Caddisflies of the Yukon - Department of Biological Sciences ...

Caddisflies of the Yukon 

FRONTISPIECE. Larva and case of Sphagnophylax meiops Wiggins and Winchester (Limnephilidae). This monotypic 

genus is a phylogenetic and geographic relict, known only from transient tundra pools in East Beringia. Length of 

mature larva 14 mm and of case 17 mm. 

787

Caddisflies (Trichoptera) of the Yukon, 

with Analysis of the Beringian and 

Holarctic Species of North America 

GLENN B. WIGGINS and CHARLES R. PARKER 

Centre for Biodiversity and Conservation Biology (formerly Department of Entomology) 

Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada M5S 2C6; 

and Department of Zoology, University of Toronto 

Current address of C.R. Parker: 

U.S. Geological Survey Biological Resources Division, Great Smokies Field Station 

1314 Cherokee Orchard Road, Gatlinburg, Tennessee 37738, U.S.A. 

Abstract. The Trichoptera recorded from the Yukon Territory now number 145 species, constituting 11 per cent 

of the North American fauna north of Mexico. Present distribution known for each species in the Yukon is outlined, 

and biological information at familial and generic levels is briefly summarized. For biogeographic analysis, evidence 

bearing on the distribution of the species is considered under 4 categories. Members of category I are wholly Nearctic 

in distribution (98 species, 68 per cent of Yukon Trichoptera) and, in the absence of evidence to the contrary, are 

considered to have repopulated the Yukon and other northern areas from glacial refugia to the south of the Laurentide 

and Cordilleran ice sheets of Wisconsinan time. 

Species of category II are Holarctic, and are now more or less widely distributed in Eurasia and northern North 

America (28 species, about 18 per cent of Yukon Trichoptera). These species could have passed the last glacial 

period in unglaciated Beringia, or to the south of the ice, or in both areas. 

Category III is composed of Palaearctic species which, from evidence available, are now confined in North 

America mainly to unglaciated Beringia or somewhat beyond (13 species, about 10 per cent of Yukon Trichoptera). 

Several represent a paradox of Beringian distribution—widely distributed Palaearctic species, evidently successful 

colonizers when they entered North America but, with retreat of the ice, have not extended their Nearctic range. 

Geological and biological factors underlying this paradox are discussed. Two Palaearctic species are recorded from 

North America for the first time: Rhyacophila mongolica Schmid, Arefina and Levanidova and Limnephilus diphyes 

McLachlan. 

Category IV comprises 8 species (including 2 additional species expected from the Yukon), about 4 per cent of 

the fauna, known mainly from the Yukon or from adjacent areas of Alaska or the Northwest Territories; these 

species are considered to be Beringian endemics or glacial relicts. Finally, because almost all of the Holarctic 

Trichoptera now recognized in North America are reviewed in the foregoing groups, the remaining Holarctic species 

that do not occur in Beringia are considered briefly in a fifth category, although they have not been recorded from 

the Yukon and most do not appear to be species of far northern latitudes. The origin of the Trichoptera of Greenland 

is also discussed. 

Ecological factors influencing the northern penetration of Yukon and Beringian Trichoptera are considered with 

an analysis of lotic and lentic-dwelling species through a latitudinal gradient of 49° to 70°N—from the southern 

border of British Columbia to the Arctic coastline of the Yukon. At latitude 60°N, the southern boundary of the 

Yukon, diversity has declined by almost 50 per cent from levels obtaining in British Columbia, 49° through 60°N. 

The main depletion occurs in the Spicipalpia and filter-feeding Annulipalpia; case-making caddisflies of the 

Integripalpia show less reduction. Similar trends are continued through the Yukon from 60° to 70°N, where species 

diversity in the Trichoptera declines by another 59 per cent. Although most North American Trichoptera occur in 

running waters, there is a marked reduction of species in these habitats with increasing latitude. Of 60 species 

recorded in the Yukon north of the Arctic Circle (67° – 70°N), 81 per cent are Integripalpia with case-making larvae 

living mainly in lentic habitats. Factors underlying the decline of lotic species, and the proportional increase of 

lentic species at higher latitudes are considered. Trichoptera of lentic habitats were much more successful in crossing 

the Bering land bridge than were species dependent on lotic waters. 

Taxonomic changes resulting from this study include suppression of Grammotaulius subborealis Schmid as a 

junior subjective synonym of G. alascensis Schmid. The status of Limnephilus fumosus Banks is clarified as a 

species distinct from Limnephilus santanus Ross, and a lectotype is designated for L. fumosus; L. isobela Nimmo 

is recognized as a junior subjective synonym of L. fumosus Banks. Goera radissonica Harper and Méthot, described 

from northern Quebec, is recognized as a junior subjective synonym of Goera tungusensis Martynov, originally 

described from Siberia. A morphological variant of Ceraclea nigronervosa (Retzius) is described. The distributional 

pp. 787 – 866 in H.V. Danks and J.A. Downes (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), 

Ottawa. 1034 pp. © 1997

Caddisflies of the Yukon 789 

and taxonomic status of Mystacides interjectus (Banks) and M. sepulchralis (Walker) is reviewed and clarified. 

This study provides a taxonomic and conceptual framework for further investigation of the Holarctic Trichoptera. 

Résumé. Les trichoptères (Trichoptera) du Yukon: inventaire et analyse des espèces béringiennes et holarctiques 

d’Amérique du Nord. Les trichoptères du Yukon comptent maintenant 145 espèces connues, soit 11 pourcent de la 

faune nord-américaine au nord du Mexique. On trouvera ici un aperçu de la répartition de chacune de ces espèces 

au Yukon et un sommaire de certains aspects de la biologie des familles et des genres. Une analyse biogéographique 

de la répartition des espèces a donné lieu à quatre grandes catégories: les membres de la catégorie I ont une répartition 

essentiellement néarctique (98 espèces, 68% des trichoptères du Yukon) et, jusqu’à preuve du contraire, sont 

estimées avoir recolonisé le Yukon et les autres zones nordiques à partir de refuges glaciaires situés au sud des 

glaciations laurentidiennes et cordillériennes au cours du Wisconsinien. 

Les espèces de la catégorie II sont holarctiques et sont maintenant généralement bien répandues en Eurasie et 

dans le nord de l’Amérique du Nord (28 espèces, environ 18% des trichoptères du Yukon). Ces espèces ont 

probablement passé la dernière période glaciaire dans la partie non englacée de la Béringie, ou alors au sud des 

glaces, ou ont occupé les deux endroits. 

La catégorie III se compose d’espèces paléarctiques qui semblent confinées, en Amérique du Nord, à la partie 

non englacée de la Béringie ou un peu au-delà (13 espèces, environ 10% des trichoptères du Yukon). Plusieurs ont 

une répartition béringienne un peu énigmatique—ce sont des espèces paléarctiques bien répandues qui ont colonisé 

l’Amérique du Nord avec succès, mais qui, au retrait des glaces, ne se sont pas répandues davantage dans la zone 

néarctique. Les facteurs géologiques et biologiques qui pourraient expliquer ce paradoxe font l’objet d’une 

discussion. Deux espèces paléarctiques sont mentionnées en Amérique du Nord pour la première fois, Rhyacophila 

mongolica Schmid, Arefina et Levanidova et Limnephilus diphyes McLachlan. 

La catégorie IV comprend 8 espèces (dont 2 qui n’ont pas encore été trouvées au Yukon), soit environ 4% de 

la faune, connues surtout au Yukon ou dans les zones adjacentes en Alaska et dans les Territoires du Nord-Ouest; 

ces espèces sont considérées comme endémiques en Béringie ou comme des espèces relictes des glaciations. Enfin, 

comme la plupart des trichoptères holarctiques reconnus en Amérique du Nord appartiennent aux catégories 

précédentes, les autres espèces holarctiques qui n’ont jamais été trouvées en Béringie sont examinées brièvement 

et forment une cinquième catégorie d’espèces jamais rencontrées au Yukon et dont la plupart ne semblent pas être 

des espèces très nordiques. L’origine des trichoptères du Groenland est également examinée. 

Les facteurs écologiques qui ont pu influencer la dispersion vers le nord des trichoptères du Yukon et de la 

Béringie sont étudiés et une analyse des espèces lotiques et lénitiques présentes le long d’un gradient latitudinal du 

49 e au 70 e parallèle, du sud de la Colombie-Britannique à la côte arctique du Yukon, donne un aperçu global de la 

situation. A la latitude 60°N, le long de la frontière australe du Yukon, la diversité est déjà réduite de près de 50% 

par rapport à la situation qui prévaut en Colombie-Britannique, soit entre les latitudes 49°N et 60°N, diminution 

qui affecte surtout les Spicipalpia et les Annulipalpia filtreurs; les Integripalpia constructeurs de fourreaux sont 

encore présents en assez grand nombre. La tendance se poursuit vers le nord, entre les parallèles 60 et 70, et la 

diversité est réduite d’un autre 59%. Bien que la plupart des trichoptères nord-américains soient des espèces d’eau 

courante, le nombre de ces espèces diminue à mesure que la latitude augmente. Des 60 espèces rencontrées au 

Yukon au nord du cercle arctique (67° – 70°N), 81% sont des Integripalpia dont les larves vivent dans des fourreaux 

en milieu lénitique. Les facteurs susceptibles d’expliquer le déclin des espèces lotiques et l’augmentation proportionnelle 

des espèces lénitiques aux latitudes plus élevées sont examinés. Les trichoptères des milieux lénitiques 

semblent avoir réussi à traverser le pont continental de Bering plus facilement que les espèces des milieux lotiques. 

Cette étude a donné lieu à certains remaniements taxonomiques: Grammotaulius subborealis Schmid est 

considéré comme un synonyme subjectif récent de G. alascensis Schmid. Le statut de Limnephilus fumosus Banks 

est redéfini et l’espèce est distincte de Limnephilus santanus Ross; un lectotype a été désigné pour représenter 

L. fumosus; L. isobela Nimmo est reconnu comme un synonyme subjectif récent de L. fumosus Banks. Goera 

radissonica Harper et Méthot, décrit du nord du Québec, est reconnu comme un synonyme subjectif récent de Goera 

tungusensis Martynov d’abord trouvé en Sibérie. Une variante morphologique de Ceraclea nigronervosa (Retzius) 

est décrite. La répartition et le statut taxonomique de Mystacides interjectus (Banks) et de M. sepulchralis (Walker) 

sont révisés et clarifiés. Ce travail a permis d’établir un nouveau cadre de recherche taxonomique et conceptuel 

pour l’étude des trichoptères holarctiques. 

Table of Contents 

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 790 

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 791 

Annotated species list of the Yukon Trichoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793 

Suborder Spicipalpia 

Family Glossosomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793 

Family Hydroptilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793 

Family Rhyacophilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 794

790 G.B. Wiggins and C.R. Parker 

Table of Contents (continued) 

Suborder Annulipalpia 

Family Hydropsychidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 796 

Family Philopotamidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 797 

Family Polycentropodidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 798 

Suborder Integripalpia 

Family Apataniidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 799 

Family Brachycentridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 799 

Family Goeridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 800 

Family Lepidostomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 800 

Family Leptoceridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 801 

Family Limnephilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 803 

Family Molannidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 809 

Family Phryganeidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 809 

Family Uenoidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 811 

Taxonomic notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 811 

Biogeographic analysis of the Yukon and Beringian Trichoptera . . . . . . . . . . . . . . . . . . . . . . . . . 820 

Geological and climatic context . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 820 

Biological aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 821 

Origin of the Beringian and Holarctic Trichoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824 

I. Nearctic species widespread in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 825 

Greenland Trichoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 828 

II. Holarctic species widespread in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 830 

III. Palaearctic-East Beringian species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 839 

IV. Beringian species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 849 

V. Holarctic species not in Beringia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 852 

Ecological considerations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 854 

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 861 

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 862 

Introduction 

In this study we have undertaken to examine the Trichoptera of the Yukon Territory of 

northern Canada from a dynamic viewpoint. Several lines of biological investigation 

coincide to make the Yukon especially appropriate for such an analysis. 

Repopulation of the Yukon following the last recession of Pleistocene glaciers is of 

unusual biogeographic interest because the western half of this area was part of the 

unglaciated Beringian refugium that extended westward through central Alaska, over the 

exposed continental shelf underlying Bering Strait and adjacent coastal areas, and incorporating 

a large part of northeastern Asia. While northern North America and extensive areas 

of Europe and Asia were covered by ice during the last glacial (Wisconsinan) advance, the 

unglaciated Beringian refugium harboured cold-adapted species, enabling some of them to 

move from one continent to the other across the Bering land bridge connecting North 

America and Asia. Evidence indicates that a number of the species in the present trichopteran 

fauna of the Yukon entered North America in this way and passed the Pleistocene glacial 

periods in the Beringian refugium, while most Nearctic species were confined to the south 

of the advancing front of the Laurentide continental and Cordilleran montane glaciers. 

From an ecological viewpoint, an investigation of Yukon Trichoptera offers an opportunity 

to contrast the ecological success of a diverse group of aquatic insects at high latitudes 

with the success of the same group in more temperate parts of North America. The species 

advancing from southern refugia as the glacial ice receded had to contend with different 

habitat conditions at higher latitudes. For terrestrial insects, evidence on the ecological costs


imposed by these conditions is available from a number of groups; but for aquatic insects 

other than Chironomidae (e.g. Oliver 1968) there is little evidence or synthesis. To what 

extent did longer colder winters, shorter summers, and shorter photoperiods influence the 

success of a group of insects with wholly aquatic larvae? Trichoptera are especially 

appropriate for seeking answers to questions of this kind because of their relatively high 

diversification; apart from Chironomidae, there are more species of Trichoptera than of any 

other group of freshwater insects, and those species occupy an exceptionally wide range of 

aquatic habitats and ecological niches (Wiggins and Mackay 1978). 

Combining both biogeographic and ecological viewpoints, investigation of the Yukon 

Trichoptera reveals results in nature when related species of Nearctic and Palaearctic origins 

come together to form aquatic communities. As an ecological testing ground for these natural 

experiments, the Yukon is well suited because of its high diversity of aquatic habitats. 

Mountain ranges divide the land into several major drainage systems, giving rise to small 

rapid streams, which in turn unite to form river systems of increasing order and potential 

biological diversity. Marshes, ponds, and lakes abound, providing rich resources for aquatic 

insects adapted for life in lentic waters. The treeline, southern margin of the arctic biome, 

passes through the northern Yukon at about latitude 67°N. To the north, arctic tundra extends 

to the Arctic Ocean; on the slopes of the mountains, biotic zones range through coniferous 

forests to alpine tundra and ice-fields. Although freshwater habitats are highly diverse within 

the Yukon, the success of insects in forming aquatic communities under climatic conditions 

of high latitudes is not well understood. Therefore, examination of these fundamental 

biological issues can add to our understanding of the biology of Trichoptera; the same issues 

underlie management of aquatic systems in the Yukon Territory. 

The analysis begins, necessarily, with a survey of the species of Trichoptera known to 

occur in the Yukon. Systematic interpretation of the species has been aided by the advanced 

state of knowledge on the Trichoptera of Russia (e.g. Lepneva 1964, 1966; Martynov 1924a), 

and by the recent synthesis of aquatic insects of the Russian Far East by I.M. Levanidova 

(1982). 

Materials and Methods 

Most of the collections of Trichoptera studied were made by field parties from the 

Department of Entomology, Royal Ontario Museum (ROME), over a 4-year period 

from 1979 to 1982, and are deposited there. Collections from other institutions were also 

studied: Canadian National Collection of Insects, Ottawa (CNCI); Illinois Natural History 

Survey, Champaign, Illinois (INHS); Royal British Columbia Museum (BCPM); 

U.S. National Museum of Natural History (USNM); University of British Columbia Insect 

Collection, Vancouver (SMDV); Zoological Institute, Russian Academy of Sciences, 

St. Petersburg (ZMAS). 

All species known from the Yukon Territory are listed, including species recorded in 

the scientific literature but not represented in the material we examined—principally records 

compiled by Nimmo and Wickstrom 1984 (henceforth NW 1984). Although we have many 

larval collections, these records are included only if larvae are the single source of evidence, 

as for Allomyia, or can be identified reliably to species as in some Hydropsyche. Distributional 

records are grouped by ecogeographic regions of the Yukon (Fig. 1); the detailed list 

of our records is too long for inclusion here, but is deposited in the library of the Royal 

Ontario Museum. At this early stage in understanding the distribution of Trichoptera in the 

Yukon, the records are highly correlated with the access roads; however, some general 

patterns seem to emerge. Biological and distributional characteristics of the higher taxa are


FIG. 1. Ecogeographic regions of the Yukon Territory (after Scudder 1997); collection records of Yukon Trichoptera 

are summarized in accordance with these numbered regions. 1, Arctic Coastal Plain; 2, British Mountains; 3, Arctic 

Plateau; 4, Porcupine Plain (including Old Crow Plain, Old Crow Mts., N. Porcupine Plateau); 5, Richardson 

Mountains; 6, Eagle Plain (including S. Porcupine Plateau); 7, Peel Plateau (including Bonnet Plume Basin); 8, 

Ogilvie Mountains (including N. and S. Ogilvie Mts.); 9, Wernecke/Selwyn Mountains; 10, Yukon/Tintina 

(including Lewes Plateau, part of Klondike Plateau, and Tintina Trench); 11, Eastern Plateaus (including Stewart, 

Macmillan, and Pelly Plateaus); 12, Shakwak Trench (including Wellesley Basin); 13, Western Ranges (including 

Ruby, Nisling, and Dawson Ranges, part of Klondike Plateau); 14, Pelly Mountains; 15, Logan Mountains; 16, 

Saint Elias/Coast Mountains; 17, Southern Lakes (including Aishihik Basin, Takhini Valley, Teslin Plateau, and 

Nisutlin Plateau); 18, Cassiar Mountains; 19, Liard Plain (including Dease Plateau); 20, Hyland/Liard Plateaus.


outlined briefly to provide a broader context for the Yukon species; distributional information 

for North America is based on the manuscript for an Annotated Catalogue of the 

Trichoptera of North America North of Mexico (Wiggins and Flint in prep.), and for Europe 

on BotojAneanu and Malicky (1978). Extended comment required on the status of certain 

species is included under Taxonomic Notes. Families, genera, and species are listed alphabetically 

under the 3 suborders of Trichoptera proposed by Wiggins and Wichard (1989) 

and Frania and Wiggins (1997). The classification of families and genera follows Wiggins 

(1996). Roman numerals following the names indicate the category to which the species is 

assigned for biogeographic analysis. Arabic numbers associated with the species names 

provide a cross-reference to distributional and other data in the annotated species list. Dates 

for collections of adults are based on material examined and on published records. 

Annotated Species List of the Yukon Trichoptera 

This list records 145 species, mainly from specimens we examined, but also from 

literature records where no material was available. Additional species undoubtedly have yet 

to be found. Distributional records for the Yukon are grouped by ecogeographic regions as 

identified in Fig. 1. 

Suborder Spicipalpia 

These are the cocoon-making caddisflies, characterized by pupal cocoons of stout silk 

lacking openings of any kind for circulation of water over the pupa. They are for the most 

part inhabitants of cool running waters, although larvae of some genera of the Hydroptilidae 

are adapted to warmer lentic sites. 

Family Glossosomatidae 

Larvae occur on rocks in flowing waters where they graze on diatoms, other algae, and 

deposits of fine organic particles. The family is represented in most faunal regions of the 

world; 6 Nearctic genera are recognized, comprising approximately 75 species. 

Genus Glossosoma. Species occur through the Nearctic, Palaearctic, and Oriental faunal 

regions; of 22 North American species now known, all but 3 are confined to the western 

mountains. Three species are recorded from the Yukon, representing 2 of the 3 Nearctic 

subgenera: G. (Ripaeglossa) alascense; G. (Synafophora = Eomystra) intermedium and verdona. 

1. Glossosoma alascense Banks (I) Nearctic, western montane 

Distribution: Yukon, Alaska, south to Utah. 

Yukon records: 8, 10, 12 (ROME); 16, 19 (NW 1984). 

Biological information: Adults have been collected in the Yukon 23 June – 9 August. 

2. Glossosoma intermedium (Klapalek) (II) Holarctic, transcontinental 

Distribution: Yukon to Montana, Quebec, Tennessee; central Europe to Finland and the northern part 

of European Russia, eastward through Siberia to Chukotka and Kamchatka (Levanidova 1975, 1982). 

Yukon records: 5, 10, 12, 19 (ROME). 

Biological information: Adults have been collected in the Yukon 23 May – 9 August. 

3. Glossosoma verdona Ross (I) Nearctic, western montane 

Distribution: Yukon, Alaska, south to California and Utah. 

Yukon records: 8, 17 (SMDV); 10 (ROME). 

Biological information: Adults have been collected in the Yukon 31 May – 28 June. 

Family Hydroptilidae 

Hydroptilidae are widely distributed throughout the world with genera characteristic of 

all types of fresh waters from cold springs to lakes. Larvae feed principally on algae,


especially filamentous forms. At least 5 of the 16 North American genera are represented in 

the Yukon, each by a single species. 

Genus Agraylea. This is an Holarctic genus with 4 North American species; larvae live in 

standing waters of lakes, and areas of reduced current in streams. 

4. Agraylea cognatella McLachlan (III) Palaearctic-East Beringian 

Distribution: Yukon; northern Europe (BotojAneanu and Malicky 1978); Russian Far East (Levanidova 

1975; BotojAneanu and Levanidova 1988). 

Yukon records: 4, 5, 11 (ROME). 

Biological information: Adults have been collected in the Yukon 5 – 23 July. 

Taxonomic notes: North American specimens recorded as A. multipunctata Curtis (e.g. NW 1984, YT: 

16, 17) will have to be re-examined because that species appears to be restricted to Europe and Asia 

(Vineyard and Wiggins in prep.). 

Genus Hydroptila. Representatives of this genus occur through much of the world, including 

approximately 90 species in North America alone; larvae occur in lakes and in flowing 

waters. 

5. Hydroptila rono Ross (I) Nearctic, transcontinental 

Distribution: Yukon to California, Quebec, and Pennsylvania. 

Yukon records: 17 (ROME). 

Biological information: Adults have been collected in the Yukon 11– 26 July. 

Genus Ochrotrichia. This genus is confined to the New World, where approximately 50 

species are known in North America. Larvae occur in running-water habitats. An unidentified 

species was recorded from the Yukon. 

6. Ochrotrichia sp. 

Distribution: Unknown. 

Yukon records: 16 (NW 1984). 

Biological information: Adults have been collected in the Yukon 26 July. 

Genus Oxyethira. Occurring widely throughout the world, this genus includes approximately 

40 North American species; larvae live in beds of aquatic plants in lakes and slow 

rivers. 

7. Oxyethira araya Ross (I) Nearctic, transcontinental 

Distribution: Yukon, Minnesota to Maine; continued absence of records between Yukon and Minnesota 

raises the possibility that these populations are disjunct, perhaps reflecting their separation during 

glaciation. 


Biological information: Adults have been collected in the Yukon 20 June –12 July. 

Genus Stactobiella. This is a small Holarctic genus with 6 North American species; larvae 

live in small, rapid streams. 

8. Stactobiella delira (Ross) (I) Nearctic, transcontinental 

Distribution: Yukon to California, Maine, and Tennessee, most of North America, but not recorded 

from central or eastern Canada. 



Family Rhyacophilidae 

Larvae are confined to cool, running waters, and for the most part are predacious on 

other insects. The family occurs in all continents of the northern hemisphere except Africa. 

Only 2 genera are recognized, and both occur in North America.


Genus Rhyacophila. This is a genus of at least 500 species, the largest in the Trichoptera. 

More than 100 species occur in North America, chiefly in the western mountains; 14 species 

have been recorded in the Yukon. Evidence shows that life cycles of Rhyacophila species 

at high latitudes tend to be longer than the single year in temperate latitudes (Irons 1988). 

9. Rhyacophila alberta Banks (I) Nearctic, western montane 

Distribution: Yukon, Alaska to New Mexico. 


Biological information: Adults have been collected in the Yukon 7 August. 

10. Rhyacophila angelita Banks (I) Nearctic, disjunct 

Distribution: Yukon to California, New Mexico; Minnesota, Quebec, New Hampshire. Distributional 

records indicate that the eastern and western populations of this species may be disjunct. 

Yukon records: Nimmo (1971). 

Biological information: None available. 

11. Rhyacophila bifila Banks (I) Nearctic, western montane 

Distribution: Yukon to California. 



12. Rhyacophila brunnea Banks (I) Nearctic, transcontinental 

Distribution: Yukon to northeastern North America, south to New Mexico and California. 

Yukon records: 10, 16 (ROME); 12, 17, 19 (NW 1984). 


Taxonomic notes: The present concept for this species (Smith and Manuel 1984) subsumes 

R. acropedes Banks, R. ignorata Schmid, and R. acuminata Fields. 

13. Rhyacophila hyalinata Banks (I) Nearctic, western montane 

Distribution: Yukon, Alaska to California and New Mexico. 

Yukon records: 12, 17, 19 (NW 1984); 16 (ROME). 


14. Rhyacophila mongolica Schmid, Arefina and Levanidova (III) Palaearctic-East Beringian 

Distribution: Previously known only from Mongolia and the Russian Far East, this species is recorded 

from North America for the first time. 



Taxonomic notes: See Taxonomic Note 1. 

15. Rhyacophila narvae Navas (II) Holarctic, western montane 

Distribution: Subsumes the western Nearctic R. vepulsa Milne (Schmid 1970), extending its distribution 

from the Russian Far East to California. 


Biological information: Adults have been collected in the Yukon 25 June. 

16. Rhyacophila pellisa Ross (I) Nearctic, western montane 

Distribution: Yukon to California and Utah. 

Yukon records: 10, 16 (ROME); 19 (SMDV); 12 (NW 1984). 

Biological information: Adults have been collected in the Yukon 24 March – 28 August. 

17. Rhyacophila tucula Ross (I) Nearctic, western montane 

Distribution: Yukon, Alaska to Utah and Colorado. 


Biological information: Adults have been collected in the Yukon 10 –12 August. 

18. Rhyacophila vao Milne (I) Nearctic, western montane 

Distribution: Yukon, Alaska to Utah and Montana. 

Yukon records: 17, 19 (NW 1984). 

Biological information: Adults have been collected in the Yukon 23 July – 8 August.


19. Rhyacophila verrula Milne (I) Nearctic, western montane 

Distribution: Yukon, Alaska to California and New Mexico. 

Yukon records: 11, 17 (SMDV). 

Biological information: Adults have been collected in the Yukon 23 July – 3 August. 

20. Rhyacophila vobara Milne (I) Nearctic, western montane 

Distribution: Yukon, Alaska to Oregon and Montana. 


Biological information: Adults have been collected in the Yukon 23 June – 27 July. 

21. Rhyacophila vocala Milne (I) Nearctic, western montane 

Distribution: Yukon to Montana and California. 

Yukon records: 10 (CNCI); 16 (ROME); 17 (SMDV). 


22. Rhyacophila vofixa Milne (I) Nearctic, western montane 

Distribution: Yukon, Alaska to Utah 

Yukon records: 8 (ROME); 12, 16, 17, 19 (NW 1984). 

Biological information: Adults have been collected in the Yukon 23 July –1 August. 

Suborder Annulipalpia 

These are the retreat-making or net-spinning caddisflies whose larvae are concealed in 

fixed tubular retreats or nets on rocks, logs, and plants. Most of the larvae live in running 

waters; some construct fine-meshed nets of silk to strain suspended food materials from the 

current, others graze fine organic particles or prey on insects. Pupation occurs in open 

perforate cells, with water circulating through the cell around the pupa. 

Family Hydropsychidae 

Hydropsychids are the dominant caddisflies of running waters over much of North 

America, both in species diversity and in biomass, but the family is markedly reduced in the 

Yukon. The family is widespread throughout the world; 11 genera with approximately 150 

species are represented in North America, but only 4 genera are known in the Yukon. Larvae 

construct nets of silken meshes which filter suspended particles and insects from the current, 

the size of the mesh differing in most genera. 

Genus Arctopsyche. Species of Arctopsyche occur through much of the Holarctic and 

Oriental regions; 4 are known in North America. Larvae construct coarse-meshed filter nets 

and are primarily insectivorous. 

23. Arctopsyche grandis (Banks) (I) Nearctic, disjunct 

Distribution: Yukon to California, with a disjunct occurrence in Quebec. 



24. Arctopsyche ladogensis (Kolenati) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland and New Hampshire; a record from Utah (Baumann 

and Unzicker 1981) leaves a large gap to the Yukon in the recorded distribution of this species in 

western North America; northern Europe and Asia through Siberia to Mongolia and Kamchatka 

(Lepneva 1964), but not recorded from Chukotka (Levanidova 1982). 

Yukon records: 4, 12, 17 (ROME); 10 (NW 1984). 

Biological information: Adults have been collected in the Yukon 26 June –14 August. 

Genus Cheumatopsyche. Some 40 species of this genus are known in North America, and 

the group is widespread through most other faunal regions.


