of segment IX extended anteriorly. Apex of inferior appendages thick and complex (type species: Oligoplectrodes potanini Martynov). Brachycentrus inhabit all types of running water, but have a preference for large watercourses. Adults of some species emerge in massive numbers in May and June. They are active during the day as well as at night and may exist in such large numbers as to become a serious nuisance. The && in particular attract attention with the bright green ovigerous mass located at the extremity of the abdomen, which can be seen when the insects fly. Near some rivers, such as the St. Lawrence in the Montreal area, two species, incanus Hagen and lateralis Say, have successive flight periods, i.e., the first in May and the second in June (Corbet et al., 1966). 171

Genus Micrasema McLachlan

Micrasema McLachlan, 1876, p. 259 Type species by original designation: Oligoplectrum morosum McLachlan Revision: Schmid, 1983 Antennae not crenulated and of equal thickness in both sexes. Maxillary palpi of % fairly long and generally extending as far as tip of 1st antennal segment. Labial palpi of % as long as maxillary palpi. In &, maxillary palpi long and slender (Figs. 730–733). Spurs: 2,2,2. Wings with a thick pilosity. Hind wings of & with a median area more heavily clothed with hairs on the underside. Hind wings generally more or less triangular, with reduced anal area. Venation (Figs. 734, 735): in front wings, R1 not sinuate at level of pterostigma. Forks I, II, III, and V present in % and I, II, III, IV, and V present in &, with FII petiolate. A2 partly lost, with the result that there are only 2 anal cells. In hind wings, forks I and V present in both sexes. M simple or divided in % and always divided in &. % genitalia (Figs. 736–740): segment IX very short dorsally, but elongated laterally. Preanal appendages triangular, fairly small, contiguous at base, slightly divergent, and membranous at base. Segment X developed into bifid plate provided with short spines at extreme tips and bearing 2 small basal median lobes also equipped with spines. Inferior appendages 1- or 2-segmented, contiguous to segment IX for short distance, without pronounced lower basal angle and with apical edge straight. Phallotheca generally curved downward. Endotheca not readily eversible, not very long, and often with faintly developed spiny framework. & genitalia (Figs. 741, 742): only sternite VIII developed for purpose of holding eggs, simple and very wide. Segment X with base only concave on underside. Vaginal apparatus very simple. The genus Micrasema has a Holarctic and Oriental distribution and is represented in North America by about 20 species, only 7 of which have been reported in our area. The genus is divided into two species groups: the rusticum group, confined to the eastern and central part of the country, and the gelidum group, found in western Canada and containing one circumboreal species, gelidum McLachlan. In the rusticum group, the front tarsi of the & are modified and the claws on the 3 legs are larger than in the % (Figs. 728, 729). The inferior appendages of the % are narrow and single-segmented (Fig. 736). In the gelidum group, the legs display only very slight sexual dimorphism. The inferior appendages of the % are wider and 2-segmented: the middle of the upper margin forms an indentation delimited by 2 lobes. The 2nd segment, which is inserted in this indentation, is very reduced, but easily recognizable and usually claw-shaped (Fig. 740). Larvae inhabit running water and occasionally very small 172

streams. Females keep their egg mass for awhile before depositing it. The hairs on the underside of the hind wings then adhere to the egg mass. Genus Amiocentrus Ross

Brachycentrus (Amiocentrus) Ross, 1938, p. 177 Type species by original designation: Brachycentrus aspilus Ross Amiocentrus Wiggins, 1965, p. 1097 Antennae slightly thicker in % than in & and weakly crenulated on inferior face. Male maxillary palpi quite short, with both 1st articles fused to each other (Figs. 743, 744) and barely reaching base of scape. Male labial palpi thick and twice longer than maxillary palpi. In &, maxillary palpi very long and thin. Spurs: 2,2,2. Wings covered with dense, appressed pilosity. Shape of wings rather similar to Brachycentrus; anal area of hind wings well developed. Venation rather similar to Micrasema (Fig. 745). In front wings, discoidal cell small, FII narrow basally and petiolated, and FIII pointed. A2 precociously ending on A1, but 2nd anal cell is distinctly longer than in Micrasema. In posterior wings, SR 4-branched as in Brachycentrus and M 2-branched as in Micrasema. % genitalia (Figs. 746–749): segment IX short dorsally and ventrally, but elongated laterally. Preanal appendages subovoid, seen laterally and dorsally, distinct from each other, but united by median membranous zone. Segment X in obtuse and simple lobe, seen laterally; from above, it looks triangular, with lateral edges thickened. Inferior appendages 2-segmented, appearing simple, slender, and bifid laterally; seen posteriorly, they look massive, complex, and of semicircular shape. Second segment small. Phallic apparatus as in Micrasema. & genitalia (Fig. 750): sternite VIII with sclerotized frame and median longitudinal suture. Segment X subrectangular, broader than long, and ending in 2 small lobes. Vulvar scale simple and bilobed. Vaginal apparatus simple. Considering adult characters only, Amiocentrus is not a well-defined genus. The only known species would normally be placed in the genus Micrasema, but Wiggins (1965) showed that the larvae have quite distinct generic characters. The genus Amiocentrus contains only one species, aspilus Ross, with a wide distribution from British Columbia to Nevada and from Colorado to Montana. The larvae live in slow moving parts of large rivers. 173

Family Sericostomatidae Stephens Sericostomatidae Stephens, 1836, p. 148 Type genus: Sericostoma Latreille The Sericostomatidae constitute a medium-sized family distributed in the Holarctic and Oriental regions and the temperate zones of the Southern Hemisphere. The Nearctic species were originally classified into a certain number of genera that were all reduced to synonymy with Sericostoma (Ross, 1944, p. 266), since they were based exclusively on secondary sexual characters. In 1974, Ross and Wallace revised the classification of these species and once again placed them in a greater number of genera and subgenera than necessary, since the characters defining these taxa are not significant beyond the specific level. Adult characters show that only Gumaga Tsuda and Agarodes Banks are valid, and in neither case is there any justification for further division into subgenera. Only Agarodes is represented in our area. Genus Agarodes Banks

Agarodes Banks, 1899, p. 217 Monobasic type species: Agarodes griseus Banks Psiloneura Banks, 1914, p. 264 Type species by original designation: Psiloneura moesta Banks = Notidobia distincta Ulmer Fattigia Ross and Wallace, 1974, p. 46 Type species by original designation: Fattigia pelle Ross Revision: Ross and Scott, 1974 Like most of the Sericostomatidae, the genus Agarodes displays strong sexual dimorphism in the antennae and maxillary palpi. Eyes very hairy and more prominent in % than in &. Vertex with 2 pairs of warts in % and 3 pairs in &. First antennal segments greatly enlarged in % and equipped with dense pilosity thickened on outer surface. These segments are contiguous and cover a variable portion of vertex posteriorly (Fig. 751). They bear glandular organs on inner surface. In &, 1st antennal segments slightly thickened and simple. Antennae stout. Maxillary palpi of % short and very thick, contiguous, raised masklike against face, and 174

apparently single-segmented. Inner surface has lobes and erectile membranous areas densely covered with golden hairs and serves a glandular function (Figs. 751, 752). Labial palpi of % very large and in all likelihood assuming tactile functions of maxillary palpi, which have taken over other functions. Maxillary palpi of & comprising 5 subequal segments (Figs. 753, 754). Legs stout, densely covered with scaly hairs, and bearing numerous black spines. Spurs: 2,2,4. Wings densely clothed with appressed pilosity and bluntly rounded at tips. Hind wings quite a bit shorter and slightly narrower than front wings. Venation (Fig. 755) similar in both sexes, almost complete, with forks I, II, III, and V present in front wings and I, II, and V in hind wings. In front wings, RS and M lie close together, and subradial and thyridial cells very wide. FI likewise very wide. Discoidal cell very small, narrow, and in contact with FI for almost its entire length. Crossvein R1–R2 present and right at base of R2. Median cell open. Cu2 terminates at Cu1b. Only 2 anal veins forming single cell. In hind wings, discoidal cell also small, and crossvein R1–R2 absent. Only 3 anal veins and 1st one incomplete. % genitalia (Figs. 756–758): segment IX stout, especially laterally. Segment X seems to consist of 2 contiguous branches, but a comparison of the Canadian species with other foreign forms shows that these branches are actually the intermediate appendages and that segment X is reduced to the laterobasal portions of these branches and is closely and completely integrated with these structures. Preanal appendages small, ovoid, and free. Inferior appendages single-segmented, in form of long, upward-slanting ovals with pebbled texture. Inner basal portion forms 2 large and more or less forked appendages joined at base. Phallic apparatus situated very high in abdomen, under intermediate appendages and without sclerotized connections with neighboring parts. It is composed of long tubular phallotheca extended virtually without discontinuity by short, spine-free endotheca without distinct phallotremal sclerite. & genitalia (Figs. 759–761): sternite VIII completely free and capable of folding down and forming large concavity for temporarily holding egg mass. Segment IX consists of triangular tergite, clearly distinct from segment X, and has lateral concavities. Sternite IX folded down posteriorly and in shape of 2 large blunt parts. Segment X developed into horizontal plate provided with appendages that cannot be readily distinguished from plate. Anal opening under rim of segment X. Vaginal opening in narrow slit situated between ventral parts of segment IX. A long and complex vaginal vestibule leading to fairly complex vaginal apparatus. Ross and Wallace (1974) divided the genus Agarodes into two subgenera, Psiloneura Banks and Agarodes s. str. Each of these subgenera is represented in Canada by a single species. The difference between these subgenera is so slight that I consider them as mere species groups. 175

The genus Agarodes is exclusively Nearctic in distribution, containing 10 species, two of which are Canadian and found in the eastern part of the country: distinctus Ulmer and griseus Banks. In the former species, the inner lobes of the inferior appendages are simple and blunt, whereas in griseus, these lobes form slender, forked branches. Larvae of these species live in cold, clear streams and in the sandy portions of rivers and lakes.

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Family Helicopsychidae Ulmer Helicopsychinae Ulmer, 1906, p. 104 Type genus: Helicopsyche von Siebold Helicopsychidae sensu Ross, 1944, p. 266 This family is represented in Canada by only a single genus. Genus Helicopsyche von Siebold

Helicospyche von Siebold, 1856, p. 38 Type species designated by Flint, 1964: Helicopsyche shuttleworthi von Siebold Head short and very wide. Eyes very large and globular in % and slightly less protuberant in &. Vertex with 2 very large median warts. In %, 2 other pedunculate warts at base of antennae. Ocelli absent. First antennal segment as long as head. In %, it is thick and slightly bulging, with inner surface erectile and covered with dense and very fine golden hairs (Fig. 762). In &, this segment more slender and unspecialized. Maxillary palpi fairly long, strongly covered with erect hairs, and 2-segmented in %, but 5-segmented in & (Figs. 763, 764). First segment long, slender, and curved at base in both sexes. Legs densely covered with scaly pilosity and provided with numerous, but tiny black spines. Abdomen covered with dense pilosity, not only on tergites and sternites, but also on pleurites. Gland of sternite V absent in both sexes. Sternites II to IV with a coarse, sclerotized reticulation. Hemogill system comprising simple, long, extremely fine tubes very difficult to observe. Sternite VI with median point in both sexes. Spurs: 1,2,4. Wings covered with dense pilosity. Front wings of moderate width, forming blunt ellipses at tips. Hind wings smaller, slightly narrower, and fairly pointed. Costal margin forming angle and bearing row of hooks on basal half. Venation (Fig. 765) similar in both sexes and considerably simplified, especially in hind wings. Forks I, III, IV, and V present in front wings and I and V in hind wings. Ocellar spots absent. In front wings, discoidal and thyridial cells very large. M2 and M3 fused to each other at base. A1+2 and A3 not confluent. In hind wings, discoidal cell absent, RS 3-branched, M2 2-branched, and all crossveins absent. Two anal veins. 177

% genitalia (Figs. 766–768): segment IX stout ventrally and particularly laterally. Segment X a tripartite horizontal piece, but comparison of borealis with other species shows that segment X actually is only the median part and that the parts completely integrated into this structure on either side in reality are the intermediate appendages. The whole unit resembles a horizontal roof that is rounded and indented at the extremity and equipped with short, thick spinules. Preanal appendages in small free knobs. Inferior appendages single-segmented, large, wide, upward-slanting, with denticulate and strongly hairy sides; inner basal portion forms knob surmounted by tuft of short, stout spines. Phallic apparatus consisting of phallotheca and endotheca. Former long, evenly shaped tube positioned very high under segment X, without any sclerotized connection with inferior appendages, and slightly swollen at tip. Endotheca small and spine-free. & genitalia (Figs. 769, 770): segment IX small and separated from sternite VIII by large membranous area. Posterior surface exhibits very little sclerotization and forms 2 large plates. Segment X reduced, in shape of horizontal plate equipped with 2 firmly fused appendages. In all likelihood, there is no external anal opening. Vaginal opening very small and situated between membranous folds. Vaginal apparatus simple and located at end of wide vestibule reinforced by 2 sclerotized ribs. The genus Helicopsyche is fairly large and virtually cosmopolitan with tropical affinities. It does not occur in most of the cold regions of the Northern Hemisphere. It is well known for the spiral-shaped cases of the larvae, which give the insects the appearance of tiny gastropods. Eight species live in the Nearctic region and of these, borealis Hagen is the most common. This species is of very widespread distribution, extending from Mexico to Nova Scotia, Alberta, and the Northwest Territories. It occasionally inhabits lakes and is found in all types of running water, exhibiting a preference for clear, fast-flowing streams.

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Family Beraeidae Wallengren Beraeidae Wallengren, 1891, p. 111 Type genus: Beraea Stephens The Beraeidae constitute a small family of Holarctic distribution. It contains only a single Nearctic genus. Genus Beraea Stephens

Beraea Stephens, 1836, p. 158 Type species designated by Fischer, 1970: Thya maurus Curtis Head short and wide. Eyes small with large ommatidia. Vertex strongly raised in middle (Fig. 771), with 3 pairs of warts. Ocelli absent. Maxillary palpi similar in both sexes, with each of 5 successive segments slightly longer than preceding one. First antennal segment long but not very convex. Spurs: 2,2,4. Legs with few sparse black spines. Abdominal hemogill system apparently absent. Internal gland of abdominal sternite V absent in % and tiny and not externally protuberant in &. Sternite VII with cone-shaped median point in both sexes. Wings strongly pubescent, in smooth ellipses. Both pairs are equally wide and similar in both sexes. Venation (Figs. 772, 773) greatly reduced and slightly different in both sexes. No crossveins and consequently no closed cells. In front wings, forks IV and V present in % and II, IV, and V in &. Discoidal cell absent. RS forked with 2 branches in % and 3 branches in &. One anal vein in %, but 2 in &. In hind wings, Sc, discoidal cell, and all apical forks absent. RS and M forked at base of wing in % and near middle of wing in &. Cu1 simple. Only 1 anal vein. In %, base of RS in front wings and of M in hind wings thickened and seta-bearing although not in same manner in front and hind wings. % genitalia (Figs. 774–776): segment IX very stout, especially laterally. Segment X indistinct from segment IX dorsally, deeply indented, and reduced to 2 semimembranous vertical lobes. Preanal appendages free, small, and ovoid. Intermediate appendages in shape of 2 long, slender spines parallel to segment X. Inferior appendages single-segmented, but complicated in shape, forming number of lobes and branches that are conspicuous from caudal view and joined together by inner basal plate. Phallic apparatus short, thick, and consists only of phallotheca and endotheca; 179

the latter not readily erectile, comprises number of pleated membranous lobes, and bears 2 long, slender spines. & genitalia (Figs. 777, 778): segment IX small, barely distinct from segment X, separated ventrally from sternite VIII, which is convex and strongly pubescent, by large membranous space and forms 2 wide parts with blunt bifid ends. Segment X considerably reduced, in horizontal plate, with 2 barely distinct appendages arising on lateral sides. Vulvar scale membranous, simple, and slightly depressed in middle. Vaginal apparatus without vestibule, very large, fairly simple, and constricted laterally at midlength. The genus Beraea is European and Nearctic in distribution. On our continent there are three species: nigritta Banks, gorteba Ross, and fontana Wiggins; the third is probably a synonym of the second. They are confined to the eastern part of the continent and are all very uncommon. They have rarely been captured. Only one, fontana, has been reported in southern Ontario, where it lives near springs in a peat bog.

