Phylogenetic Analysis of the Nearctic Androprosopa Mik (Diptera: Thaumaleidae) with an emphasis on the western species A Dissertation Presented for the Master of Science Degree The University of Tennessee, Knoxville Robert John Pivar December 2015 © 2015 by Robert John Pivar All rights reserved. Acknowledgments Abstract The family Thaumaleidae is an uncommonly encountered and understudied family. This thesis attempts to illustrate the diversity of the Thaumaleidae within North America and determine relationships between them. The Nearctic fauna of Androprosopa Mik is revised to include thirty-two species, five of which are described as new to science. Descriptions of the adult males of A. apache, A. arnaudi, A. boussi, A. magnipelvim, A. rainierensis and A. sierra are presented. Redescriptions of all remaining species are provided, as well as genitalic illustrations and updated distribution maps for each species. A diagnostic key to all Nearctic Thaumaleidae is also provided. Phylogenetic studies were conducted to determine the relationships between the eastern and western Nearctic Androprosopa, and amongst the western species. Fresh, DNA quality material was gathered for all Nearctic species except for 6, which eluded capture. Molecular sequence data from two nuclear protein-coding genes, Big Zinc Finger 2 (BZF2) and Molybdenum Cofactor Sulfurase (MCS), were used to test relationships between individual species and species groups. Analyses using maximum parsimony, maximum likelihood and Bayesian inference methods were implemented. Resulting phylogenies supported the monophyly of the western Nearctic species, however relationships between the western and eastern species remain unresolved. Analyses had difficulty placing the eastern species as closer relatives to the western fauna versus the Palearctic fauna. Six new species groups within the western Nearctic fauna are proposed herein. Table of Contents List of Tables List of Figures 25 CHAPTER 1: GENERAL INTRODUCTION AND LITERATURE REVIEW The Importance of Diptera The order Diptera (the true flies) is one of the four megadiverse insect orders, containing over 160 000 described species and comprising 10-15% of known animal species (Brown 2005; Courtney et al. 2009; Marshall 2012; Woodley et al. 2009; Yeates et al. 2007). Despite these numbers, experts believe that this represents only a small fraction of the diversity of the order, and that the actual number of species may exceed one million (Hammond 1992). Flies are found worldwide on every continent and occupy, if not dominate, a wide variety of niches (Brown 2009; Marshall 2012). It is due to their ability to occupy seemingly endless niches, ability to disperse, small size and relative scarcity of researchers that Dipteran systematic research remains a discipline with vast opportunity for discovery. Whether it is a negative or positive interaction, flies have a strong impact on humankind. No other group of animals has as much impact on human health than flies, mainly due to their ability to act as a vector for many diseases, including malaria (Hall & Gerhardt 2009; Marshall 2012). Another way in which flies have a negative impact on humans is as a plant pest. This is due to pathogen transmission and by being in direct competition for human food and forest crops (Curran 1965; Marshall 2012). Many species, however, play important positive roles for humankind, such as important pollinators, biological control, medical or forensic applications and the decomposition of plant and animal matter (Curran 1965; Brown 2009). Decades of extensive research have provided valuable knowledge on the economic importance of flies and this knowledge base continues to grow. However, in order to fully understand what pest is being dealt with, or the ecology of a certain pollinator for example, we must attempt to realize the diversity and evolutionary relationships of Diptera. These two areas of study are only beginning to be understood for many Dipteran taxa and continued systematic research will contribute to the knowledge base and reveal insight into how various species function within ecosystem processes. Biology of Thaumaleidae The Thaumaleidae, or madicolous midges, is a family of Diptera that is specialized for living in a specific type of aquatic environment. The Thaumaleidae are small (2-5mm long), stocky flies that are typically brown or black, though some may be dark orange or shiny yellow. They are found worldwide on all continents except Antarctica (Vaillant 1977; Sinclair & Saigusa 2002). As mentioned earlier, they are specialized to a very specific aquatic habitat, which as their common name suggests, is the madicolous habitat (Vaillant 1956; Sinclair & Marshall 1987; Sinclair & Stuckenberg 1995). These habitats are characterized by thin films (2mm or less) of flowing water over various substrates (e.g. rock, mud or moss) and are also known as hygropetric zones (Vaillant 1956; Mackie 2004). The larvae must be in films of no more than 1mm in depth, so that the dorsum of the larva remains out of the water, which prevents drowning (Vaillant 1956; McLellan 1988). Examples of such habitats include rock-face seepages (either natural or man-made), cascading streams, splash zones near waterfalls, and boulders in fast moving streams (Sinclair 2000). The morphological adaptations which permit thaumaleid larvae to both breathe and stay anchored within the flow of water are the presence of an unpaired prothoracic proleg, an unpaired anal proleg, as well a pair of short respiratory tubes on the prothorax (Theischinger 2004). In addition, the larval integument has been noted to be hydrophobic (Vaillant 1956; Hinton 1958), potentially as a result of the layered, reticulated ultrastructure of the dorsal cuticle visible in scanning electron microscope images taken by Boussy et al. (1994). When reacting to disturbances, larvae will exhibit characteristic sidewinding behaviour, in an attempt to evade predators. As larvae, Thaumaleidae feed on diatoms and other detritus that accumulate on the substrate (Hinton 1958; Sinclair 1990; Arnaud & Boussy 1994). Pupation occurs in damp moss, debris or mud that the larvae seek out near the madicolous habitat (Vaillant 1959). Adults are secretive and are infrequently seen. They are poor fliers and never fly far from the larval habitat. They may be seen resting on riparian vegetation or directly on the madicolous substrate or crevices therein (personal observation). The adult diet is unknown. Due to the restricted habitat and weak flight dispersal ability of the adults, the Thaumaleidae are a rarely encountered and understudied group. Trapping does not typically yield favourable results: they are not attracted to any coloured trap, nor are they active fliers, therefore traps designed around flight interception prove ineffective. However, success has been achieved (albeit rare) by placing malaise traps directly above or in front of suitable habitat. New specialized traps may prove useful for collecting as well, such as the madicolous emergent trap developed by Shimabukuro et al. (2015). The best and most efficient method for collecting Thaumaleidae is to sweep riparian vegetation near the madicolous substrate. When swept, thaumaleids can often be found at the bottom of the net with their wings folded downwards across a transverse line of weakness near the apex of the subcosta (Theischinger 1986). If sweeping is unsuccessful, inspect the madicolous substrate for resting adults on moss, in cracks and crevices, or on the rock face itself, where they may be aspirated. Larvae may simply be plucked from the substrate using a pair of forceps. Since the Thaumaleidae are rarely encountered and thusly under collected, much taxonomic work is needed on this family. In order to learn more about the behaviour, ecology and biodiversity of the group, as well as madicolous habitats in general, taxonomic studies are required. These are critical to providing stable classifications within the family Thaumaleidae and will serve as a foundation for future ecological studies. Classification of Thaumaleidae The family Thaumaleidae is a fairly easy group to recognize. Their general habitus, with a stocky build and somewhat humpbacked appearance, resembles that of the Simuliidae. The following characters are used to identify Thaumaleidae: uniform wing venation with six or seven veins reaching the wing margin, absence of ocelli, palpus five-segmented and short, and slender antennae that are roughly identical in length to the head (McAlpine 1981). Thaumaleids are generally brownish grey to black in colour, however there are a small number of species that exhibit orange or yellow colouration (Edwards 1930; Dyar & Shannon 1924; Arnaud & Boussy 1994). Genitalic characters are the main way to distinguish between species, as other structural characters are notably uniform and differences of colour are usually indeterminate (Edwards 1929). Thaumaleidae belong to the paraphyletic group Nematocera (Courtney et al. 2009; Pape et al. 2011), alternatively known as the short-horned flies or lower Diptera. Early studies trying to determine the systematic position of the Thaumaleidae resulted in different schools of thought. One proposed classification system was that thaumaleids were most closely related to the Psychodidae, based on the transverse line of weakness near the base of the wing, resulting in the characteristic folded wings in death that only these two families share (Bezzi 1913; Edwards 1929). The competing theory to the Thaumaleidae + Psychodidae notion was that thaumaleids are more closely related to the Simuliidae and Ceratopogonidae. Saunders (1923) discussed this, as well as Crampton (1926), the latter studying the pleural sclerites and how they are similar to those of the Simuliidae and Ceratopogonidae. Edwards did note in his 1929 paper that the hypopygium of thaumaleids is similar to that of ceratopogonids and simuliids, as well as one genus of psycodid. Later studies have provided strong evidence to support the Thaumaleidae + Simuliidae as sister families. Hennig (1973) recognized four infraorders of Diptera, including the Culicomorpha, in which the