25. Cheumatopsyche campyla Ross (I) Nearctic, transcontinental 

Distribution: Yukon to California, Newfoundland, Alabama. 



26. Cheumatopsyche sp. female (not C. campyla) 

Distribution: Unknown. 



Genus Hydropsyche. This is the dominant North American genus of the family with more 

than 70 Nearctic species, but only 5 are represented in the Yukon. 

27. Hydropsyche alhedra Ross (I) Nearctic, transcontinental 

Distribution: Yukon to Massachusetts and North Carolina. 



28. Hydropsyche alternans (Walker) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska and British Columbia to Newfoundland and Massachusetts. 

Yukon records: 4, 10 (NW 1984); 16, 17 (ROME). 


29. Hydropsyche amblis Ross (I) Nearctic, western montane 

Distribution: Yukon to Oregon. 

Yukon records: 17, 19 (ROME). 

Biological information: Adults have been collected in the Yukon 23 May –17 June. 

30. Hydropsyche cockerelli Banks (I) Nearctic, western montane 


Yukon records: 17 (SMDV). 


31. Hydropsyche oslari Banks (I) Nearctic, western montane 


Yukon records: 10 (ROME, SMDV). 


Genus Parapsyche. Species of Parapsyche are widely distributed through the northern 

hemisphere; 7 species are known in North America, mainly in the western mountains. Larvae 

live in small, cold streams and, like Arctopsyche, construct filter nets of coarse meshes. 

32. Parapsyche elsis Milne (I) Nearctic, western montane 

Distribution: Yukon, Alaska to California. 

Yukon records: 10, 12 (NW 1984); 16, 19 (ROME). 


Family Philopotamidae 

Larvae live in amorphous tubular silken nets of very small mesh which strain out fine 

particulate organic matter carried by the current. Wormaldia is the only one of the 3 North 

American genera recorded from the Yukon, although Dolophilodes is known from Alaska 

(Nimmo 1986). 

Genus Wormaldia. This genus is widely distributed in both the northern and southern 

hemispheres. We have Wormaldia larvae from the Yukon (10; ROME) and from the 

Northwest Territories.


33. Wormaldia gabriella (Banks) (I) Nearctic, disjunct 

Distribution: Yukon to California; Quebec. Records from the Hudson Bay drainage of northern 

Quebec (Roy and Harper 1979) indicate a major disjunction from the general western range of this 

species. 

Yukon records: A record for this species from the northern Yukon (NW 1984) was attributed to Schmid 

(1982); Yukon was not included among 9 provinces and states listed by Schmid but a Yukon record 

does appear on a distribution map. This species was recorded by Winchester (1984) from the area of 

Inuvik, Northwest Territories (68°31.2′N 135°54.2′W), close to the Arctic coast and adjacent to the Yukon. 

Family Polycentropodidae 

This is an important cosmopolitan family of retreat-making caddisflies with 6 genera 

and more than 70 species in North America; 2 genera occur in the Yukon. Most larvae are 

predacious and construct a variety of silken retreats and capture-nets. 

Genus Neureclipsis. This is a small Holarctic genus with 6 North American species. Larvae 

live in slow currents, concealed in voluminous sack-like silken nets that filter suspended 

particles. 

34. Neureclipsis bimaculata (Linnaeus) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland, Illinois; Europe through Siberia to Kamchatka. 



Genus Polycentropus. This is the largest genus in the family, widely distributed through the 

world, and with more than 40 species in North America. Larvae of different species live in 

lotic and lentic habitats and also in bog ponds and temporary pools, confirming a broad 

ecological tolerance for the genus. 

35. Polycentropus aureolus (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland and Ohio. 



36. Polycentropus flavus (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland, Illinois and California. 

Yukon records: 4, 5, 10, 12, 17 (ROME, SMDV); 11, 16 (NW 1984). 


37. Polycentropus remotus Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland and Kentucky. 

Yukon records: 4, 10, 12 (ROME, SMDV). 


38. Polycentropus smithae Denning (I) Nearctic, transcontinental 

Distribution: Yukon, British Columbia, Quebec, New Hampshire. 



39. Polycentropus weedi Blickle and Morse (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland, New Hampshire. 



Suborder Integripalpia 

These are the case-making caddisflies whose larvae construct portable tubular cases of 

plant or mineral materials fastened together with silk. In contrast to the Annulipalpia with 

fixed retreats, these larvae move with their cases in search of food. Larvae in most families


are detritivores, although some are algal grazers or predators. For pupation, the larval case 

is sealed with perforate silk at each end, allowing water to circulate directly over the pupa. 

Family Apataniidae 

Five North American genera are assigned to this family and 2 of them occur in the 

Yukon. 

Genus Allomyia. Most of the species known in this genus are confined to cold mountain 

streams of western North America; a few are known also in the Far East of Russia. Larvae 

graze diatoms and fine organic particles from rocks. 

40. Allomyia sp. 

This record is based on one larval collection from the Yukon (10 ROME) which cannot be 

identified to species. 

Genus Apatania. Seventeen species of Apatania are known in North America, and many 

others occur in the Palaearctic and Oriental regions. Larvae live in cool waters, usually 

streams but also lakes at higher latitudes, where they scrape diatoms and other algae from 

rocks (e.g. Irons 1988). 

41. Apatania crymophila McLachlan (II) Holarctic, northwestern and central 

Distribution: Yukon, Alaska, Manitoba; northern Europe and Asia. 

Yukon records: 4, 8, 10, 12, 16 (ROME, SMDV). 

Biological information: Adults have been collected in the Yukon 26 May –12 August. 

42. Apatania stigmatella (Zetterstedt) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland; northern Europe through Siberia to Chukotka, Kamchatka 

and the Amur basin (Levanidova 1982). 

Yukon records: 8, 10, 11, 12, 16, 17 (ROME, SMDV, CNCI). 


43. Apatania zonella (Zetterstedt) (II) Holarctic, transcontinental 

Distribution: Ellesmere Is. (Northwest Territories), Yukon, Alaska, Quebec, Minnesota; Greenland; 

through northern Europe and Asia to the Amur basin (I.M. Levanidova, pers. comm.). 



Family Brachycentridae 

This is a small family of the northern hemisphere with 5 genera and about 30 species 

in North America; larvae live mainly in flowing water. 

Genus Brachycentrus. Larvae of Brachycentrus species live in larger and, on the whole, 

warmer rivers and streams than do those of other genera in the family. Larvae feed on 

suspended particles from the current and graze periphytic algae. 

44. Brachycentrus americanus (Banks) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to California, Quebec, Massachusetts; Siberia, Mongolia, Japan. 



Genus Micrasema. Larvae are confined to small cold streams, where they graze algae and 

moss from rocks. 

45. Micrasema gelidum McLachlan (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Wisconsin and Quebec; northern Europe and Asia. 

Yukon records: 1 (SMDV); 4, 8, 10 (ROME); 12 (NW 1984). 


Taxonomic notes: M. kluane Ross and Morse is a junior synonym (BotojAneanu 1988).


46. Micrasema bactro Ross (I) Nearctic, western montane 

Distribution: Yukon to Utah. 



Family Goeridae 

This is a small family widely distributed in the northern hemisphere, but extended to 

tropical latitudes in Asia and even to southern Africa. There are 4 North American genera 

comprising about a dozen species. Larvae of most species live in cool running water, and 

some in the water-saturated muck of spring seepage. 

Genus Goera. This is the largest genus in the family with 6 species in North America. Larvae 

that are known live in streams and graze diatoms and organic particles from rocks. 

47. Goera tungusensis Martynov (II) Holarctic 

Distribution: Northwest Territories, Quebec; Siberia. 

Yukon records: This species is included provisionally in the Yukon fauna on the basis of a single 

female collected in the Northwest Territories, very close to the Yukon border (Midway L., 67°14′N 

135°26′W, 8 July85, SMDV). This female is similar to, but not entirely identical with, the female of 

G. tungusensis from Siberia. 


Family Lepidostomatidae 

This family is widely distributed in the northern hemisphere, with some 70 Nearctic 

species. Two genera are recognized in North America, Theliopsyche and Lepidostoma, with 

most species assigned to the latter; 4 subgenera have been proposed for species in Lepidostoma, 

3 of them represented in the Yukon fauna. No Holarctic species are known. Larvae of 

most species live in cool, running water, where they are important detritivores; some 

Lepidostoma larvae live in the littoral zone of lakes. 

Genus Lepidostoma. This genus is widely represented in North America and Eurasia. 

48. Lepidostoma cascadense (Milne) (I) Nearctic, western montane 




49. Lepidostoma cinereum Banks (I) Nearctic, transcontinental, 

western montane 

Distribution: Yukon, Alaska to California and Utah, to Newfoundland and Maine. 

Yukon records: 10, 14 (ROME); 11 (NW 1984). 


Taxonomic notes: Lepidostoma strophe Ross is a junior synonym. 

50. Lepidostoma pluviale (Milne) (I) Nearctic, western montane 




Taxonomic notes: Lepidostoma veleda Denning is a junior synonym. 

51. Lepidostoma roafi (Milne) (I) Nearctic, western montane 




52. Lepidostoma stigma Banks (I) Nearctic, western montane 



Biological information: Adults have been collected in the Yukon 24 July.


53. Lepidostoma unicolor (Banks) (I) Nearctic, transcontinental, 

western montane 

Distribution: Yukon to Quebec, California, and Arizona. 


Biological information: Adults have been collected in the Yukon 4 – 9 August. 

Family Leptoceridae 

The Leptoceridae are a large family represented on all continents. In North America 

there are approximately 100 species assigned to 8 genera, 5 of which are represented in the 

Yukon fauna. Larvae live in lakes, marshes, and slow rivers, feeding on organic particles 

and aquatic plants, or on insects. 

Genus Ceraclea. Of some 36 North American species in this genus, 4 are known in the 

Yukon. Larvae live in large rivers and the littoral zone of lakes where some species feed on 

colonies of freshwater sponges (Resh et al. 1976). 

54. Ceraclea annulicornis (Stephens) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to California, Newfoundland and Kentucky; northern Europe and Asiatic 

Russia to the Amur region and Japan (Lepneva 1966). 



55. Ceraclea cancellata (Betten) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland, south to Florida and Arizona. 



56. Ceraclea nigronervosa (Retzius) (II) Holarctic 

Distribution: Yukon, Alaska, British Columbia and Wyoming; northern Europe and Asia. 

Yukon records: 4, 10, 12, 17 (ROME, SMDV). 



57. Ceraclea resurgens (Walker) (I) Nearctic, transcontinental 

Distribution: Yukon, Oregon to Maine and Louisiana. 



Genus Mystacides. This is an Holarctic and Oriental genus with 3 widely distributed North 

American species; larvae live in standing or slowly moving water, where they are mainly 

predacious. 

58. Mystacides alafimbriata Hill-Griffin (I) Nearctic, western 

Distribution: Yukon, Alaska to California and Mexico. 

Yukon records: 10, 17 (ROME); 12 (NW 1984). 



59. Mystacides interjectus (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Quebec, Massachusetts and Ohio. 




60. Mystacides sepulchralis (Walker) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California, Newfoundland and Georgia. 

Yukon records: 4, 5, 8, 10, 12, 14, 16, 17, 19 (ROME, SMDV). 


Taxonomic notes: See Taxonomic Note 4.


Genus Oecetis. This is a genus of worldwide distribution, with approximately 20 Nearctic 

species; larvae are predacious and live in both lentic and lotic waters. 

61. Oecetis immobilis (Hagen) (I) Nearctic, transcontinental 

Distribution: Yukon, Montana to Maine, Ohio. 



62. Oecetis inconspicua (Walker) (I) Nearctic, transcontinental; 

Neotropical 

Distribution: Yukon, Alaska to California, Newfoundland, Florida, Texas; Mexico, Bahamas, Cuba, 

Venezuela. 

Yukon records: 4, 10, 12, 14, 15, 17, 19 (ROME, SMDV). 


63. Oecetis ochracea (Curtis) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to California, Tennessee; central and northern Europe (Malicky 1988), 

through Siberia to Chukotka and Kamchatka, and Mongolia (Levanidova 1982). 

Yukon records: 4, 10, 12, 16 (ROME). 


Genus Triaenodes. This is a widely distributed genus with about 25 Nearctic species; larvae 

occur in lentic and lotic habitats, where they feed on plant materials. 

64. Triaenodes baris (Ross) (I) Nearctic, transcontinental 

Distribution: Yukon to Wisconsin, Maine. 



65. Triaenodes tardus Milne (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Arizona, Maine, Tennessee. 



Genus Ylodes. Some 15 species are now assigned to this Holarctic genus, most of them from 

northern Asia. All 4 species known in North America occur in the Yukon. Larvae live in 

lentic waters. 

66. Ylodes frontalis (Banks) (I) Nearctic, western and central 

Distribution: Yukon, Alaska to California, Saskatchewan, South Dakota. 



67. Ylodes kaszabi (Schmid) (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska, Northwest Territories; Mongolia (Schmid 1965b). 



68. Ylodes reuteri (McLachlan) (II) Holarctic 

Distribution: Yukon to Colorado and Manitoba; through much of Europe (BotojAneanu and Malicky 

1978), Caucasus (Martynov 1909), Egypt and Saudi Arabia (L. BotojAneanu, pers. comm.), Siberia 

(I.M. Levanidova, pers. comm.) to Mongolia (Mey and Dulmaa 1985). 



Taxonomic notes: Triaenodes griseus Banks is a junior synonym (Manuel and Nimmo 1984). 

69. Ylodes schmidi Manuel and Nimmo (IV) East Beringian 

Distribution: Known only from the Yukon. 

Yukon records: 17 (CNCI) (Manuel and Nimmo 1984). 

Biological information: Adults have been collected in the Yukon 24 July.


Family Limnephilidae 

This is the largest family of Trichoptera in North America with some 230 species in 39 

genera. Limnephilidae are the dominant and most diverse group in northern latitudes, and 

constitute more than half of the species of Trichoptera in the Yukon. Larvae occur in all 

types of aquatic habitats, and are largely detritivorous. 

Genus Anabolia. This is an Holarctic genus with 5 North American species widely 

distributed over the northern part of the continent. Larvae are detritivores in slow streams, 

marshes, and temporary pools. 

70. Anabolia bimaculata (Walker) (I) Nearctic, transcontinental 

Distribution: Yukon to Utah, Newfoundland, New Hampshire. 



Genus Arctopora. This is a small, Holarctic genus of 3 species (Fig. 19). Larvae of at least 

A. pulchella live in temporary pools, and probably marshy sites generally. 

71. Arctopora pulchella (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland and New Hampshire. 

Yukon records: 4, 8, 11, 12, 14, 16, 17 (ROME, SMDV); 10 (NW 1984). 


72. Arctopora trimaculata (Zetterstedt) (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska; northern Europe (BotojAneanu and Malicky 1978) and Asia from 

Scandinavia through Siberia to the Amur region and Sakhalin (Schmid 1952). 

Yukon records: 4, 5 (ROME, SMDV). 


Genus Asynarchus. Species of Asynarchus occur over the northern half of North America, 

and several of them also occur in Eurasia. Larvae live in streams, ponds, and temporary 

pools, and are probably detritivores. 

73. Asynarchus aldinus (Ross) (I) Nearctic, western 


Yukon records: 10, 16, 17 (NW 1984). 


74. Asynarchus iteratus McLachlan (II) Holarctic, northwestern and central 

Distribution: Yukon, Alaska to Manitoba (Churchill: Lehmkuhl and Kerst 1979); northern Asia to 

Kamchatka (Lepneva 1966). 

Yukon records: 4 (CNCI); 5, 10 (ROME). 


75. Asynarchus lapponicus (Zetterstedt) (II) Holarctic, transcontinental 

Distribution: Yukon to Newfoundland and Maine; northern and central Europe (Malicky 1988, fig. 11) 

through Siberia to Chukotka (Levanidova 1982). 

Yukon records: 1 (SMDV); 2 (CNCI); 8 (ROME). 


76. Asynarchus montanus Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland and Utah. 

Yukon records: 4, 6, 8, 10, 11, 12, 14, 16, 17, 19 (ROME, SMDV). 


77. Asynarchus mutatus (Hagen) (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland. 


Biological information: Adults have been collected in the Yukon 29 July – 6 August.


Genus Chyranda. The single species of Chyranda is widely distributed in small, cold 

streams in northern and western montane regions of North America. Larvae are detritivorous 

(e.g. Irons 1988). 

78. Chyranda centralis (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California and Quebec. 

Yukon records: 8, 11, 14, 16, 17 (ROME, SMDV); 12 (NW 1984). 


Genus Clistoronia. Clistoronia are confined to western North America, where 4 species are 

known. Larvae live in small lakes and ponds, where they are detritivorous. 

79. Clistoronia magnifica (Banks) (I) Nearctic, western montane 


Yukon records: 11, 12, 16, 17 (CNCI, ROME, SMDV). 


Genus Dicosmoecus. Four species of Dicosmoecus inhabit running waters in western 

montane North America, and 2 others occur in eastern Asia (Wiggins and Richardson 1982). 

The 2 Yukon species feed mainly on vascular plant materials and insects. 

80. Dicosmoecus atripes (Hagen) (I) Nearctic, western montane 

Distribution: Yukon, Alaska to California, New Mexico. 

Yukon records: 8, 10, 12, 14, 16 (ROME); 17, 19 (CNCI). 

Biological information: Adults have been collected in the Yukon 23 June –18 August. Dippers, or 

water ouzels, representing a small family of passeriform birds (Cinclidae) frequenting streams, feed 

on larvae of this species in Alaska (Ellis 1978b). 

81. Dicosmoecus obscuripennis Banks (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska; Russian Far East (Nagayasu and Ito 1993). 

Yukon records: 4, 8, 10, 12 (ROME); 17 (CNCI). 


Genus Ecclisomyia. This is a small Holarctic genus confined to montane areas of western 

North America and eastern Asia; larvae live in small streams or rocky lake shores, and feed 

on diatoms and plant detritus (Irons 1988). 

82. Ecclisomyia conspersa Banks (I) Nearctic, western montane 


Yukon records: 8, 10, 12, 14, 16, 17 (CNCI, ROME, SMDV); 19 (NW 1984). 

Biological information: Adults have been collected in the Yukon 26 May –17 August. 

Genus Glyphopsyche. This is a Nearctic genus of 2 species; larvae of G. irrorata live in 

accumulations of plant materials in slowly flowing waters and in marshes, where they are 

probably detritivorous. 

83. Glyphopsyche irrorata (Fabricius) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California and Newfoundland. 


Biological information: Adults have been collected in the Yukon 26 May. 

Genus Grammotaulius. This is a widespread Holarctic genus of northern and montane 

ponds and slow streams. Larvae are probably detritivorous. 

84. Grammotaulius alascensis Schmid (IV) East Beringian 

Distribution: Yukon, Alaska, Northwest Territories. 

Yukon records: 2 (NW 1984); 4, 8, 10 (ROME, SMDV). 


Taxonomic notes: Grammotaulius subborealis Schmid (1964) is a new synonym. See Taxonomic Note 6.


85. Grammotaulius interrogationis (Zetterstedt) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland; Greenland. 



86. Grammotaulius signatipennis McLachlan (III) Palaearctic-East Beringian 

Distribution: Yukon; northern Eurasia from Sweden and Poland to Kamchatka (Schmid 1950a; 

Levanidova 1982). 

Yukon records: 11 (CNCI). 

Biological information: Adults have been collected in the Yukon 4 September. 


Genus Grensia. A single circumpolar species is assigned to this genus; larvae live mainly 

in tundra lakes and ponds. 

87. Grensia praeterita (Walker) (III) Holarctic, far northern 

Distribution: Yukon, Alaska, Northwest Territories; Greenland; far northern Eurasia. 



Genus Hesperophylax. Seven species are recognized in the Nearctic genus Hesperophylax, 

and they inhabit an unusually wide range of habitats from springs to rivers and lakes (Parker 

and Wiggins 1985). Larvae feed mainly on detritus. 

88. Hesperophylax designatus (Walker) (I) Nearctic, transcontinental 

Distribution: Yukon to California, Newfoundland, and Illinois. 



Genus Lenarchus. This is a northern Holarctic genus in which 5 of the 9 Nearctic species 

have been recorded from the Yukon. Larvae live in lentic sites—small lakes, marshes, and 

temporary pools, especially at higher elevations and latitudes, and feed on organic debris. 

89. Lenarchus crassus (Banks) (I) Nearctic, transcontinental, northern 


Yukon records: 4, 8, 14 (ROME); 12 (CNCI). 


90. Lenarchus expansus Martynov (IV) Palaearctic-East Beringian 

Distribution: Yukon, Alaska; Siberia and the Russian Far East (I.M. Levanidova, pers. comm). 

Yukon records: 1, 8, 10 (CNCI); 4 (ROME). 


91. Lenarchus fautini (Denning) (I) Nearctic, western 


Yukon records: 8, 12 (ROME); 14, 15 (SMDV). 


92. Lenarchus keratus Ross (I) Nearctic, transcontinental 

Distribution: Yukon to Michigan, Quebec. 



93. Lenarchus vastus (Hagen) (I) Nearctic, western 


Yukon records: 8, 10, 11, 12, 14, 16, 17 (ROME). 

Biological information: Adults have been collected in the Yukon 11 June – 2 September. 

Genus Limnephilus. This is the dominant trichopteran genus of northern latitudes of the 

globe, and includes approximately 85 Nearctic species (Ruiter 1995); about 27 species are


known in the Yukon, constituting 20 per cent of the Trichoptera. Larvae range widely in 

habitat, although most live in standing waters where they are major detritivores. 

94. Limnephilus argenteus Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland. 

Yukon records: 4, 6, 8, 10, 17 (ROME); 11, 16 (SMDV). 


95. Limnephilus canadensis Banks (I) Nearctic, transcontinental 

Distribution: Yukon to Maine, Oregon. 



96. Limnephilus diphyes McLachlan (III) Palaearctic-East Beringian 

Distribution: Previously known from Scandinavia through northwestern Siberia (BotojAneanu and 

Malicky 1978) to the Amur district and Kamchatka (I.M. Levanidova, pers. comm), this species is 

recorded from North America for the first time. 

Yukon records: 5, 8, 10 (ROME); 11, 17 (SMDV). 



97. Limnephilus dispar McLachlan (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Colorado, Newfoundland; northern and central Europe (Malicky 1988, 

fig. 15), Siberia (Martynov 1914, 1924a), the Amur basin and Kamchatka (Levanidova 1982). 


Biological information: Adults have been collected in the Yukon 31 May –18 July. 

Taxonomic notes: L. minusculus (Banks) is a junior synonym (Malicky 1979). 

98. Limnephilus externus Hagen (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to California, Newfoundland; northern Europe, Siberia, China (Fischer 

1968). 

Yukon records: 4, 8, 10, 11, 12, 14 (ROME); 16, 17 (SMDV). 

Biological information: Adults have been collected in the Yukon 21 July – 2 September. 

99. Limnephilus extractus Walker (I) Nearctic, transcontinental 

Distribution: Yukon to Utah, Quebec. 

Yukon records: 4, 10, 16, 17 (ROME); 12 (SMDV). 


100. Limnephilus femoralis Kirby (II) Holarctic, transcontinental 

Distribution: Yukon to Washington and Maine; Greenland (Mosely 1929); northern Europe, Kamchatka 

(Ulmer 1927) but not Siberia. 

Yukon records: 4 (SMDV); 17 (ROME). 


101. Limnephilus fenestratus (Zetterstedt) (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska, Northwest Territories; northern Europe, Iceland (Gislason 1981), 

Mongolia (Mey and Dulmaa 1985), Bering Island (I.M. Levanidova, pers. comm.) (Fig. 21). 

Yukon records: 4, 8 (ROME); 12 (CNCI, SMDV). 


102. Limnephilus fumosus Banks (IV) East Beringian 

Distribution: Yukon, Alaska, Northwest Territories. 



Taxonomic notes: This species has been confused with L. santanus Ross; Limnephilus isobela Nimmo 

(1991) is a new synonym. See Taxonomic Note 8. 

103. Limnephilus hageni Banks (I) Nearctic, transcontinental 

Distribution: Yukon, British Columbia to Quebec. 

Yukon records: 4, 10, 11, 12, 14, 15, 19 (ROME); 16 (NW 1984).



104. Limnephilus hyalinus Hagen (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Maine, Colorado. 

Yukon records: 10 (ROME, CNCI); 16 (NW 1984); 17 (SMDV). 


105. Limnephilus infernalis (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Maine. 



106. Limnephilus janus Ross (I) Nearctic, transcontinental 

Distribution: Yukon to Colorado, Maine. 



107. Limnephilus kennicotti Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Oregon to Newfoundland; Greenland (Fig. 21). 

Yukon records: 2 (ROME); 10 (CNCI); 12 (NW 1984); 16, 19 (SMDV). 


Taxonomic notes: See under L. fenestratus (category III). 

108. Limnephilus nigriceps (Zetterstedt) (II) Holarctic, northern 

Distribution: Yukon, Alaska to Manitoba; Europe and Siberia to Kamchatka (Lepneva 1966). 

Yukon records: 8 (SMDV); 10 (CNCI); 12, 16 (NW 1984). 


109. Limnephilus pallens Banks (IV) Nearctic 

Distribution: Yukon, Alaska, Northwest Territories, Michigan (Ruiter 1995). 

Yukon records: 1 (SMDV); 2 (CNCI); 16 (NW 1984). 


110. Limnephilus partitus Walker (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland. 

Yukon records: 4 (SMDV); 10, 12, 16 (ROME); 17 (CNCI). 


111. Limnephilus parvulus (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland, Maine. 


Biological information: Adults have been collected in the Yukon 31 May – 25 June. 

112. Limnephilus perpusillus Walker (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland, south to Colorado. 



113. Limnephilus picturatus McLachlan (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Colorado, east to Quebec; Greenland; northern Europe, Iceland 

(Gislason 1981) through Siberia to Kamchatka, the Amur region, Sakhalin, and Chukotka (Levanidova 

1982). 

Yukon records: 1, 16 (NW 1984); 2 (CNCI); 5, 6, 12 (ROME); 4, 8, 14, 19 (ROME, SMDV); 17 

(SMDV). 


114. Limnephilus rhombicus (Linnaeus) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Colorado, east to Newfoundland, West Virginia, Ohio; Greenland; 

Europe (Malicky 1988, fig. 18), through Turkestan (Schmid 1955), Siberia, Mongolia to Kamchatka 

(Lepneva 1966; Levanidova 1982). 

Yukon records: 4, 10, 12 (ROME); 17 (INHS). 

Biological information: Adults have been collected in the Yukon 16 June – 30 July.


115. Limnephilus sansoni Banks (I) Nearctic, western montane 

Distribution: Yukon, Alaska to Colorado. 

Yukon records: 6, 8, 10, 11, 12 (ROME); 11, 16 (ROME, SMDV); 17 (CNCI). 


116. Limnephilus secludens Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California, Quebec. 

Yukon records: 10, 12, 17 (CNCI, ROME, SMDV). 


117. Limnephilus sericeus (Say) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to Colorado, Newfoundland to Ohio; Europe (Malicky 1988, fig. 16), 

Siberia to Kamchatka, Japan (Lepneva 1966). 

Yukon records: 10, 14, 16 (ROME); 12, 17 (SMDV). 

Biological information: Adults have been collected in the Yukon 2 – 31 August. 

118. Limnephilus stigma Curtis (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska (Flint 1964); Europe (BotojAneanu and Malicky 1978), through Siberia 

to Kamchatka and the eastern extremity of Russia (Lepneva 1966; Levanidova 1975). 

Yukon records: 4, 12, 17 (SMDV); 8, 10, 11 (ROME). 


Taxonomic notes: L. indivisus (Figs. 26, 27), the similar sister species, has not been confirmed for the 

Yukon; see under L. stigma (III). 

119. Limnephilus sublunatus Provancher (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland, Colorado. 


Biological information: Adults have been collected in the Yukon 3 –10 August. 

120. Limnephilus tarsalis (Banks) (I) Nearctic, western 

Distribution: Yukon, Alaska, Northwest Territories, Alberta, British Columbia, Wyoming, Colorado. 



Taxonomic notes: L. alvatus Denning is a junior synonym (Ruiter 1995). 

Genus Nemotaulius. This is a small Holarctic genus with a single species in North America 

distributed across the northern part of the continent. Larvae live in lentic waters. 

121. Nemotaulius hostilis (Hagen) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland, West Virginia, Colorado. 



Genus Onocosmoecus. Onocosmoecus is a small Holarctic genus of 2 species (Wiggins and 

Richardson 1987); larvae live in cool waters of slow streams and lake margins where they 

are detritivores. 

122. Onocosmoecus unicolor (Banks) (II) Holarctic, transcontinental 

Distribution: Yukon, Alaska to California, Colorado, Michigan, Newfoundland; eastern Siberia and 

Kamchatka to the Kurile Islands. 

Yukon records: 6, 8, 10, 11, 12, 14, 16, 17, 19 (ROME, SMDV). 