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Family Leptoceridae Leach Leptoceridae Leach, 1915, p. 136 Type genus: Leptocerus Leach Elongate, slender, and slight insects. %% larger than && and often with wider hind wings. Head short and wide (Fig. 779) with eyes occasionally globular and very large, particularly in %. Ocelli absent. Vertex strongly convex, generally with 3 pairs of warts, and concave behind antennae to allow latter to flatten backward against head and body. Antennae very fine, over twice length of front wings in % and 1.5 times as long in &. First antennal segment convex, slightly shorter than head. Subsequent segments 2 to 3 times longer than thick. Maxillary palpi 5-segmented in both sexes, very long, and densely covered with erect hairs. Second segment always very long and last segment fairly short and partially unsclerotized (Fig. 781). Pronotum very short, concave in middle, and partly covered by mesonotum. Legs clothed with a very dense, flattened, scaly pilosity, and bearing weakly developed black spines. Claws very small. Front claws of % occasionally asymmetrical (Fig. 780). Spurs: 0,2,2 or 1,2,2 or 2,2,2. Abdominal hemogill system generally well developed (Figs. 782, 826). Gland of sternite V absent. Wings long, narrow, and densely clothed with appressed woolly hairs. Front wings with elliptical tips and partially sclerotized anal angle. Hind wings as wide as front wings or wider, occasionally exhibiting slight sexual dimorphism. Venation fairly homogeneous in series of genera and fairly considerably simplified. FI and FV alone present on both pairs of wings, with one exception. In front wings, crossveins Sc-R1 always present and R1–R2+3 occasionally present. Discoidal cell always closed and very elongate. Median cell open or absent and thyridial cell closed and very variable in length. RS forks twice, producing R2, R,3 and R4+5, whereas M forks once to yield M1+2 and M3+4. Basal trunk of M absent in Triaenodes. Cu2 meets wing margin and is joined to Cu1b by crossvein. A1+2+3 ends at Cu2. Second anal cell very long. In hind wings, discoidal cell always open, and RS and M as in front wings. Their bases are occasionally evanescent. Three or 4 anal veins. % genitalia fairly varied within family, with slight polyphyletic tendency to asymmetry. Segment IX well developed all around and occasionally slightly reduced dorsally. Apicolateral edge above inferior appendages generally forms inwardly folded strip connected to winglike structures of 181

phallotheca. Framework of segment X difficult to interpret with any degree of certainty. Two large, ear- or cone-shaped preanal appendages free or integrated with segment X. They are strongly pubescent, and there is no question as to their identity. Between these appendages are 2 spurs or 2 paired branches or 1 unpaired roof-like lobe. These parts are probably the intermediate appendages, which are either free or secondarily fused together, rather than the body of segment X. In Ylodes and Triaenodes, segment X forms a single piece and coexists with intermediate appendages paired or fused into single branch. Inferior appendages rather small, joined at base by inner basal plate, generally single-segmented, complex in form, and occasionally accompanied by upper basal branch. Phallic apparatus highly variable in development, either primitive and complete in structure, with phallotheca, endotheca, aedeagus, and parameres, or else reduced to phallotheca and spine-bearing endotheca. It may also be obscurely structured. It is situated fairly high above inferior appendages, with sclerotized connections with lateral sides of segment IX or with base of inferior appendages and even occasionally with both. & genitalia: segment IX continuous all around, but short dorsally and laterally. Laterally, developed into 2 oval, valve-like parts that allow insect to keep egg mass for period of time before depositing. Ventrally, segment IX elongate, occasionally continuous with sternite VIII, and forms flat or concave and occasionally carinate surface consisting of several plates designed to hold egg mass. Segment X small, in shape of short truncated tube with anal opening at the extremity (except in Ceraclea), and accompanied by 2 lateral appendages free or secondarily integrated with segment. Vulvar scale variable in size (absent in Nectopsyche). Supragenital plate absent. Vaginal opening gaping, occasionally forming large vaginal chamber proportional in size and complexity to development of phallic apparatus. Vaginal apparatus simple or complex. The Leptoceridae constitute one of the largest trichopteran families. It is cosmopolitan, but is particularly well represented in tropical regions, although a fair number of lineages occur in the Holarctic area. The larvae inhabit primarily lakes and large rivers. Adults are occasionally very abundant and at times fly in swarms late in the afternoon or at dusk and have a very wide geographical range. The family is divided into two subfamilies: the Triplectidinae, which occur mainly in the Southern Hemisphere, and the Leptocerinae, which are very widely distributed in the Northern Hemisphere. Only the latter subfamily is represented in Canada. 1a Thyridial cell of front wings and FV of hind wings absent (Fig. 801) . . . . . 2 1b Thyridial cell of front wings and FV of hind wings present (Fig. 783) . . . . 3 2a Inferior appendages of % forming wide, indented ovals (Fig. 807). Appendages of & free (Figs. 809, 810) . . . . . . . . . . . . . . . . . . . . . . . . Ylodes, p. 187

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2b Inferior appendages of % slender and 2-branched (Fig. 802). Appendages of & fused with segment X (Fig. 805, 806). . . . . . . . . . . . Triaenodes, p. 186 3a Discoidal cell of front wings very long and thyridial cell short. Former begins basally of latter and ends distally after it (Fig. 783) . . . . Ceraclea, p. 183 3b Thyridial cell of front wings very long and begins well basally of discoidal cell (Fig. 793) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a M1+2 of front wings a straight extension of M (Fig. 820) . . Oecetis, p. 190 4b M1+2 of front wings not a straight extension of M (Fig. 811) . . . . . . . . . . . 5 5a RS and M of hind wings evanescent (Figs. 827, 828). Very fragile, white or very lightly colored and delicately shaded species. . Nectopsyche, p. 191 5b All apical cells of front wings sessile (Fig. 793). Species generally bluish black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mystacides, p. 184 5c Base of RS of hind wings evanescent. In front wings, M1+2 and M3+4 fork after anastomosis (Fig. 811) . . . . . . . . . . . . . . . . . . . . . . . . . Setodes, p. 188 5d Base of RS on the hind wings normal. In front wings, M1+2 and M3+4 fork at level of anastomosis (Fig. 816) . . . . . . . . . . . . . . . . . Leptocerus, p. 189

Genus Ceraclea Stephens

Ceraclea Stephens, 1829, p. 28 Monobasic type species: Phryganea nervosa Fourcroy = Ceraclea nigronervosa Retzius Revision: Morse, 1975 In %, eyes variable in size, occasionally huge and sometimes contiguous on vertex. Spurs: 2,2,2. Front claws of % asymmetrical (Fig. 780). Abdominal hemogill system highly developed and composed of arborescent branches not only on pleurites but also between sternites (Fig. 782). Front wings with bluntly rounded tips. Hind wings distinctly wider than front wings in % and barely wider in &. Venation displaying slight sexual dimorphism (Figs. 783, 784). In front wings, forks I and V present in % and I, III, and V in &, with M 2-branched in % and 3-branched in &. FI and M fork petiolate. Discoidal cell very long and, proximally and distally, extending beyond short thyridial cell. In hind wings, base of RS not evanescent. Four anal veins. % genitalia (Figs. 785–789): segment IX fairly evenly short all around. Preanal appendages large, free, ear-shaped, horizontal, and fused to each other at base for variable distance. Segment X virtually absent. Intermediate appendages fused into unpaired, horizontal roof-like plate with entire or cleft extremity and occasionally accompanied by 2 lateral spurs. Inferior appendages 2-segmented, with 1st article long, slender, upward-slanting, and simple or complex in shape. Lower basal angle may be 183

stretched to variable degree, while inner side of article displays level of complexity characteristic of species. Inner basal portion stretched into protuberance that acts as support for phallic apparatus. Second article small, fairly strongly sclerotized, spur-shaped, and always inserted below apical lobe of 1st article and before tip of latter. Phallic apparatus small, sometimes simplified, sometimes complete, i.e., with phallotheca, endotheca, aedeagus, and parameres and with inner sclerotized connection with middle of lateral sides of segment IX, but not with base of inferior appendages. & genitalia (Figs. 790–792): segment IX very elongate ventrally but not continuous with sternite VIII. Ventral side flat and may be entire or cleft by simple, longitudinal or else V-shaped furrow, isolating 2 or 3 plates. Valves medium-sized or small. Appendages triangular, small, and free. Segment X also small, cone-shaped, but closed at extremity, with result that there is no anal opening. Vulvar scale simple, 2- or 3-lobed. Vaginal apparatus simple, flat, with or without a dorsal membranous vaginal chamber. The genus Ceraclea was for a long time confused with the genus Athripsodes. Morse (1975) isolated it from the latter and divided it into three subgenera: Pseudoleptocerus Ulmer (African) and Athripsodina Kimmins and Ceraclea Stephens. The latter two occur in the Holarctic and Oriental regions and are represented in our area. For the purposes of this text, we consider these two subgenera as species groups, since only a small number of genital characters in the % distinguish them. In Ceraclea s. str., the phallotheca is very short and forms an apicoventral plate. There is a pair of parameres developed into short branches and a small, but clearly distinct aedeagus (Fig. 789). In Athripsodina, the phallotheca is longer, less sclerotized, without a ventral plate, and cleft vertically at the tip into 2 paired lobes. The aedeagus is reduced, barely distinct from the endotheca that has lost the parameres, but through the phenomenon of vicarianism, it bears 1, 2, or 3 slender spines that are probably secondary endothecal developments (Fig. 788). Ceraclea contains 36 Nearctic species, 6 of which are found in Canada. Virtually all of them have a widespread distribution. The Canadian species inhabit all types of lotic and lentic biotopes, showing preference for large lakes and rivers.

Genus Mystacides Berthold

Mystacides Berthold, 1827, p. 437 Monobasic type species: Phryganea nigra Linnaeus Revision: Yamamoto and Wiggins, 1964 184

Eyes of % very large, globular, reddish, and widely separated. Top of head and thorax smooth and shiny. Spurs: 0,2,2. Abdominal hemogill system well developed. Front wings very bluntly rounded at extremity, with strong venation, and barely narrower than hind wings. In repose, extremities frequently bent toward body along line beginning at notch on costal margin and extending across anastomosis to arculus. Venation (Fig. 793): in front wings, apical cells particularly short and all sessile. Strongly slanting anastomosis. Thyridial cell particularly long and beginning close to base of wing. In hind wings, costal margin forming distinct angle before tip. Base of RS evanescent. Four anal veins. % genitalia (Figs. 794–797): segment IX very short dorsally, but particularly elongate apico-ventrally where it forms a more or less extensively bifid protuberance acting as lock notch for inferior appendages. Segment X virtually absent and merged with the base of the intermediate appendages. Preanal appendages in shape of long, horizontal, slender, and free rods. Intermediate appendages forming 2 or 3 long, strongly sclerotized, symmetrical or asymmetrical spines. Inferior appendages single-jointed, small, oval-shaped in side view, equipped with sharp, lower, subbasal heel-like protuberance and rounded, inner, subapical lobe or carina. Phallic apparatus medium-sized, with inner sclerotized connection with base of inferior appendages, but not with margin of segment IX. It comprises downward-curving phallotheca, occasionally equipped with upper median branch, and faintly developed and spine-free endotheca. & genitalia (Figs. 798, 799): segment IX stout and robust, not extended anteroventrally, but continuous with sternite VIII. Huge valves, developed into large callipers. Segment X in bluntly truncated tube. Appendages similar in shape to those of %, forming long, free, horizontal rods. Vulvar scale strong and bilobed. Base of valves hollowed out into deep, sclerotized vaginal chamber above fairly complex vaginal apparatus. Whole unit appears very complicated in ventral view. The genus Mystacides is small and of Holarctic and Oriental distribution. There are three species in our area. In interjecta Banks (transcontinental, from Maine to Alaska), the front wings are brown, the intermediate appendages are symmetrical, and the apicoventral protuberance of segment IX slightly bifid. In alafimbriata Hill-Griffin (western, from Mexico to Alaska), the front wings are black with a bluish metallic sheen, the intermediate appendages are asymmetrical, and the apicoventral protuberance of segment IX has a deep V-shaped notch. In sepulchralis Walker (transcontinental, from Newfoundland to Alaska), the wings have the same bluish-black coloring, and one of the intermediate appendages is divided, with the result that there are 3 asymmetrical spines on segment X. The apicoventral protuberance of segment IX has a deep U-shaped notch. 185

These last two species display reverse symmetry, a phenomenon that is unique among the Trichoptera: the 2 or 3 spines making up the intermediate appendages are twisted either to the left or to the right in equal proportions within any given population (Figs. 796, 797). Unlike many other caddisflies, adults of Mystacides are active only during the day. Early in the morning or late in the afternoon, when the sun’s rays are oblique and the air is still, they can very often be observed in large numbers skimming above the water’s surface along the shores of lakes and ponds. These flights consist almost exclusively of %%. The insects become inactive after nightfall and are not attracted by lights. Genus Triaenodes McLachlan

Triaenodes McLachlan, 1865, p. 110 Type species designated by Ross, 1944: Leptocerus bicolor Curtis Triaenodella Mosely, 1932, p. 308 Type species by original designation: Triaenodella chelifera Mosely Triaena McLachlan, 1865, p. 34 Type species designated by Fischer, 1965: Leptocerus bicolor Curtis First antennal segment especially long. In %, inner dorsal portion bears a large, oval, movable scale protecting 2 stout brushes of hairs that may fan out (Fig. 800). Legs slender. Spurs: 1,2,2. Front claws in % very small and slightly asymmetrical. Abdominal hemogill system well developed. In &, 4th and 5th abdominal pleurites with area of sclerotized black striations. Front wings fairly broad, with elliptical tips, not any narrower than hind wings and clothed with fine pilosity of dark gold and pale yellow hairs creating simple design. Venation (Fig. 801): in front wings, crossvein R1–R2+3 absent. Discoidal cell large and wide. FI petiolate. M stem completely absent, with the result that there is no thyridial cell. M1+2 and M3+4 are therefore stemless, and their bases constitute part of the anastomosis, which is basically straight. In hind wings, M forks virtually at same point as RS. Cu1 not forked, and FV absent. Three anal veins. % genitalia (Figs. 802–804): segment IX well developed all around, particularly ventrally. Segment X small, inconspicuous, membranous, often asymmetrical, and occasionally tiny. Preanal appendages free, developed into slender, horizontal rods. Intermediate appendages forming free, paired spines (injustus group) or fused into faintly sclerotized median lobe (marginatus group) or even lost (aba). Inferior appendages small, probably 2-segmented, horizontal, and complex in shape and with sharp apex, which is stretched to some extent. On upper surface, a large spine-bearing knob, which may be 2nd segment completely fused with 1st. Base of inferior 186

appendages forms a very long, slender, paired spine pointing forward, then backward and downward in broad curve, paralleling curve of phallic apparatus, and assuming role of parameres. Phallic apparatus very large and very stout, downward-curving, and comprising sclerotized phallotheca extended without discontinuity by large, spine-free, strongly erectile, and forked membranous endotheca. The base of the phallotheca is no longer connected directly to the inferior appendages, but the winglike structures joining it to the apical margins of segment IX are developed into 2 bands that are connected to both segment IX and the base of the inferior appendages. These bands are strongly developed, wedge-shaped, semi-external, and border the apical margins of segment IX with a 2nd ridge (“b” on Fig. 802). & genitalia (Figs. 805, 806): segment IX stout, not continuous ventrally with sternite VIII but has 2 furrows. Valves large and subcircular. Segment X forming short, stout, truncated tube with completely laterally integrated appendages. Spacious vaginal chamber with sclerotized walls and very complex relief at base of valves. Vulvar scale large and thick, with complicated relief and shape. The classification of the genus Triaenodes is unsatisfactory and needs to be revised on a worldwide scale. For the purposes of this text, I have accepted the validity of the genus Ylodes Milne, which most authors confuse with the genus Triaenodes. The latter, as defined here, actually corresponds to Triaenodella Mosely, but I have not used this name since I am still not sure of its status or validity. The genus Triaenodes is large and ubiquitous, with tropical affinities. It contains 23 Nearctic species, 12 of which are found in Canada. They live in lotic and lentic biotopes and produce swimming larvae that remain in submerged aquatic vegetation. Genus Ylodes Milne