Thaumaleidae are strongly nested. Subsequent studies have agreed with this hypothesis and further placed the Thaumaleidae within the superfamily Chironomoidea, together with the Simuliidae, Ceratopogonidae and Chironomidae (Wood & Borkent 1989; Oosterbroek & Courtney 1995; Sinclair et al. 2007). Wood and Borkent (1989) listed two morphological characters that support the inclusion of the Thaumaleidae in the Chironomoidea: presence of a prothoracic proleg and abdominal spiracles with posterior flaps modified into cylindrical procerci. A recent monograph by Borkent (2012) used pupal characters to propose a new superfamily known as the Simulioidea (Ceratopogonidae + Thaumaleidae + Simuliidae) where thaumaleids and simuliids are sisters. The characteristics that Borkent used to support the Thaumaleidae + Simuliidae group are as follows: egg swollen dorsally, larval stage with at least six instars, short pupal and adult antennae, and wing with fork of Cu situated less than 0.25 the total length of the wing. Molecular study of the phylogeny of Diptera has also supported a Thaumaleidae + Simuliidae clade. Pawlowski et al. (1996) were the first to recover this grouping by using partial 28S ribosomal RNA gene sequences resulting in 100% bootstrap support for this clade, which was also supported by Moulton (2000). Bertone et al. (2008) used four nuclear markers (28S rDNA, CAD, TPI and PGD) to resolve relationships among the lower Diptera. This resulted in the novel superfamily Simulioidea, consisting of Thaumaleidae + Simuliidae, again with 100% bootstrap support for this topology. This differs from the Simulioidea that Borkent (2012) proposed using morphology, in that he also includes the Ceratopogonidae. More recently, Wiegmann et al. (2011) used a combination of 14 nuclear loci and complete mitochondrial genomes to provide a phylogenomic estimate of fly relationships, again recovering the Thaumaleidae + Simuliidae clade with >95% bootstrap support. Thaumaleidae currently contains approximately 185 described species worldwide, classified within seven genera. The family is most speciose in the Palearctic region, while the Neotropical and Afrotropical fauna are the least well known with six and three species described, respectively. Compared to the Simuliidae (~2,120 spp.), thaumaleids are much less speciose, perhaps due to the vertebrate blood feeding habits of adult black flies (Borkent 2012). There is one described extinct species, Mesothaumalea fossilis Kovalev, that was discovered in fossil deposits in Transbaicalia, east of Lake Baikal in Siberia, Russia (Kovalev 1989). These deposits contain fossils from the Upper Jurassic/Early Cretaceous, which date back to 145mya. There is a simuliid fossil from the Jurassic period (176mya), which likely indicates that thaumaleids were present as well (Borkent 2012). Where the Thaumaleidae originated from and how they radiated is still unknown. Ruthe described the first thaumaleid in 1831 when he described Thaumalea testacea Ruthe at which time the family was known as Orphnephilidae. Bezzi proposed the use of the name Thaumaleidae in 1913. Bezzi’s paper was the first detailed study of the Thaumaleidae, providing an extensive bibliography, discussing previous studies, as well describing new species. The next large revision of the family was Edwards' 1929 monograph. Edwards recognized the usefulness of terminalia in species identification (both male and female). He also revised all known species at the time and placed them in four genera: Androprosopa Mik, Austrothaumalea Tonnoir, Thaumalea Ruthe and Trichothaumalea Edwards. Vaillant and Schmid described most of the Palearctic fauna, while other scientists described much smaller numbers. The Australasian species belong to three genera (Austrothaumalea, Niphta Theischinger and Oterere McLellan) and are well known, particularly because of work from Theischinger (1986), McLellan (1988) and Sinclair (2000, 2008a, 2008b). The Neotropics and Afrotropics still require much study. Only six species are known from the Neotropics, all confined to southern Chile and Argentina and sharing the same genera as those in Australasia (Edwards 1930; Schmid 1970; McLellan 1988). Only three Afrotropical species have been described. They are all in the genus Afrothaumalea Stuckenberg and found in South Africa (Stuckenberg 1960; Sinclair & Stuckenberg 1995; Sinclair 2015). It is expected that focused collecting in appropriate microhabitats will continue to yield new species worldwide; particularly within the latter two understudied regions. The Nearctic region consists of three genera and 29 described species: Androprosopa, Thaumalea and Trichothaumalea. The earliest taxonomic work on the Nearctic Thaumaleidae dates back to Bezzi’s 1913 paper in which he described the first Nearctic species, Androprosopa americana (Bezzi). There have been numerous Androprosopa discoveries since then. These will be discussed later in this paper. The genus Trichothaumalea is rare and difficult to predict where it will be collected. When collected, it seems to be in small