Genus Philarctus. Several species of Philarctus occur in Asia but only one in North 

America, from Manitoba westward; larvae live in slow streams and small ponds where they 

are probably detritivores. 

123. Philarctus quaeris (Milne) (I) Nearctic, western and north-central 

Distribution: Yukon, British Columbia to Manitoba, Colorado. 

Yukon records: 17 (ROME, SMDV).



Genus Psychoglypha. This is an important Nearctic genus wholly confined to western 

montane areas except for one northern and transcontinental species, P. subborealis. Larvae 

are confined to cool running waters and are detritivorous. 

124. Psychoglypha alascensis (Banks) (I) Nearctic, western montane 




125. Psychoglypha subborealis (Banks) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California, New Hampshire, Newfoundland. 


Biological information: Adults have been collected in the Yukon 26 April – 24 July. 

Genus Sphagnophylax. Sphagnophylax is a relict genus of arctic tundra in the Yukon and 

adjacent Northwest Territories (Wiggins and Winchester 1984). Larvae of the single species 

live in transient tundra pools and feed on plant materials. 

126. Sphagnophylax meiops Wiggins and Winchester (IV) East Beringian 

Distribution: Yukon, Northwest Territories. 



Family Molannidae 

The Molannidae are a small family mainly confined to the northern hemisphere; larvae 

are omnivorous, living in lentic or slowly flowing waters. 

Genus Molanna. This is an Holarctic and Oriental genus with 6 North American species. 

Only one, M. flavicornis, is transcontinental; the others are eastern. 

127. Molanna flavicornis Banks (II) Holarctic, transcontinental 

Distribution: Yukon to Nova Scotia, New York, disjunct populations in Colorado; Eurasia. 



Taxonomic notes: Designation of this species as Holarctic hinges on the proposal by Fuller (1987) that 

it is conspecific with the Eurasian M. albicans Zett. 

Genus Molannodes. Molannodes is an Holarctic genus with one species widely distributed 

across northern Europe and Asia, and highly localized in northern North America; a second 

species occurs in Japan. 

128. Molannodes tinctus Zetterstedt (II) Holarctic 

Distribution: Yukon, Alaska, Northwest Territories, Saskatchewan, Ontario; northern Europe, Asia. 



Family Phryganeidae 

The Phryganeidae are an Holarctic and Oriental family of 74 species, well represented 

throughout northern North America (Wiggins in press). Larvae live in lakes and marshes or 

in slow rivers and streams; they are omnivorous to largely predacious in feeding. The largest 

caddisflies belong to this family. 

Genus Agrypnia. This is an Holarctic genus with 10 North American species, mainly 

transcontinental and northern; all but one of the 10 are known in the Yukon. Larvae occur 

in marshes, lakes and slow rivers, and are mainly predacious.


129. Agrypnia colorata Hagen (II) Holarctic, transcontinental 

Distribution: Yukon, Utah to Newfoundland; Siberia to Mongolia, Finland. 



130. Agrypnia deflata (Milne) (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Colorado, Newfoundland. 

Yukon records: 4, 12 (ROME, SMDV); 6, 10, 16 (ROME); 17 (CNCI). 


131. Agrypnia glacialis Hagen (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California, North Dakota, Newfoundland; Greenland. 



132. Agrypnia improba (Hagen) (I) Nearctic, transcontinental 

Distribution: Yukon, Oregon to Newfoundland, North Carolina. 

Yukon records: 10, 12, 15, 17 (ROME, SMDV); 19 (CNCI). 


133. Agrypnia macdunnoughi (Milne) (I) Nearctic, transcontinental 


Yukon records: 10, 17 (ROME, SMDV); 19 (NW 1984). 


134. Agrypnia obsoleta (Hagen) (III) Palaearctic-East Beringian 

Distribution: Yukon and British Columbia; Europe and Asia. 



135. Agrypnia pagetana Curtis (II) Holarctic 

Distribution: Yukon, Alaska, British Columbia through the Northwest Territories to Manitoba; 

northern and central Europe to northern Italy, Bulgaria, Caucasus (BotojAneanu and Malicky 1978). 

Yukon records: 2, 4, 10, 11, 12, 19 (ROME, SMDV). 


136. Agrypnia sahlbergi (McLachlan) (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska, and British Columbia; northern Scandinavia, Asia. 



137. Agrypnia straminea Hagen (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland and Colorado, Illinois, North Dakota. 

Yukon records: 8, 10, 11, 12, 15, 16, 17, 19 (ROME, SMDV). 


Genus Banksiola. This is a Nearctic genus of 4 eastern species, and one common and 

widespread transcontinental species (Wiggins 1956). Larvae live in lentic waters. 

138. Banksiola crotchi Banks (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to California, Newfoundland, Ohio. 



Genus Oligotricha. This is a small Eurasian genus with a single species extending into 

northwestern North America (Wiggins and Kuwayama 1971). Larvae live in standing waters. 

139. Oligotricha lapponica (Hagen) (III) Palaearctic-East Beringian 

Distribution: Yukon, Alaska; northern Europe, Asia. 


Biological information: Adults have been collected in the Yukon 6 – 25 July.


Genus Phryganea. This is an Holarctic genus with 2 North American species living in lentic 

waters. One of them, P. cinerea, is common throughout northern and montane areas. 

140. Phryganea cinerea Walker (I) Nearctic, transcontinental 

Distribution: Yukon, Alaska to Newfoundland and California, North Dakota, Ohio. 

Yukon records: 4, 10, 12, 17 (ROME); 17 (NW 1984). 


Genus Ptilostomis. This Nearctic genus of 4 species includes 2 with transcontinental 

distributions; in addition to P. semifasciata, P. ocellifera (Walker) has been recorded from 

Alaska and British Columbia (Liard R. Hot Springs Prov. Pk., 59°26′N 126°04′W, 8.vi.80, 

ROME), and probably occurs in the Yukon. Larvae are predacious for the most part, and 

live in all types of aquatic habitats, including temporary pools. 

141. Ptilostomis semifasciata (Say) (I) Nearctic, transcontinental 

Distribution: Yukon to Newfoundland, Virginia. 



Family Uenoidae 

This is a family of the northern hemisphere with 5 North American genera, all inhabiting 

rapid streams (Vineyard and Wiggins 1988). Two genera are recorded from the Yukon, and 

one other, Neophylax, is represented in Alaska. Larvae graze algae and fine organic particles 

on rocks. 

Genus Neothremma. This genus is restricted to western montane North America where 7 

species are known (Wiggins and Wisseman 1992). Larvae occur in rapid streams, and we 

have collected them at several Yukon sites. 

142. Neothremma didactyla Ross (I) Nearctic, western montane 

Distribution: Yukon to Oregon. 



Genus Oligophlebodes. This is a Nearctic genus of 7 western species; larvae are confined 

to turbulent mountain streams. 

143. Oligophlebodes ruthae Ross (I) Nearctic, western montane 




144. Oligophlebodes sierra Ross (I) Nearctic, western montane 




145. Oligophlebodes zelti Nimmo (I) Nearctic, western montane 

Distribution: Yukon to Montana. 



Taxonomic Notes 

Note 1. Rhyacophila mongolica Schmid, Arefina and Levanidova (14) 

Among a series of 5 pharate adults collected by a Royal Ontario Museum field party in 

the Yukon (30 mi W Old Crow, Sunaghun Cr., 16.vii. 1981, ROME #810565), one well 

developed male (Fig. 2) is considered conspecific with specimens of R. mongolica (Schmid


FIGS. 2 – 3. 2, Rhyacophila mongolica Schmid, Arefina and Levanidova (14) (Rhyacophilidae), male genitalia: a, 

lateral; b, segment X, caudal (YT: ROME 810565); 3, Rhyacophila sibirica McLachlan, segment X, caudal (Russia, 

Siberia: ROME). 

et al. 1993); our material has been compared with specimens provided by these authors from 

Russia. This is the first North American record for this species, otherwise widely distributed 

in the Russian Far East and Mongolia. The pharate male differs from its sister species R. 

sibirica McLachlan in details of segment X (cf. Figs. 2b, 3). 

Note 2. Goera tungusensis Martynov (47) 

Goera radissonica Harper and Méthot 1975, new synonymy 

During this study, we found that Goera radissonica Harper and Méthot (1975) from 

northern Quebec is a junior subjective synonym of Goera tungusensis Martynov (1909) from 

Siberia, confirmed through comparison by W.K. Gall of the holotype male of G. radissonica 

with specimens of G. tungusensis obtained from the Zoological Institute, St. Petersburg. 

Note 3. Ceraclea nigronervosa (Retzius) (56) 

We found 2 forms of adults of this species in the Yukon and Beringian collections 

studied. In both forms the dark veins of the forewing contrast strongly against the membrane, 

but the wing membrane is reddish brown in one and gray to colourless in the other. The 

gray-winged form is inferred to be typical for the species, in accordance with McLachlan’s 

(1877) description which applies to specimens from northern and central Europe; we have 

similar material from Finland. However, the brown-winged form shows several distinctive 

characters in the male genitalia as described below. 

Male genitalia (brown-winged form, Fig. 4). Segment IX in ventral view approximately 

twice as wide as long (width less than twice length in gray-winged form, Fig. 6); basal 

segment of inferior appendages extended into a prominent posteroventral process (little 

extended in gray-winged form). Segment X with superior appendages long and tapered 

apically (shorter and not tapered in gray-winged form); segment X with each ventrolateral 

lobe bearing short stout setae basally (stout setae lacking in gray-winged form). Phallus with 

apical membranous endotheca spherical in caudal view (less expanded in gray-winged form). 

Female genitalia. The female genitalic structure is variable and no characters concordant 

with the males were found; characters illustrated for each form (Figs. 5, 7) can occur in both


FIGS. 4 – 7. Ceraclea nigronervosa (Retzius) (56) (Leptoceridae). 4, Brown-winged form (YT: 810525d ROME), 

male genitalia: a, lateral; b, dorsal; c, caudal; d, ventral; e; phallus, lateral; 5, Brown-winged form (YT: 810525d 

ROME), female genitalia: a, lateral; b, dorsal; 6, Gray-winged form (YT: 810522b ROME), male genitalia: a, lateral; 

b, dorsal; c, caudal; d, ventral; e, phallus, lateral; 7, Gray-winged form (Russia: Tunguska, Yenisei R., ROME), 

female genitalia: a, lateral; b, dorsal.


brown- and gray-winged specimens. Thus females can be identified only by the brown 

membrane of the forewings and by association with males. 

In North America material of the brown-winged form was examined from the Yukon 

(10, 12, 17; ROME, SMDV), Alaska, and British Columbia; the gray-winged form was 

collected only in the Yukon (4, 10, 12; ROME). Adult specimens were collected along lakes 

and streams and on the banks of the Yukon River. However, we received a series of the 

brown-winged form collected near St. Petersburg, Russia, by V.D. Ivanov; genitalic structure 

of males is entirely consistent with our Beringian variant (Fig. 4). Consequently, we infer 

that two morphs may occur sympatrically over much of the range attributed to C. nigronervosa, 

and perhaps the two differ in preferred habitat or life cycle. The issue has to be resolved 

by a worker with access to populations in Europe and Asia. 

Note 4. Mystacides sepulchralis (Walker) (60) 

Although this species is common and widely distributed over much of North America 

including the Yukon and Alaska (Yamamoto and Wiggins 1964), its status beyond North 

America requires clarification. Holarctic distribution was attributed to M. sepulchralis by 

Yamamoto and Ross (1966), even while recognizing as valid the Siberian species M. bifidus 

Martynov (1924b); supporting evidence was not given, and we know of none apart from a 

record from Siberia attributed initially to M. sepulchralis (Martynov 1910) but later assigned 

to M. bifidus (Martynov 1935). 

There is considerable variation in genitalic characters of M. sepulchralis. Through the 

assistance of V.D. Ivanov, we have obtained information on genitalic characters of the 

holotype of M. bifidus in the Zoological Institute, St. Petersburg; and we have examined 

other specimens, indicating that there is at least some variation in this species, too. The two 

forms are very close, and clearly are sister taxa; but Martynov affirmed his view (Betten and 

Mosely 1940) that the Siberian M. bifidus Martynov and the North American M. sepulchralis 

(Walker) are separate species. We follow that interpretation here because small morphological 

differences in other species are used here as the basis for inferring separation during 

glacial periods. Detailed analysis of the variation in populations within each continent might 

reveal evidence of a Beringian interchange; whether such differences would confirm status 

as species or as intraspecific variants would emerge from the analysis. For the present, we 

interpret M. sepulchralis as a strictly North American species. 

Larvae of M. bifidus, for which we have no information, could be relevant to this 

question. In recent years, distinctive larvae in some populations of M. sepulchralis from 

eastern North America have been found with head markings of discrete spots similar to those 

in M. alafimbriata, and quite unlike the largely black heads characteristic of most sepulchralis 

populations in North America (cf. figs. 3 and 4, Yamamoto and Wiggins 1964). 

Association of these larvae with adults confirms that they are M. sepulchralis. This larval 

dimorphism could have been established in populations isolated during the glacial period to 

the southeast of the continental ice mass, although it does not appear to be reflected in 

characters of the adults. 

The sister species of M. sepulchralis/bifidus is M. alafimbriata, confined to western 

North America including the Yukon and Alaska (Yamamoto and Wiggins 1964). Mystacides 

alafimbriata and sepulchralis are only slightly different morphologically; and although there 

appears to be little overlap in their ranges, the two have been taken in the same collections 

in a few localities, all in the Yukon and Alaska. Possibly M. alafimbriata was derived from 

populations isolated to the south of the Cordilleran glacier in the mountains of western North 

America. Outlying montane glaciers along the Rocky and Cascade Mountain ranges served


FIGS. 8 – 9. 8, Mystacides sibiricus Martynov (Leptoceridae) (Russia, Lake Baikal: ROME), male genitalia, caudal 

with detail at apex of phallus, lateral; 9, Mystacides interjectus (Banks) (59) (YT: ROME 795058), male genitalia, 

caudal with detail of apex of phallus, lateral. 

to isolate refugia in the coniferous forest zone south of the continental ice sheet and in the 

southern coastal refugium (Kavanaugh 1988, fig. 37). 

Note 5. Mystacides interjectus (Banks) (59) 

This species is widely distributed across the northern half of North America (Yamamoto 

and Wiggins 1964, as longicornis), including the Yukon and Alaska. In distinguishing 

between the North American M. interjectus (Banks) and the Eurasian M. longicornis 

(Linnaeus), Yamamoto and Ross (1966) treated M. sibiricus Martynov (1935) as a junior 

synonym of M. interjectus (Banks 1914), thereby apparently extending the range of the North 

American form into Asia. This interpretation was followed by Mey and Dulmaa (1985) in 

identifying specimens from Mongolia as M. interjectus. However, illustrations of the male 

genitalia of M. sibiricus by Martynov (1935, figs. 28 – 30) show that this species belongs not 

to the interjectus lineage, but to the longicornis complex. Our interpretation is consistent 

with a male from Lake Baikal identified as M. sibiricus (Fig. 8) in the Zoological Institute, 

St. Petersburg; it is confirmed from examination of the syntype series of approximately 100 

specimens in the Zoological Institute by V.D. Ivanov, who has designated a lectotype male 

of Mystacides sibiricus Martynov (Ivan Lake, 45 km NW of Chita City, 15.vii.1925, 

Vinogradov leg.). Whether Mystacides sibiricus Martynov and longicornis Linnaeus are 

distinct species will have to be determined by examination of specimens over a wide 

geographic range in Eurasia; currently, we know of no evidence to support the occurrence 

of M. interjectus in the Palaearctic region. 

Distinction between M. interjectus and longicornis was based by Yamamoto and Ross 

(1966) on the structure of segment X; but we believe that the two are best distinguished by 

the structure of the male inferior appendages in caudal aspect. In M. interjectus (Fig. 9) the 

concave ventral face of the inferior appendage is conspicuously narrowed in caudal aspect; 

in M. longicornis and M. sibiricus, the concave ventral face is wide, and rather uniformly 

so throughout. Moreover, the phallotheca in M. longicornis and M. sibiricus terminates 

apically in a pair of stout sclerotized hooks; these hooks are scarcely developed in M. interjectus. 

Over the whole of its range in North America, M. interjectus is somewhat variable 

in genitalic morphology, but we find no indication of regional differences in the Yukon, 

Alaska, or elsewhere to indicate that populations were separated during glaciation.


The longicornis species group comprising M. nigra, longicornis, and interjectus 

(Yamamoto and Ross 1966) is apparently of Palaearctic origin. Separation between M. longicornis 

and interjectus was probably by vicariant speciation of a continuous circumboreal 

ancestral range brought about by separation of Asia and North America. 

Note 6. Grammotaulius alascensis Schmid (84) 

Grammotaulius subborealis Schmid 1964, new synonymy 

These 2 taxa, together with G. signatipennis McLachlan, constitute a complex in which 

identification is rendered all the more difficult by the fact that no explicit diagnosis has been 

proposed to distinguish one from another. Because a series of Yukon specimens (Dempster 

Hwy. km 105, nr. Blackstone R., 30.vii.1979, ROME #791183, 19 males) shows characters 

intermediate between those illustrated for males of G. alascensis and G. subborealis (Schmid 

1964, figs. 6 –13), the latter name is placed here as a junior subjective synonym under 

G. alascensis which has page priority. In the characters illustrated by Schmid for the females, 

the pair of slender membranous lobes arising from the posterodorsal margin of segment 

IX + X was interpreted inconsistently (Schmid 1964, figs. 4, 5): in G. alascensis the slender 

lobe is shown as an appendage of a sclerotized rounded lobe at the base of the anal tube; in 

G. subborealis the slender lobe is shown correctly as a median structure separate from the 

sclerotized lobe. These paired membranous lobes are actually eversible, and appear in 

genitalic preparations of Grammotaulius and many other Limnephilidae in variable conditions 

of eversion; in Schmid’s figures of G. signatipennis (1950a, figs. 52 – 54) they do not 

appear at all, as is often the case, although the lobes are extended in other cleared specimens 

of this species. 

In this emended sense, G. alascensis appears to be the North American sister species of 

G. signatipennis. Our material of G. signatipennis is not extensive, but the principal 

distinguishing character of the males is evidently the posteroventral edge of the superior 

appendages, which is very heavily sclerotized, darkened, and rather linear in G. alascensis, 

but more rounded and less sclerotized in G. signatipennis. In some specimens of the latter 

species, the posteroventral margin of the superior appendage is somewhat more heavily 

sclerotized than illustrated (Schmid 1950a, fig. 50). The intermediate appendages in our 

North American material show considerable variation, as was illustrated by Schmid. We 

have seen a specimen of G. alascensis from the Hudson Bay coastline of the Northwest 

Territories (Eskimo Point, 30.ix.1939, CNCI). From all of this, G. alascensis can be regarded 

as a highly variable North American species. We do not have sufficient material to assess 

variation for a consistent character separating females of these 2 species. 

We are advised by O.S. Flint that the Alaskan specimens attributed to G. signatipennis 

McLachlan (Flint 1964: Nunivak Is., USNM) are in fact G. alascensis Schmid. However, 

G. signatipennis must remain as part of the North American fauna because we have found 

a single male from the Yukon (Dawson, Moose Cr., CNCI) in which the genitalic structure 

is typical for this species. The forewings show the typical markings only faintly, which we 

attribute to the rather teneral condition of the specimen. 

Note 7. Limnephilus diphyes McLachlan (96) 

This species is recorded here for the first time from North America; diagnostic 

characters in genitalic morphology are illustrated (Figs. 10, 11; and also by Malicky 1983, 

pp. 191 and 200). The original description by McLachlan (1880) provides a good account 

of the general morphology. Limnephilus diphyes appears to belong to the sitchensis group 

(Schmid 1955), and can be distinguished from the other species by the prominent notch in 

the superior appendages of the male.


FIGS. 10 –11. Limnephilus diphyes McLachlan (96) (Limnephilidae) (YT: ROME 800105a). 10, Male genitalia: a, 

lateral; b, ventral; c, phallus, lateral; 11, Female genitalia: a, lateral; b, ventral. 

Note 8. Limnephilus fumosus Banks (102) 

Limnephilus isobela Nimmo 1991, new synonymy 

Information available in the taxonomic literature is inadequate for the distinction of 

L. fumosus Banks from its very similar sister species L. santanus Ross. Banks (1900) based 

L. fumosus on a series of 3 male and 2 female specimens from Alaska; diagnostic characters 

for this species were not clearly described, and Banks’ assertion that L. fumosus also occurred 

in Washington probably stemmed from failure to distinguish between L. fumosus and the 

more southerly sister taxon later recognized as L. santanus. In the original description of 

L. santanus from Oregon, Ross (1949) gave an explicit diagnosis only for the female. To 

establish the identity of these 2 species, we studied the type material of L. fumosus in the 

U.S. National Museum of Natural History, and of L. santanus in the Illinois Natural History 

Survey; genitalic morphology of both sexes is illustrated here. We also designate Limnephi-


FIGS. 12 –13. Limnephilus fumosus Banks (102) (Limnephilidae) (AK: USNM). 12, Lectotype male, genitalia: a, 

lateral; b, dorsal; c, caudal; d, phallus, lateral; e, phallus, dorsal; 13, Paratype female, genitalia: a, lateral; b, ventral. 

lus isobela Nimmo (1991) as a junior subjective synonym of L. fumosus Banks; the type 

locality given for L. isobela Nimmo is Isobel Pass (= Isabel or Isabell Pass) mi. 206, 

Richardson Highway, Yukon, but the Richardson Highway and Isabel Pass are within 

Alaska. 

Males of the 2 species can be separated by the parameres, which are bifid apically in 

L. fumosus (Fig. 12) but undivided in L. santanus (Fig. 14). The females are distinguished 

by the structure of segment X; in L. fumosus segment X is produced as a pair of heavily 

sclerotized blade-like projections (Fig. 13), broad apically in ventral aspect with the posterior 

margin broadly U-shaped. In females of L. santanus (Fig. 15) segment X terminates in a pair 

of elongate slender processes, and in ventral aspect is narrowed apically. In addition to 

genitalic differences, L. fumosus and L. santanus differ in the colour of the forewings: 

Limnephilus fumosus is uniformly brown with conspicuous light areas, much as illustrated 

by Banks (1900, fig. 10), whereas L. santanus is brown with light speckling widely 

distributed over the wing, in addition to the large light areas along the chord and at the apex, 

as described by Ross (1949).


FIGS. 14 –15. Limnephilus santanus Ross (Limnephilidae) (OR: INHS). 14, Holotype male, genitalia: a, lateral; b, 

dorsal; c, caudal; d, phallus, lateral; e, phallus, dorsal; 15, Allotype female, genitalia: a, lateral; b, ventral. 

Ross (1949) stated that the type of L. fumosus is a female; however, a type has not been 

designated. Therefore, we select the following male from the type series as the lectotype of 

Limnephilus fumosus (Banks, 1900): LECTOTYPE male: Alaska, Berg Bay, June 10 ’99; 

Harriman Expedition ’99, T. Kincaid, collector; #192, USNM Paratype (sensu cotype) 

No. 5262. 

Records confirmed for Limnephilus fumosus indicate that this species is known from 

Alaska, Yukon, and the Northwest Territories (NT: Midway L., 67°14′N 135°26′W, 

8.vii.1985, SMDV). A series in the ROME identified as L. santanus was cited by Nimmo 

and Wickstrom (1984: YT. Mirror Creek, Alaska Hwy. mile 1209, 28.vi.1958, ROME); 

these specimens have been re-examined in light of the information now available for these 

2 species, and they are L. fumosus. Nimmo’s list of Alaskan Trichoptera (1986) includes 

L. fumosus based on earlier records from the literature, and also L. santanus based on material


deposited in the CNCI; we have been unable to locate this recent material in the CNCI, but 

we suspect that it belongs to L. fumosus. Consequently, occurrence of L. santanus in Alaska 

remains subject to confirmation. Nimmo and Scudder (1983) record L. santanus from British 

Columbia; we have been unable to locate their material in the CNCI, but the record should 

be confirmed in light of this new information. 

The morphological distinction between these 2 species is rather slight but seems 

consistent; as in a number of close congeneric species in the Limnephilidae, the structural 

distinction between them appears to have arisen in the parameres of the males and in small, 

presumably reciprocal, changes in the genitalic structure of the females. 

Biogeographic Analysis of the Yukon and Holarctic Trichoptera 

Here we seek some understanding of the sources from which Trichoptera have come to 

occupy the Yukon. The number of species of Trichoptera known in the Yukon Territory 

stands now at 145, constituting 11 per cent of the Nearctic Trichoptera known north of 

Mexico, and doubtless more have yet to be recorded. The history of the Yukon trichopteran 

fauna is analyzed under 4 categories of species; several other species from adjacent Alaska 

or the Northwest Territories, although not recorded from the Yukon, are relevant to this 

analysis and are included (designated by †). The remaining Holarctic species that have not 

been recorded in Beringia are brought together for comparison in a fifth category; and 

consequently all North American species of Trichoptera that also occur in Europe or Asia 

are treated here in some manner. 

Category I. Nearctic species widespread in North America 

II. Holarctic species widespread in North America 

III. Palaearctic-East Beringian species 

IV. Beringian species 

V. Holarctic species not in Beringia 

Relevant evidence is outlined for the species discussed, and although it is incomplete 

and often speculative in interpretation, a taxonomic and conceptual context is established 

from which more focussed questions can be identified. Phylogenetic analysis of the genera 

for the species concerned, although an important asset, is well beyond the scope of this study. 

In addition to the investigation of Yukon Trichoptera, our objective in this study is an initial 

outline of issues relating to the origin of Holarctic Trichoptera in North America. Particular 

attention has been given to intraspecific morphological variation in the Holarctic and 

amphiberingian species, in an attempt to detect differences in populations that could be 

indicative of geographic disjunction during glacial periods. The broad scope of this study 

has precluded special efforts to enlarge series of the variable species to statistically significant 

levels; our observations, based on the material available, are to be regarded as an 

indication of the potential for more exacting analysis of intraspecific variation in certain 

species. In the absence of evidence to the contrary, we have assumed that these differences 

have a genetic basis, but the possibility cannot be excluded that ecological factors could be 

involved (e.g. Danks 1981: 360). 

Geological and Climatic Context. For much of the past 65 million years of the Cenozoic 

era, overland connections between the major continents at the northern end of the globe 

permitted interchange of animals and plants (Matthews 1979a, fig. 2.6), establishing the 

ancestry of the present biota. Continental crust connecting North America with Asia stood


above the sea for the first 40 million years of the Cenozoic era during the Palaeocene (65 – 58 

million years B.P.), Eocene (58 – 37 ma B.P.), and Oligocene (37 – 24 ma B.P.) epochs, 

permitting open biotic interchange between the 2 continents (Stanley 1986). The effective 

barrier to biotic interchange through early Cenozoic epochs was the Turgai Strait and perhaps 

the Cherskiy Seaway subdividing Siberia on a north-south axis (Matthews 1979a, fig. 2.2). 

The Bering land bridge connecting North America and Asia was inundated by the sea at 

intervals during the last 25 million years of the Cenozoic, through the Miocene (24 – 5 ma 

B.P.) and Pliocene (5 –1.8 ma B.P.) epochs (Stanley 1986); flooding during the mid-Pliocene 

some 3 ma B.P. closed the Bering land bridge until lowered sea level again exposed it during 

the major Pleistocene glacial advances. 

Combined with these intercontinental connections, warm moist climatic conditions 

prevailed at arctic latitudes through the Palaeocene to the Eocene. Cooler and drier 

climates followed in late Eocene, probably caused by changes of geography, ocean currents, 

volcanism, and variations in Earth’s solar orbit. This cooling trend continued through the 

Oligocene, Miocene, and Pliocene until marked decline in temperature led to establishment 

of an arctic climate in the north, with glaciers forming about 2 ma B.P. (Stanley 1986). 

Exposure of the Bering land bridge (Fig. 16) resulted from declining sea levels as large 

volumes of water were bound up in the glacial ice that covered much of northern North 

America, Asia, and Europe. A number of major glacial advances followed in the Pleistocene, 

each advance receding to some extent during a subsequent interglacial period. The last major 

Pleistocene glacial advance in North America, the Wisconsinan, began approximately 

100 000 years ago and reached its maximum about 18 000 years ago, before beginning the 

present period of glacial recession. 

Land connections between northeastern North America and Europe had disappeared by 

mid-Miocene time, some 15 million years ago. Prior to that, in early Cenozoic epochs before 

sea-floor spreading between North America and Europe had separated the land masses, there 

were at least 2 principal overland connections (Matthews 1979a, fig. 2.3): the De Geer bridge 

interconnecting through northern Greenland; and the Greenland – Faeroes bridge through 

Iceland. 

Biological Aspects. As a result of this geological and climatic history, the present fauna of 

North America has been derived over millions of years when intercontinental movement 

across northern land connections was favoured by warm to mild climates. When ocean 

barriers arose between the continents, and northern climates cooled, the North American 

biota was isolated from that of Europe, followed intermittently by separation from Asia. The 

Pleistocene epoch of the past 1.8 million years provided an opportunity once again for biotic 

interchange between North America and Asia. This time, however, harsh climates prevailed, 

and the largely unglaciated Bering land bridge connecting Nearctic (East) and Palaearctic 

(West) Beringia permitted only cold-adapted species to pass from one continent to the other. 