Ylodes Milne, 1934, p. 11 Type species by original designation: Triaenodes grisea Banks This genus is very closely related to Triaenodes and could be a subgenus of the latter, particularly with respect to the venation that is identical in both genera. The distinguishing features are: stouter legs, with front claws of % exhibiting greater asymmetry, 1st antennal segment of % without any neoformations and front wings uniformly grey-brown instead of gold with yellow markings, and abdominal pleurites of & without dark striations. % genitalia (Figs. 807, 808): segment IX clearly divided into lateral and ventral parts. Segment X well developed, forming an elongated oval, symmetrical roof above phallic apparatus. Preanal appendages free and 187

also in elongated oval shape. Intermediate appendages fused into fairly short unpaired lobe. Inferior appendages forming large ovals concave on inner surface and notched at lower apical margin. Basal branch small, not parallel to phallic apparatus, concealed in concavity created by appendages, and accompanied by knob near base. Phallic apparatus similar to that of Triaenodes, but less stout, membranous and erectile at base, and accompanied by unpaired branch arising from phallotheca, very long and slender and situated dorsally. It probably plays the role of the paramere, which in Triaenodes has been taken over by the basal branches of the inferior appendages. Apparatus without sclerotized connection to either inferior appendages or margin of segment IX. & genitalia (Figs. 809, 810) fairly similar to those of Triaenodes, but segment IX less stout laterally, concave ventrally, and continuous with sternite VIII. Valves smaller. Segment X larger, with free, triangular appendages. Vulvar scale simple and 2-lobed. No complex vaginal chamber at inner base of valves. Long, simple vaginal apparatus. The genus Ylodes is medium-sized, of Holarctic distribution, and is particularly well represented in Central Asia. It contains only four Nearctic species that are represented in our area: reuteri McLachlan (Alberta, British Columbia), in which the inner branches of the inferior appendages are short and blunt, frontalis Banks (Manitoba, Yukon Territory), in which they are longer and more slender, kaszabi Schmid (Alaska, Yukon Territory), and schmidi Manuel and Nimmo (Yukon Territory), in which they are thick. Larvae live in ponds. Genus Setodes Rambur

Setodes Rambur, 1842, p. 515 Type species designated by Milne, 1934: Setodes punctella Rambur = Phryganea viridis Fourcroy Revision: Holzenthal, 1982 Spurs: 0,2,2. Front claws of % thin and symmetrical. Abdominal hemogill system poorly developed. Wings long, narrow, and fairly pointed, both pairs equally wide, but hind wings with very long fringes. Venation (Fig. 811): in front wings, discoidal cell short. FI and M fork petiolate. In hind wings, costal margin forming a fairly distinct angle, RS base evanescent, FV short and 3 anal veins. % genitalia (Figs. 812, 813): segment IX short dorsally and long ventrally. Segment X forming a horizontal, bifid roof and ending in 2 sharp, more or less elongate and occasionally asymmetrical points. Preanal appendages long and free or completely integrated with segment X. Intermediate appendages absent. Inferior appendages single-segmented, fairly large, bifid or trifid. Phallic apparatus complete; phallotheca and endotheca 188

reduced to very short basal parts. Aedeagus robust, very large, and broadly bent downward; spermatic duct opens dorsally well before apex of organ. Parameres very long and thin. A sclerotized connection between phallotheca, lateral margin of segment IX, and inferior appendages. & genitalia massive, very complex, and highly sclerotized (Figs. 814, 815). Segment IX stout all around, forming large, raised ventral plate not continuous with sternite VIII. Valves large and spatulate. Segment X consisting of large, oval, horizontal, and prominent roof with tiny subanal plate. Appendages completely integrated with base of segment X. Vulvar scale bulky, bilobed, concave on upper surface, and completely contained within space between valves. Spacious vaginal chamber with semimembranous walls occurs at base of valves. Vaginal apparatus narrow and very long. The genus Setodes is large and found on every continent, except in Neotropical America. It is particularly richly represented in the tropics where many species have extraordinary genitalia. There are only nine Nearctic species, only three of which have been reported in this country and are confined to Quebec and Ontario. In oligius Ross and guttatus Banks, the preanal appendages are fused with segment X, and the inferior appendages are short. In incertus Walker, the preanal appendages are free, and the inferior appendages long and slender. Adults are very delicate, with front wings the color of burnt gold with silver speckles. Larvae live in lakes and gently flowing water. Genus Leptocerus Leach

Leptocerus Leach, 1915, p. 136 Monobasic type species: Phryganea interrupta Fabricius Ymymia Milne, 1934, p. 16 Monobasic type species: Setodes americanus Banks Spurs: 0,2,2. Front claws of % tiny and symmetrical. Wings very narrow, with hind wings slightly narrower than front wings. Both pairs have pointed tips. Venation (Fig. 816): in front wings, discoidal cell long and M fork sessile. In hind wings, RS not evanescent at base, M fork also sessile, and 3 anal veins. % genitalia (Fig. 817): segment IX fairly uniformly elongate all around. Segment X reduced and completely integrated with dorsal portion of segment IX, but extended by 2 sharp branches. Preanal appendages completely fused with bulk of segments IX and X. Intermediate appendages absent. Inferior appendages single-segmented, very large and wide, horizontal, concave on inner surface, with slight thickening provided with stout spines at base of this surface. Phallic apparatus stout, horizontal, and not deeply invaginated within abdomen. Phallotheca and endotheca reduced to very short basal parts. Aedeagus robust and of complex shape. Parameres 189

absent. Long vertical band provides sclerotized connection between phallotheca and base of lateral edge of segment IX, above base of inferior appendages. & genitalia (Figs. 818, 819): segment IX with a pebbled area containing irregular, shallow depressions in the middle of the lateral sides. Ventrally, it is not continuous with sternite VIII. Valves in the shape of large, elongate ovals. Segment X and its appendages completely integrated with the dorsal portion of segment IX, forming a bulky, horizontal, roof-like structure. Membranous subanal plate present. Vulvar scale membranous and with an elongate oval shape. Vaginal chamber absent and vaginal apparatus wide and fairly complex. The genus Leptocerus is large and cosmopolitan in distribution, although not found in South America. It is especially well represented in the Old World tropics where some species have the most extraordinary genitalia to be found among the Trichoptera. There is only a single Nearctic species: americanus Banks, which is widely distributed in central and eastern Canada. The larvae are swimmers and occur in the submerged aquatic vegetation growing in still water. Genus Oecetis McLachlan

Oecetis McLachlan, 1877, p. 329 Type species designated by Ross, 1944: Leptocerus ochraceus Curtis Oecetina Banks, 1899, p. 215 Type species by original designation: Oecetis incerta Provancher nec Walker = Leptocerus inconspicuus Walker Oecetodes Ulmer, 1907, p. 144 Type species designated by Milne, 1934: Setodes avarus Banks Friga Milne, 1934, p. 16 Type species by original designation: Setodes immobilis Hagen Yrula Milne, 1934, p. 17 Type species by original designation: Oecetina fumosa Banks = Setodes cinerascens Hagen Setodina Banks, 1907, p. 130 Type species by original designation: Setodina parva Banks Quaria Milne, 1934, p. 17 Monobasic type species: Oecetis scala Milne Palpi especially long, Front claws of % symmetrical. Spurs: 1,2,2. Wings long and narrow, with front wings fairly pointed, and hind wings slightly wider than front wings. Venation (Fig. 820): in front wings, 190

discoidal cell long. M1+2 generally in direct line with M and, as a result, there seems to be only a single median vein, but M3+4 is present and seems to arise from Cu1a. Anastomosis fairly variable in design, forming straight or slanting line, or in tiered or zigzag pattern. In hind wings, FI tiny, bases of RS and M not evanescent. Three anal veins. % genitalia simple and only slightly prominent (Figs. 821–823): segment IX short all around, with posterior lateral angle occasionally stretched. Preanal appendages developed into slightly prominent, rounded, ear-shaped structures. Segment X forming simple oval roof, or else, possibly lost and replaced with intermediate appendages that are fused together to form this oval part. Inferior appendages single-segmented, rather small, forming simple callipers, with lower subbasal angle and occasionally 2nd, also subbasal but upper angle. Phallic apparatus extremely variable in development. It comprises phallotheca, forming protruding lower apical angle, and more or less developed endotheca bearing from 6 spines (persimilis) to a single, asymmetrically twisted spine (inconspicua). Aedeagus and parameres lost. & genitalia barely prominent (Figs. 824, 825). Segment IX short all around and ventrally continuous with sternite VIII, forming slightly concave surface, occasionally reinforced with sclerotized strips, to hold egg mass temporarily. Medium-sized and distinctly concave valves. Segment X in short truncated tube. Appendages developed into faintly protruding lobes. Vulvar scale small and membranous. Internal vaginal chamber missing, except in species with strongly developed endotheca in % (persimilis). Vaginal apparatus small and simple. The genus Oecetis is large, cosmopolitan, and well represented on every continent. It contains 20 Nearctic species, a dozen of which live in Canada. They all have a very widespread distribution and inhabit mainly lakes and the major rivers where they occasionally are very abundant locally. A certain number of genera, including some of the ones mentioned above, were originally part of the genus Oecetis but were established as separate genera on the basis of details relating to venation and the shape of the inferior appendages of the %. Later on, they have all been reassembled under the same generic heading. I have adopted this approach here but am not sure that it is justified in every instance. It may be too radical a solution, and some of the genera may constitute natural generic or subgeneric lines. Only a revision of the genus on a worldwide scale will allow the problem to be solved. Genus Nectopsyche Müller

Nectopsyche Müller, 1879, p. 40 Monobasic type species: Setodes gemma Müller 191

Leptocella Banks, 1899, p. 214 Type species by original designation: Mystacides uwarowii Kolenati = Leptocerus albidus Walker & smaller than %. Top of head and thorax and base of antennae covered with dense, scaly pilosity. Eyes larger in % than in &. First antennal segment considerably thickened. Labial palpi strongly reduced in both sexes. Spurs: 0,2,2. Front claws of % fairly poorly developed and symmetrical. Middle femora and posterior femora and tibiae with fringe of long, fine hairs in % only. Abdominal hemogill system well developed (Fig. 826). Front wings elliptical at apex in both sexes. Hind wings of % in wide triangles, with anal area strongly developed. In &, hind wings barely wider than front wings. Venation similar in both sexes in spite of dimorphism in wing size and shape (Figs. 827, 828). In front wings, crossvein R1–R2+3 present. Discoidal cell wide and not very long, thyridial cell also not very elongate, and M fork petiolate. In hind wings, RS and M 2-branched. RS and M almost completely evanescent over virtually their entire length, with extremely thin extremities. Four anal veins. % genitalia considerably modified (Fig. 829). Segment IX short and completely interrupted ventrally. Segment X virtually absent and unusual in the disposition of its armature. Preanal appendages free, forming long, very prominent, horizontal rods. Intermediate appendages very low, between inferior appendages, forming deeply bifid plate and creating large concave space between themselves and preanal appendages. Inferior appendages situated below lower lateral angles of segment IX; single-segmented and upward-slanting and of complex structure, denticulate margins, and constantly strongly sclerotized inner apical lobe; fused together ventrally and forming 2 paired and slender appendages or 1 large unpaired plate. Two branches curved upward and backward, possibly assuming role of parameres, arise from base of inner portion of inferior appendages. Phallic apparatus large and positioned very low, between inferior appendages. Short phallotheca and large, complex endotheca; latter strongly erectile, spine-free, and equipped with large sclerotized ventral plate, which provides protection when structure is retracted; it ends in cylinder with small apical phallotremal sclerite. Aedeagus and parameres lost. & genitalia greatly simplified (Figs. 830, 831). Segments IX and X firmly fused into large part that appears triangular from side; reduced to narrow band ventrally, with break in middle. Appendages free, long, and slender. Valves in form of simple lobes, under another slender point. Vulvar scale and vaginal chamber absent. Anal and vaginal openings in concave, membranous space between appendages and valves. A long cylindrical, membranous vaginal vestibule ends in simple vaginal apparatus. 192

The genus Nectopsyche is fairly large and among the most specialized in the family. It is exclusively New World in distribution, but found primarily in Neotropical America. Half a dozen species have come as far north as Canada. Most are distributed in the central, eastern, and northeastern parts of the continent. They have a very wide ecological tolerance and occur in lakes, large and small rivers, and even marshes. Nectopsyche are the most exotic looking of the various Canadian Leptoceridae. They can be recognized by their slim structure and white or gold coloring, which gives them a delicate, exquisite, and almost immaterial appearance.

193

Family Odontoceridae Wallengren Odontoceridae Wallengren, 1891, p. 12 Type genus: Odontocerum Leach Antennae distinctly longer than front wings, with 1st segment strongly thickened. Ocelli absent. Maxillary palpi 5-segmented in both sexes. Legs with weakly developed black spines. Middle tibiae as long as femora, with preapical spurs before middle of the tibia length. Spurs: 2,4,4. Gland of abdominal sternite V and hemogill system absent. Wings displaying fairly pronounced sexual dimorphism in shape and venation. In front wings, discoidal cell present and closed, and median cell open or absent in both sexes. Thyridial cell absent in % since M is missing, but present in &. Forks I, II, V or I, V present in % and I, II, III, V in &. Cu2 ending at Cu1a or occasionally absent in %. Behind A, an additional postanal vein (PA). In hind wings, discoidal cell closed. Forks I or I, II, V present in % and I, II, III, V or I, II, V in &. Three or 4 anal veins. % genitalia: segment IX well developed. Segment X more or less distinct from segment IX, a simple or complex roof-like structure with bifid extremity. Preanal appendages free, large, and elongate. Intermediate appendages present or absent. Inferior appendages consisting of 2 simple segments. Phallic apparatus situated fairly high in abdomen, without any sclerotized connections with segment X or inferior appendages. It comprises tubular phallotheca, membranous, spine-free endotheca, and short, partially unsclerotized aedeagus containing large phallotremal sclerite. Parameres absent. & genitalia: segment IX whole and fairly stout; ventrally, it consists of 3 flat or slightly concave plates framing vaginal opening and designed to hold egg mass temporarily. Segment X very small and reduced to thin border. Appendages large, ovoid, and faintly prominent. Supragenital plate and vulvar scale absent. Anal and vaginal openings situated in membranous area. Vaginal apparatus fairly simple, at end of long, more or less sclerotized vestibule. The Odontoceridae constitute a small, virtually cosmopolitan family with tropical affinities. It is heterogenous and still poorly defined. The six genera found in the Nearctic region have been classified into two subfamilies, the Odontocerinae and the Pseudogoerinae. The two Canadian genera, 194

Psilotreta Banks and Marilia Müller belong to the former. They differ greatly from each other and represent very unequal stages of specialization. 1a Discoidal cell on both pairs of wings long and narrow, joined by FI for long distance (Fig. 834). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psilotreta, p. 195 1b Discoidal cell wider and joined by FI for short distance only (Fig. 845). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marilia, p. 196

Genus Psilotreta Banks

Psilotreta Banks, 1899, p. 213 Monobasic type species: Psilotreta frontalis Banks Astoplectron Banks, 1914, p. 264 Type species by original designation: Heteroplectron borealis Provancher = Psilotreta indecisa Walker Revision: Parker and Wiggins, 1987 Head very short and wide. Eyes slightly larger in % than in &. Vertex with 3 pairs of warts (Fig. 832). Face flat or slightly concave. Antennae slightly longer than front wings and fairly thick. Maxillary palpi covered with strong pilosity of erect hairs, with first 2 segments short and last 2 distinctly thinner (Fig. 833). Legs stout. Spurs: 2,4,4. Wings covered with a dense pilosity. Two pairs subequal in width, but slightly wider in & than in %. Venation with distinct sexual dimorphism (Figs. 834, 835). In both sexes, discoidal cell on both pairs of wings long, tapered, situated in middle of wing, and joined for long distance by FI. Crossveins Sc-R1 and R1–R2+3 present. FII with long petiole. In front wings in both sexes, R1 meets wing margin near R2, Cu2 present and ends at Cu1b. One of anal veins missing, with the result that there is only 1 anal cell. Postanal vein very long. In front wings of %, only FI, FII, and FV present. M without a base, with the result that there is no thyridial cell, represented by single apical branch, separating from R4+5. FV especially wide. In front wings of &, forks I, II, III, and V present. M 3-branched, forming long, narrow thyridial cell. FV wide. In hind wings of %, only forks I, II, and V present and M 2-branched. Three anal veins. In hind wings of &, forks I, II, III, and V present and M 3-branched. Four anal veins. % genitalia (Figs. 836–838): segment IX stout and distinctly extended anteriorly in middle of lateral sides. Dorsally, a long lobe overhanging segment X and framed by 2 hairy lateral thickenings. Segment X complex, truncated, with anal opening at extremity, and ending in 2 winglike structures or 2 points. Preanal appendages free and forming large, elongate earlike lobes. Intermediate appendages present, inserted on sides of 195

segment X, in shape of curved or looped spurs. Inferior appendages 2-segmented, stout, subhorizontal, and simple in shape. Second article smaller than 1st, inserted at apex of latter, and provided with short, stout spines. Phallic apparatus large and very elongate. Aedeagus with very large phallotremal sclerite. & genitalia (Figs. 839–841): segment IX massive and open ventrally, forming 3 posterior plates in more or less rounded triangles. Appendages of segment X ovoid and barely prominent. Vaginal vestibule complex and sclerotized. The genus Psilotreta is small and of Oriental and Nearctic distribution. It contains six Nearctic species, only four of which are represented in our area: indecisa Walker, rufa Hagen, frontalis Banks, and labida Ross. They are closely related and can be distinguished primarily by the shape of segment X and the intermediate appendages. They are confined to the eastern part of the country and inhabit the calmer stretches of the running water. Psilotreta originated in the Oriental region, where the greatest number, and the most primitive species occur (Schmid 1989, p. 110). Genus Marilia Müller

Marilia Müller, 1880, p. 127 Type species designated by Mosely and Kimmins, 1953: Marilia major Müller Head, thorax, palpi, and legs clothed with a slightly scaly pilosity. Eyes very large, globular, reddish, and almost touching on vertex that is concave and without warts in % (Fig. 842). In &, head distinctly smaller, eyes also smaller, and vertex with 3 pairs of warts (Fig. 843). Antennae very fine, longer than front wings, with 1st segment convex. Maxillary palpi with all the segments long. Mesonotum strongly convex with scutellum very large and ogival. Front and hind legs reduced in size, but middle legs are very long, with tibia and femur of equal length and preapical spurs inserted at point corresponding to 1st third of tibia (Fig. 844). Spurs: 2,4,4. Wings exhibiting strong sexual dimorphism in their shape and venation (Figs. 845, 846). Front wings narrow and distinctly truncated at apex, but to greater degree in % than in &. Hind wings forming broad triangles, with very long brush of hairs on anal lobe in %. In &, hind wings barely wider than front wings and without brush of hairs. Venation: in front wings, R1 ends on R2 and pterostigma clearly marked in %. Forks I and V present in % and I, II, III, and V in &. In %, base of M completely absent, with the result that there is no thyridial cell. M 2-branched. In &, base of M present, but it coalesces with Cu1 at early stage, with the result that the 196

thyridial cell is present, but very narrow. M 3-branched. Cu2 absent in %, but present and very faint in &. Single anal vein. No basal cells. Postanal vein present. In hind wings, SC, RS, R2+3, and R2 close to C, and closer in % than in &. R2 very short and ending at an early stage at R1 in both sexes. FI only present in % and FI and FV in &. M single-branched in % and 2-branched in &. Three anal veins in both sexes. % genitalia very simple (Figs. 847–849). Segment IX evenly elongate all around, not forming lobe above segment X, continuous with latter, and without visible suture. Segment X forming a simple roof with long cleft at extremity. Preanal appendages forming long rods. Intermediate appendages lost. Inferior appendages comprising 2 subcylindrical articles, 2nd fairly reduced and bearing fine apical spinules. Phallic apparatus small, downward-curving, with aedeagus barely distinct from phallotheca and provided with small phallotremal sclerite. & genitalia (Figs. 850, 851): segment IX strongly indented in middle of lateral sides, and forming ventrally 2 large subrectangular plates framing vaginal opening. Appendages very large, triangular, barely separate from segment X at base, and slightly prominent. Segment X thin and without external anal opening. Vaginal apparatus simple. Vaginal vestibule simple and membranous. The genus Marilia is medium-sized and found primarily in the Oriental, Australian, and Neotropical regions. In North America, there are two species that inhabit the southern United States, one of which, flexuosa Ulmer, has been recorded in southern Ontario.

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Family Calamoceratidae Ulmer Calamoceratidae Ulmer, 1905, p. 106 Type genus: Calamoceras Brauer The Calamoceratidae are a small family that is found on every continent. It contains large, heavy-bodied, and very pubescent insects. There are only three Nearctic genera, one of which is represented in Canada. Genus Heteroplectron McLachlan

Heteroplectron McLachlan, 1871, p. 123 Monobasic type species: Heteroplectron californicum McLachlan Eyes fairly small. Vertex with 5 pairs of warts. Ocelli absent. Antennae slender, slightly longer than front wings in % and of equal length in &, and covered with a pilosity forming erect brush at apex of each segment, giving a crenulated appearance. First segment very short and vestigial. Second segment large and subglobular. Fourth and 5th segments fused together. Segments IV to X with median crescent-shaped cross stripe (Fig. 852). Maxillary palpi 5-segmented in both sexes, covered with dense, erect setae; first 2 segments long and last 2 fairly slender. Legs with faintly developed black spines. Spurs: 2,4,4. Hind tibiae and tarsi of % with posterior margin bearing very long fringe of fine hairs. Gland of abdominal sternite V absent. Hemogill system composed of 3 long, slender, simple tubes per segment (Fig. 859). Wings uniformly brown-black and densely clothed with fine hairs. Front wings forming long, obliquely stretched triangles; hind wings in semicircles and much shorter than front wings. Venation complete and without sexual dimorphism (Fig. 853), with forks I, II, III, IV, and V present in front wings and I, II, III, and V in hind wings. In front wings, crossvein Sc-R1 present. Discoidal cell halfway along length of wing, small and narrow. Median and thyridial cells large and closed. All forks sessile. In hind wings, crossvein Sc-R1 present and R1 ending at R2. Discoidal cell small and open. FII and FIII petiolate. M forking at early stage at level of RS. Cu1 with large fringe of long black hairs. There are 3 anal veins. % genitalia (Figs. 854–856): segment IX stout all around, with an area clothed with long setae on lateral sides. Segment X forming simple, 198

stretched, bifid roof-like structure covered with small spinules. Preanal appendages free and fairly large. Intermediate appendages absent. Inferior appendages 2-segmented, with 1st articles thick, divergent, and upwardslanting and densely covered with tubercles on inner surface. Second article forming very short cylinder also spinulose at tip. Phallic apparatus situated fairly high, under segment X, and without sclerotized connection with latter or inferior appendages. It consists of tubular phallotheca continuous with membranous, spine-free endotheca, and very short, partially unsclerotized aedeagus. Parameres absent. & genitalia (Figs. 857, 858): last tergites and sternites small, with correspondingly larger pleurites. Sternite VIII short, transverse, cleft in middle of posterior margin and ciliate. Segment IX small, stout, and folding onto ventral side of abdomen where it forms 2 large, partially striated plates creating concave surface for temporarily holding egg mass. Segment X vestigial, developed into simple trapezoidal plate with appendages, semi-oval in shape, lying against dorsal surface. Supragenital plate and vulvar scale absent. Vaginal apparatus simple, with long, partially sclerotized vestibule. The genus Heteroplectron is small, exclusively New World in distribution, and contains only two Canadian species: californicum McLachlan, found along the Pacific coast as far north as British Columbia, where it is common, and americanum Banks, which occurs in the eastern part of the continent as far north as Quebec, where it apparently is very uncommon. Larvae of these species inhabit cold, running water.

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Family Molannidae Wallengren Molannidae Wallengren, 1891, p. 116 Type genus: Molanna Curtis Head short and wide, with eyes hairy and slightly more globular in % than in &. Ocelli absent. Antennae thick, with 1st segment slightly convex. Maxillary palpi 5-segmented in both sexes, densely clothed with erect hairs; segments differing slightly in length. Spurs: 2,4,4. Legs with numerous short, black spines. Abdominal hemogill system well developed and comprising half dozen simple tubes per segment. Internal gland of sternite V present in & alone and very small (Fig. 862). Wings fairly narrow, both pairs equal in width, fairly bluntly rounded at tips, and densely clothed with fine hairs. Venation fairly complete, with or without sexual dimorphism, but with variable secondary rearrangement of vein forks, with the result that most of forks are no longer recognizable as such. In front wings, discoidal cell absent. RS 2- or 3-branched, M 3or 4-branched, depending on sex, and Cu1, single- or 2-branched. In hind wings, discoidal and median cells absent, RS 3-branched, M 2-branched, and Cu1 simple. Three anal veins. % genitalia: segment IX unevenly short all around. Segment X present or virtually absent. Preanal appendages large, free, and elongate. Intermediate appendages stout and curved downward. Inferior appendages single-segmented, forming callipers of more or less complex design, and joined at base by inner plate. Phallic apparatus immediately above inferior appendages and comprising more or less long phallotheca and variably developed endotheca poorly equipped with spines. Aedeagus and parameres absent. & genitalia: sternite VIII large and surrounding lower part of segment IX. Latter very short when viewed from side, but posteriorly, folding backward, forming flat, tripartite surface for temporarily holding egg mass. Segment X small, barely distinct from segment IX, and bearing 2 pubescent, poorly differentiated appendages. Anal and vaginal openings close to each other and located in more or less complex membranous area. Supragenital plate and vulvar scale absent. Vaginal apparatus rather simple. The Molannidae constitute a small family of Holarctic and Oriental distribution. The family contains only three genera, two of which exist in 200

Canada: Molanna Curtis and Molannodes McLachlan. They represent two related but unequal stages of specialization. The species live in lotic as well as lentic biotopes, but have a decided preference for lakes. 1a Front wings long and narrow. Venation different in both sexes. RS in front wings 2-branched and Cu1a and Cu1b distinct (Figs. 863, 864) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molanna, p. 201 1b Front wings wider and rounded. Venation similar in both sexes. RS in front wings 3-branched and Cu1a and Cu1b fused to each other (Fig. 871) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molannodes, p. 202

Genus Molanna Curtis

Molanna Curtis, 1834, p. 214 Monobasic type species: Molanna angustata Curtis Vertex with 4 pairs of warts (Fig. 860). Maxillary palpi with modified 2nd or 3rd segment in % only and specialized pilosity in both sexes (Fig. 861). Spurs: 2,4,4. Pronotum with pair of large, simple, transverse warts. Wings long and narrow, slightly narrower and larger in & than in %. Venation (Figs. 863, 864) slightly different in both sexes. In front wings RS 2-branched in both sexes. M 3-branched in % and 4-branched in &. Discoidal and median cells absent. Crossvein Cu1-Cu2 situated medially. Cu2 evanescent at extremity, but more strongly so in % than in &. Two short anal cells. A1+2+3 very long, extending to Cu1. In hind wings, venation considerably simplified in musetta Betten which has long row of modified hairs. In other species, RS 3-branched, M 2-branched, and Cu1 simple. % genitalia (Figs. 865–867): segment IX not reduced in height and not enclosed within sternite VIII. Segment X virtually absent and reduced to 2 sclerites lining genital cavity. Preanal appendages large, slightly downward-curving, and calliper-shaped. Intermediate appendages large, stout, contiguous, always distinctly downward-curving, and provided with short, thick setae. Inferior appendages in fairly slender callipers, at base with large inner knob equipped with spines. Phallotheca in a long slender tube and without sclerotized connection to base of inferior appendages. Endotheca small, simple, equipped with few processes, and ending in large phallotremal sclerite. & genitalia very wide (Figs. 868, 869). Segment X barely distinct from its appendages, which form simple hairy protuberances. Posterior portion of segment IX forming large, simple, tripartite plate. Anal and vaginal openings in narrow, simple, membranous area. Vaginal vestibule fairly long and slightly sclerotized. Vaginal apparatus small and simple. The genus Molanna is small, Holarctic and Oriental in distribution, represented in Canada by five closely related species that can be 201

distinguished primarily by the shape of the intermediate appendages. They live near lakes and occasionally in running water and have a very widespread geographical distribution. They are easy to recognize due to the fact that in repose the adults curve their wings around their bodies, and the longitudinal axis of the latter is at an angle to the substrate. This is undoubtedly a case of mimicry in which the insects look like short branch segments. In Greek, phryganion means dry twig and led to the French word Phrygane, synonym of caddisfly. Genus Molannodes McLachlan

Molannodes McLachlan, 1866, p. 178 Monobasic type species: Molannodes zelleri McLachlan = Phryganea tincta Zetterstedt Vertex with 3 pairs of warts, lateral median ones forming large lunule. Spurs: 2,4,4. Pronotum with 2 pairs of rounded warts. In %, median pair is transformed into 2 erectile, wing-shaped membranous organs bearing long fringe of golden setae (Fig. 870). Wings wider than in Molanna. Venation (Fig. 871) similar in both sexes. In front wings, RS 3-branched and M 4-branched. Discoidal cell absent. Median cell open. Crossvein Cu1-Cu2 situated anteriorly. Cu2 complete to end. First anal cell very short and 2nd anal cell very long, since A1+2+3 short and ends at wing margin. Hind wings like those of Molanna. % genitalia (Figs. 872–875): segment IX reduced in height and invaginated within sternite VIII, which is enlarged and forms few apical carinae and concavities. Segment X clearly differentiated, forming horizontal, quadrangular, roof-like structure. Preanal appendages large, fairly long, and indented. Intermediate appendages developed into bifid spurs, far apart and curved downward. Inferior appendages in short, simple callipers with denticulate inner margin. Phallotheca forming short cone-like structure with lateral wings joined to base of inferior appendages. Endotheca very large, complex, consisting of stout, paired and unpaired lobes, and equipped with tubercles and short spines. & genitalia (Figs. 876, 877): segment X well differentiated and distinct from subcircular appendages. Ventroposterior portion of segment IX forming complex unit of plates and membranous folds, including false vulvar scale and gaping vaginal vestibule. Vaginal apparatus large. The genus Molannodes contains only a single Nearctic species, tinctus Zetterstedt, which is also widely distributed in the Palaearctic region. It has been reported from western Alaska and the Northwest Territories. It inhabits the same biotopes as Molanna. 202

List of Abbreviations Used in the Figures a. Anal opening of the Integripalpia a.ph. Phallic apparatus app. Appendages of segment X in the & Integripalpia b. Sclerotized band joining the phallotheca and the base of the inferior appendages of many Leptocerinae c. Cerci of the & Annulipalpia c.v. Vaginal chamber éc. éd. end. ép. ext.