Formation of the glaciers was governed by the configuration of land, sea, and mountains—relationships 

that differ between Asia and North America. Over much of northeastern 

Asia glaciers were confined to higher mountains (Fig. 16), leaving most of that area 

unglaciated throughout the Pleistocene epoch. In North America, by contrast, the Laurentide 

continental glacier and the Cordilleran glacier of the western mountain ranges flowed 

together to the east and south of the Yukon (Fig. 16); and with each glacial advance the 

unglaciated Nearctic Beringian refugium was closed off as the advancing ice eliminated 

animals and plants in its path, reducing the levels of species-packing in Beringian communities. 

During periods of interglacial recession, contact may have been possible between the


FIG. 16. Approximate boundaries of unglaciated Beringia during Wisconsinan glacial maximum; based on Ager 

(1982), Lafontaine and Wood (1988). Separation between Laurentide and Cordilleran glaciers indicates location 

of postulated intermittent ice-free corridor. 

Beringian refugium, and perhaps other glacial refugia, with the main body of the North 

American biota to the south of the glaciated areas. An ice-free corridor was opened between 

the Laurentide and Cordilleran glaciers during periods of interglacial recession, connecting 

Beringia with unglaciated areas to the south for extensive periods (e.g. Reeves 1973; Pielou 

1991). This corridor would have held major freshwater drainage systems arising from glacial 

melting, but whether the climate sustained suitable habitats for aquatic insects such as 

Trichoptera is an open question. To provide food for aquatic insects, habitats must support 

plant materials of both allochthonous and autochthonous origin. However, at least one 

species of Trichoptera now exists in the rigorous climate of Ellesmere Island, indicating that 

the ice-free corridor may have supported some species during the Pleistocene. Should that 

have happened, Beringian species including those of Palaearctic origin could have passed 

to the south of the Laurentide and Cordilleran glaciers during Pleistocene interglacial 

periods. A Nahanni glacial refuge for freshwater organisms in the southeastern Yukon is 

supported by genetic analysis of lake whitefish stocks (Foote et al. 1992), adding support to 

the possibility of survival of aquatic insects in the corridor area during glaciation. But if the 

ice-free corridor did function as an interglacial passage for Trichoptera, it must have been a 

two-way corridor; and the ecological resistance of expanding Nearctic communities in the 

south to dispersing Palaearctic species could have been a constraint to the southbound 

movement of species from Beringia (e.g. Vermeij 1991). 

Exposure of the Bering land bridge during Pleistocene glaciations connected quite 

different refugial areas in Nearctic and Palaearctic Beringia (Fig. 16). Palaearctic Beringia


was much larger than Nearctic Beringia and also extended to lower latitudes; therefore, it 

would have supported more species of plants and animals, including Trichoptera (see under 

Ecological Considerations). By contrast, glaciation in North America isolated the Alaska – 

Yukon peninsula, largely if not entirely closing off exchange of species with lower latitudes 

(above). Consequently, the 2 species pools must have been imbalanced when the Bering land 

bridge was exposed. We infer that whatever dispersal followed the opening of the Bering 

land bridge would necessarily have been weighted in favour of a Palaearctic-to-Nearctic 

movement; in a group-to-group comparison, not only would there have been more species 

in Palaearctic Beringia, but also there probably would have been more vacant ecological 

niches in Nearctic Beringia because of extinctions caused by glaciation. For mammalian 

species, predominantly eastward dispersal across the Bering land bridge has been inferred 

from evidence for the Pleistocene, but movement at about the same level in each direction 

during the Pliocene (Vermeij 1991). These conclusions are consistent with the concept that 

dissimilar patterns of glaciation in North America and Asia are correlated with asymmetric 

interchange of species. Moreover, global analysis of past biotic interchanges leads to the 

general conclusion that the resistance of a biota to invasion is reduced by previous extinction 

of species in that biota (Vermeij 1991). 

Superimposed upon these general factors influencing the interchange of animals and 

plants between North America and Asia are the special requirements of aquatic insects. Only 

the Odonata have the capacity for active dispersal over longer distances between the aquatic 

habitats of larvae. Adult Trichoptera are subject at best to passive dispersal by strong winds; 

even then, ovipositing females would have to be relocated in the vicinity of appropriate 

aquatic habitats. Dispersal by wind over the narrow sea barrier of the present-day Bering 

Strait seems at least as likely as dispersal across long distances overland; but there is little 

supporting evidence for successful dispersal across Bering Strait, nor is there evidence for 

post-Pleistocene intercontinental interchange of Trichoptera by way of the Aleutian Island 

chain (e.g. Karlstrom and Ball 1969). Even so, several species of Trichoptera evidently did 

reach Greenland by dispersal at high latitudes over sea barriers from distant glacial refugia 

(see category I, Greenland Trichoptera). Beringia during much of the Pleistocene is depicted 

as arid and steppe-like with the muskeg and wet tundra, now so typical, greatly diminished 

in extent (Schweger et al. 1982). Thus, Nearctic Beringia, the corridor between Asia and 

North America, would seem to have provided a different mix of habitats and probably more 

restricted habitats for aquatic insects than are now available. Moreover, the glacier-fed water 

courses were probably similar to glacial streams of today that support few aquatic insects 

because they carry heavy loads of suspended glacial debris which are deposited to form 

unstable and largely unproductive substrates, shifted by braided channels as the volume of 

meltwater fluctuates. Unglaciated upland parts of Beringia, however, probably had streams 

that were similar to present-day upland streams in tundra regions. These streams would have 

joined the lowland river systems that traversed the exposed coastal shelf (Fig. 16), forming 

a network of freshwater systems that is believed to have facilitated an exchange of fish 

species between Asia and North America (Lindsey and McPhail 1986). Because of the high 

biological diversity of Trichoptera, however, the availability of freshwater habitats across 

Beringia is only one aspect of the issue; another aspect is the extent to which these freshwater 

habitats coincided with the requirements of the species that had access to them. These issues 

are explored under Ecological Considerations. 

We infer that the Bering land bridge of Pleistocene time would have been a difficult 

passage for many aquatic insects because the cold dry climate was not well suited to dispersal 

of the adult insects, and because freshwater habitats would have been localized, unsuitable,


or different in other ways from those of the present. More favourable conditions for the 

movement of aquatic insects between Asia and North America would seem to have been 

available during the millions of years that the 2 continents were connected in Pliocene time 

and earlier, before the land connection between the 2 continents was inundated in the 

mid-Pliocene, some 3 million years ago. 

Origin of the Beringian and Holarctic Trichoptera. The origin of Beringian and Holarctic 

species can be interpreted in several ways. Some species could have had an Holarctic 

distribution pre-dating the separation of North America from Asia in Pliocene time some 

3 ma B.P. Thus, some Holarctic species (category II) could have been confined to the south 

of the advancing ice in unglaciated parts of the continent, giving rise to intraspecific variants 

or even sibling species which now can be detected; disjunct populations of some of these 

species may have been isolated in unglaciated Nearctic Beringia by the converging Laurentide 

and Cordilleran ice masses. Palaearctic-East Beringian species (category III) may never 

have occurred much beyond their present range in North America, or, if they did, were 

overtaken by the advancing ice and confined as glacial relicts to the Beringian refugium 

where they remain. Some others could be Nearctic species that dispersed from Beringia to 

Asia during the past 1.8 million years of the Pleistocene. For most of the Beringian Holarctic 

species, the evidence now available is not sufficient to interpret clearly which of the 

possibilities seems most likely. However, a pattern of vicariant distribution of sister species 

in North America and Eurasia often emerges, dating perhaps from the Pliocene or earlier, 

and augmented in some cases by apparent later dispersal of the Palaearctic form to North 

America, presumably by way of the Bering land bridge during the Pleistocene. 

The fundamental issues in piecing together the origin of the Beringian Trichoptera are 

the age of the species and their rate of structural change over time. This information is 

unknown, but clearly we are dealing with species that differ from their closest living relative 

(sister species) by a wide range of distinguishing characters, and thus could and probably 

do represent species of different ages. Some Beringian species pairs are virtually sibling 

species, e.g. Limnephilus fenestratus (101) and L. kennicotti (107). The small, but consistent, 

morphological distinctions between some pairs of sister species could have arisen in disjunct 

populations along the southern margin of the North American glaciers. Others are well 

differentiated although clearly sister species, e.g. Agrypnia deflata (130) and A. obsoleta 

(134), which probably had a vicariant origin arising from an earlier intercontinental subdivision 

of the range of a common circumboreal ancestor. Flooding of the land connection 

between Asia and North America in the mid-Pliocene, some 3 million years ago, or earlier, 

offers possible origins for dichotomies of this sort, although disjunctions between some 

conspecific North American and Asian populations could have been imposed by cold climate 

even before the land connection was overrun by the sea. At the far end of the scale for 

the age of trichopteran taxa inferred from morphological divergence is Sphagnophylax 

meiops (126) (Frontispiece, Fig. 28), known only from unglaciated Nearctic Beringia or 

marginally beyond, and which appears to have lost the power of flight. The monotypic genus 

Sphagnophylax is the only genus-level taxon known in Trichoptera that is confined to 

Beringia, and is clearly both a phylogenetic and a geographic relict (Wiggins and Winchester 

1984). 

Placing these speciation events in geological time with any assurance is not yet possible 

for Trichoptera. Inferring the relative times of origin of species from levels of morphological 

divergence achieves some consistency in interpreting the history of a number of species of 

disparate age, and is employed in this study because few other clues are available; but we


recognize that if correct for some species, this morphological guideline may be illusory for 

others. One indication that the species did not diverge at uniform rates is provided by 

members of category V; several of these are widespread Holarctic species that must have 

passed between Asia and North America at some time before the onset of harsh Pleistocene 

climates because there is little evidence that they are adapted now to far northern latitudes. 

Secondly, fossil evidence of known age is rarely available for species of Trichoptera, but in 

Coleoptera where it is available, extant species of the north were found to be unexpectedly 

old and clearly in existence well before Pleistocene time, with some identifiable in late 

Miocene 5.7 ma B.P. (e.g. Downes and Kavanaugh 1988). A very slow rate of evolutionary 

change in some Trichoptera is demonstrated by the small structural differences between a 

Baltic amber species of Lype (Psychomyiidae) and a Recent European species (Ross 1958, 

fig. 1); the age of the amber species is placed at upper Eocene—some 50 ma B.P. Perhaps 

this is an isolated and atypical example, but for insect species generally an age of 25 million 

years has been estimated (e.g. Briggs 1966). Comprehensive analysis of the fossil record 

reveals that insects show very low rates of extinction; among vertebrates, by contrast, few 

living species are more than a million years old (Labandeira and Sepkoski 1993). The age 

of species and the time required for their origin are different concepts; but inferences 

reflecting the rate of evolution for insect species must be tempered by the evidence that at 

least 760 species of Drosophilidae have evolved on the Hawaiian Islands since their origins 

6 million to 500 000 years ago (Kaneshiro 1993). 

Finally, there is the larger question of the age of the cold, far northern biomes 

themselves. Tundra develops in regions that are too cold for the growth of trees, and arctic 

tundra is believed to have arisen in the north as disjunct areas of ecological regression in late 

Miocene forests (Matthews 1979b; Danks 1981). The Seward Peninsula of Alaska, just 

below the Arctic Circle (67°N lat.), was forested up to about 5.7 million years ago in late 

Miocene time (Hopkins et al. 1971). Palaeobotanical evidence indicates that the forest-tundra 

ecotone probably lay just to the north of the Seward Peninsula at that time, suggesting that 

tundra probably covered most of the North American Arctic Archipelago by 6 ma B.P. 

(Matthews 1979b). Thus, tundra may be a relatively young biome in the north, and adaptation 

of tundra species a rather recent phenomenon. Some arctic tundra Trichoptera may have been 

derived from alpine tundra habitats, which in Beringia would have dated from the time that 

the high mountains of the western Cordillera and of northeastern Asia reached elevations 

which precluded the growth of trees. Evidence from fossil insects suggests, however, that 

species may have adapted to arctic tundra conditions from lineages occurring in the boreal 

forest (Matthews 1979a; Danks 1981). Because of their diversity and broad ecological 

penetration, Trichoptera could be an appropriate group for testing these hypotheses. Phylogenetic 

analysis in the highly diverse boreomontane genus Limnephilus would be a promising 

avenue of investigation to reveal the habitats of the phyletic ancestors of the present 

inhabitants of arctic and alpine tundra. However, a good deal more basic information about 

the biology, distribution, taxonomy, and phylogenetic relationships of the species of Limnephilus 

will be required. Trichoptera established in pre-Pleistocene arctic tundra might have 

found glacial refuge in alpine tundra of accessible mountain ranges, or in disjunct lowland 

tundra refugia along the southern margin of the continental glacier. 

I. Nearctic Species Widespread in North America 

This is the largest category of the Yukon Trichoptera—98 species comprising 68 per 

cent of the total fauna. None of these species is known from Europe or Asia, and their 

generally widespread occurrence now at more southerly latitudes in North America suggests


that they passed the last glaciation south of the ice or in other refugia under climatic 

conditions less severe than those in Beringia, and returned to northern latitudes as the glaciers 

receded. The possibility cannot be excluded that some of these species could have passed 

the last glacial period in East Beringia, failed to cross the Bering land bridge to Asia, and 

dispersed southward after recession of the glaciers. Most of these strictly Nearctic caddisfly 

species which now occur in the Yukon probably originated before the glaciation of the past 

2 million years, although some may be the product of isolation in separate refugial areas to 

the south of the glaciers, e.g. the dichotomy between Mystacides alafimbriata and M. sepulchralis. 

Numbers in parentheses following the species names provide a cross-reference to 

distributional and other data in the Annotated List of Yukon Trichoptera. 

Spicipalpia 

Glossosomatidae 

Glossosoma alascense Banks (1) 

Glossosoma verdona Ross (3) 

Hydroptilidae 

Hydroptila rono Ross (5) 

Ochrotrichia sp. (6) 

Oxyethira araya Ross (7) 

Stactobiella delira (Ross) (8) 

Rhyacophilidae 

Rhyacophila alberta Banks (9) 

Rhyacophila angelita Banks (10) 

Rhyacophila bifila Banks (11) 

Rhyacophila brunnea Banks (12) 

Rhyacophila hyalinata Banks (13) 

Rhyacophila pellisa Ross (16) 

Rhyacophila tucula Ross (17) 

Rhyacophila vao Milne (18) 

Rhyacophila verrula Milne (19) 

Rhyacophila vobara Milne (20) 

Rhyacophila vocala Milne (21) 

Rhyacophila vofixa Milne (22) 

Annulipalpia 

Hydropsychidae 

Arctopsyche grandis (Banks) (23) 

Cheumatopsyche campyla Ross (25) 

Cheumatopsyche sp. (26) 

Hydropsyche alhedra Ross (27) 

Hydropsyche alternans (Walker) (28) 

Hydropsyche amblis Ross (29) 

Hydropsyche cockerelli Banks (30) 

Hydropsyche oslari Banks (31) 

Parapsyche elsis Milne (32) 

Philopotamidae 

Wormaldia gabriella (Banks) (33) 

Polycentropodidae 

Polycentropus aureolus (Banks) (35) 

Polycentropus flavus (Banks) (36) 

Polycentropus remotus Banks (37) 

Polycentropus smithae Denning (38) 

Polycentropus weedi Blickle and Morse (39) 

Integripalpia 

Apataniidae 

Allomyia sp. (40) 

Brachycentridae 

Micrasema bactro Ross (46) 

Lepidostomatidae 

Lepidostoma cascadense (Milne) (48) 

Lepidostoma cinereum Banks (49) 

Lepidostoma pluviale (Milne) (50) 

Lepidostoma roafi (Milne) (51) 

Lepidostoma stigma Banks (52) 

Lepidostoma unicolor (Banks) (53) 

Leptoceridae 

Ceraclea cancellata (Betten) (55) 

Ceraclea resurgens (Walker) (57) 

Mystacides alafimbriata Hill-Griffin (58) 

Mystacides interjectus (Banks) (59) 

Mystacides sepulchralis (Walker) (60) 

Oecetis immobilis (Hagen) (61) 

Oecetis inconspicua (Walker) (62) 

Triaenodes baris (Ross) (64) 

Triaenodes tardus Milne (65) 

Ylodes frontalis (Banks) (66) 

Limnephilidae 

Anabolia bimaculata (Walker) (70) 

Arctopora pulchella (Banks) (71) 

Asynarchus aldinus (Ross) (73) 

Asynarchus montanus Banks (76) 

Asynarchus mutatus (Hagen) (77) 

Chyranda centralis (Banks) (78) 

Clistoronia magnifica (Banks) (79) 

Dicosmoecus atripes (Hagen) (80) 

Ecclisomyia conspersa Banks (82) 

Glyphopsyche irrorata (Fabricius) (83) 

Grammotaulius interrogationis 

(Zetterstedt) (85) 

Hesperophylax designatus (Walker) (88) 

Lenarchus crassus (Banks) (89) 

Lenarchus fautini (Denning) (91) 

Lenarchus keratus Ross (92) 

Lenarchus vastus (Hagen) (93) 

Limnephilus argenteus Banks (94) 

Limnephilus canadensis Banks (95)

Limnephilus extractus Walker (99) 

Limnephilus hageni Banks (103) 

Limnephilus hyalinus Hagen (104) 

Limnephilus infernalis (Banks) (105) 

Limnephilus janus Ross (106) 

Limnephilus kennicotti (Banks) (107) 

Limnephilus partitus Walker (110) 

Limnephilus parvulus (Banks) (111) 

Limnephilus perpusillus Walker (112) 

Limnephilus sansoni Banks (115) 

Limnephilus secludens Banks (116) 

Limnephilus sublunatus Provancher (119) 

Limnephilus tarsalis (Banks) (120) 

Nemotaulius hostilis (Hagen) (121) 

Philarctus quaeris (Milne) (123) 

Psychoglypha alascensis (Banks) (124) 


Psychoglypha subborealis (Banks) (125) 

Phryganeidae 

Agrypnia deflata (Milne) (130) 

Agrypnia glacialis Hagen (131) 

Agrypnia improba (Hagen) (132) 

Agrypnia macdunnoughi (Milne) (133) 

Agrypnia straminea Hagen (137) 

Banksiola crotchi Banks (138) 

Phryganea cinerea Walker (140) 

Ptilostomis semifasciata (Say) (141) 

Uenoidae 

Neothremma didactyla Ross (142) 

Oligophlebodes ruthae Ross (143) 

Oligophlebodes sierra Ross (144) 

Oligophlebodes zelti Nimmo (145) 

These Nearctic Trichoptera from glacial refuges to the south are of 2 broad groups— 

western montane species and transcontinental species. The western montane species are 

mainly members of the Spicipalpia and Annulipalpia whose larvae live in lotic habitats 

characteristic of mountainous terrain. This group is particularly diverse southward through 

Oregon to California and Colorado, but markedly less successful to the north (Table 1 

below): Glossosomatidae, Rhyacophilidae, Hydropsychidae, Philopotamidae, Polycentropodidae, 

Brachycentridae (Micrasema), Lepidostomatidae, Limnephilidae (Chyranda, 

Discosmoecus, Ecclisomyia, Psychoglypha), and Uenoidae. Figures for declining species 

diversity at higher latitudes (Table 1) indicate that the lotic species of the western montane 

Trichoptera have populated the glaciated terrain of the Yukon with limited success. Lentic 

species of western montane origin in a few genera of the Limnephilidae (Asynarchus, 

Clistoronia, Lenarchus, Limnephilus) and the Leptoceridae (Mystacides, Ylodes) have 

extended farther north in the Yukon than have the lotic species of montane origin. 

Transcontinental species are inferred to have passed glaciation along a much wider ice 

front because overall they are now widely distributed in North America. They are, for the 

most part, species of lentic waters and slowly moving streams. These species are primarily 

case-makers of the Integripalpia; their diversity also declines with increasing latitude 

(Table 1), but not as much as in the Spicipalpia and Annulipalpia where most species live 

in lotic waters. Transcontinental Trichoptera occur to a large extent across the boreal forest 

biome but their lake and marsh habitats also extend through deciduous forests and grasslands. 

Larvae of these species, and of Trichoptera generally, have little or no specific relationship 

with vascular plants, and feed for the most part on the fungi colonizing plant detritus and on 

algae, or on other insects. In general, the effect of low temperature on aquatic habitats seems 

to be the major constraint to repopulation of the Yukon by species in both groups of Category 

I Trichoptera; this topic is explored further under Ecological Considerations. 

There is an interesting recurrence in northern Quebec of disjunct populations of several 

species which otherwise appear to be confined to western or northwestern North America 

(Roy and Harper 1979; Harper 1989): Arctopsyche grandis (23) (Hydropsychidae), Wormaldia 

gabriella (33) (Philopotamidae), and Goera tungusensis (47) (Goeridae). The first 

2 are Nearctic species (category I) which probably passed the Wisconsinan glaciation south 

of the ice, and now range broadly in western montane areas from the Yukon southward. 

Goera tungusensis is Holarctic (category II) and may also have passed the last glacial period


along the southern margin of the ice, or it may have entered North America for the first time 

during the Pleistocene. This shared pattern of postglacial distribution might indicate that 

each of these species passed the Wisconsinan glaciation as disjunct eastern populations; 

larvae of each of them usually occur in cool lotic habitats. 

Greenland Trichoptera. Species of Trichoptera occurring in both North America and 

Greenland pose relevant distributional questions arising from glaciation because most of 

them occur in the Yukon as well. Some are Holarctic species, and some are strictly Nearctic, 

but the fact that all of them occur in Greenland raises the possibility that they illustrate some 

general principles of high-latitude distribution. Since the insect fauna of Greenland includes 

few if any endemic species, and was derived to a large extent by postglacial dispersal from 

North America (Downes 1966), we infer that these Nearctic species of Trichoptera reached 

Greenland after recession of the glaciers. Occurrence of a Palaearctic species, Limnephilus 

griseus (Linnaeus), in Greenland (Fristrup 1942) raises a further possibility that some of 

the Holarctic species might have reached Greenland from Europe; records for a second 

Palaearctic limnephilid from Greenland, Halesus radiatus (Curt.), are believed to be in error 

(N.P. Kristensen, pers. comm.). Ice rafting from western Norway has been postulated as a 

sweepstakes route for the repopulation of Iceland and other North Atlantic islands by insects 

about 10 000 years ago (Buckland et al. 1986). Although adult Trichoptera are generally 

short-lived, an adult diapause in species of Limnephilus adapted to transient waters could 

prolong the life span, perhaps enabling these species to endure such a passage; their egg 

masses embedded in gelatinous matrix are resistant to desiccation (Wiggins 1973). All of 

the Greenland Trichoptera are case-making species of the Integripalpia whose larvae live in 

lentic waters or in slow currents of streams. 

Common patterns in the far northern distribution of Lepidoptera and of several other 

groups of insects reveal 2 elements in the fauna of Greenland (Downes 1966). Species of 

the high arctic occurring on adjacent Ellesmere Island are very largely shared with northern 

Greenland. In the Trichoptera, only 2 species of the Greenland fauna can be considered 

high-arctic forms—Apatania zonella (43) and Grensia praeterita (87). The first occurs on 

Ellesmere Island (Corbet 1966) where it is the most northerly species of Trichoptera known 

in North America, although it is an Holarctic species and is recorded also from Yukon to 

Quebec; Grensia praeterita is a circumpolar species recorded mainly above treeline although 

not from Ellesmere Island. The high-arctic species are held to have reached Greenland by 

way of a far northern route across the Arctic Archipelago (Downes 1966), raising the 

possibility that they could have passed the Wisconsinan glaciation in a Peary Land refugium 

along the northern edge of Greenland (e.g. Danks 1981, fig. 5). The relatively small number 

of high-arctic Trichoptera indicates that these insects are not well adapted to the rigours of 

existence at high latitudes. 

The second element in the Greenland insect fauna comprises species of more southerly 

distribution that are of low-arctic or boreal origin (Downes 1966). Most of the Greenland 

Trichoptera appear to be of this group because they range widely across the boreal forest 

and often farther to the south. These species are likely to be postglacial dispersants to 

Greenland, although probably by way of the Arctic Archipelago and Baffin Island. It appears 

that most of them moved into deglaciated North America from refuges in the central part of 

the continent, finally reaching Greenland. 

Evidence relevant to category I species in Greenland is outlined below, and for other 

species under the category indicated; 2 of the Nearctic species known from Greenland have 

not been recorded from the Yukon (†).

Apataniidae 

Apatania zonella (Zetterstedt) (II, 43) 

Limnephilidae 

Grammotaulius interrogationis (Zetterstedt) (I, 85) 

The type locality of this transcontinental Nearctic species is in Greenland (Mosely 1929; 

Schmid 1950a). Genitalic structures of males showed marked variation both within and 

between populations, but we found no congruent pattern. Schmid (1950a) considered 

G. interrogationis to be the sister species of 3 others (2 Palaearctic, 1 Nearctic). 

Grensia praeterita (Walker) (III, 87) 


Limnephilus femoralis Kirby (II, 100); L. picturatus McLachlan (II, 113); L. rhombicus 

(Linnaeus) (II, 114) 

These 3 widely distributed Holarctic species could have reached Greenland from North 

America, or from Europe as did the Palaearctic species L. griseus. In Yukon and other 

Nearctic material of L. rhombicus there is substantial variation in both male and female 

genitalic structure, but we found no geographic correlation to indicate separation of populations 

by glacial barriers. Congruent variation found in L. picturatus was analyzed (II). 

Limnephilius kennicotti Banks (I, 107) 

This Nearctic species is discussed under L. fenestratus (III), its very similar sister 

species. 

Limnephilus moestus Banks (†) 

The range of this Nearctic species extends from the Northwest Territories to Colorado, 

east to Newfoundland and West Virginia, and to Greenland (Forsslund 1932). It has not been 

recorded from the Yukon or Alaska. The species was assigned to the sitchensis group of 6 

Nearctic species (Schmid 1955). Few species of Nearctic Trichoptera have the broad 

latitudinal range from West Virginia to Greenland shared by L. moestus and the following 

species L. ornatus. 

Limnephilus ornatus Banks (†) 

This species is known from Alaska [Naknek (CNCI); Parks Hwy. (ROME 821118); 

Kodiak Is. (ROME); Popof Is. (Banks 1900)], and from Alberta and Montana to Newfoundland, 

south to West Virginia. Although recorded from Alaska, L. ornatus is not known from 

the Yukon or from British Columbia (Nimmo and Scudder 1978, 1983), nor is it known 

elsewhere in the western montane states; this is a large gap for a conspicuous species and 

suggests a disjunct population in Alaska. The species has been recorded from Greenland 

(Mosely 1932), but not from northern Asia; a record from Japan (Ulmer 1907), never 

confirmed, was rejected by Schmid (1965a). If L. ornatus passed the last glacial period in 

the Beringian refugium, there is no evidence that it spread to Asia. No close relatives of 

L. ornatus were indicated in the classification by Schmid (1955). 

Phryganeidae 

Agrypnia glacialis Hagen (I, 131) 

This transcontinental Nearctic species is the sister of the Palaearctic A. picta Kolenati, 

widely distributed through northern Europe and Asia to Kamchatka. There is no evidence 

that either species has moved across the Bering land bridge, in contrast to the congeneric 

species pair A. deflata and obsoleta (see category III). However, there are records for 

A. glacialis from Greenland, and the Eurasian sister species A. picta occurs as far west as


Britain and Iceland. We infer that this evidence indicates vicariant speciation from a 

circumboreal ancestor. Outlying records of A. glacialis in Idaho, Utah and California suggest 

relict montane populations from the time when the species occurred south of the Pleistocene 

glacial ice. In these populations, and also in the Greenland material, variations in male 

genitalic structure are more common than in the populations of northern North America 

(Wiggins in press). 

II. Holarctic Species Widespread in North America 

Each of these species is now widely distributed in both North America and Eurasia; 28 

of them are known in the Yukon, constituting about 18 per cent of the Trichoptera. Many 

are now transcontinental, and these broad Nearctic ranges seem more likely to follow from 

glacial refuge along the southern margin of the ice. Thus, a number of these Holarctic species 

probably occurred in North America before the Pleistocene, and probably have been on this 

continent for at least the past 3 million years, before the connection between Asia and North 

America was overrun by the sea in mid-Pliocene time. However, southward movement of 

any of these species from Beringia through the ice-free corridor between the Laurentide and 

Cordilleran glaciers might have occurred during interglacial periods of the Pleistocene (see 

Biological Aspects), and is difficult to detect. For some category II species, sister relationships 

between Nearctic and Palaearctic species pairs suggest origin from common circumboreal 

ancestors, and perhaps arose from intercontinental vicariance in the Pliocene or 

earlier. Evidence indicates that for the most part species involved in recent dispersal moved 

from Asia to North America (see Biogeographic Analysis). A few species, e.g. Brachycentrus 

americanus and Onocosmoecus unicolor, appear to have originated in North America 

and dispersed to Asia across the Bering land bridge. 