Vulvar scale Aedeagus Endotheca Spurs Outer branches of segment X of the Limnephilidae and the Goeridae

g. Phallic apparatus guide of the Leptoceridae and Tinodes inf. Inferior appendages and lower branches of segment X of the Limnephilidae int. Intermediate appendages and inner branches of segment X of the Limnephilidae l.m.d.IX Dorsal median lobe of segment IX of the Goeridae oc. Ocelli o.a.v. Anovaginal opening of the & Annulipalpia o.v. Vaginal opening of the & Integripalpia PA Postanal vein of the Odontoceridae par. Parameres ph. Phallotheca phcr. Phallocrypt Pl. Labial palpi Plm. Middle plate of segment X of the Neophylacinae Pm. Maxillary palpi pr. Preanal appendages s.a Subanal plate and anal sclerites of the Rhyacophilidae Scl. Lateral sclerites of segment X of the Limnephilinae and Pseudostenophylacinae 203

s.ph. Phallotremal sclerite spr. Supragenital plate of the & Integripalpia sup. Superior appendages of the Limnephilidae t.c. Cephalic warts v. Valves of the & Leptoceridae vag. Vaginal apparatus vest. Vaginal vestibule VII, VIII, IX, X, XI, VIIIt., VIIIst., IXt., IXst. Refer to the last abdominal segments and their tergites and sternites. 1, 2 First and second segments of the inferior appendages, antennae or maxillary palpi.

204

Figs. 1–7. 1–3, Rhyacophila acropedes: 1, head, lateral; 2, head, dorsal; 3, venation. 4–7, Rh. vocala: 4, front leg; 5, % genitalia, lateral, with the right inferior appendage removed to show the structure of the phallic apparatus; 6, & genitalia, lateral; 7, vaginal apparatus, dorsal.

205

Figs. 8–19. 8–10, Himalopsyche phryganea: 8, % genitalia, lateral; 9, preanal appendages, dorsal; 10, & genitalia, lateral. 11, Palaeagapetus celsus, head and pronotum, dorsal (Ross 1944). 12, Glossosoma intermedium, head and pronotum, dorsal (Ross 1944). 13–16, Palaeagapetus celsus: 13, % genitalia, lateral; 14, % genitalia, ventral; 15, % genitalia, dorsal (Ross 1944); 16, vaginal apparatus; 17, Palaeagapetus finisorientis, venation (Botosaneanu and Levanidova 1987). 18–19, Culoptila cantha: 18, % genitalia, lateral; 19, & genitalia, ventral.

206

Figs. 20–31. 20–23, Glossosoma lividum: 20, maxillary and labial palpi; 21, front leg; 22, venation; 23, apical spurs of middle leg. 24, Gl. hoodi: % genitalia, lateral. 25, Gl. alascense: % genitalia, lateral. 26–29, Gl. lividum: 26, % genitalia, lateral; 27, ventral plate of sternite VI; 28, & genitalia, lateral; 29, vaginal apparatus, dorsal. 30–31, Gl. penitum: 30, % genitalia, lateral; 31, segment IX, ventral.

207

Figs. 32–41. 32–36, Agapetus hessi: 32, venation; 33, % genitalia, lateral; 34, inferior appendage, dorsal; 35, sternites V and VI; 36, & genitalia, lateral. 37, Protoptila erotica: venation (Ross 1944). 38–39, Pr. tenebrosa: 38, % genitalia, lateral; 39, sternite VIII, ventral. 40, Pr. maculata: & genitalia, ventral. 41, Glossosoma hoodi: venation.

208

Figs. 42–56. 42–51, Agraylea multipunctata: 42, head of the %, dorsal; 43, maxillary palpi; 44, venation; 45, sternite VII; 46, sternite V; 47, % genitalia, lateral; 48, segment IX and inferior appendages, ventral; 49, & genitalia, lateral; 50, & genitalia, ventral; 51, vaginal apparatus, ventral. 52–56, Ithytrichia clavata: 52, venation; 53, % genitalia, lateral; 54, % genitalia, ventral; 55, & segment VII, ventral (Ross 1944); 56, vaginal apparatus, ventral.

209

Figs. 57–68. 57, Hydroptila ampoda: % head, dorsal; the right posterior cephalic tubercle is flattened against the head in repose; on the left, it is raised, as the erectile organ is turgescent (drawn by A. Lutes). 58–62, H. spatulata: 58, venation; 59, % genitalia, lateral; 60, segment IX and inferior appendages, ventral; 61, phallic apparatus; 62, & segment VII and vaginal apparatus, ventral. 63–68, Orthotrichia cristata: 63, venation; 64, sternite VII, lateral; 65, % genitalia, lateral; 66, % genitalia, ventral; 67, phallic apparatus; 68, & segment VII, ventral.

210

Figs. 69–79. 69–74, Stactobiella delira: 69, venation; 70, % genitalia, lateral; 71, % genitalia, ventral; 72, phallic apparatus; 73, & segment VII, ventral; 74, vaginal apparatus, ventral. 75–79, Oxyethira serrata: 75, venation; 76, % genitalia, lateral; 77, % genitalia, ventral; 78, phallic apparatus; 79, & genitalia, lateral.

211

Figs. 80–91. 80–86, Leucotrichia pictipes: 80, venation; 81, pronotum and tegulae; 82, base of antennae; 83, % genitalia, lateral; 84, % genitalia, ventral; 85, apex of phallic apparatus; 86, & segment VII and vaginal apparatus, ventral. 87–91, Neotrichia okapa: 87, venation; 88, % genitalia, lateral; 89, % genitalia, ventral; 90, phallic apparatus. 91, N. halia: & segment VII, ventral.

212

Figs. 92–101. 92–97, Ochrotrichia tarsalis: 92, venation; 93, % genitalia, lateral; 94, % genitalia, dorsal; 95, aedeagus; 96, & segment VII, lateral; 97, vaginal apparatus, ventral. 98–101, Mayatrichia ayama: 98, venation; 99, % genitalia, lateral; 100, % genitalia, ventral; 101, vaginal apparatus, ventral (Ross 1944).

213

Figs. 102–109. 102–103, Chimarra obscura: 102, head, lateral; 103, head, dorsal. 104–109, Dolophilodes novusamericanus: 104, venation; 105, % genitalia, lateral; 106, segments IX and X, dorsal; 107, inferior appendage, ventral; 108, & genitalia, lateral; 109, vaginal appartus, lateral.

214

Figs. 110–122. 110, Wormaldia occidea: radial sector of front wing. 111–115, W. gabriella: 111, complete venation; 112, % genitalia, lateral; 113, segments IX and X, dorsal; 114, & genitalia, lateral; 115, vaginal apparatus, lateral. 116–122, Chimarra obscura: 116, venation; 117, % genitalia, lateral; 118, inferior appendages, ventral; 119, phallic apparatus, lateral, with endotheca completely extruded; 120, phallic apparatus, lateral, of another specimen, with endotheca invaginated; 121, & genitalia, lateral; 122, vaginal apparatus, lateral.

215

Figs. 123–136. 123–132, Arctopsyche ladogensis: 123, base of % antenna; 124, & middle leg; 125, abdominal hemogill system; 126, % venation; 127, & venation; 128, % genitalia, lateral; 129, segments IX and X, dorsal; 130, inferior appendage, ventral; 131, & genitalia, lateral; 132, & genitalia, ventral. 133–136, Parapsyche elsis: 133, % genitalia, lateral; 134, inferior appendage, ventral; 135, segments IX and X, dorsal; 136, & segments X and XI, lateral.

216

Figs. 137–143. Macrostemum zebratum: 137, % maxillary palpus; 138, head, dorsal; 139, venation; 140, % genitalia, lateral; 141, segments IX and X, dorsal; 142, & genitalia, lateral; 143, sternite VIII, ventral.

217

Figs. 144–156. 144–153, Hydropsyche recurvata: 144, % maxillary palpus; 145, % front claws; 146, % sternite V; 147, & sternite V; 148, venation; 149, % genitalia, lateral; 150, segments IX and X, dorsal; 151, phallic apparatus, dorsal; 152, & genitalia, lateral; 153, sternite VIII, ventral. 154–156, Potamyia flava: 154, venation; 155, % genitalia, lateral; 156, segments IX and X, dorsal.

218

Figs. 157–170. 157–162, Cheumatopsyche campyla: 157, venation; 158, % genitalia, lateral; 159, segment X, caudal; 160, phallic apparatus, dorsal; 161, & genitalia, lateral; 162, & genitalia, ventral. 163–170, Diplectrona modesta: 163, base of antenna; 164, % internal gland and filament of sternite V, dorsal; 165, & internal gland and filament of sternite V, lateral; 166, venation; 167, % genitalia, lateral; 168, segments IX and X, dorsal; 169, & genitalia, lateral; 170, & genitalia, ventral.

219

Figs. 171–177, Aphropsyche doringa: 171, venation (Ross 1944); 172, % genitalia, lateral; 173, inferior appendage, ventral; 174, phallic apparatus, lateral; 175, phallic apparatus, dorsal; 176, & genitalia, latera; 177, & genitalia, ventral (Nimmo 1987).

220

Figs. 178–192. 178, Neureclipsis bimaculata: % sternite V of the. 179, N. crepuscularis: venation. 180–181, N. bimaculata: 180, % genitalia, lateral; 181, inferior appendage, ventral. 182–184, N. crepuscularis: 182, phallic apparatus, lateral; 183, & genitalia, lateral; 184, & genitalia, ventral. 185, Polycentropus interruptus: maxillary palpus. 186, P. variegatus: hemogill system of segment IV. 187–192, P. interruptus: 187, % venation; 188, & hind wing; 189, % sternite V; 190, % genitalia, lateral with the left inferior appendage removed; 191, & genitalia, lateral; 192, & genitalia, ventral.

221

Figs. 193–198. Paranyctiophylax moestus: 193, venation; 194, % genitalia, lateral; 195. inferior appendage, caudal; 196, phallic apparatus, lateral; 197, & genitalia, lateral; 198, & genitalia, ventral.

222

Figs. 199–209. 199–204, Cyrnellus fraternus: 199, maxillary palpus; 200, venation; 201, % genitalia, lateral; 202, inferior appendage, ventral; 203, & genitalia, lateral; 204, & genitalia, ventral. 205–206, Cernotina calcea: 205, maxillary palpus; 206, venation. 207–209, C. pallida: 207, % genitalia, lateral; 208, & genitalia, lateral; 209, & genitalia, ventral.

223

Figs. 210–223. 210–216, Phylocentropus placidus: 210, % head, dorsal; 211, maxillary and labial palpi; 212, hemogill system of segment III; 213, venation; 214, % genitalia, lateral; 215, inferior appendage, ventral; 216, segment X, ventral. 217, Ph. lucidus: segment X, ventral. 218–219, Ph. placidus: 218, & genitalia, lateral; 219, & genitalia, ventral. 220–223, Lype diversa: 220, venation; 221, % genitalia, lateral; 222, inferior appendage, ventral; 223, & genitalia, lateral.

224

Figs. 224–233. 224, Tinodes cascadius: venation. 225–229, T. provo: 225, % genitalia, lateral; 226, intermediate appendages, dorsal; 227, inferior appendage, ventral; 228, & genitalia, lateral; 229, & genitalia, ventral. 230–233, Psychomyia flavida: 230, maxillary palpus; 231, venation; 232, % genitalia, lateral, 233, & genitalia, ventral.

225

Figs. 234–240. Dicosmoecus gilvipes: 234, venation; 235, head, dorsal view; 236, % genitalia, lateral; 237, % genitalia, dorsal; 238, phallic apparatus, dorsal; 239, & genitalia, lateral; 240, & genitalia, ventral.

226

Figs. 241–245. Onocosmoecus unicolor: 241, % genitalia, lateral; 242, % genitalia, dorsal; 243, phallic apparatus, dorsal; 244, & genitalia, lateral; 245, & genitalia, ventral.

227

Figs. 246–252, Eocosmoecus schmidi: 246, % genitalia, lateral; 247, % genitalia, dorsal; 248, % genitalia, ventral; 249, phallic apparatus, lateral; 250, phallic apparatus, dorsal; 251, & genitalia, lateral; 252, & genitalia, ventral (Wiggins et al. 1989).

228

Figs. 253–260. Allocosmoecus partitus: 253, % maxillary palpus; 254, & maxillary palpus; 255, % genitalia, lateral; 256, % genitalia, dorsal; 257, segment IX and inferior appendage, ventral; 258, phallic apparatus, dorsal; 259, & genitalia, lateral; 260, & genitalia, ventral.

229

Figs. 261–265. Amphicosmoecus canax: 261, % genitalia, lateral; 262, inferior appendage, ventral; 263, phallic apparatus, dorsal; 264, & genitalia, lateral; 265, & genitalia, ventral.

230

Figs. 266–271. Pedomoecus sierra: 266, venation; 267, % genitalia, lateral; 268, % genitalia, caudal; 269, phallic apparatus, lateral; 270, & genitalia, lateral; 271, & genitalia, ventral.

231

Figs. 272–278. 272, Allomyia bifosa: venation. 273–278, A. tripunctata: 273, % genitalia, lateral; 274, % genitalia, ventral; 275, segments IX and X, dorsal; 276, phallic apparatus, lateral; 277, & genitalia, lateral; 278, & genitalia, ventral.

232

Figs. 279–286, Manophylax annulatus: 279, % front wing; 280, & both wings; 281, % genitalia, lateral; 282, % genitalia, dorsal; 283, phallic apparatus, lateral, 284, phallic apparatus, ventral; 285, & genitalia, lateral; 286, & genitalia, ventral (Wiggins et al. 1973).

233

Figs. 287–293. Cryptochia pilosa: 287, venation; 288, % genitalia, lateral; 289, % genitalia, dorsal; 290, segment X, caudal; 291, phallic apparatus, lateral; 292, & genitalia, lateral; 293, & genitalia, ventral.

234

Figs. 294–300. Rossiana montana: 294, % maxillary palpus; 295, venation; 296, % genitalia, lateral, with the left inferior appendage removed; 297, % genitalia, dorsal; 298, phallic apparatus, lateral; 299, & genitalia, lateral; 300, & genitalia, ventral.

235

Figs. 301–307. Ecclisomyia conspersa: 301, front wing venation; 302, % genitalia, lateral; 303, segments IX and X, dorsal; 304, inferior appendage, caudal; 305, phallic apparatus, dorsal; 306, & genitalia, ventral; 307, & genitalia, lateral.

236

Figs. 308–313. Ecclisocosmoecus scylla: 308, % front wing venation; 309, % genitalia, lateral; 310, % genitalia, caudal; 311, phallic apparatus, dorsal; 312, & genitalia, lateral; 313, & genitalia, ventral.

237

Figs. 314–320. Ironoquia punctatissima: 314, % genitalia, lateral; 315, % genitalia, caudal; 316, phallic apparatus, dorsal; 317, inferior appendage, caudal; 318, & genitalia, lateral; 319, & genitalia, dorsal; 320, & genitalia, caudal.

238

Figs. 321–326. Apatania zonella: 321, % venation; 322, % genitalia, lateral; 323, segments IX and X, dorsal; 324, phallic apparatus, lateral; 325, & genitalia, lateral; 326, &, ventral.

239

Figs. 327–331. Pseudostenophylax sparsus sparsus: 327, % genitalia, lateral; 328, % genitalia, caudal; 329, phallic apparatus, ventral; 330, & genitalia, lateral; 331, & genitalia, ventral.

240

Figs. 332–338. 332, Limnephilus submonilifer: % front leg. 333–338, L. indivisus: 333, venation; 334, % genitalia, lateral; 335, % genitalia, caudal; 336, phallic apparatus, lateral; 337, & genitalia, lateral; 338, & genitalia, ventral.

241

Figs. 339–352. 339–345, Grammotaulius interrogationis: 339, % genitalia, lateral; 340, % genitalia, caudal; 341, segments IX and X, dorsal; 342, paramere, lateral; 343, & genitalia, lateral; 344, & genitalia, ventral; 345, & genitalia, dorsal. 346–352, Nemotaulius hostilis: 346, % genitalia, lateral; 347, % genitalia, caudal; 348, segments IX and X, dorsal; 349, paramere, lateral; 350, & genitalia, lateral; 351, & genitalia, caudal; 352, & genitalia, dorsal.

242

Figs. 353–362. 353–357, Leptophylax gracilis: 353, venation; 354, % genitalia, lateral; 355, phallic apparatus, lateral; 356, & genitalia, lateral; 357, & genitalia, ventral. 358–362, Asynarchus batchawana: 358, % genitalia, lateral; 359, % genitalia, caudal; 360, phallic apparatus, lateral; 361, & genitalia, lateral; 362, & genitalia, ventral.

243

Figs. 363–378. 363–369, Anabolia bimaculata: 363, % genitalia, lateral; 364, % genitalia, dorsal; 365, % genitalia, caudal; 366, paramere, lateral; 367, & genitalia, lateral; 368, & genitalia, ventral; 369, & genitalia, dorsal. 370–378, Halesochila taylori: 370, % genitalia, lateral; 371, % genitalia, caudal; 372, % genitalia, dorsal; 373, apex of the aedeagus, dorsal; 374–375, apex of paramere, seen from two different angles; 376, & genitalia, dorsal; 377, & genitalia, ventral; 378, & genitalia, lateral.

244

Figs. 379–389. 379–385, Arctopora pulchella: 379, % genitalia, lateral; 380, % genitalia, dorsal; 381, % genitalia, caudal; 382, paramere, lateral; 383, & genitalia, lateral; 384, & genitalia, dorsal; 385, & genitalia, ventral. 386–389, Lenarchus (Prolenarchus) keratus: 386, % genitalia, lateral; 387, % genitalia, dorsal; 388, % genitalia, caudal; 389, phallic apparatus, dorsal.

245

Figs. 390–402. 390–396, Lenarchus (Lenarchus) rho: 390, % genitalia, lateral; 391, % genitalia, dorsal; 392, % genitalia, caudal; 393, paramere, lateral; 394, & genitalia, lateral; 395, & genitalia, dorsal; 396, & genitalia, ventral. 397–402, Lenarchus (Paralenarchus) vastus: 397, % genitalia, lateral; 398, % genitalia, dorsal; 399, % genitalia, caudal; 400, & genitalia, lateral; 401, & genitalia, dorsal; 402, & genitalia, ventral.

246

Figs. 403–408. Philarctus quaeris: 403, % front leg; 404, % genitalia, lateral; 405, % genitalia, dorsal; 406, phallic apparatus, lateral; 407, & genitalia, lateral; 408, & genitalia, caudal.

247

Figs. 409–418. 409–413, Clistoronia (Clistoronia) magnifica: 409, % genitalia, lateral; 410, % genitalia, caudal; 411, phallic apparatus, lateral; 412, & genitalia, lateral; 413, & genitalia, caudal. 414–418, Clistoronia (Clistoroniella) flavicollis: 414, % genitalia, lateral; 415, % genitalia, caudal; 416, phallic apparatus, lateral; 417, & genitalia, lateral; 418, & genitalia, caudal.

248

Figs. 419–425. Platycentropus radiatus: 419, apical spur of % hind legs; 420, front wing venation; 421, % genitalia, lateral; 422, % genitalia, dorsal; 423, phallic apparatus, lateral; 424, & genitalia, lateral; 425, & genitalia, caudal.

249

Figs. 426–432. Hydatophylax argus: 426, % genitalia, lateral; 427, % genitalia, dorsal; 428, % genitalia, caudal; 429, phallic apparatus, lateral; 430, & genitalia, lateral; 431, & genitalia, dorsal; 432, & genitalia, ventral.

250

Figs. 433–444. 433–439, Pycnopsyche subfasciata: 433, % genitalia, lateral; 434, % genitalia, caudal; 435, % genitalia, dorsal; 436, phallic apparatus, lateral; 437, & genitalia, lateral; 438, & genitalia, caudal; 439, % abdominal sternite V. 440–442, Philocasca thor: 440, % genitalia, lateral; 441, % genitalia, dorsal; 442, phallic apparatus, dorsal. 443–444, Ph. demita: 443, & genitalia, lateral; 444, & genitalia, caudal (Wiggins et al. 1968).

251

Figs. 445–454, Sphagnophylax meiops: 445, % genitalia, lateral; 446, % genitalia, dorsal; 447, % genitalia, caudal; 448, % genitalia, ventral; 449, phallic apparatus, dorsal; 450, phallic apparatus lateral; 451, venation; 452, & genitalia, lateral; 453, & genitalia, dorsal; 454, & genitalia, ventral (Wiggins et al. 1984).

252

Figs. 455–466. 455–459, Hesperophylax incisus: 455, % genitalia, lateral; 456, % genitalia, caudal; 457, phallic apparatus, dorsal; 458, & genitalia, caudal; 459, & genitalia, lateral. 460–466, Chyranda centralis: 460, % head, lateral; 461, % maxillary and labial palpi; 462, % genitalia, lateral; 463, % genitalia, caudal; 464, phallic apparatus, dorsal; 465, & genitalia, lateral; 466, & genitalia, caudal.

253

Figs. 467–471. Clostoeca disjuncta: 467, % genitalia, lateral; 468, % genitalia, caudal; 469, phallic apparatus, lateral; 470, & genitalia, lateral; 471, & genitalia, caudal.

254

Figs. 472–484. 472–478, Frenesia missa: 472, % genitalia, lateral; 473, % genitalia, dorsal; 474, % genitalia, caudal; 475, phallic apparatus, dorsal; 476, & genitalia, lateral; 477, & genitalia, dorsal; 478, & genitalia, caudal. 479–484, Glyphopsyche irrorata: 479, % genitalia, lateral; 480, % genitalia, dorsal; 481, % genitalia, caudal; 482, & genitalia, lateral; 483, & genitalia, dorsal; 484, & genitalia, ventral.

255

Figs. 485–500. 485–488, Chilostigmodes areolatus: 485, % genitalia, lateral; 486, % genitalia, dorsal; 487, % genitalia, caudal; 488, phallic apparatus, dorsal. 489–495, Grensia praeterita: 489, % genitalia, lateral; 490, % genitalia, dorsal; 491, % genitalia, caudal; 492, phallic apparatus, dorsal; 493, & genitalia, lateral; 494, & genitalia, dorsal; 495, & genitalia, caudal. 496–500, Chilostigma itascae: 496, % genitalia, lateral; 497, % genitalia, dorsal; 498, phallic apparatus, lateral; 499, & genitalia, ventral; 500, & genitalia, lateral (Wiggins 1975).

256

Figs. 501–509. 501–506, Psychoglypha alascensis: 501, % genitalia, lateral; 502, % genitalia, dorsal; 503, segment IX and inferior appendages, ventral; 504, phallic apparatus, lateral; 505, & genitalia, ventral; 506, & genitalia, lateral. 507, Frenesia missa, pterostigma and anastomosis of front wing. 508, Grensia praeterita, pterostigma and anastomosis of front wing. 509, Chilostigmodes areolatus, pterostigma and anastomosis of front wing.

257

Figs. 510–516. Desmona mono: 510, % genitalia, lateral; 511, % genitalia, dorsal; 512, % genitalia, ventral; 513, phallic apparatus, lateral; 514, phallic apparatus, dorsal; 515, & genitalia, lateral; 516, & genitalia, ventral (Wiggins et al. 1990).

258

Figs. 517–527. 517, Homophylax baldur: % venation of the base of the front wing. 518, H. andax: % abdominal sternite V, with the internal gland. 519, H. flavipennis: & abdominal sternite V, with the internal gland. 520, H. baldur: % venation. 521–522, H. andax: 521, % hind wing venation. 522, & hind wing venation. 523–525, H. baldur: 523, % genitalia, lateral; 524, % genitalia, dorsal; 525, phallic apparatus, lateral. 526–527, H. crotchi: 526, & genitalia, lateral; 527, & genitalia, caudal.

259

Figs. 528–534. Phanocelia canadensis: 528, venation; 529, % genitalia, lateral; 530, % genitalia, dorsal; 531, % genitalia, caudal; 532, phallic apparatus, dorsal; 533, & genitalia, lateral; 534, & genitalia, caudal.

260

Figs. 535–543. 535, Neophylax oligius: % venation. 536, N. oligius: & hind wing venation. 537, N. concinnus: & hind wing venation. 538, N. occidentis: % hind spurs. 539, N. rickeri: % hind spurs. 540–543, N. occidentis: 540, % genitalia, lateral; 541, % genitalia, caudal; 542, & genitalia, lateral; 543, & genitalia, caudal.

261

Figs. 544–548. 544, Oligophlebodes minutus: % venation. 545–546, O. mostbento: 545, % genitalia, lateral; 546, segment IX and inferior appendages, ventral. 547–548, O. sierra: 547, & genitalia, lateral; 548, & genitalia, ventral.

262

Figs. 549–554. 549–550, Neothremma didactyla: 549, & venation; 550, % hind wing venation. 551–552, N. alicia: 551, % genitalia, lateral; 552, inferior appendages, ventral. 553–554, N. didactyla: 553, & genitalia, lateral; 554, & genitalia, ventral.

263

Figs. 555–561. Farula jewetti: 555, & venation; 556, % hind wing venation; 557, % genitalia, lateral; 558, segments IX and X, dorsal; 559, inferior appendages, ventral; 560, & genitalia, ventral; 561, & genitalia, lateral.

264

Figs. 562–572, Sericostria surdickae: 562, venation; 563, head, dorsal; 564, % maxillary palpus; 565, % genitalia, lateral; 566, % genitalia, dorsal; 567, phallic apparatus, lateral and dorsal; 568, % genitalia, ventral; 569, % genitalia, caudal; 570, & genitalia, lateral; 571, & genitalia, ventral; 572, & genitalia, dorsal (Wiggins et al. 1985).

265

Figs. 573–583, Goera calcarata: 573, % head, dorsal; 574, % maxillary palpus in erection; 575, & maxillary palpus; 576, % venation; 577, % points of sternite VI; 578, % genitalia, lateral; 579, % genitalia, dorsal; 580, segment IX and inferior appendage, ventral; 581, phallic apparatus, lateral; 582, & genitalia, lateral; 583, & genitalia, ventral.

266

Figs. 584–591, Goeracea genota: 584, % maxillary palpus; 585, % labial palpus; 586, venation; 587, % genitalia, lateral; 588, % genitalia, dorsal; 589, segment IX and inferior appendage, ventral; 590, & genitalia, lateral; 591, & genitalia, ventral.

267

Figs. 592–600, Goereilla baumanni: 592, % maxillary palpus; 593, % labial palpus; 594, venation; 595, % genitalia, lateral; 596, % genitalia, dorsal; 597, segment IX and inferior appendage, ventral; 598, phallic apparatus, lateral; 599, & genitalia, lateral; 600, & genitalia, ventral (Wiggins 1974).

268

Figs. 601–605, Lepania cascada: 601, venation (Wiggins 1973); 602, % genitalia, lateral; 603, % genitalia, dorsal; 604, & genitalia, lateral; 605, & genitalia, ventral.

269

Figs. 606–618. 606–607, L. (Lepidostoma) togatum: 606, % head, dorsal; 607, & first article of antenna. 608, L. (Nosopus) cascadense, % first two segments of antenna. 609–617, L. (Lepidostoma) togatum, 609, median tibia; 610, % venation; 611, & venation; 612, % genitalia, lateral; 613, % segments IX and X, dorsal; 614, inferior appendage, ventral; 615, phallic apparatus, lateral; 616, & genitalia, lateral; 617, & genitalia, ventral; 618, L. (Lepidostoma) cascadense, phallic apparatus, lateral.

270

Figs. 619–627. 619–620, L. (Nosopus) podager: 619, % venation; 620, & venation; 620a, L. (Lepidostoma) latipenne, spermatheca. 621–627, L. (Lepidostoma) americanum: 621, % genitalia, lateral; 622, segments IX and X, dorsal; 623, inferior appendage, ventral; 624, phallic apparatus, lateral; 625, phallic apparatus, dorsal; 626, & genitalia, lateral; 627, vaginal apparatus (Weaver 1988).

271

Figs. 628–636, L. (Mormomyia) vernale: 628, % venation; 629, & venation; 630, % genitalia, lateral; 631, segments IX and X, dorsal; 632, inferior appendages, dorsal; 633, phallic apparatus, lateral; 634, phallic apparatus, dorsal; 635, & genitalia, lateral; 636, vaginal apparatus, dorsal (Weaver 1988).

272

Figs. 637–647, L. (Neodinarthrum) pluviale: 637, % venation; 638, & venation; 639, % genitalia, lateral; 640, segment IX and X, dorsal; 641, % X segment, caudal; 642, inferior appendages, ventral; 643, phallic apparatus, lateral; 644, & genitalia, lateral; 645, & genitalia, ventral; 646, vaginal apparatus, ventral; 647, vaginal apparatus, lateral (Weaver 1988).

273

Fig. 648–657. 648, Theliopsyche grisea: % head, dorsal. 649, L. (Neodinarthrum) pluviale: % venation. 650, L. (Nosopus) cascadense: % front wing venation. 651–657, Th. grisea: 651, % venation; 652, & of hind wing venation; 653, % sternite VIII, ventral; 654, % genitalia, lateral; 655, segment X, dorsal; 656, inferior appendage, ventral; 657, & genitalia, ventral.

274

Figs. 658–667. 658–661, Oligostomis ocelligera: 658, % genitalia, lateral; 659, segment IX and inferior appendage, ventral; 660, phallic apparatus, lateral; 661, & genitalia, ventral. 662–667, Hagenella canadensis: 662, % venation; 663, % genitalia, lateral; 664, segments IX and X, dorsal; 665, segment IX and inferior appendage, ventral; 666, phallic apparatus, lateral; 667, & genitalia, ventral.

275

Figs. 668–672. Ptilostomis semifasciata: 668, % genitalia, lateral; 669, segments IX and X, dorsal; 670, segment IX and inferior appendage, ventral; 671, phallic apparatus, lateral; 672, & genitalia, ventral.

276

Figs. 673–678. 673–675, Oligotricha lapponica: 673, % genitalia, lateral; 674, inferior appendage, caudal; 675, & genitalia, ventral. 676–678, Banksiola crotchi: 676, % genitalia, lateral; 677, inferior appendage, ventral; 678, & genitalia, ventral.

277

Figs. 679–686. 679–682, Fabria inornata: 679, % genitalia, lateral; 680, inferior appendage, ventral; 681, phallic apparatus, lateral; 682, & genitalia, ventral. 683–686, Agrypnia vestita: 683, % genitalia, lateral; 684, inferior appendage, ventral; 685, phallic apparatus, lateral; 686, & genitalia, ventral.

278

Figs. 687–699. Phryganea cinerea: 687, % head, dorsal; 688, base of antenna; 689, % maxillary palpus; 690, & maxillary palpus; 691, sternite V with internal gland; 692, % venation; 693, & venation; 694, % genitalia, lateral; 695, % genitalia, dorsal; 696, phallic apparatus, lateral; 697, & genitalia, lateral; 698, & genitalia, ventral; 699, abdominal hemogill system of second and third segments.

279

Figs. 700–707. Beothukus complicatus: 700, % venation; 701 % genitalia, lateral; 702, % genitalia, dorsal; 703, % genitalia, caudal; 704, phallic apparatus, lateral; 705, & genitalia, lateral; 706, & genitalia, ventral; 707, vulvar scale (Wiggins et al. 1989).

280

Figs. 708–714. Eobrachycentrus gelidae: 708, % venation; 709, & venation; 710, % maxillary palpus; 711, % genitalia, lateral; 712, % genitalia, dorsal; 713, & genitalia, lateral; 714, & genitalia, ventral.

281

Figs. 715–727. Brachycentrus numerosus: 715, % head, dorsal; 716, % labial palpus; 717, % maxillary palpus; 718, & maxillary palpus; 719, % middle tibia; 720, % venation; 721, & venation; 722, % genitalia, lateral; 723, segment IX and preanal appendages, dorsal; 724, segment X (or intermediate appendages), dorsal; 725, segment IX and inferior appendage, caudal; 726, & genitalia, lateral; 727, & genitalia, ventral.