Evidence from intraspecific morphological variation suggests that some Nearctic species 

were divided, with populations confined to the south of the advancing glacial front while 

disjunct conspecific populations were restricted to Beringia: e.g. Brachycentrus americanus 

(44), Micrasema gelidum (45), Oecetis ochracea (63), Ylodes reuteri (68), Limnephilus 

picturatus (113), Onocosmoecus unicolor (122), and Agrypnia colorata (129). For the 

remaining species of category II, evidence available now is inadequate to support any 

inference about their origin and direction of dispersal. 

Although several of these species now occur in arctic tundra habitats, most of them are 

also widely distributed in forest and grassland biomes at more temperate latitudes. Consequently, 

these species do not comply with the general pattern in some other groups that the 

arctic and treeline species were derived from Beringia, while the boreal and subarctic fauna 

moved northward from refugia below the glacial front. The most northerly species of 

category II are Asynarchus iteratus (74) and Apatania zonella (43). 

Two species (†) known from Alaska or the Northwest Territories are also considered 

under this category; although not yet recorded from the Yukon, they probably occur there. 

Spicipalpia 


Glossosoma intermedium (Klapalek) 

Hydroptilidae 

Oxyethira ecornuta Morton (†) 


Rhyacophila narvae Navas 

Annulipalpia 


Arctopsyche ladogensis (Kolenati) 


Neureclipsis bimaculata McLachlan

Integripalpia 

Apataniidae 

Apatania crymophila McLachlan 

Apatania stigmatella (Zetterstedt) 

Apatania zonella (Zetterstedt) 


Brachycentrus americanus (Banks) 

Micrasema gelidum McLachlan 

Goeridae 

Goera tungusensis Martynov 

Leptoceridae 

Ceraclea annulicornis (Stephens) 

Ceraclea excisa (Morton) (†) 

Ceraclea nigronervosa (Retzius) 

Oecetis ochracea (Curtis) 

Ylodes reuteri (McLachlan) 


Limnephilidae 

Asynarchus iteratus McLachlan 

Asynarchus lapponicus (Zetterstedt) 

Limnephilus dispar McLachlan 

Limnephilus externus Hagen 

Limnephilus femoralis Kirby 

Limnephilus nigriceps (Zetterstedt) 

Limnephilus picturatus McLachlan 

Limnephilus rhombicus (Linnaeus) 

Limnephilus sericeus (Say) 

Onocosmoecus unicolor (Banks) 

Molannidae 

Molanna flavicornis Banks 

Molannodes tinctus Zetterstedt 

Phryganeidae 

Agrypnia colorata Hagen 

Agrypnia pagetana Curtis 


Glossosoma intermedium (Klapalek) (2) 

This is the only Holarctic species of Glossosoma, and the only transcontinental species 

in North America. According to Ross (1956, charts 31, 43), the sister species of G. intermedium 

is G. verdona Ross; he inferred that the two originated in North America and 

intermedium spread to Eurasia by way of the Bering land bridge. If this dispersal occurred 

during the Pleistocene, G. intermedium would be one of the few species that moved from 

North America to Asia. However, by this interpretation, some populations of G. intermedium 

probably would have been confined south of the ice, accounting for the present transcontinental 

distribution. This sister-group relationship opens a second possibility that G. intermedium 

is the Eurasian vicariant from a common Holarctic ancestor; this species might then 

have reached North America by way of the Bering land bridge. Occurrence of G. intermedium 

in the northern Yukon (Fig. 1, region 5) indicates its adaptation to far northern 

conditions; and if G. intermedium reached North America during the Pleistocene glaciation, 

its present transcontinental distribution would have been achieved after glacial recession. In 

any event, occurrence of the 2 sister species in the Yukon invites ecological comparison (e.g. 

Irons 1988), leading perhaps to some understanding of factors underlying the wide distribution 

of G. intermedium. Ross considered the subgenus G. (Synafophora = Eomystra), to which 

G. intermedium belongs, to be the biological analogue of the sibirica group of Rhyacophila—species 

of both groups tolerant of low gradient streams and sharing the dispersal 

capability of ecological generalists which enabled some to reach eastern North America. 

Hydroptilidae 

Oxyethira ecornuta Morton (†) 

One of the few Holarctic species known in this family, O. ecornuta occurs through 

northern Europe (BotojAneanu and Malicky 1978) to the Far East of Russia (BotojAneanu 

and Levanidova 1988); our collections include specimens from Alaska, and although not yet 

recorded from the Yukon, this species probably occurs there. There is material from Ontario 

in the ROME collection, suggesting a disjunct distribution and the possibility that this 

species at least in part passed the last glaciation south of the continental glacier. The species 

was placed in the flavicornis group by Kelley (1984), and was believed to be the sister species 

of O. flavicornis (Pictet) from Europe; both are said to occur in lentic habitats.



Rhyacophila narvae Navas (15) 

This species is a member of the sibirica group, and was proposed as the sister taxon of 

R. transquilla Tsuda from Japan (Ross 1956; Schmid 1970). Origin of Rhyacophila narvae 

in Asia was proposed by these authors. If dispersal to North America occurred during the 

Pleistocene, river systems of the Bering land bridge must have provided larval habitats of 

cool, fast-flowing water required by the larvae (Lepneva 1964). If dispersal of R. narvae to 

North America occurred before the Pliocene separation of the 2 continents, this Holarctic 

species has resisted the vicariant subdivision postulated for some others. 


Arctopsyche ladogensis (Kolenati) (24) 

This is the only Holarctic species in the genus and, of 4 species of Arctopsyche in North 

America, it is the only one with a transcontinental distribution. It is the sister species of 

A. amurensis Martynov from the Amur region of Russia (Schmid 1968). If the 2 species 

originated in Asia, A. ladogensis could have reached North America during the Pleistocene 

or before by way of the Bering land bridge. However, vicariant speciation of a circumboreal 

ancestor following separation of North America and Asia, perhaps in the Pliocene, is also a 

possible explanation for the sister-group relationship. In this interpretation, A. ladogensis 

would be a Nearctic species confined at least in part to the Beringian refugium, and dispersing 

to Asia across the land bridge; a disjunct population in Utah (Baumann and Unzicker 1981) 

is consistent with Nearctic origin of A. ladogensis and its occurrence both north and south 

of the ice. We found no consistent morphological differences between Nearctic and 

Palaearctic representatives, but have not examined specimens from Utah. Larvae live in 

strong currents of rivers and larger streams (Lepneva 1964), and occurrence of this species 

in the far north of the Yukon (Fig. 1, region 4) indicates that it is suited for trans-Beringian 

dispersal. 


Neureclipsis bimaculata (Linnaeus) (34) 

This is the only Holarctic species of Neureclipsis, and the only transcontinental species 

in North America. There are morphological differences between Nearctic and Palaearctic 

specimens which could reflect distinctions at the level of populations or even species, but 

the general background variation has yet to be assessed over broad geographic areas. These 

differences suggest the possibility of vicariant subdivision of a circumboreal ancestor when 

North America and Asia were separated during the Pliocene. Transcontinental distribution 

across central North America suggests that, at least in part, this species occurred to the south 

of the ice during glaciation. 

Apataniidae 

Apatania crymophila McLachlan (41); A. stigmatella (Zetterstedt) (42); A. zonella (Zetterstedt) 

(43) 

These are the only Holarctic species known in Apatania, and all 3 could have passed 

the last glaciation in Beringia. However, transcontinental penetration of North America 

seems likely to have been achieved through postglacial dispersal from refugial areas south 

of the ice mass, implying that at least A. stigmatella and zonella occurred in North America 

before the Pleistocene, and thus probably before the Pliocene separation between North 

America and Asia. Apatania crymophila was proposed as the sister species of the European 

A. wallengreni McLachlan, and A. stigmatella the sister species of A. shoshone Banks in


western North America (Schmid 1954a). Apatania zonella shows rather distant relationship 

with other species (Schmid 1954a), and is parthenogenetic (Corbet 1966). Recorded 

from Ellesmere Island and Greenland, A. zonella is the most northerly of Nearctic 

Trichoptera. 


Brachycentrus americanus Banks (44) 

This species occurs through northern and western North America, and its range has been 

extended to include Siberia, Mongolia, and Japan with recognition of Oligoplectrodes 

potanini Martynov as a junior subjective synonym (Schmid 1983); it has been recorded also 

from Kamchatka, but not Chukotka (Levanidova 1982). This is the only Nearctic species 

assigned to the subgenus B. (Oligoplectrodes) (Flint 1984); 2 other species occur in 

Asia—B. kozlovi Martynov and B. punctatus Forsslund. From illustrations of the male 

genitalia of these species, it appears that the 2 Asian taxa are sister species, together 

constituting the sister lineage of B. americanus. 

We have examined specimens from much of the range in North America [159 #, 

220 !] and also material from Siberia, Mongolia, and Japan; a unique characteristic of 

B. americanus is the high variability of tibial spurs in North American populations. The 

typical and plesiomorphic tibial spur complement for all Brachycentrus species is 2,3,3; but 

in populations of B. americanus in Asia and Alaska tibial spurs are largely consistent at 2,2,3 

(Fig. 17). Most Yukon specimens also have spur counts of 2,2,3; although approximately 

20 per cent of those examined have 2,2,2, and occasionally the full complement of 2,3,3, 

with some spurs reduced in size. Regional variation representing all 3 spur conditions of 

B. americanus was found in varying proportions over much of North America to the south; 

in Manitoba, northern Ontario, and in Utah and Colorado the plesiomorphic 2,3,3 condition 

occurred in most specimens examined. Assuming genetic control for this variation, the 

parsimonious interpretation for the pattern of character distribution is that B. americanus 

arose in North America where the plesiomorphic spur count of 2,3,3 occurs. Uniformity in 

northeastern Asia for the reduced spur condition of 2,2,3 indicates a pervasive founder effect 

from an apomorphic ancestor that probably came from North America. In Nearctic Beringian 

populations isolated by encroaching glaciers, tibial spurs could have stabilized at 2,2,3, with 

colonizers from these populations entering Asia by way of rivers of the Bering land bridge. 

Occurrence of B. americanus in the northern Yukon (Fig. 1, region 4: Porcupine Plain) 

indicates that it is adapted to conditions that would have been available in trans-Beringian 

dispersal during the Pleistocene. The main Nearctic body of B. americanus south of the 

continental ice mass would have been characterized by plesiomorphic tibial spurs of 2,3,3 

(Fig. 17). Distribution for B. americanus in North America (Flint 1984, fig. 23) suggests 

that disjunct populations occur in the northeast, in Wisconsin, Minnesota and Michigan, and 

in the western mountains. These disjunctions may be maintained now by the Great Lakes 

and by extensive areas where the cool streams required by larvae of this species are lacking. 

The disjunctions could be a consequence of separate refugia during glacial advances, 

restricting gene flow and enabling differing numbers of tibial spurs to become established 

in different areas. Following retreat of the ice, renewed contact between the Nearctic 

populations could have led to the mixture of spur counts against a strong background of the 

plesiomorphic 2,3,3 that now characterizes populations of B. americanus in North America 

(Fig. 17). The high incidence of the reduced 2,2,3 condition in western North America might 

also be a result of southward dispersal of Beringian populations. Based on life-history data 

from the interior of Alaska, a life cycle of 2 years was inferred (Irons 1988).


FIG. 17. Approximate distribution of character states for tibial spurs in Brachycentrus americanus (Banks) (44) 

(Brachycentridae).


Micrasema gelidum McLachlan (45) 

An intensive taxonomic analysis of this species complex was carried out by 

BotojAneanu (1988), who concluded from a study of male genitalic morphology that 

M. gelidum represents a polytypic superspecies of 6 prospecies, allopatrically distributed 

over a large part of northern Eurasia and North America. Three of the prospecies are 

represented in the Yukon and Alaska and in the Far East of Russia: M. (gelidum) gentile 

McLachlan (junior synonym M. scissum McLachlan, Kimmins and Denning 1951); 

M. (gelidum) extremum BotojAneanu; and M. (gelidum) gelidum McLachlan which also 

extends across the whole of northern Eurasia to Norway (junior synonyms: M. kluane Ross 

and Morse 1973; M. subscissum Martynov; M. sibiricum Martynov). Because M. gelidum 

is a member of a Nearctic group comprising M. bactro Ross, M. diteris Ross, M. onisca 

Ross, and M. sprulesi Ross, its origin was believed to have been in western North America 

(BotojAneanu 1988); and the mosaic of prospecies constituting M. gelidum was attributed 

to the isolation of populations during Pleistocene glacial periods and their recolonization 

following glacial retreat. Complexity within the superspecies M. gelidum was interpreted to 

indicate that glacial intervals preceding the last Wisconsinan period were effective in 

isolating populations for periods sufficient to effect morphological divergence. One form in 

the complex, M. (gelidum) canusa BotojAneanu (1988), occurs in central North America, 

and probably was isolated along the southern margin of the glacial ice; but the more northerly 

Nearctic forms were inferred to have passed periods of glacial advance in the Beringian 

refugium. As with species of Apatania, larval habitats for members of the gelidum complex 

broaden with increasing latitude—those in the most southerly localities living in spring 

streams, but northward habitats include rivers and also lakes in the most northerly localities 

(BotojAneanu 1988). Consequently larval habitat probably would not have been a 

deterrent to dispersal of the northern populations across the Bering land bridge during the 

Pleistocene, an inference further supported by occurrence of M. gelidum on the Arctic coastal 

plain (Fig. 1, region 1). 

Goeridae 

Goera tungusensis Martynov (47) 

The range of this species known originally from Siberia must now be extended to 

northern Quebec, the Northwest Territories, and almost certainly the Yukon (see Taxonomic 

Note 2). We also have from Alaska a pharate male that may represent a morphological 

variant of this species, or perhaps an unnamed sister taxon. If Pleistocene dispersal of 

G. tungusensis across the Bering land bridge to North America is indicated, the species 

could have extended eastward in North America after withdrawal of the glaciers. However, 

the widely scattered North American records suggest the possibility of repopulation 

of deglaciated areas from south of the ice where a pre-Pliocene circumboreal 

ancestor took refuge, and from which a disjunct Alaskan population has begun to 

diverge. 

Leptoceridae 

Ceraclea annulicornis (Stephens) (54) 

Phylogenetic analysis (Yang and Morse 1988) indicated that C. annulicornis is one of 

an unresolved group of 4 species—2 in China, and C. ruthae (Flint) from eastern North 

America. Species that are now transcontinental in North America, such as C. annulicornis, 

probably passed the Pleistocene to the south of the ice mass and if so, would have occurred 

in North America before glaciation.


Ceraclea excisa (Morton) (†) 

Although not known from the Yukon, this species probably occurs there because it is 

recorded from Alaska (Milne 1934; Morse 1975), Wisconsin, and Michigan to Quebec and 

Massachusetts; and it extends through northern Europe and Siberia to the Amur region 

(Lepneva 1966). It is considered to be the sister taxon of the group of 4 species that includes 

C. annulicornis (Yang and Morse 1988); its widespread occurrence in North America leads to 

the same interpretation as for the preceding species. 

Ceraclea nigronervosa (Retzius) (56) 

Although most species of the nigronervosa group are confined to North America (Morse 

1975), C. nigronervosa is the sole Holarctic species. Larvae live in large rivers, where they 

feed on colonies of freshwater sponge (Resh 1976; Solem and Resh 1981). In addition to 

typical nigronervosa specimens that are consistent with most others examined from Europe 

and Asia, our North American material reveals a brown-winged variant with distinctive male 

genitalic characters (Fig. 4) which is also represented in Europe (Taxonomic Note 3). Thus 

the 2 forms appear to be broadly sympatric and could have reached North America during 

the Pleistocene by way of unglaciated Beringia, and spread southward following glacial 

retreat. This interpretation would require that sponges, the food of the larvae, occurred in 

waters of the Bering land bridge; and this is feasible because C. nigronervosa now occurs 

in the far northern Yukon (Fig. 1, region 4). However, this species might have been in North 

America before the Pliocene separation from Asia because the range of C. nigronervosa now 

extends to Wyoming. 

Oecetis ochracea (Curtis) (63) 

This is the only Holarctic species in Oecetis, and no phylogenetic analysis of the genus 

has been made. The subspecies O. o. carri Milne (1934) was segregated to discriminate 

between North American and European populations by genitalic characters. We found no 

evidence to support this interpretation; variation in genitalic characters was evident, but 

showed no geographic correlation. However, specimens from northern Europe and the 

Yukon are darker brown in colour overall than those from other parts of North America. 

Although slight, this indication of genetic continuity in O. ochracea through Nearctic 

Beringia and Europe suggests that the light-coloured Nearctic populations were derived from 

ancestors forced to the south of the continental glacier; if so, this species probably was 

present in North America before the Pleistocene glaciation. 

Ylodes reuteri (McLachlan) (68) 

In discussing the Nearctic range of this species, Ross (1965, as Triaenodes griseus) 

alluded to morphological variants in the Rocky Mountains, the western part of the Northwest 

Territories, and the Hudson Bay area. Highly variable populations in southern Saskatchewan 

were interpreted as an intergrading blend from all 3 areas—an example of caddisfly species 

that passed the last glaciation in discrete areas in the Rocky Mountains, in eastern North 

America, and in Beringia. If so, Y. reuteri or its ancestor must have occupied North America 

early enough, before Asia and North America were separated in the Pliocene, to have become 

established to the south of the glaciers. 

Limnephilidae 

Asynarchus iteratus McLachlan (74) 

This species was assigned to the lapponicus group (Schmid 1954b), and was considered 

to be so close to A. aldinus (Ross) from Alberta that the two could be geographic variants 

of the same species. The close similarity suggests that separation between them might have


been brought about by segregation of a portion of the Nearctic population of A. iteratus to 

the south of the advancing glaciers, giving rise to A. aldinus, while A. iteratus became 

confined to Beringia as a glacial relict. If that were so, A. iteratus or its ancestor probably 

reached North America from Asia before the land bridge between the 2 continents was 

overrun by the sea during the Pliocene. 

Asynarchus lapponicus (Zetterstedt) (75) 

Variation in the superior and intermediate appendages of the males showed no coherent 

pattern correlated with geographic distribution. Among the species assigned to the lapponicus 

group (Schmid 1954b), the sister species of A. lapponicus would be the northern North 

American species A. montanus Banks. Prominent morphological differences in the genitalia 

of the 2 species suggest that they have been separate for some time, perhaps through 

inter-continental vicariant speciation, with A. lapponicus dispersing from Asia to North 

America across the Bering land bridge during the Pleistocene. Disjunct distribution between 

northern and central populations of this species in Europe (Malicky 1988, fig. 11) suggests 

that it was widely distributed in Europe and Asia before the glacial advances of the 

Pleistocene. If A. lapponicus was confined to Beringia during Pleistocene glaciation, it has 

spread widely in North America following recession of the ice, in contrast to some other 

Nearctic Beringian species (see category III). The larval habitat for A. lapponicus is littoral 

areas of lakes, tundra pools, and slow streams (Winchester 1984), and the species is 

univoltine at the latitude of Tuktoyaktuk, Northwest Territories (69°29′N). 

Limnephilus dispar McLachlan (97); L. externus Hagen (98); L. femoralis Kirby (100); 

L. nigriceps (Zetterstedt) (108); L. picturatus McLachlan (113); L. rhombicus (Linnaeus) 

(114); L. sericeus (Say) (117). 

Phylogenetic relationships within the large genus Limnephilus have not been investigated 

in sufficient depth to support inferences about the geographic origin of the species. 

All of these species have a transcontinental distribution in North America, and probably 

occurred on this continent before glaciation in the Pleistocene. We found morphological 

variation in several of the species, but a congruent pattern was evident only in L. picturatus. 

Comparison of specimens of L. picturatus from all parts of the range in North America 

[64 #, 85 !] with Eurasian material indicates genetic continuity between Nearctic Beringia 

and Eurasia, and restriction in gene flow between Beringian populations and those from 

other parts of North America. This argues for isolation of Nearctic populations of L. picturatus 

south of the glaciers while Palaearctic immigrants of the species entered Beringia. 

Accordingly, L. picturatus probably would have occurred in North America before the 

Pleistocene, and its Holarctic distribution would pre-date the Pliocene separation between 

North America and Asia. On melting of the glaciers, the more southerly Nearctic populations 

appear to have been more successful in colonizing deglaciated territory than have the 

Beringian populations. Analysis of Greenland populations for these variable characters 

might shed light on their origin. 

Thus, the patch of stout setae on the ventral surface of the hindwing R2 in males is well 

developed in all Eurasian specimens examined and in about 95 per cent of Beringian 

specimens, but equally well developed in only about one third of other North American 

specimens. The pterostigma is light in colour in 90 per cent of Beringian and Palaearctic 

specimens examined, but light in about 50 per cent of other North American specimens. 

Inferior appendages in the male genitalia are triangular in dorsal aspect in about 86 per 

cent of Nearctic Beringian specimens; about 4% of specimens from other parts of North 

America are similar in this character, where the predominant condition is for parallel dorsal


margins on the inferior appendages. Eurasian specimens examined are, however, about 

evenly divided between these 2 conditions. In the parameres of the male genitalia of Nearctic 

Beringian specimens there are more than 10 long, very fine, setae on the dorsal preapical 

lobe and the setae on the mesal surface extend anterad of the dorsal process. In specimens 

from other parts of North America there are fewer than 10 setae on the dorsal lobe of the 

parameres and the setae are coarser, as they are also in Eurasian specimens examined; but 

the fine setae on the mesal surface in Eurasian material extend anterad of the dorsal lobe as 

in Beringian specimens. 

In females, the dorsal margin of segment IX is truncate in 98 per cent of Nearctic 

Beringian specimens examined and also in Eurasian specimens, differing from the rounded 

condition found in 87 per cent of North American populations generally. Segment X in dorsal 

aspect is bifid in only about 17% of Nearctic Beringian females examined, but in 90% of 

specimens from elsewhere in North America; in most Nearctic Beringian specimens the apex 

of X is blunt or pointed rather than bifid. 

This species is univoltine at Tuktoyaktuk, Northwest Territories, where larvae live in 

tundra ponds (Winchester 1984). 

Onocosmoecus unicolor (Banks) (122) 

It seems likely that Onocosmoecus is part of a complex of dicosmoecine genera that 

arose in North America (Wiggins and Flint in prep.). Marked variation in this species is 

confirmed by no fewer than 6 synonyms, but analysis of specimens from many localities 

revealed no congruent geographic pattern (Wiggins and Richardson 1987). We infer that 

some populations passed the Pleistocene glaciation south of the ice in North America because 

the only other species known in the genus, O. sequoiae Wiggins and Richardson, is confined 

to the Sierra Nevada Mountains of California and may have originated there as a glacial 

disjunct. Widespread occurrence of variable populations of O. unicolor in North America 

from British Columbia to Newfoundland indicates that other populations of this species 

extended across a broad front to the south of the glaciers, and that O. unicolor occurred in 

North America before Pleistocene glaciation began. This leads to the further possibility that 

other populations of O. unicolor in the Beringian refugium dispersed to eastern Asia across 

the Bering land bridge during the Pleistocene glaciation. Larvae of this species live in cool 

waters of slow streams and the littoral zone of lakes, habitats that would have been readily 

available on the Bering land bridge; larvae in streams in interior Alaska fed entirely on plant 

detritus (Irons 1988). 

Molannidae 

Molanna flavicornis Banks (127) 

This is the only North American species of Molanna with a transcontinental distribution. 

It is very similar to and is perhaps identical with the Eurasian Molanna albicans Zett., 

resulting in a circumboreal range through northern Eurasia and North America (Fuller 1987). 

The albicans group of several species evidently arose in Eurasia and only this single Nearctic 

extension now exists. Montane populations in Colorado are interpreted as glacial relicts, 

indicating that this species ranged widely to the south of North American glaciers during the 

Pleistocene, and thus its Holarctic distribution would have been established before Asia and 

North America were separated in the Pliocene. Larvae live on the bottom of cool lakes. 

Molannodes tinctus Zetterstedt (128) 

Collections of this northern Eurasian species in Alaska and the Yukon have led to the 

entrenched view that it was a glacial relict confined to the Beringian refugium. In recent


years, however, isolated collections of the species have been made in Saskatchewan and 

northern Ontario (Fuller 1987). Based on the broad distribution of M. tinctus in northern 

Europe and Asia, and on phylogenetic evidence for the origin of Molannodes in Asia (Fuller 

1987), an eastward dispersal by way of the Bering land bridge is inferred. Evidence is 

equivocal as to whether dispersal occurred in Pleistocene time or earlier. 

Phryganeidae 

Agrypnia colorata Hagen (129) 

Material from Asia has been referred by most authors to A. principalis (Martynov), but 

this name is a junior subjective synonym of A. colorata Hagen (Wiggins in press). All 

Beringian specimens examined and all from the Palaearctic portion of the range are dark in 

colour on the thorax, legs, and wings; and all have fully developed tibial spurs of 2,4,4, 

typical for the family Phryganeidae (Fig. 18). The single exception in Palaearctic material 

that we examined is a light-coloured female from the Keriya River area of China (Sinkiang 

Prov.; ZMAS), although the tibial spurs were fully developed. In contrast, most North 

American specimens taken outside Beringia are light in colour on the thorax, legs, and wings, 

and have the tibial spurs reduced in some way. The pattern of reduction is variable—some 

spurs may be absent, others reduced to tiny knobs; and in any pair of spurs, one might be 

reduced or lacking and the other normal; opposite members of a pair of legs often have 

different spur conditions. Exceptions occur in series from Fort McPherson, Northwest 

Territories and Kamloops, British Columbia (ROME), in which specimens are dark as in 

Beringian and Palaearctic material, but the tibial spurs are variable as in most North 

American populations. No congruent variation was found in genitalic characters. [Specimens 

examined: N. Am.—58 #, 55 !] 

The sister species of A. colorata is A. legendrei (Navas) known only from China. We 

interpret the light-coloured A. colorata to be a vicariant Nearctic form derived from dark 

ancestral stock with unmodified spurs following separation of Asia from North America, 

perhaps in Pliocene time. Nearctic populations would have been isolated to the south of the 

Pleistocene glaciers when light colour and unstable tibial spurs could have been established; 

disjunct montane populations of the light-coloured form in Wyoming and Utah are consistent 

with this interpretation (Wiggins in press). The plesiomorphic dark Palaearctic form with 

stable tibial spurs would have dispersed to Nearctic Beringia across the land bridge. 

Following retreat of the ice, contact between the 2 forms in North America could have given 

rise to intergrading populations such as those near Kamloops and Fort McPherson. 

Agrypnia pagetana Curtis (135) 

No close relative of this species is known (Wiggins in press). We infer that A. pagetana 

is a Eurasian species that reached North America across the Bering land bridge, perhaps 

during the Pleistocene, and dispersed through northern North America following recession 

of the glaciers. It has not been recorded east of Hudson Bay. Populations of A. pagetana in 

Europe range farther south to more temperate climates than in North America. Larvae of this 

species live in small tundra ponds and slow-flowing streams; they are univoltine at 

Tuktoyaktuk, Northwest Territories (lat. 69°29′N) (Winchester 1984). 

III. Palaearctic-East Beringian Species 

Because of their broad Eurasian range and restricted North American distribution, these 

species are inferred to be of Palaearctic origin. Dispersal to North America across the Bering 

land bridge during the Pleistocene seems the most likely route for a number of these species. 

The 13 species in category III constitute about 10 per cent of the Yukon fauna.


FIG. 18. Global distribution of variant characters in Agrypnia colorata Hagen (129) (Phryganeidae). Dark-coloured 

adults with tibial spurs constant at 2,4,4 occur throughout northern Eurasia and Beringia; light-coloured adults with 

tibial spurs variably reduced occur through most of North America outside Beringia; intermediate dark specimens 

with variable tibial spurs occur in northwestern North America between the two areas outlined. 

Failure of these East Beringian species to broaden their postglacial Nearctic range 

perhaps could be attributed to competition for resources from species advancing from the 

south which were better adapted to the freshwater habitats of the deglaciated terrain, or in 

any case from species that reached the new habitats first. As discussed previously, closure 

of Beringia by conjunction of the Laurentide and Cordilleran ice sheets probably would have 

caused some extinction in Nearctic Beringia; consequently a wider range of ecologically 

coordinate species from the much larger, and biologically more diverse, West Beringia 

would have encountered reduced competition when they dispersed to East Beringia. But this 

advantage of the Palaearctic species might well have been less effective in postglacial time 

when Nearctic species were reassembled in more tightly packed communities. For these or

other reasons, most Palaearctic-East Beringian species of category III have functioned 

neither as aggressive colonists nor ecological generalists in North America; but quite clear 

is the contrast with Europe and Asia where most of them are widely distributed, and 

apparently are successful generalists. It is one of the striking paradoxes of Beringian distributions 

that wide-ranging, and evidently competitively successful, Eurasian species remain 

confined to their East Beringian outpost, evidently unable to disperse much beyond their 

former glacial refuge. This distributional paradox can be added to the productivity paradox 

(e.g. Hopkins et al. 1982) as significant questions about the biological history of Beringia. 