282

Figs. 728–742. 728–739, Micrasema rusticum: 728, % front leg; 729, & front leg; 730, % maxillary palpus; 731, % labial palpus; 732, & maxillary palpus; 733, & labial palpus; 734, % venation; 735, & venation; 736, % genitalia, lateral; 737, segment IX and preanal appendages, dorsal; 738, segment X (or intermediate appendages), dorsal; 739, inferior appendage, caudal. 740, M. bactro: % inferior appendage, lateral. 741–742, M. rusticum: 741, & genitalia, lateral; 742, & genitalia, ventral.

283

Figs. 743–750. Amiocentrus aspillus: 743, % maxillary palpus; 744, % labial palpus; 745, & front wing anal area and hind wing; 746, % genitalia, lateral; 747, segments IX and X, dorsal; 748, inferior appendage, ventral; 749, & segment X, dorsal; 750, & genialia, ventral.

284

Figs. 751–761. Agarodes distinctus: 751, % head, lateral; 752, % maxillary palpus, inner side; 753, & maxillary palpus; 754, & labial palpus; 755, % venation; 756, % genitalia, lateral; 757, segments IX and X, dorsal; 758, segment IX and inferior appendage, ventral; 759, & genitalia, lateral; 760, & genitalia, dorsal; 761, & genitalia, ventral.

285

Figs. 762–770. Helicopsyche borealis: 762, % head, dorsal; 763, % maxillary palpus; 764, & maxillary palpus; 765, % venation; 766, % genitalia, lateral, with right inferior appendage removed to show inside; 767, % genitalia, dorsal; 768, segment IX and inferior appendage, ventral; 769, & genitalia, lateral; 770, & genitalia, ventral.

286

Figs. 771–778. Beraea fontana: 771, % head, lateral; 772, % venation; 773, & venation; 774, % genitalia, lateral; 775, % genitalia, ventral; 776, phallic apparatus, ventral; 777, & genitalia, lateral; 778, & genitalia, ventral.

287

Figs. 779–792. 779–782, Ceraclea annulicornis: 779, % head, dorsal; 780, front claws; 781, maxillary palpus. 782, C. submaculata: abdominal hemogill system of segments III and IV. 783–788, C. annulicornis: 783, % venation; 784, & venation; 785, % genitalia, lateral; 786, segments IX and X, dorsal; 787, inferior appendage, caudal; 788, phallic apparatus, lateral. 789, C. erratica: phallic apparatus, lateral. 790–792, C. annulicornis: 790, & genitalia, lateral; 791, & genitalia, dorsal; 792, & genitalia, ventral.

288

Figs. 793–799. Mystacides sepulchralis: 793, venation; 794, % genitalia, lateral; 795, segment IX, ventral; 796–797, intermediate appendages of 2 specimens, dorsal; 798, & genitalia, lateral; 799, & genitalia, ventral.

289

Figs. 800–810. 800–804, Triaenodes marginatus: 800, % first antennal segment; 801, venation; 802, % genitalia, lateral; 803, preanal and intermediate appendages, dorsal; 804, inferior appendage, ventral; 805–806, Tr. injustus: 805, & genitalia, lateral; 806, & genitalia, ventral. 807–810, Ylodes griseus: 807, % genitalia, lateral; 808, segments IX and X, dorsal; 809, & genitalia, lateral; 810, & genitalia, ventral.

290

Figs. 811–819. 811–815, Setodes oligius: 811, venation; 812, % genitalia, lateral; 813, inferior appendage, ventral; 814, & genitalia, lateral; 815, & genitalia, ventral. 816–819, Leptocerus americanus: 816, venation; 817, % genitalia, lateral; 818, & genitalia, lateral; 819, & genitalia, ventral.

291

Figs. 820–825. Oecetis inconspicua: 820, venation; 821, % genitalia, lateral; 822, % genitalia, dorsal; 823, segment IX and inferior appendage, ventral; 824, & genitalia, lateral; 825, & genitalia, ventral.

292

Figs. 826–831. 826–829, Nectopsyche exquisita: 826, hemogill system of segment III; 827, % venation; 828, & venation; 829, % genitalia, lateral. 830–831, Nectopsyche albida: 830, & genitalia, lateral; 831, & genitalia, ventral.

293

Figs. 832–841. Psilotreta indecisa: 832, % head, dorsal; 833, % maxillary palpus; 834, % venation; 835, & venation; 836, % genitalia, lateral; 837, segments IX and X, dorsal; 838, phallic apparatus, lateral; 839, & genitalia, lateral; 840, & genitalia, dorsal; 841, & genitalia, ventral.

294

Figs. 842–851. Marilia flexuosa: 842, % head, dorsal; 843, & head, dorsal; 844, % middle leg; 845, % venation; 846, & venation; 847, % genitalia, lateral; 848, segments IX and X, dorsal; 849, segment IX and inferior appendage, ventral; 850, & genitalia, lateral; 851, & genitalia, ventral.

295

Figs. 852–859. Heteroplectron californicum: 852, % head, dorsal; 853, venation; 854, % genitalia, lateral; 855, % genitalia, dorsal; 856, phallic apparatus, lateral; 857, & genitalia, lateral; 858, & genitalia, ventral; 859, abdominal hemogill system of segment V.

296

Figs. 860–869. Molanna flavicornis: 860, % head, dorsal; 861, % maxillary palpus; 862, abdominal hemogill system and internal gland of segment V; 863, % venation; 864, & venation; 865, % genitalia, lateral; 866, segment IX and inferior appendage, ventral; 867, phallic apparatus, lateral; 868, & genitalia, lateral; 869, & genitalia, ventral.

297

Figs. 870–877. Molannodes tinctus: 870, % prothoracic tubercles in erection; 871, % venation; 872, % genitalia, lateral; 873, segments IX and X, dorsal; 874, sternite VIII, segment IX and inferior appendage, ventral; 875, phallic apparatus, lateral; 876, & genitalia, lateral; 877, % genitalia, ventral.

298

Glossary acuminate Tapering to a long point. anastomosis A network of crossveins located at the apical third of both pairs of wings. androconia Elongated scent scales. apodemal Refers to an internal extension of the base of an appendage or segment. callosity Thickening of the membrane of the anal area of the front wing of Glossosoma. campodeiform Refers to a type of larva in which the head forms a very obtuse angle with the body axis. crenulation Transverse ridges. eruciform Refers to a type of larva in which the head forms a right angle with the body axis. eurytherm An organism that tolerates a wide range of temperature; adj. eurythermal. genitalia Latin, plural noun, designating all the external genital parts. habitus General appearance of an insect. hemogills Thread-like organs located on the abdominal pleurites and designed to oxygenate the blood. hydropetric A species living on a stone wall, under a thin film of water. inerm A neologism from Latin inermis; unarmed, a part without spines or teeth. labial palpi Small articulated organs inserted on the labium. lentic Refers to running water, rivers, torrents, and streams. lotic Refers to still and stagnant water, lakes, ponds, and marshes. maxillary palpi Small articulated organs inserted on the maxillae. moniliform Refers to antennae in which the segments are subglobular and not longer than wide. neoformation A specialization of a pre-existing form. ocelli Simple eyes, of which there are 3. They are situated on the vertex. ogival In the shape of an ogive, an arch with a pointed apex. ommatidia Simple eyes, which together form the compound eyes. pedunculate Synonym for petiolate. petiolate Having a petiole. Used for a wing fork that is joined to a cross vein by a stalk. sessile Without a petiole or a peduncle. Refers to a wing fork that begins at its connection with a cross vein. spurs Strongly thickened spines on the tibiae. 299

stenotherm An organism that does not withstand large changes in temperature; adj. stenothermal. sternite Lower part of an abdominal or thoracic segment. tegulae Mesothoracic scales protecting the base of the front wings. tergite Upper part of an abdominal segment. venation Arrangement of veins on the wings, providing a stiff framework for the wings (see Fig. 41 for an explanation of the veins). vertex Upper part of the head, located between the eyes. vicariism Replacement of a lost part by a new one. wing forks Forking of the radial, median and cubital veins, numbered from I to V (see Fig. 41).

300

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Wiggins, G.B. 1965. Additions and revisions to the genera of North American caddis-flies of the family Brachycentridae with special reference to the larval stages, Can. Entomol., 97: 1089–1106. Wiggins, G.B. 1973a. New systematic data for the North American caddisfly genera Lepania, Goeracea and Goerita, Life Sci. Contr., R. Ont. Mus., Toronto, 91: 33 p. Wiggins, G.B. 1973b. Contributions to the systematics of the caddisfly family Limnephilidae I, Life Sci. Contr., R. Ont. Mus., Toronto, 94: 32 p. Wiggins, G.B. 1974. Contributions to the systematics of the caddisfly family Limnephilidae III. The genus Goereilla, First Int. Symp. Trichoptera, Junk, Dordrecht: 7–19. Wiggins, G.B. 1975. Contibutions to the systematics of the caddisfly family Limnephilidae II, Can. Entomol. 107: 325–336. Wiggins, G.B. 1996. The larvae of the North American caddisfly genera, 2nd edition, University of Toronto Press, 457 p. Wiggins, G.B. and N.H. Anderson. 1968. Contributions to the systematics of the caddisfly genera Pseudostenophylax and Philocasca, with special reference to the immature stages, Can. J. Zool., 46: 61–75. Wiggins, G.B. and D.S. Larson. 1989. Systematics and biology for a new Nearctic genus in the caddisfly family Phryganeidae, Can. J. Zool., 67(6): 1550–1556. Wiggins, G.B. and J.S. Richardson. 1982. Revision and synopsis of the caddisfly genus Dicosmoecus, Aquatic Insects, 4(4): 181–217. Wiggins, G.B. and J.S. Richardson. 1986. Revision of the Onocosmoecus unicolor group, Psyche, 93(3–4): 187–216. Wiggins, G.B. and J.S. Richardson. 1989. Biosystematics of Eocosmoecus, a new Nearctic caddisfly genus, J. North Amer. Benth. Soc., 8(4): 355–369. Wiggins, G.B. and N.N. Winchester. 1984. A remarkable new caddisfly genus from northwest North America, Can. J. Zool., 62(9): 1853–1858. Wiggins, G.B. and R.W. Wisseman. 1990. Revision of the North American caddisfly genus Desmona, Ann. Entomol. Soc. Amer., 83(2): 155–161. Wiggins, G.B., J.S. Weaver and J.D. Unzicker. 1985. Revision of the caddisfly family Uenoidae, Can. Entomol., 117: 763–800. Yamamoto, T. and G.B. Wiggins. 1964. A comparative study of the North American species of the caddisfly genus Mystacides, Can. J. Zool., 42: 1105–1126.

310

Index Acrocentropus 68 Acronopsyche 136 pilosa 136 Adicrophleps 13 Agapetinae 9, 21, 22, 26, 30 Agapetus 4, 9, 26, 27 fuscipes 26 hessi 208 Agarodes 13, 174–176 distinctus 176, 285 griseus 174, 176 Agraylea 9, 33, 34 costello 34 multipunctata 34, 209 saltesa 34 sexmaculata 33 Agrypnia 13, 157, 158, 164, 165 glacialis 164 pagetana 164 vestita 278 Alepomyia 151 bryanti 151, 152 Alepomyiodes 152 Algonquina 98 Alisotrichia 9 Allegophylax 118 Alleodes 160 Allocosmoecus 11, 77, 81, 82 partitus 81, 82, 229 Allomyia 11, 76–78, 84, 85 bifosa 85, 232 tripunctata 232 Allophylax 90, 91 Amiocentrus 13, 169, 173 aspilus 173, 284 Amphicosmoecus 11, 77, 78, 82, 83 canax 83, 230 Anabolia 11, 98, 104, 105 bimaculata 105, 244 clathrata 160 consocia 105

311

ozburni 105 sordida 105 Anabolina 98 diversa 98 Anagapetini 23 Anagapetus 9, 23, 24 bernea 24 debilis 23, 24 hoodi 24 Anchitrichia 9 Anisocentropus 14 Anisogamus disjunctus 123 Annulipalpia 6, 7, 9 Anseriglossa 9, 23, 25, 26 penitum 26 Anticyra phaeopha 70 Aopsyche 154 corona 154 Apatania 11, 91–93 canadensis 132 tripunctata 84 wallengreni 92 zonella 93, 239 Apataniinae 11, 74, 75, 91, 147 Aphelocheira flavomaculata 57 ladogensis 48 Aphropsyche 58 aprilis 58 doringa 220 Apolopsyche 98 Aragodes 174–176 distinctus 176 griseus 174, 176 Arcadopsyche 152 prominens 152 Arctopora 11, 97, 107, 108 pulchella 108, 245 salmon 108 trimaculata 107, 108

Arctopsyche 9, 47–50 apicalus 49 grandis 49 inermis 49 ladogensis 48, 49, 216 Arctopsychidae 4, 10, 16, 47, 48, 70 Astenophylax 117 Astoplectron 195 Asynarchus 11, 98, 103, 104 alascensis 164 batchawana 104, 243 bicornis 109 centralis 122 flavicollis 115 fusorius 103 montanus 104 mutatus 104 productus 108, 110 Athripsodes 184 Athripsodina 13, 184 Athripsodini 13 Atomyia 151 modesta 151 Atomyiodes 152 bispinosus 152 Atopsyche 10 Austrotinodes 11 Banksiola 13, 157–159, 162, 163 crotchi 163, 277 dossuaria 162 Beothukus 13, 158, 166, 167 complicatus 167, 280 Beraea 13, 179, 180 fontana 180, 287 gorteba 180 maculata 28 nigritta 180 Beraeidae 13, 16, 179 Brachycentridae 4, 13, 16, 17, 168, 169 Brachycentrinae 168 Brachycentrus 13, 168–171, 173 americanus 170 aspilus 173 incanus 171 lateralis 171 nigrisoma 171 numerosus 282 subnubilus 170 Caborius 91 Calamoceras 198 Calamoceratidae 4, 14, 16, 198

Centromacronema 10 Ceraclea 13, 182–184 annulicornis 288 erratica 288 nigronervosa 183 submaculata 288 Ceratopsyche 54 Cernotina 10, 61, 65, 66 calcea 65, 223 pallida 66, 223 spicata 66 Chaetopterygini 95 Chaetopterygopsis parvula 90 Cheumatopsyche 10, 54–57 campyla 57, 219 speciosa 57 Chilostigma 12, 97, 129 itascae 129, 256 sieboldi 128 Chilostigmini 12, 95 Chilostigmodes 12, 97, 126, 127 areolatus 127, 256, 257 forcipatus 126 Chimarra 10, 45, 46 aterrima 46 feria 46 obscura 46, 214, 215 socia 46 Chimarrinae 10, 42, 43, 45 Chyranda 12, 97, 122, 123 centralis 123, 253 Clistoronia 11, 97, 113–115 flavicollis 248 formosa 114 magnifica 114, 248 Clistoroniella 11, 114, 115 flavicollis 115 Clostoeca 12, 98, 123, 124 disjuncta 124, 254 sperryae 123 Clymene 36 aegerfasciella 36 Cochliopsyche 13 Cryptochia 11, 76, 78, 86, 87 furcata 87 pilosa 87, 234 Culoptila 9, 28, 29 aluca 29 cantha 29, 206 Cyllene 39 minutisimella 39

312

Cyrnellus 10, 61, 65 fraternus 65, 223 minimus 65 Dasystegia 164 Desmona 12, 97, 130, 131 bethula 130, 131 mono 131, 258 Dibusa 10 Dicosmoecinae 11, 74–77, 95 Dicosmoecus 11, 76–80, 82 atripes 79 canax 82 gilvipes 79, 226 obscuripennis 79 tristis 80 Dinarthrodes 149 Dinarthrum 149 Diplectrona 10, 57, 58 doringa 58 felix 57 modesta 58, 219 Diplectroninae 10, 52, 57 Dipseudopsidae 67 Dipseudopsis 67 Doloclanes 10 Dolophiliella 44 gabriella 44 Dolophilodes 10, 43, 44 distinctus 43 novusamericanus 214 ornatus 43 Dolophilus 44 copiosus 44 Drusinus 93 frontalis 81 uniformis 93 Ecclisocosmoecus 11, 77, 78, 89, 90 scylla 90, 237 spinosus 89 Ecclisomyia 11, 77, 78, 88, 89, 91 complicata 166 conspersa 88, 89, 236 maculosa 89 Ecnomidae 11, 67 Eobrachycentrus 13, 168–170 gelidae 169, 170, 281 Eocosmoecus 11, 77, 78, 81 frontalis 81 schmidi 81, 228 Eodinarthrum 149 Eomystra 25 dulkejti 25

313

Eremopsyche 151 frontalis 151 Eustenace 118 Fabria 13, 158, 159, 163, 164 inornata 164, 278 Farula 12, 135, 139, 140 jewetti 264 rainieri 139, 140 Fattigia 174 pelle 174 Flura 151 Frenesia 12, 96, 124, 125 difficilis 125 missa 125, 255, 257 Friga 190 Fumonta 10 Glossosoma 9, 21, 22, 24–26 alascense 207 boltoni 22 hoodi 207, 208 intermedium 206 lividum 207 parvulum 24 penitum 25, 207 Glossosomatidae 7, 9, 15, 21, 30 Glossosomatinae 9, 21, 22 Glyphopsyche 12, 96, 125, 126 bryanti 125 irrorata 125, 126, 255 missouri 126 Goera 12, 142–144 calcarata 144, 266 fuscula 144 stylata 144 tungusensis 144 Goeracea 12, 143–145 genota 145, 267 oregona 145 Goereilla 12, 146 baumanni 146, 268 Goeridae 4, 12, 14, 17, 142, 147 Goerinae 12, 142, 143 Goerita 12 genota 144 Grammotaulius 11, 96, 100, 101 betteni 100 brevilinea 101 interrogationis 100, 242 Grensia 12, 97, 127, 128 praeterita 127, 128, 256, 257 Gumaga 13, 174

Hagenella 13, 157–160 canadensis 160, 275 Halesochila 11, 96, 105, 107 taylori 106, 107, 244 Halesus maculipennis 115 magnificus 113 minutus 137 punctatissimus 90 Helicopsyche 13, 177, 178 borealis 178, 286 shuttleworthi 177 Helicopsychidae 4, 13, 16, 177 Helicopsychinae 177 Hesperophylax 11, 97, 121, 122 alaskensis 122 consimilis 122 designatus 122 incisus 253 occidentalis 122 Heteroplectron 14, 198, 199 americanum 199 borealis 195 californicum 198, 199, 296 Himalopsyche 9, 19, 20 phryganea 20, 206 Holocentropus 63 placidus 67 Homophylax 12, 96, 131, 132 acutus 132 andax 132, 259 baldur 132, 259 crotchi 132, 259 flavipennis 131, 132, 259 Homoplectra 10, 57–59 alseae 58 doringa 59 Hyalopsyche 67 Hyalopsychidae 4, 11, 15, 67 Hydatophylax 12, 96, 117–120 argus 117, 118, 250 hesperus 118 variabilis 118 victor 118 Hydrobiosidae 10 Hydropsyche 10, 51, 54–57 bronta 54 cinerea 54 flavomaculata 62 hyalinata 52 instabilis 54 lepida 56

occipitalis 44 recurvata 55, 218 Hydropsychidae 4, 10, 16, 48, 51, 52, 70 Hydropsychinae 10, 49, 52–54 Hydropsychoidea 6, 7, 10, 64, 157 Hydropsychoides 10 Hydroptila 9, 30, 33, 35 ampoda 210 angustella 36 costalis 37 spatulata 210 tineoides 35 Hydroptilidae 2, 7, 9, 14, 15, 27, 30–33 Hydroptilinae 9, 30–32 Hydroptilini 9 Imania 84 sichotalinensis 84 Integripalpia 6–8, 11 Ironoquia 11, 76, 78, 90, 91 parvula 91 punctatissima 91, 238 Ithytrichia 10, 33–35 clavata 35, 209 confusa 39 lamellaris 34 Jenortha 152 canadensis 152 Jyrvia 164 Klapalekia 25 Lenarchulus 107 Lenarchus 11, 12, 97, 108–112 crassus 111 expansus 111 keratus 110, 245 productus 111 rho 111, 246 vastus 246 Lepania 12, 145–147 cascada 147, 269 Lepaniinae 12, 143, 145 Lepaniini 145 Lepidostoma 12, 148–150, 154 americanum 271 cascadense 270, 274 latipenne 151, 271 pluviale 273, 274 podager 271 squamulosum 150 togatum 151, 270

314

vernale 272 wisconsinensis 152 Lepidostomatidae 4, 12, 17, 148, 149 Lepidostomatinae 12, 148 Leptocella 192 Leptoceridae 4, 13, 16, 181, 182, 193 Leptocerinae 13, 182 Leptocerini 13 Leptoceroidea 8, 13 Leptocerus 13, 181, 183, 189, 190 albidus 192 americanus 190, 291 bicolor 186 inconspicuus 190 ochraceus 190 Leptonema 10 Leptophylax 12, 96, 102, 103 gracilis 102, 103, 243 Leucotrichia 9, 33, 38, 39 melleopicta 38 pictipes 39, 212 Leucotrichiini 9 Limnephila scabripennis 118 Limnephilidae 4, 5, 11, 15, 74, 75, 98, 135 Limnephilinae 11, 75, 76, 90, 94, 95, 120 Limnephilini 11, 95 Limnephiloidea 8, 11 Limnephilus 12, 74, 97–100, 104, 111, 113, 121 difficilis 124 dispar 98 indistinctus 115 indivisus 241 nervosus 104 praeteritus 127 submonilifer 98, 241 vastus 111 Limnophilus 98 Lipoglossa 23, 24 Lype 11, 69–71 diversa 71, 224 Macronema 10 flavum 55 Macrostemum 10, 52, 53 zebratum 53, 217 Macrotaulius 12, 102 Madeophylax 86 altus 86

315

Manophylax 11, 77, 78, 85, 86 altus 86 annulatus 85, 86, 233 Marilia 14, 195–197 flexuosa 197, 295 major 196 Matrioptila 9 Mayatrichia 9, 33, 40, 41 ayama 40, 41, 213 Metrichia 9 Mexipsyche 10 Micrasema 13, 169, 172, 173 bactro 283 gelidum 172 rusticum 172, 283 Microtrichoptera 31 Molanna 14, 200–202 angustata 201 flavicornis 297 musetta 201 Molannidae 4, 14, 16, 200 Molannodes 14, 201, 202 tinctus 202, 298 zelleri 202 Monocosmoecus 82 Monophylax 130 mono 130 Mormomyia 12, 150, 152 togata 150 vernalis 152, 272 Mormonia 150 gracilicornis 150 Moselyana 11, 77 Muroglossa 23 Mystacides 13, 183–186 alafimbriata 185 interjecta 185 sepulchralis 185, 289 uwarowii 192 Mystacidini 13 Mystrophora 24 intermedium 24 Mystrophorella 25 Namamyia 14 Nectopsyche 13, 182, 183, 191, 193 albida 293 exquisita 293 Nectopsychini 13 Nemotaulius 12, 96, 101, 102 hostilis 101, 102, 242 Neodinarthrum 12, 150, 153 aporna 154

pluviale 154, 273, 274 rayneri 154 Neophryganea 166 Neophylacinae 75, 134, 135 Neophylax 12, 134–138 concinnus 136, 261 occidentis 261 oligius 261 rickeri 261 Neothremma 12, 135, 138–140 alicia 138, 139, 263 didactyla 139, 263 Neotrichia 9, 33, 39 collata 39 halia 212 okapa 212 Neotrichiini 9 Nepaloptila 28 Nerophilus 14 Neureclipsis 10, 61, 62 bimaculata 62, 221 crepuscularis 62, 221 valida 62 Neuronella 13, 161 Neuronia augustipennis 161 canadensis 160 inornata 163 vestita 164 Neuropsyche 151 tibialis 151 Nosopus 12, 150, 151 cascadense 270, 274 podager 151, 271 Notidobia distincta 174 Notiopsyche 150 carolina 150 latipennis 150 Nyctiophylax 65 Ochrotrichia 9, 33, 39, 40 insularis 39 potomus 40 stylata 40 tarsalis 40, 213 Ochrotrichiini 9 Odontoceridae 4, 14, 16, 194 Odontocerinae 14, 194 Odontocerum 194 Oecetina 190 fumosa 190 Oecetini 13

Oecetis 13, 183, 190, 191 incerta 190 inconspicusa 191, 292 persimilis 191 scala 190 Oecetodes 190 Oestropsinae 10, 52 Oestropsis 52 Olemira 151 americana 151 mexicana 152 pluviale 153 Oligophlebodes 12, 135, 137, 138 coloradensis 137 minutus 262 mostbento 262 sierra 262 Oligophorae 74, 75 Oligoplectrodes 13, 170, 171 potanini 170, 171 Oligoplectrum morosum 172 Oligopsyche 150 Oligostomis 13, 158–160 ocelligera 160, 275 pardalis 160 Oligotricha 13, 158, 159, 162 chloroneura 162 lapponica 162, 277 Onocosmoecus 11, 76–78, 80, 81 unicolor 80, 227 Oropsyche 10 Orthotrichia 10, 33, 36 aegerfasciella 36 cristata 36, 210 curta 36 Orthotrichiini 10 Oxyethira 9, 33, 37, 38 serrata 211 Paduniella 11 Paduniellinae 11, 70 Palaeagapetus 9, 30, 31 celsus 31, 206 finisorientis 206 guppyi 31 nearcticus 31 rotundatus 30 Parachiona pilosa 86 Paragapetus 44 moestus 44 Paralenarchus 11, 109, 111, 112

316

Paranyctiophylax 10, 60, 61, 64 kisoenis 64 moestus 222 Parapsyche 10, 48–50 almota 50 apicalis 50 elsis 50, 216 Parthina 14 Paucicalcaria 9 Pedomoecus 11, 77, 78, 83, 84 sierra 83, 84, 231 Phanocelia 12, 96, 133 canadensis 133, 260 Phanopsyche 152 grisea 152 Philarctus 12, 96, 112, 113 bergrothi 112 przewalskii 113 quaeris 113, 247 Philocasca 12, 120, 121 alba 121 antennata 121 banksi 121 demita 120, 121, 251 thor 121, 251 Philopotamidae 4, 15, 42 Philopotaminae 10, 42, 43 Philopotamus 42, 43 Phryganea 13, 156–158, 165, 166 arga 117 bimaculata 61 cinerea 166, 279 dossuaria 162 grandis 165 hirta 150 interrogationis 100 interrupta 189 irrorata 125 lapponica 103 marginata 45 nervosa 183 nigra 184 obsoleta 164 pilosa 143 radiata 115 reticulata 159 rhombica 98 sayi 166 semifasciata 161 striata 162 subfasciata 118 tincta 202

317

trimaculata 107 vestita 92 viridis 188 waeneri 71 Phryganeidae 4, 13, 15, 156, 157 Phryganeinae 13, 157 Phryganoidea 8, 13 Phryganomyia 164 Phylloicus 14 Phylocentropus 11, 67, 68 carolinus 68 lucidus 68, 224 placidus 68, 224 Platycentropus 12, 96, 115, 116 amicus 116 indistinctus 116 radiatus 116, 249 Platyphylax occidentalis 121 Plectrocnemia 63 Plectropsyche 10 Polycentropodidae 4, 10, 15, 60, 61, 67, 69, 70 Polycentropus 10, 60–63 aureolus 63 cinereus 63 confusus 63 interruptus 63, 221 irroratus 62 lucidus 68 remotus 63 variegatus 63, 221 Polyphorae 74, 75 Polyplectropus 10 Polytrichia 39 Potamyia 10, 54–56 flava 56, 218 Pristosilo 150 canadensis 150 Prolenarchus 12, 109, 110 keratus 110 Prophryganea 164 principalis 164 Protoptila 9, 27–29 erotica 208 maculata 208 tenebrosa 208 Protoptilinae 9, 21, 22, 27 Pseudogoera 14 Pseudogoerinae 14, 194 Pseudoleptocerus 184 Pseudoneureclipsinae 61

Pseudostenophylacinae 11, 75, 76, 93 Pseudostenophylax 11, 93, 94 edwardsi 94 fumosus 93 sparsus sparsus 94, 240 Psiloneura 13, 174, 175 moesta 174 Psilotreta 14, 195, 196 frontalis 195, 196 indecisa 195, 196, 294 labida 196 rufa 196 Psychoglypha 12, 97, 121, 129, 130 alascensis 257 avigo 129 bella 129 mono 130 subborealis 130 Psychomyia 11, 69, 70, 72, 73 annulicornis 72 flavida 72, 73, 225 lumina 73 nomada 73 pusilla 72 Psychomyiidae 11, 15, 61, 67–70 Psychomyiinae 11, 70 Psychoronia 12 Ptilocolepinae 9, 30 Ptilocolepus 30 Ptilostomis 13, 158, 159, 161 angustipennis 161 kovalevskii 161 ocellifera 161 postica 161 semifasciata 161, 276 Pycnopsyche 12, 97, 118–120 guttifer 119 subfasciata 119, 251 Quaria 190 Quisilo 154 Quissa 72 Radema 92 Rheophylax 9, 98 Rhyacophila 9, 18–20, 22, 23, 32 acropedes 205 tibetana 20 vocala 205 vulgaris 19 Rhyacophilidae 4, 7, 9, 15, 18, 19, 21 Rhyacophiloidea 6, 7, 9 Rhyacophylax 10

Ripaeglossa 9, 23, 24 Rossiana 11, 77, 78, 87, 88 montana 87, 88, 235 Sericostoma 174 Sericostomatidae 4, 13, 16, 174 Sericostomatoidea 8, 13 Sericostriata 12, 135, 140, 141 surdickae 140, 141, 265 Setodes 14, 183, 188, 189 americanus 189 avarus 190 cinerascens 190 gemma 191 guttatus 189 immobilis 190 incertus 189 oligius 189, 291 punctella 188 Setodina 190 parva 190 Setodini 14 Silo griseus 154 Sisko 10 Smicridea 10 Sphagnophylax 12, 96, 119, 120 meiops 119, 120, 252 Sphinctogaster 13, 170, 171 lateralis 171 lutescens 170 Stactobia palmata 36 ulmeri 36 Stactobiella 10, 33, 36, 37 delira 37, 211 palmata 37 Stactobiini 10 Stenophylacini 12, 95 Stenophylax infumatus 117 limbatus 118 minusculus 98 palatus 78 parvula 98 Sychnothrix 13 Synafophora 9, 23–25 intermedium 25 lividum 25 minutum 24 nigrior 25 verdona 25 Tascobia 36

318

Thamastes dipneumus 4 Theliopsyche 13, 149, 154, 155 grisea 155, 274 parva 154, 155 Theliopsychinae 13, 154 Thremmatinae 135 Thya maurus 179 Tinodes 11, 69–72 cascadius 225 lurida 71 provo 72, 225 Toluaca 28 Trentonius 44 distinctus 44 Triaena 186 Triaenodella 186, 187 chelifera 186 Triaenodes 14, 181–183, 186–188 aba 186 grisea 187 injustus 186, 290 marginatus 186, 290

319

Triaenodini 14 Triplectides 14 Triplectidinae 14, 182 Uenoa 134 Uenoidae 12, 14, 75, 134, 135 Uenoinae 134 Wormaldia 10, 43–45 gabriella 45, 215 occidea 215 Xiphocentronidae 67 Ylodes 14, 182, 187, 188 frontalis 188 griseus 290 kaszabi 188 reuteri 188 schmidi 188 Ymymia 189 Yphria 13 Yphriinae 13, 157 Yrula 190 Zaporota 98 pallens 98 Zumatrichia 9