This issue focusses on species of category III because they may differ from category II 

essentially in lacking the competitive ecological edge required in new communities; 

Palaearctic species having that competitive edge now meet the distributional criterion of 

category II. 

For several of them (Ylodes kaszabi, Arctopora trimaculata, Grammotaulius signatipennis, 

Limnephilus stigma, and Agrypnia obsoleta), sister-group relationships suggest 

intercontinental vicariance, perhaps during the Pliocene or earlier, followed later by dispersal 

of the Palaearctic form to East Beringia, probably across the Pleistocene land bridge. 

In Agraylea cognatella and Limnephilus fenestratus, morphological similarity to their 

respective Nearctic sister species is so close that dichotomy during the Pleistocene could be 

reasonably inferred. Subdivision of the range by glacial ice seems likely, with the Nearctic 

sister species originating in isolation to the south of the glaciers. 

Spicipalpia 

Hydroptilidae 

Agraylea cognatella McLachlan 


Rhyacophila mongolica Schmid, Arefina 

and Levanidova 

Integripalpia 

Leptoceridae 

Ylodes kaszabi Schmid 

Hydroptilidae 

Agraylea cognatella McLachlan (4) 

The Palaearctic range of A. cognatella appears to be circumscribed by A. multipunctata 

Curtis which is widely distributed through Europe and western Asia (BotojAneanu and 

Levanidova 1988); in North America, A. cognatella is evidently circumscribed to the south 

of Beringia, not by A. multipunctata as has been the traditional interpretation (e.g. Ross 

1944), but by a sibling species A. fraterna Banks apparently now widely distributed on this 

continent (Vineyard and Wiggins in prep.). It seems reasonable to infer from present 

evidence that A. cognatella is a Palaearctic-East Beringian species now confined in North 

America to the refugium, and that A. fraterna passed the glacial period to the south of the 

ice where it may have arisen. 


Rhyacophila mongolica Schmid, Arefina and Levanidova (14) 


Limnephilidae 

Arctopora trimaculata (Zetterstedt) 

Dicosmoecus obscuripennis Banks 

Grammotaulius signatipennis McLachlan 

Grensia praeterita (Walker) 

Limnephilus diphyes McLachlan 

Limnephilus fenestratus (Zetterstedt) 

Limnephilus stigma Curtis 

Phryganeidae 

Agrypnia obsoleta (Hagen) 

Agrypnia sahlbergi (McLachlan) 

Oligotricha lapponica (Hagen)


FIG. 19. Distribution of Arctopora species (71, 72) (Limnephilidae); symbols superimposed indicate occurrence of 

both species at the same locality. 

In phylogenetic analyses of Rhyacophila, some 25 species from eastern and western 

North America, Siberia, Japan, and Europe have been assigned to the sibirica species group 

(Ross 1956; Schmid 1970). This group forms the major component of Rhyacophila in 

Palaearctic Asia, and includes the only 2 Holarctic members of the genus known to 

date—R. narvae and R. mongolica (Taxonomic Note 1). Rhyacophila mongolica is considered 

the sister species of R. sibirica McLachlan (Schmid et al. 1993). Among 14 species of 

Rhyacophila occurring in the Yukon, R. mongolica (Fig. 2) is recorded much farther north 

than any of the others (Fig. 1, region 4), indicating that the species was ecologically adapted 

for dispersal to North America across the Bering land bridge during the Pleistocene. 

Leptoceridae 

Ylodes kaszabi (Schmid) (67) 

The genus Ylodes appears to have originated in north-central Asia (Manuel and Nimmo 

1984), and Y. kaszabi occurs in Mongolia. A sister-species relationship with the Nearctic 

Y. schmidi suggests the possibility of intercontinental vicariant origin during the Pliocene 

or perhaps earlier; thus Y. kaszabi is inferred to have been a Palaearctic species that later 

reached North America by way of the Bering land bridge during the Pleistocene. 

Limnephilidae 

Arctopora trimaculata (Zetterstedt) (72)


In addition to A. trimaculata, 2 North American sister species are recognized in 

Arctopora: A. pulchella (Banks) (71), transcontinental from Yukon and Alaska to Newfoundland 

and New Hampshire; and A. salmon (Smith) in Idaho (Fig. 19). The evidence 

suggests vicariant speciation from a circumboreal common ancestor, followed by derivation 

of the North American species pair from subdivision of the range, possibly by glacial ice. 

This interpretation leads further to Pleistocene dispersal of A. trimaculata from Asia to North 

America across the Bering bridge; but the species remained confined to East Beringia 

following deglaciation, perhaps because A. pulchella was better adapted for colonizing the 

new habitats. In any case, the 2 species are now geographically sympatric in East Beringia 

(Fig. 19). 

Dicosmoecus obscuripennis Banks (81) 

Dicosmoecus obscuripennis is the only member of the Palaearctic palatus species group 

recorded from North America (Wiggins and Richardson 1982). Study of additional collections 

provided by I.M. Levanidova (Vladivostok) revealed that this species is actually 

widespread in eastern Russia—a situation further investigated by Nagayasu and Ito (1993); 

Asian specimens examined are larger, the length of male forewings 20 – 25 mm, in contrast 

to 18 – 20 mm for North American specimens. If this species reached North America from 

Asia during the Pleistocene, passage over the Bering land bridge might not have been wholly 

dependent upon the clear running waters to which larvae of species of Dicosmoecus are 

usually confined because larvae of D. obscuripennis have been found along lake margins in 

the Russian Far East (I.M. Levanidova, pers. comm.). After retreat of the glaciers, D. obscuripennis 

appears not to have penetrated North American habitats much beyond its 

glacial-age Beringian range. Possibly a limiting factor in North America is the widespread 

occurrence of D. atripes (80), an ecologically vigorous species of the western montane region 

ranging from California to Yukon and Alaska, which appears to have reached Beringia from 

the south following glacial recession (Fig. 20). 

Grammotaulius signatipennis McLachlan (86) 

This is the sister species of G. alascensis Schmid recorded from northern North America 

(Taxonomic Note 6). From the evidence available, G. signatipennis is probably a Palaearctic 

species, extended in range to North America across the Bering land bridge during the 

Pleistocene. 

Grensia praeterita (Walker) (87) 

This is one of the very few truly arctic species of Trichoptera, and it probably occurred 

in Beringia, at least in part, during glaciation. In contrast to most of the other species assigned 

to category III, G. praeterita has extended its postglacial range considerably beyond Beringia. 

Evidently, the species does not occur on the mainland of North America east of Hudson 

Bay (Harper 1989), indicating that its dispersal to Greenland was probably by way of the 

Arctic Archipelago. Comparison of Yukon specimens with material from the Northwest 

Territories and Russia revealed no morphological differences, although the latter were 

slightly larger. 

Limnephilus diphyes McLachlan (96) 

This species is recorded from North America for the first time (Taxonomic Note 7). Its 

affinities are obscure, but since it has not been collected south of Alaska and the Yukon, the 

evidence suggests that L. diphyes is a Palaearctic species that reached North America during 

the Pleistocene and has not spread beyond East Beringia. Larvae of L. diphyes live in 

sphagnum bog pools (Johansson et al. 1991).


FIG. 20. Distribution of Dicosmoecus atripes (Hagen) (80) and D. obscuripennis Banks (81) (Limnephilidae); the 

latter is widely distributed in eastern Asia.


FIG. 21. Distribution of the sister species Limnephilus kennicotti Banks (107) and L. fenestratus (Zetterstedt) (101) 

(Limnephilidae); symbols superimposed indicate occurrence of both species at the same locality. 

Limnephilus fenestratus (Zetterstedt) (101) 

The very similar sister taxon of L. fenestratus is L. kennicotti (107) (category I), which 

occurs widely through northern North America and in Greenland (Fig. 21). We have 

identified both taxa in separate collections from the Yukon (Kluane and Firth River). The 

two are most readily separable by male genitalic structure: in L. fenestratus the intermediate 

appendages are relatively short and do not extend beyond the superior appendages in lateral 

aspect, but are longer in L. kennicotti and do extend beyond the superior appendages in lateral 

aspect. Diagnostic characters for males and females of both species were illustrated by 

Kimmins and Denning (1951, figs. 10, 11, as L. miser McLachlan which is a junior subjective 

synonym of L. fenestratus proposed by Schmid 1955; and figs. 12, 13, as L. moselyi Kimmins 

and Denning from Greenland which is a junior subjective synonym of L. kennicotti proposed 

by Ross and Merkley 1952). 

We presume that records of L. fenestratus from Greenland by Fristrup (1942), repeated 

by Gislason (1981), were based on Mosely’s (1929) misidentification of L. kennicotti as 

L. miser (= fenestratus); a series of specimens we examined from Greenland (Zoologisk 

Museum, Copenhagen, per N.P. Kristensen) proved to be L. kennicotti. From the Yukon, 

Nimmo and Wickstrom (1984) do not record L. fenestratus, but provide several records for


L. kennicotti; we have confirmed that their material from Kluane National Park and Firth 

River is L. kennicotti, but that from Burwash Landing is really L. fenestratus. In the Yukon 

material examined, we found some indication that male genitalic characters intergrade 

between these 2 taxa. Moreover, putative distinction between females (Kimmins and 

Denning 1951) has proven unreliable because of variation in genitalic characters, culminating 

in our material in 2 females collected with a male of L. fenestratus (Yukon: Old Crow 

Flats, ROME 810587a); one female shows characters attributed to L. kennicotti, the other to 

L. fenestratus. Further evidence on variation is required before the status of these taxa can 

be confirmed. [North American specimens examined: L. fenestratus 26 #, 29 !; L. kennicotti 

18 #, 32 !]. 

Whether or not these 2 forms are distinguished as species, their distributional relationships 

are informative in a Beringian context. If the very close morphological similarity 

between the 2 taxa is interpreted to indicate separation during the Pleistocene, L. kennicotti 

could have diverged as a disjunct population along the southern margin of the North 

American glaciers. By this interpretation, L. fenestratus would have been the Holarctic 

ancestor, and its present East Beringian population would be a glacial relict. As the ice 

receded, widespread recolonization by L. kennicotti over northern North America evidently 

led to its dispersal to Greenland, approaching the western limit of its ancestral stock 

represented by L. fenestratus in Iceland (Fig. 21). Judging from material we have examined, 

L. fenestratus has extended its Nearctic range little if at all beyond unglaciated Beringia; 

L. kennicotti has shown marked capacity for colonizing deglaciated habitats, evidently far 

into the northern Yukon where the 2 forms are now apparently sympatric. Some intergradation 

in the morphological characters distinguishing the two (see above) suggests that this 

postglacial sympatry has yet to reach some equilibrium. 

Limnephilus stigma Curtis (118) 

Records for the close sister species L. indivisus Walker, now widespread over much of 

North America, approach the Nearctic limit of L. stigma in the Northwest Territories (Fort 

Smith, CNCI) and northern British Columbia (Alaska Hwy. km 359, Prophet R. Prov. Park, 

ROME). Specimens from Kluane in the Yukon were identified as L. indivisus by Nimmo 

and Wickstrom (1984), but our examination of that same material indicates that they are 

L. stigma. In the continued absence of intermediates, we infer that the 2 species do not 

hybridize. It has long been recognized that L. stigma (Figs. 22 – 25) is very similar morphologically 

to the widespread North American species L. indivisus Walker (e.g. Betten and 

Mosely 1940). Males of L. indivisus are distinguished by the much longer intermediate 

appendages and by the ventral gap in the mesal dentation of the superior appendages 

(Fig. 26); females of L. indivisus are distinguished by the narrow and deeply incised apex 

of segment X, especially evident in ventral aspect (Fig. 27). 

Material of L. stigma from Yukon and Alaska [26 #, 24 !] differs slightly in genitalic 

characters from specimens examined from Europe and northern Asia [18 #, 16 !]. In East 

Beringian males (Fig. 24b), the peripheral dentate ridge on the superior appendages bears a 

large point underlying the intermediate appendages which is lacking from European specimens 

(Fig. 22b). Males in our Nearctic material of L. stigma bear a prominent sclerotized 

point on each paramere (Fig. 24d); this sclerotized point is lacking in specimens we examined 

from Europe (Fig. 22c; and Malicky 1983, p. 188), but is present in specimens examined 

from Kamchatka. In East Beringian females (Fig. 25), the apex of segment X is more 

narrowly tapered than in Eurasian material (Fig. 23), especially in ventral aspect.


FIGS. 22 – 25. Limnephilus stigma Curtis (111) (Limnephilidae). 22, Male genitalia of specimen from Europe: a, 

lateral; b, caudal; c, phallus, lateral; 23, Female genitalia of specimen from Europe: a, lateral; b, dorsal; c, ventral; 

24, Male genitalia of specimen from Yukon: a, lateral; b, caudal; c, superior appendage, mesal; d, phallus, tip of 

paramere; 25, Female genitalia of specimen from Yukon: a, lateral; b, dorsal; c, ventral.


FIGS. 26 – 27. Limnephilus indivisus Walker (Limnephilidae), specimens from Ontario. 26, Male genitalia: a, lateral; 

b, caudal; c, superior appendage, mesal; d, phallus with detail of tip of paramere; 27, Female genitalia: a, lateral; 

b, dorsal; c, ventral. 

The sister species could have arisen through vicariant speciation of the circumboreal 

range of a common Pliocene ancestor, giving rise to L. stigma in Eurasia and L. indivisus in 

North America; during the last glacial advance, L. indivisus probably would have been 

restricted to the south of the ice, while L. stigma could have entered the East Beringian 

refugium but did not extend its range. Larvae of these species inhabit small marshy water 

bodies, including temporary pools (Wiggins 1973). 

Phryganeidae 

Agrypnia obsoleta (Hagen) (134) 

We have material of this widely distributed Eurasian species from the northern Yukon. 

Its Nearctic sister species, A. deflata (Milne) (130), is common over much of northern and 

western montane North America; although designated as a subspecies A. obsoleta deflata 

by some authors (e.g. Milne 1934; Fischer 1964; Nimmo and Wickstrom 1984), the evidence 

does not support this interpretation (Wiggins in press). Specimens from British Columbia 

identified as A. obsoleta (Nimmo and Scudder 1983: Glacier Nat. Park, 1# 1!) have been 

examined, and are A. deflata. 

Two successive events seem to have been involved: vicariant subdivision of the 

circumboreal ancestor from which the sister species A. obsoleta and deflata were derived in 

Eurasia and North America respectively, perhaps during the Pliocene or earlier; and 

subsequent Pleistocene dispersal of A. obsoleta to East Beringia during a recent glacial 

advance, perhaps with A. deflata restricted to the south of the Laurentide and Cordilleran 

glaciers. Outlying montane populations of A. deflata, for example in Colorado (Wiggins in


press), can be interpreted as relicts from this time. With recession of the ice, A. deflata now 

occurs in northern and montane North America to the Yukon and Alaska, but evidently 

A. obsoleta has dispersed little beyond Beringia. 

Agrypnia sahlbergi (McLachlan) (136) 

This species is widespread from Scandinavia through northern and eastern Asia. It is 

recorded here from the Yukon and also from Alaska, and recently from British Columbia 

(Nimmo and Scudder 1978). We infer that it reached this continent by way of the Bering 

land bridge during the Pleistocene, and evidently has dispersed southward to a limited extent. 

Oligotricha lapponica (Hagen) (139) 

All 5 species of Oligotricha are Palaearctic (Wiggins and Kuwayama 1971), but only 

O. lapponica extends to North America where records are confined to the Yukon and Alaska. 

From this evidence, we infer a Eurasian origin for O. lapponica, and it probably reached 

North America across the Bering land bridge during the Pleistocene. The forewings usually 

bear reticulate dark markings on a lighter background (Wiggins and Kuwayama 1971), but 

in some populations from northern Lapland of Sweden and Finland the wings are uniform 

dark brown—O. lapponica var. hyperborea (Forsslund 1933). Some specimens we have 

seen from East Beringia also have uniform dark brown wings, others have normal reticulate 

wings. The brown-winged form of O. lapponica in Europe and in East Beringia probably 

arose from independent changes in the genotype. This condition suggests a parallel with 

melanic forms that occur in some butterflies, apparently as they approach their ecological 

limits (Downes 1966); but in one species of the genus, O. striata (Linnaeus) of Europe, the 

wings are consistently uniform dark brown (Wiggins and Kuwayama 1971). 

This species is a good example of the Beringian distributional paradox—a widespread 

and successful species throughout Eurasia, but evidently still circumscribed in North 

America within the borders of unglaciated Beringia (cf. Asynarchus lapponicus (75), 

category II). Biotic factors responsible for this situation could include the closely related 

phryganeid Banksiola crotchi (Banks) (138)—a common and locally abundant transcontinental 

Nearctic species that is clearly aggressive ecologically (Wiggins 1956); Oligotricha 

and Banksiola are sister genera (Wiggins in press). Larvae of both genera live in shallow 

waters of lakes, marshes, and slow streams; consequently, suitable habitats that might 

otherwise accommodate O. lapponica in North America beyond Beringia are likely to have 

been pre-empted by B. crotchi or possibly by some other species in the Phryganeidae. 

Coupled with this is the probability of some genetic drift in the populations isolated in 

Nearctic Beringia, perhaps altering their competitive potential; some change in the genome 

can be inferred from reappearance of the uniform brown forewings. 

IV. Beringian Species 

These species share distributional and biological characteristics indicating that they 

were confined to Beringia during the Pleistocene glacial period. Five are Nearctic species 

recorded from the Yukon, Alaska, or the Northwest Territories. On present evidence these 

species appear to be East Beringian endemics or glacial relicts from a broader pre-Pleistocene 

range in North America. Two species, Lenarchus expansus and Asynarchus innuitorum, 

occur in both Nearctic and Palaearctic Beringia, but are not known elsewhere. Most of these 

species appear to be confined to arctic or alpine tundra, and are consistent with the general 

pattern in other groups in which the arctic species were derived from Beringia. One Nearctic 

species, Limnephilus pallens, is assigned provisionally to category IV because some records


are anomalous with the general pattern. Two species (†) not yet recorded from the Yukon 

may occur there. 

Integripalpia 

Leptoceridae 

Ylodes schmidi Manuel and Nimmo 

Apataniidae 

Allomyia picoides (Ross) (†) 

Limnephilidae 

Asynarchus innuitorum (Nimmo) (†) 

Grammotaulius alascensis Schmid 

Lenarchus expansus Martynov 

Limnephilus fumosus Banks 

Limnephilus pallens Banks 

Sphagnophylax meiops Wiggins and 

Winchester 

Leptoceridae 

Ylodes schmidi Manuel and Nimmo (69) 

This species is known only from the type locality in the Yukon, and was proposed as 

the sister species of Y. kaszabi (67) (Manuel and Nimmo 1984). Under that species (category 

III), it was suggested that Y. schmidi may have originated as the Nearctic vicariant from a 

common ancestor that occurred in both northwestern North America and eastern Asia. 

Apataniidae 

Allomyia picoides (Ross) (†) 

This species has not been recorded outside of Alaska (Katmai; Ross 1950); although its 

range may or may not extend to the Yukon, it can be considered a Nearctic (East) Beringian 

endemic or relict. 

Limnephilidae 

Asynarchus innuitorum (Nimmo) (†) 

Although this species has not been recorded yet from the Yukon, it was described from 

a tundra stream in the vicinity of Tuktoyaktuk, Northwest Territories, a short distance from 

the Yukon border (Fig. 1) (Winchester 1984: Limnephilus species A); and it has been found 

also across the Bering Strait in Chukotka (A.P. Nimmo, pers. comm.). Assigned originally 

to Limnephilus, this species was transferred to Asynarchus by Ruiter (1995:23-24). On the 

basis of existing information, we infer that A. innuitorum is a Beringian endemic, and 

probably also a glacial relict. 

Grammotaulius alascensis Schmid (84) 

This species appears to be the Nearctic sister species of the originally Palaearctic 

G. signatipennis (86) (category III). Present evidence suggests that G. alascensis passed 

glaciation in East Beringia, and dispersed eastward to Hudson Bay as the ice receded (See 

Taxonomic Note 6). 

Lenarchus expansus Martynov (90) 

Records for this species indicate that it is known only from the Beringian refugium in 

both Asia (Schmid 1952: Kolyma delta, Jana plains) and North America (Alaska, see also 

Nimmo 1986; Yukon, see above). In West Beringian specimens the forewings are light tan 

in colour with only faintly contrasting markings (Martynov 1914); but in all except one 

specimen (YT: Firth R.; CNCI) in the Nearctic material that we have seen, the dark markings 

of the forewing contrast strongly with the light base colour. Diversification in colour is 

probably a consequence of the isolation of the 2 populations. Since we found no morphological 

differences, the amphi-Beringian range could have been achieved by way of the


Pleistocene Bering land bridge. The species was assigned to the subgenus L. (Lenarchus) 

(Schmid 1952), the only Holarctic member of the group of 6 Nearctic and Palaearctic species. 

Although L. expansus is an amphi-Beringian species, its origin in either Asia or North 

America remains unresolved. 

Larvae of most Lenarchus live in standing waters, but those of L. expansus have been 

found in water-saturated tundra sod (MacLean and Pitelka 1971). Under these conditions, 

restriction of this species to Beringia is noteworthy because, although wet tundra habitat 

seems scarcely limiting now, it is thought to have been rare in Pleistocene Beringia, when 

much drier conditions prevailed (Schweger et al. 1982). Consequently, wet tundra must have 

been available in Beringia throughout the Pleistocene, at least in localized patches. (See also 

Sphagnophylax meiops.) 

Limnephilus fumosus Banks (102) 

Present information on distribution of the two very similar North American sister 

species, L. fumosus and L. santanus, suggests that they may be the products of populations 

that became disjunct during the formation of glaciers. If Limnephilus fumosus is confined to 

northern latitudes, it probably passed the Pleistocene glacial period in Beringia, although 

there is no evidence for dispersal to Asia across the Bering land bridge. Limnephilus santanus 

is known from Oregon and probably occurs in Washington (Banks 1900). Oregon was not 

covered by ice, but lies at the southern edge of the maximum extent of glaciation; thus records 

from British Columbia are critical in interpreting the postglacial history of these 2 species 

(see Taxonomic Note 8). All of this indicates that L. fumosus is probably a glacial relict 

confined to East Beringia. 

Limnephilus pallens Banks (109) 

This species was assigned to the asiaticus group (Schmid 1955) with 3 other North 

American and 8 Eurasian species. Judging from the male genitalic morphology (Nimmo 

1991), L. pallens appears to be more closely related to several of the Eurasian species, 

especially L. tricalcaratus Mosely (1936) from Tibet, than to North American species. 

Larvae of this species live in tundra ponds (Lehmkuhl and Kerst 1979), and thus its 

adaptation to far northern conditions during Pleistocene glaciation seems assured. If this 

species passed the glacial period in a northern refugium such as East Beringia, evidently it 

did not become established in West Beringia. If L. pallens is a Nearctic Beringian endemic 

species, it must have dispersed eastward to Hudson Bay (Lehmkuhl and Kerst 1979: Rankin 

Inlet) following glacial recession. However, a recent record for L. pallens from the Michigan 

shore of Lake Huron (Ruiter 1995) raises the possibility that this eastern extension could 

have been derived from the southern margin of the glaciers, as the ice retreated. Because of 

these discrepancies, assignment of Limnephilus pallens to category IV is provisional. 

Sphagnophylax meiops Wiggins and Winchester (126) 

This species (Frontispiece, Fig. 28) has been collected in the Yukon and in adjacent 

parts of the Northwest Territories (Aklavik, Tuktoyaktuk), but is otherwise unknown. 

Sphagnophylax is a monotypic genus, which on morphological grounds appears to represent 

an aberrant lineage in the limnephiline tribe Limnephilini (Winchester et al. 1993); and 

Sphagnophylax is the only trichopteran genus known that is confined to Beringia. Consequently, 

S. meiops would be both a phylogenetic and a geographic relict, apparently 

preserved from extinction only by the unglaciated Beringian refugium. The larval habitat of 

wet moss in transient tundra pools is significant because larvae of some other phylogenetically 

relict Trichoptera also occur in wet edaphic sites; a low incidence of competitors and


FIG. 28. Adult of Sphagnophylax meiops Wiggins and Winchester (126) (Limnephilidae). Brachyptery and 

anomalous venation suggest that this species is probably flightless, and perhaps was preserved from extinction only 

by the unglaciated Beringian refugium. Forewing length 4 mm. (From Canadian Journal of Zoology) 

predators in these sites may be a factor in the survival of relict Trichoptera (Wiggins 1984). 

Occurrence of this species in wet tundra is also interesting because the Beringian refugium 

for at least part of the Pleistocene is inferred to have been a region of low precipitation 

throughout the year and a predominantly dry upland region with little muskeg where 

Sphagnum was absent (Schweger et al. 1982). Survival of S. meiops, and also of Lenarchus 

expansus (see above) which evidently lives under similar conditions, indicates that areas of 

wet tundra persisted in Beringia. 

Reduced wings (Fig. 28) and anomalous venation suggest that this species has limited 

ability to fly and perhaps is even flightless; and the eyes are unusually small for adult 

Trichoptera. Similar modifications occur in a number of arctic Lepidoptera (Downes 1964). 

V. Holarctic Species Not in Beringia 

Because categories II, III and IV include almost all of the North American Trichoptera 

now recognized as occurring in Europe and Asia as well, it is useful for comparison to add 

here the few remaining Holarctic species. Six species are assigned to category V, and none 

has been recorded from the Yukon; most of them probably dispersed from one continent to 

the other under more equable conditions of climate some time before the separation of Asia 

and North America in the Pliocene. With Holarctic distribution established long ago, these 

species appear now to be disjunct relicts, still resistant to cladogenetic divergence.


Hydroptilidae 

Ithytrichia clavata Morton 

This species is widely distributed in North America from British Columbia to California, 

through Texas, Kansas, Oklahoma to Quebec and New Hampshire; it is widespread through 

northern and western Europe (L. BotojAneanu, pers. comm.). It is not now a far northern 

species in either North America or Europe, and would seem unlikely to have dispersed across 

the Bering land bridge during a glacial climate regime. Moreover, the larvae live in running 

water, a habitat used by only very few successful Pleistocene dispersants. This evidence 

suggests that Holarctic distribution was achieved before the Quaternary glacial periods 

occurred, and perhaps the Nearctic and Palaearctic forms have diverged to some extent not 

yet recognized, as in the Agraylea multipunctata complex (Vineyard and Wiggins in prep.). 

In any event, the name I. clavata was based initially on populations in New York (Ithaca), 

and further taxonomic resolution is focussed on European populations. 

Oxyethira mirabilis Morton 

The range of this species in northern Europe has been extended to eastern Canada with 

recognition of O. barnstoni Harper as a junior subjective synonym (Kelley 1984). The 

pattern of distribution, if now adequately understood, suggests a north Atlantic dispersal. It 

is the only European representative in the aeola group of O. (Oxytrichia), which is otherwise 

entirely confined to North and South America (Kelley 1984). 


Polycentropus picicornis Stephens 

This species occurs through most of Europe (BotojAneanu and Malicky 1978), Siberia 

and Mongolia to Kamchatka (Lepneva 1964; Mey and Dulmaa 1985). It is known locally in 

North America from the Northwest Territories to New Hampshire and may yet be recorded 

from the Yukon. Larvae live in small bodies of standing water and in slow currents of rivers 

(Lepneva 1964). This pattern of northern distribution suggests that P. picicornis could have 

passed between Asia and North America more recently than other species of category V, 

possibly during the Pleistocene. 

Psychomyiidae 

Psychomyia flavida Hagen 

This species was described from North America where it is transcontinental and 

abundant in running waters. The Asian P. composita Martynov was proposed as a junior 

subjective synonym of P. flavida (Schmid 1965b); consequently, a wide distribution through 

Siberia, Mongolia, and North America now has to be attributed to P. flavida. In this context, 

the absence of records of any Psychomyia from the Russian Far East (e.g. Levanidova 1982) 

is of interest. In North America, P. flavida has not been recorded in the extreme northwest, 

but only to the edge of treeline at Churchill, Manitoba (Lehmkuhl and Kerst 1979), and 

ranges from British Columbia to California and Nova Scotia to North Carolina. Thus, 

whatever its place of origin, dispersal of P. flavida across the Bering land connection between 

North America and Asia probably occurred in a climate more moderate than the Pleistocene 

glacial periods. 

Limnephilidae 

Grammotaulius betteni Hill-Griffin 

Known originally from Oregon, this species has also been recorded from China (Schmid 

1950a). Larvae live in slow streams and small marshy ponds (Hill-Griffin 1912), some of


which are probably temporary (Wiggins 1977); however, the species is not part of the 

northern Nearctic fauna. 

Hydatophylax variabilis Martynov 

Widely distributed through northern Eurasia from Sweden to Kamchatka (Schmid 

1950b) and Chukotka (Levanidova 1982) at the eastern extremity of Siberia, this species has 

also been collected at several localities in southeastern Alaska. It has not been recorded from 

the Yukon or elsewhere in North America. The species is part of an Asian complex (Schmid 

1950b), and on present evidence represents dispersal across the land connection between 

North America and Asia—either in Pleistocene or Pliocene time. This is the only Holarctic 

trichopteran known to occur in the coastal extension of southeastern Alaska; several Nearctic 

species appear to have moved northward to this area after retreat of the glaciers, but are not 

known elsewhere in Alaska or in the Yukon. This pattern of distribution might reflect a 

coastal glacial refugium for aquatic insects (e.g. Kavanaugh 1988), but streams with ample 

organic detritus would be required for survival of the larvae (e.g. Irons 1988). 

Ecological Considerations 

Aquatic insects have major roles in the cycling of nutrients and energy which underlies 

the natural productivity of freshwater systems. At temperate latitudes, aquatic insect larvae 

are diverse and abundant; and in individual systems, Trichoptera are usually high in both 

diversity and abundance in relation to other aquatic insects (Wiggins and Mackay 1978). 

However, our analysis has shown that species diversity in Trichoptera shows a marked 

decline at higher latitudes. Because the fauna is now fairly well defined, the Yukon 

Trichoptera provide a promising focus for inquiry into the exploitation of aquatic habitats 

by caddisflies at higher latitudes and into the biological factors responsible for this latitudinal 

decline in diversity. 

We examine these questions by contrasting patterns of geographic distribution and the 

use of resources in different behavioural and ecological groups of Yukon Trichoptera with 

the same groups at more temperate latitudes in North America. Considering first the generic 

level, approximately 150 genera are recognized in North American Trichoptera—case-makers, 

retreat-makers, and cocoon-making species combined (Wiggins 1996). Comparison 

between genera of running-water (lotic) and standing-water (lentic) forms in the North 

American Trichoptera yields a ratio for lotic genera to lentic genera of about 4:1. This is a 

valid biological comparison because the genus is an ecological as well as a morphological 

unit for Trichoptera (Wiggins and Mackay 1978); with few exceptions, habitats for species 

at this broad level of discrimination are consistent within genera of Trichoptera. A ratio of 

4:1 demonstrates that in North America overall there are, by a substantial margin, more 

ecological niches accessible to Trichoptera in running waters than in standing waters. 

A similar comparison between lotic and lentic genera for the 51 genera of Trichoptera 

recorded from the Yukon yields a ratio of approximately 1:1. The contrast between 4:1 for 

North America and 1:1 for the Yukon substantiates a marked decline in lotic-dwelling taxa 

with increasing latitude. This decline could be attributed either to reduction in the resources 

available to lotic Trichoptera, or to diminishing ability of Trichoptera to exploit lotic niches 

at higher latitudes; this issue is best approached at the species level. 

To examine the decline in diversity of Trichoptera at the species level in the north, the 

Yukon fauna of 145 species can be contrasted with that of British Columbia immediately to 

the south, where 279 species have been recorded (Nimmo and Scudder 1978, 1983). In broad


TABLE 1. Numbers of species in families of Trichoptera: data for Yukon from this study; for British Columbia from 

Nimmo and Scudder (1978, 1983), and for Alaska from Nimmo (1986), with taxonomic emendations. 

Family British Columbia 

lat. 49° to 60°N 

Alaska Yukon 

lat. 60° to 70°N lat. 67° to 70°N 

Spicipalpia 70 23 22 4 

Glossosomatidae 13 4 3 1 

Hydroptilidae 14 4 5 2 

Rhyacophilidae 43 15 14 1 

Annulipalpia 41 14 17 7 

Hydropsychidae 21 7 10 2 

Philopotamidae 7 1 1 1 

Polycentropodidae 13 6 6 4 

Integripalpia 168 95 106 49 

Apataniidae 7 5 4 2 

Brachycentridae 5 4 3 2 

Calamoceratidae 1 - - - 

Goeridae 1 1 1 1 

Lepidostomatidae 11 3 6 - 

Leptoceridae 22 12 16 7 

Limnephilidae 98 57 57 26 

Molannidae 1 2 2 2 

Phryganeidae 14 8 13 9 

Rossianidae 1 - - - 

Uenoidae 8 3 4 - 

Totals 279 132 145 60 

terms, these figures demonstrate that from latitude 49°N on the southern border of British 

Columbia to 60°N at the southern border of the Yukon, the trichopteran fauna decreases by 

134 species—approximately 50 per cent (Table 1). An adjunct to this database is available 

from a list of the Trichoptera of Alaska (Nimmo 1986). These 3 adjacent areas have similar 

montane topography and all have been affected by glaciation to some extent, either directly 

or indirectly. 

To extend the analysis, the northern terminus of this latitudinal gradient in the Yukon 

can be represented by ecogeographic regions 1 through 5 (Fig. 1), from approximately 

latitude 67°N at the Arctic Circle to 70°N, extending through treeline and coastal tundra to 

the Arctic shoreline. Sixty species of Trichoptera are recorded within that area (Table 1), 

although others will likely still be found. Consequently, the Yukon Trichoptera fauna 

declines from 145 species to approximately 60 species, about 59 per cent, through the 

latitudinal gradient of 60° to 70°N. 

Numbers of species in each family of Trichoptera compiled for these areas are summarized 

in Table 1. Through approximately the same latitudinal range, total numbers of species 

in the faunas of Yukon and Alaska are close, as are the figures for all 3 suborders. The 

trichopteran fauna of British Columbia is approximately twice that of either Yukon or 

Alaska; and within the overall latitudinal range of 49° to 70°N the fauna decreases from 279 

species to 60—approximately 78 per cent overall. Changes in proportions of the 3 major 

groups of Trichoptera in these regional faunas are shown in Table 2; there is a general decline 

in proportions of species of the Spicipalpia and Annulipalpia, and an increase in the 

proportion of species of the case-making Integripalpia.


TABLE 2. Percentages of the regional trichopteran faunas in major groups, based on data from Table 1. 

Major group British Columbia Alaska Yukon 

lat. 49° to 60°N lat. 60° to 70°N lat. 67° to 70°N 

Spicipalpia 

(mainly carnivores and 

algal feeders in lotic 

habitats) 

25% 17% 15% 7% 

Annulipalpia 

(fixed retreats in lotic 

habitats) 

15% 11% 10% 12% 

Integripalpia 

(portable cases, mainly in 

lentic habitats) 

60% 72% 75% 81% 

100% 100% 100% 100% 

In far northern parts of the Yukon from latitude 67° to 70°N (Table 2), the Spicipalpia 

decline substantially from their representation over the Yukon and Alaska as a whole, due 

mainly to a drastic reduction in the lotic and predatory Rhyacophilidae. The Annulipalpia 

maintain about the same proportion as for the Yukon generally, with a large decline in 

filter-feeding lotic Hydropsychidae, but a small reduction in the Polycentropodidae which 

are mainly predacious species tolerant of slow currents and lentic waters. Correspondingly, 

the proportion of case-making Integripalpia in the far northern trichopteran fauna increases 

somewhat to 81 per cent (Table 2). Species of the Limnephilidae decline from 57 in the 

Yukon generally to 26 in the far northern section (Table 1). The genus Limnephilus is 

represented in the Yukon overall by 27 species, but declines to 15 in the extreme northern 

part. Limnephilid species of the far northern part of the Yukon occur almost entirely in 

standing waters or slow currents in streams of low gradient; Dicosmoecus obscuripennis 

(81) is perhaps the sole exception, and occurs largely in lotic waters. 

Reduction in species of the Spicipalpia and Annulipalpia in the Yukon and Alaska 

(Table 1) translates biologically into a decrease of Trichoptera in running-water habitats at 

these latitudes, consistent with the generic analysis. Each of the 3 Spicipalpian families 

represented is substantially reduced (Table 1); and to whatever extent trophic factors are 

responsible for the decline, they do differ for each family. Larvae in the Glossosomatidae 

are grazers of diatoms and fine organic particles. In the Hydroptilidae, larvae in almost all 

of the genera represented feed on filamentous algae. Larvae of most species in the Rhyacophilidae 

are believed to be predacious, although there is some evidence for specific restriction 

to certain prey groups, and a few feed on algae and vascular plant tissue; for this family a 

decline of prey organisms in lotic habitats at higher latitudes could have some influence on 

diversity. 

In the Annulipalpia through the latitudinal gradient of 49° to 70°, species decrease by 

about 50 per cent in both the Hydropsychidae and Polycentropodidae. The Philopotamidae 

show a marked decline in species by about 85 per cent, and are rare in collections from 

the Yukon; larvae in this family are filter-feeders, but consume finer organic particulates 

than do other filter-feeding Trichoptera. The fine particulate organic matter consumed by 

filter-feeders and by Annulipalpia generally is largely produced by larvae of the shredder 

guild of aquatic insects, which includes most case-making Trichoptera (Integripalpia). 

Decline in filter-feeding Hydropsychidae could be caused by a decrease in fine particulate 

organic matter carried in suspension by the current; reduced populations of lotic insects


generally could affect the supply. Part of the nutrient value of this resource is aquatic 

hyphomycete fungi growing on faecal particles of other aquatic invertebrates; the fungi are 

known to grow at low temperatures (Bärlocher and Kendrick 1973). However, studies by 

Irons et al. (1994) suggest that microbial populations are physiologically less able to maintain 

optimal metabolic rates at the cold temperatures of high latitudes. Other suspended materials 

including ultra-fine particulate matter may be involved, but the decline in filter-feeding 

Trichoptera raises the likelihood that nutritional resources from suspended particulates in 

general may be marginal for supporting Trichoptera. These factors suggest that filter-feeding 

species of the Hydropsychidae which are successful at higher latitudes may derive a larger 

proportion of their food from insect prey than do those at lower latitudes, and perhaps are 

preadapted through a tendency toward increased predation; the hydropsychid subfamily 

Arctopsychinae (Arctopsyche, Parapsyche) is such a group, and is relatively well represented 

at higher latitudes. Filter-feeding larvae of the Polycentropodidae (Polycentropus, 

Neureclipsis) also derive a larger part of their food from insect prey than do Hydropsyche 

and Cheumatopsyche (e.g. Wiggins 1977, 1996). This difference could underlie the more 

northerly Yukon distribution for polycentropodids among Annulipalpia, and is another 

question to which more detailed study of Yukon Trichoptera might be directed. 

Our analysis shows that within the latitudinal gradient of 49° to 70°N, there is a marked 

decline in lotic Trichoptera, chiefly Spicipalpia and Annulipalpia, in all of their niches of 

resource exploitation. Changing trophic resources in lotic habitats will be responsible for 

some of this decline, but physical changes in running-water habitats must be taken into 

account. The range of lotic habitats at higher latitudes does not differ significantly from that 

available to the south, and these habitats are highly diverse in the Yukon and Alaska; but 

colder temperatures influence winter survival of lotic Trichoptera in smaller streams that 

freeze to the bottom (e.g. Harper 1981), including the hyporheic zone. Larval Trichoptera 

do not overwinter in streams that freeze to the bottom in subarctic interior Alaska (Irons et 

al. 1993), but survive only in sections that do not freeze (Irons 1988). Surveys of Arctic slope 

streams show that communities of benthic invertebrates are more diverse and of higher 

density in spring streams than in tundra streams, which in turn are richer than mountain 

streams (Craig and McCart 1975). The differences were attributed largely to the perennial 

flow from groundwater in spring streams, contrasted with interrupted flow when the other 

streams are frozen to the bottom during the long winter period. Trichoptera occurred in 68 

per cent of 59 spring streams sampled, but in only 21 per cent of 98 tundra streams, and 1 

per cent of 137 mountain streams. Taxonomic refinement below the ordinal level was not 

provided, but would have been highly informative even at the family level; in any event, 

both Plecoptera and Ephemeroptera maintained relatively high occurrence of 95 to 63 per 

cent in all 3 stream types in the same survey. In the larger rivers, water would remain unfrozen 

beneath the ice. Consequently, the low extremes of temperature in rivers would not differ 

greatly from lotic waters at more southerly latitudes, although the longer duration of the ice 

cover might reduce the annual period suitable for growth. However, bottom substrates of 

large northern rivers tend to be mainly unstable shifting sediments (Barton 1986; Soluk 

1985), and inappropriate for stationary, filter-feeding annulipalpian Trichoptera. For example, 

the low number of caddisflies colonizing streams following the Mt. St. Helens eruption 

in Washington was attributed by Anderson (1992) to shifting substrata and high mobility of 

the stream bed. In a glacier-fed river in Alaska, Trichoptera were one of the last insect orders 

to appear at the progression of sampling sites from the headwaters, downstream (Slack et al. 

1979). Studying development of freshwater communities following rapid recession of a 

neoglacial ice sheet in Alaska, Milner (1987) found that Chironomidae, Ephemeroptera, and


Plecoptera colonized streams but Trichoptera had a minimal role in the formation of new 

communities. Marked decline in the proportion of Trichoptera in lotic systems of Alaska 

generally was found by Oswood (1989). Therefore, compared to their dominant role in lotic 

systems at temperate latitudes (Wiggins and Mackay 1978), Trichoptera are ill-suited to 

running waters at high latitudes, where larvae are exposed to encasement in ice, unstable 

substrates, and suspended flow. 

However, larvae of several species of case-making Trichoptera (Integripalpia: Limnephilidae, 

Phryganeidae) in a slow arctic tundra stream near Tuktoyaktuk, Northwest 

Territories (lat. 69°N), remained frozen in the ice from October through May, when they 

resumed development to complete their univoltine life cycle (Winchester 1984). Tundra 

streams support lentic species for the most part. These observations suggest that certain lentic 

Trichoptera are physiologically capable of tolerating freezing of body fluids, even though 

some evidence indicates that avoiding freezing by supercooling is unlikely for most aquatic 

insects (Oswood et al. 1991). In Norway, Solem (1981) found larvae of Agrypnia obsoleta 

(Phryganeidae) to survive enclosure in solid ice for 6 months to –11°C; laboratory experiments 

confirmed freezing resistance for A. obsoleta, but larvae of Phryganea bipunctata 

were dead after several weeks in ice. Larvae of Agrypnia obsoleta (Phryganeidae), Oecetis 

ochracea (Leptoceridae), and 2 species of Molanna (Molannidae) which survived freezing 

in ice in a Swedish river had blocked the openings of their cases, although they were not in 

prepupal or pupal stages (Olsson 1981). These observations raise the critical question 

whether the portable integripalpian case confers some physiological advantage for overwintering 

trichopteran larvae embedded in ice? In Chironomidae, larvae constructing winter 

cocoons have a higher survival rate in frozen habitats than do larvae without cocoons (Danks 

1971). Tolerance to the freezing of body fluids appears to be a requirement of Trichoptera 

living at high latitudes, but these observations further suggest that there may be behavioural 

as well as physiological components to that tolerance. Resistance to low levels of oxygen is 

another aspect of the survival of aquatic insects encased in ice (Moore and Lee 1991). 

A wholly different approach to cold winter temperatures is shown in 2 species of the 

Limnephilidae, Glyphopsyche irrorata and Psychoglypha subborealis (83, 125). Adults of 

both species collected from October through May near Juneau, Alaska by Ellis (1978a) 

became sexually mature in spring. This unusual strategy for surviving cold winter conditions 

raises the question whether larvae of these species are tolerant of freezing; larvae of 

G. irrorata occur in lentic habitats, P. subborealis in lotic. Both species are assigned to 

category I, and are inferred to have reached the Yukon from more southerly areas following 

retreat of the glaciers. 

Among the 3 suborders, there is a considerably smaller latitudinal decline in species of 

the case-making Integripalpia of about 40 per cent through 49° to 70°N (Table 1). Again, 

trophic characteristics in the families of Integripalpia are not uniform. Larvae of Apataniidae, 

Goeridae, Uenoidae, and Brachycentridae in part, feed mainly by grazing diatoms from rock 

surfaces; this is the same trophic guild to which the Glossosomatidae (Spicipalpia, see above) 

belong. All of these groups occur in lotic habitats, demonstrating that in running waters food 

resources for grazing larvae do support Trichoptera at latitudes of 60° to 70°N. 

Other groups of Integripalpia in streams at high latitudes are detritivorous: Lepidostomatidae; 

and Limnephilidae (Chyranda, Dicosmoecus, Hesperophylax, Onocosmoecus, 

Psychoglypha). As members of the functional group of shredders, larvae of these species 

feed mainly on allochthonous plant debris supporting microbial growth. This food resource 

may be limiting; the supply of detritus in a subarctic Alaskan stream was meagre compared 

with that in temperate streams (Cowan and Oswood 1984), and was believed to influence


strongly the spatial and temporal patterns of detritivores. Moreover, microbial processing of 

plant detritus in high latitude streams was held to be impeded at low temperatures (Irons 

et al. 1994). 

Although the success of case-making Trichoptera in subarctic running waters is constrained 

by ice and by low levels of allochthonous detritus, biological interactions in 

subarctic ponds and lakes appear to be rather different because it is in these lentic habitats 

that Trichoptera flourish at higher latitudes. Case-making Integripalpia constitute 75 per cent 

of the species of Yukon Trichoptera (Table 2), and more than half of these (56 per cent) live 

in lentic habitats. Since inputs of allochthonous detritus are relatively lower in ponds and 

lakes because the ratio of shoreline to water area is much lower than it is in streams, 

autochthonous plant matter seems to be the energy resource underlying the success of 

detritivorous Trichoptera in these habitats. A number of species of both submerged and 

emergent vascular aquatic plants occur in southern parts of the arctic tundra, and considerably 

more in the subarctic (e.g. Hobbie 1973; I.L. Wiggins and Thomas 1962). 

Case-making larvae in the Phryganeidae, Molannidae, and Leptoceridae (Ceraclea, 

Mystacides, Oecetis) are predacious on insects and other invertebrates; overall, these groups 

decrease rather little at higher latitudes (Table 1). 

The latitudinal gradient analyzed here reflects the scale of environmental constraints 

met by species of Trichoptera dispersing to the north in the wake of receding glacial ice. The 

gradient also reveals a pattern in their use of the resources of aquatic systems under 

conditions imposed by increasing latitude. However, the relatively smaller decline in 

case-making Integripalpia within this latitudinal gradient can also be interpreted as a 

reflection of the capacity of some species of the Limnephilidae and Phryganeidae to exploit 

lentic habitats at high latitudes. Both families appear to have originated in running waters, 

but most extant species have been derived subsequently in lentic habitats. Thus, some groups 

of Limnephilidae and Phryganeidae can be seen as preadapted through origin to low 

temperatures and northerly photoperiods. If a number of species in these families have the 

physiological capability to live under conditions of extreme cold, the failure of most other 

families to do so may reflect the warmer climatic conditions in which they originated. 

Consequently, although reduced resources in aquatic communities may account for some of 

the depletion in species diversity, most groups of Trichoptera may have evolved at more 

temperate latitudes and probably still lack the physiological capability to persist under 

relatively recent regimes of cold climates at high latitudes (see above, Origin of the Beringian 

and Holarctic Trichoptera). 

Diptera are one of the most successful insect orders in the far north, particularly the 

family Chironomidae (e.g. Oliver 1968). Because most chironomid larvae are aquatic, and 

adults feed sparingly if at all, the success of this family in the far north has been attributed 

to their independence from plants and other insects (Downes 1962). Trichoptera, however, 

are biological analogues of Chironomidae but are represented in the far north by far fewer 

species, further indicating that intrinsic limitations of Trichoptera influence the penetration 

of these insects into far northern latitudes. Larvae of Chironomidae and of Empididae survive 

freezing in Alaskan subarctic streams (Irons et al. 1993). 

Larval habitat and food must also have governed the success of Trichoptera in their 

passage across the Bering land bridge. Among the present Holarctic and Beringian Trichoptera 

are 43 lentic species but only 8 lotic species—a ratio of approximately 5 lentic : 1 

lotic—indicating that caddisflies inhabiting standing waters have been far more successful 

as colonizers bridging the gaps between larval habitats, and in moving between Asia and 

North America. During glacial maxima, Beringian species dispersing across the land bridge


would have had to cross cold, dry land of low relief (Schweger et al. 1982). The clear, stable 

and well-oxygenated streams required by some species probably would have been sparse, 

although lakes and marshes would have occurred in low-lying areas. 

A related aspect is that larvae in Apatania, Ecclisomyia, and Micrasema are mainly 

grazers confined to cool streams at lower latitudes, but in the far north also occur in lakes. 

If ice conditions at higher latitudes render stream habitats unsuitable for larvae in these 

genera (see the foregoing), their transfer to cold lakes would provide an ecological alternative, 

which in Apatania may underlie extension of A. zonella to Ellesmere Island (Lake 

Hazen 81°49′N; Corbet 1966)—the most northerly record for Trichoptera. 

Because many Holarctic and Beringian species are colonizers, it is significant that 13 

of the 51 species (25 per cent) belong to the single genus Limnephilus, a dominant group in 

standing waters through much of the northern hemisphere. Although the life histories of few 

Nearctic Limnephilus are known in detail, a good deal of the success of these species in 

standing waters is probably due to the specialized characteristics by which many of them 

successfully exploit temporary pools (Wiggins 1973): marked tendency to disperse as adults; 

eggs deposited not in water but on damp substrates in a gelatinous matrix resistant to 

desiccation; and rapid larval development. Moreover, 6 additional Beringian species belong 

to other northern limnephilid genera that also occur in transient waters—2 in Asynarchus, 2 

in Grammotaulius, and 1 in each of Arctopora and Lenarchus. Species in these genera share 

most of the same developmental characteristics found in Limnephilus (e.g. Wiggins 1973), 

and in all they account for 37 per cent of the Holarctic Trichoptera in North America. 

Adaptations to transient waters are clearly apomorphic specializations in Trichoptera (Wiggins 

et al. 1980) and occur mainly in the family Limnephilidae; transient pools are a dominant 

aquatic biotope at high latitudes. 

One characteristic of species of Limnephilidae in transient waters at temperate latitudes 

is deferral of oviposition through diapause until the height of the dry phase of the pool basins 

has passed (Wiggins 1973). Females of these species are inactive until sexual maturity is 

induced by decreasing photoperiod (Novak and Sehnal 1963); at higher elevations temperature 

was also found to be a factor in breaking diapause. At high latitudes, temperatures 

conducive to larval development are available for only a few months, and yet species of 

Limnephilidae appear to be univoltine in tundra streams and ponds near Tuktoyaktuk at 

latitude 69°29′N (e.g. Winchester 1984). These species appear to have no diapause during 

their life cycle, development proceeding until inhibited by low temperature. Development 

deferred by obligate diapause in the short growing season would be biologically costly for 

Trichoptera at high latitudes if a second year of larval growth was thereby required, 

suggesting that the ecological strategy in species of Trichoptera successful at high latitudes 

is uninterrupted larval development initiated from a low temperature threshold. At temperate 

latitudes, diapause in the life cycle of these species of Limnephilidae makes existence in 

transient pools possible by imposing a delay in development while the pool and its basin are 

dry; this delay can be accommodated because the annual growth period is still adequate for 

univoltine development. By contrast, life cycles with uninterrupted development have a 

selective advantage for rapid growth in transient waters at high latitudes, suggesting that the 

incidence of diapause in congeneric species of Limnephilidae must change between temperate 

and arctic latitudes. 

At high latitudes within the zone of continuous permafrost, basins of transient surface 

pools probably do not become desiccated by exposure to the sun, as do basins of temporary 

pools at more southerly latitudes. Although exposed surface waters in transient tundra pools 

do evaporate during the brief summer (e.g. Winchester et al. 1993), a supply of water is


assured from melting permafrost, and the meltwater is shielded from evaporation by moss 

and other plant materials; underlying substrates are not likely to become desiccated because 

the frozen permafrost below prevents the water from draining away. These are the classic 

factors producing muskeg in the far north (e.g. Pielou 1991); and this rather paradoxical 

relationship between transient tundra pools, permafrost, and insolation could underlie a 

unique type of habitat for aquatic insect larvae adapted to an arctic climate. Among the 

Trichoptera, only case-making detritivorous species occur in transient tundra pools. Three 

species, all members of the Limnephilidae, are now known in these habitats: Lenarchus 

expansus and Sphagnophylax meiops (category IV), and Asynarchus lapponicus (II) (90, 

126, 75); but to what extent do larvae of other species use water-saturated tundra habitats in 

the basins of transient tundra pools? Moreover, if because of the relatively recent origin of 

the arctic biome (see Biogeographic Analysis), time has been insufficient for the evolutionary 

potential of aquatic insects to fully exploit these tundra pools with their constant supply 

of water from permafrost, these habitats may provide an evolutionary plateau of the future. 

Perhaps it is not coincidental that Sphagnophylax meiops (Frontispiece, Fig. 28), a relict 

species on both geographic and phylogenetic grounds, and the sole trichopteran genus 

confined to Beringia, still persists only in just such an arctic tundra habitat. 

Acknowledgements 

Field work for this study was supported by a Co-operative Grant (1981– 84) from 

the Natural Sciences and Engineering Research Council of Canada to G.G.E. Scudder 

and G.B. Wiggins. Analysis of the material, including participation by C.R. Parker and 

additional field work, was supported by an NSERC Operating Grant (G5707) to G.B. 

Wiggins. Participants in ROM field parties included H.E. Frania, E.R. Fuller, R. 

Jaagumagi, and B.D. Marshall. Assistance in analysis of the collections was provided 

by J. Conn, H.E. Frania, E.R. Fuller, R. Jaagumagi, J.D. Kerr, J. Thomson-Delaney, and 

by P.W. Schefter who also identified material of Hydropsyche and Cheumatopsyche. 

Information on particular species was provided by T.J. Arefina, L. BotojAneanu, O.S. 

Flint, W.K. Gall, V.D. Ivanov, N.P. Kristensen, I.M. Levanidova, J. Lukyanchenko, H. 

Malicky, W. Mey, A.P. Nimmo, and R.N. Vineyard who also translated literature in the 

Russian language. Critical comments on the manuscript were provided by L. BotojAneanu, 

J.A. Downes, O.S. Flint, H.E. Frania, I.M. Levanidova, R.J. Mackay, J.V. 

Matthews, A.P. Nimmo, V.H. Resh, and R.A. Ring. Illustrations were prepared by P. 

Stephens-Bourgeault, except for Figs. 2 and 3 by Z. Zichmanis and 12 –15, 17 –18 by 

C.R. Parker. C. Rutland and R. Darling rendered valuable assistance in preparing the 

manuscript. Additional collections of Trichoptera were made available from the Canadian 

National Collection of Insects maintained by the Biological Resources Division of 

Agriculture Canada (F. Schmid); the Illinois Natural History Survey (J.D. Unzicker); 

the Royal British Columbia Museum (R.A. Cannings); the Spencer Museum, University 

of British Columbia (G.G.E. Scudder and S.G. Cannings); the Strickland Museum, 

University of Alberta (G.E. Ball and A.P. Nimmo); the Department of Entomology, 

United States National Museum of Natural History (O.S. Flint); D.G. Huggins of the 

Kansas State Biological Survey; and N.N. Winchester and R.A. Ring of the Department 

of Biology, University of Victoria.


References 

Ager, T.A. 1982. Vegetational history of western Alaska during the Wisconsin glacial interval and the Holocene. 

pp. 75 – 83 in D.M. Hopkins, J.V. Matthews Jr., C.E. Schweger, and S.B. Young (Eds.), Paleoecology of 

Beringia. Academic Press, New York. 489 pp. 

Anderson, N.H. 1992. Influence of disturbance on insect communities in Pacific Northwest streams. Hydrobiologia 

248:79 – 92. 

Banks, N. 1900. Neuropteroid insects. In Papers from the Harriman Alaska Expedition. Proc. Acad. Sci. Wash. 

2:465 – 473. 

______ 1914. American Trichoptera—notes and descriptions. Can. Ent. 46:149 –156, 201– 205, 252 – 258, 261– 

268. 

Bärlocher, F. and B. Kendrick. 1973. Fungi and food preferences of Gammarus pseudolimnaeus. Arch. Hydrobiol. 

72:501– 516. 

Barton, D.R. 1986. Invertebrates of the Mackenzie system. pp. 473 – 492 in B.R. Davies, and K.F. Walker (Eds.), 

The Ecology of River Systems. W. Junk, The Hague. 793 pp. 

Baumann, R.W. and J.D. Unzicker. 1981. Preliminary checklist of Utah caddisflies. Encyclia 58:25 – 29. 

Betten, C. and M.E. Mosely. 1940. The Francis Walker types of Trichoptera in the British Museum. British Museum 

(Natural History), London. 

BotojAneanu, L. 1988. A superspecies, or Formenkreis, in caddisflies: Micrasema (superspecies gelidum) McLachlan 

(Trichoptera). Populational thinking versus Hennigian fundamentalism. Riv. Idrobiol. 27:181– 210. 

BotojAneanu, L. and I.M. Levanidova. 1988. Trichoptera Hydroptilidae (Insecta) from Soviet Union Far-Eastern 

Territories. Bull. zool. Mus. Univ. Amsterdam 11:169 –176. 

BotojAneanu, L. and H. Malicky. 1978. Trichoptera. pp. 333 – 359 in J. Illies (Ed.), Limnofauna Europaea. Gustav 

Fischer Verlag, Stuttgart, New York; Swets and Zeitlinger B.V., Amsterdam. 474 pp. 

Briggs, J.C. 1966. Zoogeography and evolution. Evolution 20:282 – 289. 

Buckland, P.C., D.W. Perry, G.M. Gislason, and A.J. Dugmore. 1986. The pre-Landnám fauna of Iceland: a 

palaeontological contribution. Boreas 15:173 –184. 

Corbet, P.S. 1966. Parthenogenesis in caddisflies (Trichoptera). Can. J. Zool. 44:981– 982. 

Cowan, C.A. and M.W. Oswood. 1984. Spatial and seasonal associations of benthic macroinvertebrates and detritus 

in an Alaskan subarctic stream. Polar Biol. 3:211– 215. 

Craig, P.C. and P.J. McCart. 1975. Classification of stream types in Beaufort Sea drainages between Prudhoe Bay, 

Alaska, and the Mackenzie Delta, N.W.T., Canada. Arct. Alp. Res. 7:183 –198. 

Danks, H.V. 1971. Overwintering of some north temperate and arctic Chironomidae. II. Chironomid biology. Can. 

Ent. 103:1875 –1910. 

______ 1981. Arctic Arthropods. A review of systematics and ecology with particular reference to the North 

American fauna. Entomological Society of Canada, Ottawa. 608 pp. 

Downes, J.A. 1962. What is an arctic insect? Can. Ent. 94:143 –162. 

______ 1964. Arctic insects and their environment. Can. Ent. 96:279 – 307. 

______ 1966. The Lepidoptera of Greenland; some geographic considerations. Can. Ent. 98:1135 –1144. 

Downes, J.A. and D.H. Kavanaugh. 1988. Origins of the North American insect fauna. Introduction and commentary. 

pp. 1 –11 in J.A. Downes and D.H. Kavanaugh (Eds.), Origins of the North American Insect Fauna. 

Mem. ent. Soc. Can. 144. 168 pp. 

Ellis, R.J. 1978a. Over-winter occurrence and maturation of gonads in adult Psychoglypha subborealis (Banks) 

and Glyphopsyche irrorata (Fabricius). Pan-Pacif. Ent. 54:178 –180. 

______ 1978b. Seasonal abundance and distribution of adult caddisflies of Sashin Creek, Baranof Island, southeastern 

Alaska. Pan-Pacif. Ent. 54:199 – 206. 

Fischer, F.C.J. 1964. Trichopterorum Catalogus, vol. 5. Nederlandsche Entomologische Vereeniging, Amsterdam. 

214 pp. 

______ 1968. Trichopterorum Catalogus, vol. 9. Nederlandsche Entomologische Vereeniging, Amsterdam. 

363 pp. 

Flint, O.S., Jr. 1964. Two species of Limnephilidae new to North America. Proc. ent. Soc. Wash. 66:60. 

______ 1984. The genus Brachycentrus in North America, with a proposed phylogeny of the genera of Brachycentridae 

(Trichoptera). Smithsonian Contribs Zool. 398. 58 pp. 

Foote, C.J., J.W. Clayton, C.C. Lindsey, and R.H. Bodaly. 1992. Evolution of lake whitefish (Coregonus 

clupeaformis) in North America during the Pleistocene: evidence for a Nahanni glacial refuge race in the 

northern Cordillera region. Can. J. Fish. aquat. Sci. 49:760 – 768. 

Forsslund, K.H. 1932. Zur Kenntniss der Trichopteren Grönlands. Ent. Tidskr. 53:56 – 59. 

______ 1933. Eine neue melanistiche Neuronia - Varietät (Trich.). Ent. Tidskr. 54:36. 

Frania, H. E. and G. B. Wiggins. 1997. Analysis of morphological and behavioural evidence for the phylogeny and 

higher classification of Trichoptera. Contr. Life Sci. Div. R. Ont. Mus. 160. 

Fristrup, B. 1942. Neuroptera and Trichoptera. Zool. Iceland 3:1– 23. 

Fuller, E.R. 1987. Systematics of the caddisfly family Molannidae (Trichoptera). M.Sc. Thesis (unpublished), Dept. 

Zoology, Univ. Toronto.


Gislason, G.M. 1981. Distribution and habitat preferences of Icelandic Trichoptera. pp. 99 –109 in G.P. Moretti 

(Ed.), Proc. Third Int. Symp. on Trichoptera. Series Entomologica 20. W. Junk, The Hague. 472 pp. 

Harper, P.P. 1981. Ecology of streams at high latitudes. pp. 313 – 337 in M.A. Lock and D.D. Williams (Eds.), 

Perspectives in Running Water Ecology. Plenum, New York. 450 pp. 

______ 1989. Zoogeographical relationships of aquatic insects (Ephemeroptera, Plecoptera, and Trichoptera) from 

the eastern James Bay drainage. Can. Fld-Nat. 103:535 – 546. 

Harper, P.P. and G. Méthot. 1975. Goera radissonica n. sp., nouveau Trichoptère de la région de la baie James. 

Naturaliste can. 102:593 – 595. 

Hill-Griffin, A.L. 1912. New Oregon Trichoptera. Ent. News 23:17 – 21. 

Hobbie, J.E. 1973. Arctic limnology: a review. pp. 127 –168 in M.E. Britton (Ed.), Alaskan Arctic Tundra. Arct. 

Inst. N. Am. Tech. Pap. 25. 224 pp. 

Hopkins, D.M., J.V. Matthews, J.A. Wolfe, and M.L. Silberman. 1971. A Pliocene flora and insect fauna from the 

Bering Strait region. Palaeogeogr. Palaeoclimatol. Palaeoecol. 9:211– 231. 

Hopkins, D.M., J.V. Matthews Jr., C.E. Schweger, and S.B. Young (Eds.). 1982. Paleoecology of Beringia. 

Academic Press, New York. 489 pp. 

Irons, J.G. 1988. Life history patterns and trophic ecology of Trichoptera in two Alaskan (U.S.A.) subarctic streams. 

Can. J. Zool. 66:1258 –1265. 

Irons, J.G., L.K. Miller, and M.W. Oswood. 1993. Ecological adaptations of aquatic macroinvertebrates to 

overwintering in interior Alaska (U.S.A.) subarctic streams. Can. J. Zool. 71:98 –108. 

Irons, J.G., M.W. Oswood, R.J. Stout, and C.M. Pringle. 1994. Longitudinal patterns in leaf litter breakdown: is 

temperature really important? Freshwat. Biol. 32:401– 411. 

Johansson, A., A.N. Nilsson, and B.W. Svensson. 1991. Larval morphology, habitat and distribution of Limnephilus 

diphyes (Trichoptera, Limnephilidae). Ent. Tidskr. 112:19 – 25. 

Kaneshiro, K.Y. 1993. Introduction, colonization, and establishment of exotic insect populations: fruit flies in 

Hawaii and California. Am. Ent. 39:23 – 29. 

Karlstrom, T.N.V. and G.E. Ball (Eds.). 1969. The Kodiak Island Refugium; its Geology, Flora, Fauna and History. 

Boreal Institute, Univ. Alberta. Ryerson Press, Toronto. 262 pp. 

Kavanaugh, D.H. 1988. The insect fauna of the Pacific Northwest coast of North America: present patterns and 

affinities and their origins. pp. 125 –149 in J.A. Downes and D.H. Kavanaugh (Eds.), Origins of the North 

American Insect Fauna. Mem. ent. Soc. Can. 144. 168 pp. 

Kelley, R.W. 1984. Phylogeny, morphology and classification of the micro-caddisfly genus Oxyethira Eaton 

(Trichoptera: Hydroptilidae). Trans. Am. ent. Soc. 110:435 – 463. 

Kimmins, D.E. and D.G. Denning. 1951. The McLachlan types of North American Trichoptera in the British 

Museum. Ann. ent. Soc. Am. 44:111 –140. 

Labandeira, C.C. and J.J. Sepkoski Jr. 1993. Insect diversity and the fossil record. Science 261:310 – 315. 

Lafontaine, J.D. and D.M. Wood. 1988. A zoogeographic analysis of the Noctuidae (Lepidoptera) of Beringia, and 

some inferences about past Beringian habitats. pp. 109 –123 in J.A. Downes and D.H. Kavanaugh (Eds.), 

Origins of the North American Insect Fauna. Mem. ent. Soc. Can. 144. 168 pp. 

Lehmkuhl, D.M. and C.D. Kerst. 1979. Zoogeographical affinities and identification of central Arctic caddisflies 

(Trichoptera). Musk-Ox 25:1– 28. 

Lepneva, S.G. 1964. Fauna of the U.S.S.R.; Trichoptera, vol. 2, no. 1. Larvae and Pupae of Annulipalpia. (Original 

in Russian, translated into English, Israel Program for Scientific Translations, 1970. 638 pp.) 

______ 1966. Fauna of the U.S.S.R.; Trichoptera, vol. 2, no. 2. Larvae and Pupae of Integripalpia. (Original in 

Russian, translated into English, Israel Program for Scientific Translations, 1971. 700 pp.) 

Levanidova, I.M. 1975. The caddisflies (Trichoptera) of Kamchatka; an ecological-faunistic outline (In Russian). 

Bull. Pacif. Sci. Inst. Fish. and Oceanogr. 97:83 –114. 

______ 1982. Amphibiotic insects of the mountainous regions of the Soviet Far East (Faunistics, ecology, 

zoogeography of the Ephemeroptera, Plecoptera and Trichoptera) (In Russian). U.S.S.R. Academy of Science, 

Far Eastern Science Centre. Biology - Soils Institute. 214 pp. 

Lindsey, C.C. and J.D. McPhail. 1986. Zoogeography of fishes of the Yukon and Mackenzie basins. pp. 639 – 674 

in C.H. Hocutt and E.O. Wiley (Eds.), The Zoogeography of North American Freshwater Fishes. Wiley, New 

York. 866 pp. 

MacLean, S.F. and F.A. Pitelka. 1971. Seasonal patterns of abundance of tundra arthropods near Barrow. Arctic 

24:19 – 40. 

Malicky, H. 1979. Notes on some caddisflies (Trichoptera) from Europe and Iran. Aquat. Insects 1:3 –16. 

______ 1983. Atlas of European Trichoptera. Series Entomologica 24. W. Junk, The Hague. 298 pp. 

______ 1988. Spuren der Eiszeit in der Trichopterenfauna Europas (Insecta, Trichoptera). Riv. Idrobiol. 27(2 – 3):247 – 

297. 

Manuel, K.L. and A.P. Nimmo. 1984. The caddisfly genus Ylodes in North America (Trichoptera: Leptoceridae). 

pp. 219 – 224 in J.C. Morse (Ed.), Proc. Fourth Int. Symp. on Trichoptera. Series Entomologica 30. W. Junk, 

The Hague. 486 pp. 

Martynov, A.V. 1909. Les Trichoptères de la Sibérie et des régions adjacentes. I. Ezheg. zool. Muz. [Annls Mus. 

Zool. Acad. Imp. Sci. St. Petersbourg] 14:223 – 255.


______ 1910. Les Trichoptères de la Sibérie et des régions adjacentes. II. Ezheg. zool. Muz. [Annls Mus. Zool. Acad. 

Imp. Sci. St. Petersbourg] 15:351– 429. 

______ 1914. Les Trichoptères de la Sibérie et des régions adjacentes. IV, Limnophilinae. Ezheg. zool. Muz. [Annls 

Mus. Zool. Acad. Imp. Sci. St. Petersbourg] 19:173 – 285. 

______ 1924a. Practical entomology, vol. 5. Trichoptera (In Russian). Gosudarstvennoe Izdatelstvo, Leningrad. 

______ 1924b. Notice sur les Trichoptères de la District de Minoussinsk (In Russian). Ezheg. gosud. Muz. N.M. 

Mart’yanova [Jb. Martjanovischen Staats Mus. Minoussinsk] 2(3):62 –107. 

______ 1935. Trichoptera of the Amur region. I. Trudy zool. Inst., Leningr. [Trav. Inst. Zool. Acad. Sci. URSS] 

2:205 – 395. 

Matthews, J.V., Jr. 1979a. Tertiary and Quaternary environments: historical background for an analysis of the 

Canadian insect fauna. pp. 31– 86 in H.V. Danks (Ed.), Canada and its Insect Fauna. Mem. ent. Soc. Can. 

108. 573 pp. 

______ 1979b. Fossil beetles and the late Cenozoic history of the tundra environment. pp. 371– 378 in J. Gray and 

A.J. Boucot (Eds.), Historical Biogeography, Plate Tectonics, and the Changing Environment. Oregon State 

Univ. Press, Corvallis. 500 pp. 

McLachlan, R. 1874 –1880. A monographic revision and synopsis of the Trichoptera of the European fauna. Pt. 1, 

1874:1– 46, pls. 1– 5. Pt. 2, 1875:47 –108, pls. 6 –11. Pt. 3, 1875:109 –144, pls. 12 –15. Pt. 4, 1876:145 – 220, 

pls. 16 – 23. Pt. 5, 1876:221– 280, pls. 24 – 31, w. supplement I – XII. Pt. 6, 1877:281– 348, pls. 32 – 37., Pt. 7, 

1878:349 – 428, pls. 38 – 44. Pt. 8, 1879:429 – 500, pls. 45 – 51. Pt. 9, 1880:501– 23, w. supplement XIII – 

LXXXIV, pls. 52 – 59. 

Mey, W. and A. Dulmaa. 1985. Die Köcherfliegenfauna der Mongolei (Insecta, Trichoptera). Mitt. zool. Mus. Berl. 

61:79 –104. 

Milne, L.J. 1934. Studies in North American Trichoptera, 1. Publ. by author, Cambridge, Mass. 19 pp. 

Milner, A.M. 1987. Colonization and ecological development of new streams in Glacier Bay National Park, Alaska. 

Freshwat. Biol. 18:53 – 70. 

Moore, M.V. and R.E. Lee Jr. 1991. Surviving the big chill: overwinter strategies of aquatic and terrestrial insects. 

Am. Ent. 37:111 –118. 

Morse, J.C. 1975. A phylogeny and revision of the caddisfly genus Ceraclea (Trichoptera, Leptoceridae). Contr. 

Am. Ent. Inst. 11(2). 97 pp. 

Mosely, M.E. 1929. Trichoptera and Ephemeroptera of Greenland. Additional records made by the Oxford 

University Expedition to Kugssuk, Godthaab Fjord, W. Greenland, 1928. Ann. Mag. nat. Hist. (ser. 10) 

4:501– 509. 

______ 1932. Oxford University Greenland expedition, 1928: a correction. Ann. Mag. nat. Hist. (ser. 10) 9:573 – 74. 

______ 1936. The Indian caddis-flies (Trichoptera). Pt. IV. J. Bombay nat. Hist. Soc. 38(3):447 – 478. 

Nagayasu, Y. and T. Ito. 1993. The caddisfly genus Dicosmoecus in Asia (Trichoptera, Limnephilidae). I. Males. 

pp. 123 –127 in C. Otto (Ed.), Proc. Seventh Int. Symp. on Trichoptera. Backhuys, Leiden. 312 pp. 

Nimmo, A.P. 1971. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta and eastern British 

Columbia and their post-glacial origin. Quaest. ent. 7:3 – 234. 

______ 1986. Preliminary annotated checklist of the Trichoptera (Insecta) of Alaska. Contr. nat. Sci., Br. Columb. 

Prov. Mus. 5:1– 7. 

______ 1991. Seven new species of Limnephilus from western North America with description of female of 

L. pallens (Banks) (Trichoptera, Limnephilidae, Limnephilinae, Limnephilini). Proc. ent. Soc. Wash. 93:499 – 

508. 

Nimmo, A.P. and G.G.E. Scudder. 1978. An annotated checklist of the Trichoptera (Insecta) of British Columbia. 

Syesis 11:117 –133. 

______ 1983. Supplement to an annotated checklist of the Trichoptera (Insecta) of British Columbia. Syesis 

16:71– 83. 

Nimmo, A.P. and R. Wickstrom. 1984. Preliminary annotated checklist of the Trichoptera (Insecta) of the Yukon. 

Syesis 17:3 – 9. 

Novak, K. and F. Sehnal. 1963. The development cycle of some species of the genus Limnephilus (Trichoptera). 

$ as. % sl. Spol. ent. 60:68 – 80. 

Oliver, D.R. 1968. Adaptations of Arctic Chironomidae. Annls zool. fenn. 5:111 –118. 

Olsson, T.I. 1981. Overwintering of benthic invertebrates in ice and frozen sediment in a North Swedish river. 

Holarct. Ecol. 4:161 –166. 

Oswood, M.W. 1989. Community structure of benthic invertebrates in interior Alaskan (USA) streams and rivers. 

pp. 97 –110 in W.F. Vincent and J.C. Ellis-Evans (Eds.), High Latitude Limnology. Kluwer Academic 

Publishers, Dordrecht, Netherlands. 

Oswood, M.W., L.K. Miller, and J.G. Irons. 1991. Overwintering of freshwater benthic invertebrates. pp. 360 – 375 

in R.E. Lee Jr. and D.L. Denlinger (Eds.), Insects at Low Temperature. Chapman and Hall, New York. 513 pp. 

Parker, C.R. and G.B. Wiggins. 1985. The Nearctic caddisfly genus Hesperophylax Banks (Trichoptera: Limnephilidae). 

Can. J. Zool. 63:2443 – 2472. 

Pielou, E.C. 1991. After the Ice Age. The Return of Life to Glaciated North America. Univ. Chicago Press, Chicago. 

366 pp.


Reeves, B.O.K. 1973. The nature and age of the contact between the Laurentide and Cordilleran ice sheets in the 

western interior of North America. Arct. Alp. Res. 5:1 –16. 

Resh, V.H. 1976. The biology and immature stages of the caddisfly genus Ceraclea in eastern North America 

(Trichoptera: Leptoceridae). Ann. ent. Soc. Am. 69:1039 –1061. 

Resh, V.H., J.C. Morse, and I.D. Wallace. 1976. The evolution of the sponge feeding habit in the caddisfly genus 

Ceraclea (Trichoptera: Leptoceridae). Ann. ent. Soc. Am. 6:937 – 941. 

Ross, H.H. 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Ill. nat. Hist. Surv. 23(1). 326 pp. 

______ 1949. Descriptions of some western Limnephilidae (Trichoptera). Pan-Pacif. Ent. 25:119 –128. 

______ 1950. Synoptic notes on some Nearctic limnephilid caddisflies (Trichoptera, Limnephilidae). Am. midl. 

Nat. 43:410 – 429. 

______ 1956. Evolution and Classification of the Mountain Caddisflies. Univ. Illinois Press, Urbana. 213 pp. 

______ 1958. Affinities and origins of the northern and montane insects of western North America. pp. 231-252 in 

C.L. Hubbs (Ed.), Zoogeography. Am. Assoc. Advance. Sci. Publ. 51. 509 pp. 

______ 1965. Pleistocene events and insects. pp. 583 – 596 in D.G. Frey and H.E. Wright (Eds.), The Quaternary 

of the United States. Princeton Univ. Press, Princeton. 922 pp. 

Ross, H.H. and D.R. Merkley. 1952. An annotated key to the Nearctic males of Limnephilus (Trichoptera: 

Limnephilidae). Am. midl. Nat. 47:435 – 455. 

Ross, H.H. and J.C. Morse. 1973. Micrasema kluane, a probable stepping stone to the Arctic (Trichoptera, 

Brachycentridae). Ent. News 84:291– 293. 

Roy, D. and P.P. Harper. 1979. Liste préliminaire des Trichoptères (insectes) du Québec. Annls Soc. ent. Québec 

24:148 –172. 

Ruiter, D. E. 1995. The adult Limnephilus Leach (Trichoptera: Limnephilidae) of the New World. Bull. Ohio Biol. 

Surv. (n.s.) 11(1). 200 pp. 

Schmid, F. 1950a. Monographie du genre Grammotaulius Kolenati (Trichoptera, Limnophilidae). Revue suisse 

Zool. 57:317 – 352. 

______ 1950b. Le genre Hydatophylax Banks (Trichoptera, Limnophilidae). Mitt. Schweiz. ent. Ges. 23:265 – 296. 

______ 1952. Le groupe de Lenarchus Mart. (Trichoptera, Limnophilidae). Mitt. Schweiz. ent. Ges. 25:157 – 210. 

______ 1954a. Contribution à l’étude de la sous-famille des Apataniinae II. Tijdschr. Ent. 97:1– 74. 

______ 1954b. Le genre Asynarchus McL. Mitt. Schweiz. ent. Ges. 27:57 – 96. 

______ 1955. Contribution à l’étude des Limnophilidae (Trichoptera). Université de Lausanne, Suisse. 245 pp. 

______ 1964. Some Nearctic species of Grammotaulius Kolenati (Trichoptera, Limnephilidae). Can. Ent. 96:914 – 

917. 

______ 1965a. Quelques Trichoptères asiatiques II. Ent. Tidskr. 86:28 – 35. 

______ 1965b. Trichoptera. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei. 

Reichenbachia 7:201– 203. 

______ 1968. La Famille des Arctopsychides (Trichoptera). Mem. Soc. ent. Québec 1. 84 pp. 

______ 1970. Le Genre Rhyacophila et la Famille des Rhyacophilidae (Trichoptera). Mem. ent. Soc. Can. 66. 230 

pp. + 52 pl. 

______ 1982. Revision des Trichoptères canadiens. II. Les Glossosomatidae et Philopotamidae (Annulipalpia). 


______ 1983. Revision des Trichoptères canadiens. III. Les Hyalopsychidae, Psychomyiidae, Goeridae, Brachycentridae, 

Sericostomatidae, Helicopsychidae, Beraeidae, Odontoceridae, Calamoceratidae et Molannidae. 


Schmid, F., T.J. Arefina, and I.M. Levanidova. 1993. Contribution to the knowledge of the Rhyacophila (Trichoptera) 

of the sibirica group. Bull. Inst. r. Sci. nat. Belg., Ent. 63:161 –172. 

Schweger, C.E., J.V. Matthews, D.M. Hopkins, and S.B. Young. 1982. Paleoecology of Beringia—a synthesis. 

pp. 425 – 444 in D.M. Hopkins, J.V. Matthews Jr., C.E. Schweger, and S.B. Young (Eds.), Paleoecology of 

Beringia. Academic Press, New York. 489 pp. 

Scudder, G.G.E. 1997. Environment of the Yukon. pp. 13 – 57 in H.V. Danks and J.A. Downes (Eds.), Insects of 

the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 

Slack, K.V., J.W. Naumann, and L.J. Tilley. 1979. Benthic invertebrates in a north-flowing stream and a 

south-flowing stream, Brooks Range, Alaska. Wat. Resour. Bull. 15:108 –135. 

Smith, S.D. and K.L. Manuel. 1984. Reconsideration of the Nearctic species of the Rhyacophila acropedes subgroup 

based on adults (Trichoptera: Rhyacophilidae). pp. 369 – 374 in J.C. Morse (Ed.), Proc. Fourth Int. Symp. on 

Trichoptera. Series Entomologica 30. W. Junk, The Hague. 486 pp. 

Solem, J.O. 1981. Overwintering strategies in some Norwegian caddisflies. Summary. pp. 321– 322 in G.P. Moretti 

(Ed.), Proc. Third Int. Symp. on Trichoptera. Series Entomologica 20. W. Junk, The Hague. 472 pp. 

Solem, J.O. and V.H. Resh. 1981. Larval and pupal description, life cycle, and adult flight behaviour of the 

sponge-feeding caddisfly Ceraclea nigronervosa (Retzius) in central Norway (Trichoptera). Entomologica 

scand. 12:311– 319. 

Soluk, D.A. 1985. Macroinvertebrate abundance and production of psammophilous Chironomidae in shifting sand 

areas of a lowland river. Can. J. Fish. aquat. Sci. 42:1296 –1302. 

Stanley, S.M. 1986. Earth and Life Through Time. W.H. Freeman, New York. 690 pp.


Ulmer, G. 1907. Trichopteren, Fasc. VI(I). Collections Zoologiques du baron Edm. de Selys Longchamps. Hayez, 

Impr. des Académies, Bruxelles. 102 pp. 

______ 1927. Entomologische Ergebnisse der schwedischen Kamtchatka-Expedition 1920-1922. 11. Trichopteren 

und Ephemeropteren. Ark. Zool. 19A (8). 17 pp. 

Vermeij, G.J. 1991. When biotas meet: understanding biotic interchange. Science 253:1099 –1104. 

Vineyard, R.N. and G.B. Wiggins. 1988. Further revision of the caddisfly family Uenoidae (Trichoptera): evidence 

for inclusion of Neophylacinae and Thremmatidae. Syst. Ent. 13:361– 372. 

Wiggins, G.B. 1956. A revision of the North American caddisfly genus Banksiola (Trichoptera: Phryganeidae). 

Contr. R. Ont. Mus. Zool. Palaeont. 43. 13 pp. 

______ 1973. A contribution to the biology of caddisflies (Trichoptera) in temporary pools. Contr. Life Sci. Div. 

R. Ont. Mus. 88. 28 pp. 

______ 1977. Larvae of the North American Caddisfly Genera (Trichoptera). First edition. Univ. Toronto Press, 

Toronto. 401 pp. 

______ 1984. Trichoptera - some concepts and questions, Keynote address. pp. 1 –12 in J.C. Morse (Ed.), Proc. 

Fourth Int. Symp. on Trichoptera. Series Entomologica 30. W. Junk, The Hague. 486 pp. 

______ 1996. Larvae of the North American Caddisfly Genera (Trichoptera). Second edition, revised. Univ. 

Toronto Press, Toronto. 457 pp. 

______ in press. The Caddisfly Family Phryganeidae (Trichoptera). Univ. Toronto Press, Toronto. 

Wiggins, G.B. and S. Kuwayama. 1971. A new species of the caddisfly genus Oligotricha from northern Japan and 

Sakhalin, with a key to the adults of the genus (Trichoptera: Phryganeidae). Kontyu 39:340 – 346. 

Wiggins, G.B. and R.J. Mackay. 1978. Some relationships between systematics and trophic ecology in Nearctic 

aquatic insects with special reference to Trichoptera. Ecology 59:1211 –1220. 

Wiggins, G.B., R.J. Mackay, and I.M. Smith. 1980. Evolutionary and ecological strategies of animals in annual 

temporary pools. Arch. Hydrobiol. Suppl. 58:97 – 206. 

Wiggins, G.B. and J.S. Richardson. 1982. Revision and synopsis of the caddisfly genus Dicosmoecus (Trichoptera: 

Limnephilidae). Aquat. Insects 4:181– 217. 

______ 1987. Revision of the Onocosmoecus unicolor group (Trichoptera: Limnephilidae, Discosmoecinae). 

Psyche 93:187 – 216 (1986). 

Wiggins, G.B. and W. Wichard. 1989. Phylogeny of pupation in Trichoptera, with proposals on the origin and 

higher classification of the order. J. N. Am. benthol. Soc. 8:260 – 276. 

Wiggins, G.B. and N.N. Winchester. 1984. A remarkable new caddisfly genus from northwestern North America 

(Trichoptera, Limnephilidae, Limnephilinae). Can. J. Zool. 62:1853 –1858. 

Wiggins, G.B. and R.W. Wisseman. 1992. New North American species in the genera Neothremma and Farula, 

with hypotheses on phylogeny and biogeography (Trichoptera: Uenoidae). Can. Ent. 124:1063 –1074. 

Wiggins, I.L. and J.H. Thomas. 1962. A Flora of the Alaskan Arctic Slope. Univ. Toronto Press, Toronto. Arct. 

Inst. N. Am. Publ. 4. 425 pp. 

Winchester, N.N. 1984. Life histories and post-glacial origins of tundra caddisflies (Trichoptera) from the 

Tuktoyaktuk Peninsula, Northwest Territories. M.Sc. Thesis (unpublished), Department of Biology, Univ. 

Victoria. 

Winchester, N.N., G.B. Wiggins, and R.A. Ring. 1993. The immature stages and biology of the unusual North 

American arctic caddisfly Sphagnophylax meiops, with consideration of the phyletic relationships of the genus 

(Trichoptera: Limnephilidae). Can. J. Zool. 71:1212 –1220. 

Yamamoto, T. and H.H. Ross. 1966. A phylogenetic outline of the caddisfly genus Mystacides (Trichoptera: 

Leptoceridae). Can. Ent. 98:627 – 632. 

Yamamoto, T. and G.B. Wiggins. 1964. A comparative study of the North American species in the caddisfly genus 

Mystacides (Trichoptera: Leptoceridae). Can. J. Zool. 42:1105 –1126. 

Yang, L.-F. and J.C. Morse. 1988. Ceraclea of the People’s Republic of China (Trichoptera; Leptoceridae). Contr. 

Am. ent. Inst. 23. 69 pp.

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