numbers. Trichothaumalea has an interesting distribution with two species found in western North America, one species in eastern North American and one species in Japan, exhibiting a Trans-Pacific biogeographic vicariance (Sinclair & Saigusa 2002). The third genus found in North America is represented by one species, Thaumalea verralli Edwards. Thaumalea verralli is primarily a Palearctic species whose presence in North America gives it the first Holarctic distribution for the Thaumaleidae (Sinclair 1996). Sinclair (1996) hypothesized that T. verralli’s occurrence in North America is likely due to artificial introduction via a ship’s ballast, as it has only been collected from one of the oldest seaports in North America, located in St. John’s, Newfoundland. No phylogenies of the Thaumaleidae have been constructed to depict generic or species relationships. Sinclair has hypothesized generic relationships, indicating that Thaumalea + Androprosopa is the sister group to the remaining genera, and that the abbreviated Sc vein is a synapomorphy supporting the Trichothaumalea + Afrothaumalea + Niphta + Oterere + Austrothaumalea assemblage (pers. comm.). Previously (Sinclair 2000), the absence of an apical, spine-like gonostylar setae in Afrothaumalea, Niphta, Oterere and Austrothaumalea further supported this hypothesis, but the recent discovery of Afrothaumalea stuckenbergi Sinclair (2015) has put this hypothesis into question. A. stuckenbergi has independently derived a crown of spines on its gonostyli as opposed to 1-2 apical spines seen in Androprosopa and Thaumalea. Developing a stable phylogeny of the Thaumaleidae would be extremely useful in testing generic concepts and relationships, particularly amongst the Gondwanan genera. Genus Androprosopa Mik Androprosopa Mik is a Holarctic genus with about half of the species distributed in the Nearctic Region and the remainder in the Palearctic Region. The type species, Androprosopa larvata (Mik), was described in 1888 originally under the genus Orphnephila Haliday. Ten years later, Mik described the new genus Androprosopa and subsequently designated A. larvata as the type species (Mik 1898). After the description of A. larvata, no other species had been described as belonging to Androprosopa, rather they were described under Thaumalea or Orphnephilina Enderlein. Martinovský and Rozkošný (1976) observed that the form of the genitalia of several species of Thaumalea resembled that of Orphnephilina and proposed that eight species be reassigned to the genus Orphnephilina, including several Nearctic species. Since then, the assignment of species within Androprosopa had been problematic until Sinclair (1996) redefined the genus on the basis that gonostylar action is oblique in Androprosopa vs. the gonostylar action in the horizontal plane of Thaumalea. This resulted in the expansion of the genus to include 51 species, as well as the recognition that Orphnephilina is a junior synonym to Androprosopa as originally proposed by Wagner in 1995. Until now, there have been 52 described species of Androprosopa, with the latest description being A. zempoala Sinclair & Huerta (2010). The first taxonomic work on Nearctic Androprosopa was Bezzi’s 1913 description of the eastern species A. americana. The other two remaining eastern species are A. thornburghae (Vaillant) and A. vaillantiana Sinclair. Sinclair’s 1996 paper dealt with all the eastern Nearctic Thaumaleidae and provided excellent drawings and keys to the males, pupae and larvae. The genus is far more diverse in the western half of the Nearctic Region with 22 described species (Dyar & Shannon 1924; Garrett 1925; Schmid 1970; Brothers 1972; Arnaud & Boussy 1994; Sinclair & Huerta 2010). Arnaud & Boussy’s 1994 treatment of the group is the largest to date. They described 17 new species, as well as provided a diagnostic species key for males to all of the western Nearctic Thaumaleidae. They described females for some species, though they still pose a problem for identification, as they are difficult to associate with males. Gillespie et al. (1994) discussed the biology and taxonomy of larvae and pupae of ten species of Androprosopa found in Idaho and California. They provided descriptions and drawings of both life stages. However no key was provided and this represents less than half of the western fauna. Descriptions of the larvae and pupae of remaining species would be useful not only for identification, but also in the construction of a phylogeny to determine relationships between Nearctic species and Palearctic species. Due to the absence of a phylogeny, the relationships amongst the Nearctic species of Androprosopa remain uncertain. The eastern species are characterized by short, blunt apical gonostylar spines, tapering gonostyli and the dense setae on the inner face of the gonocoxites, and may be related to A. ericfisheri (Arnaud & Boussy) (Sinclair 1996). Larval antennal characters may also have phylogenetic use (Sinclair 1996). Also, some species have only been collected on one occasion and are known from only a few specimens (A. fusca (Garrett) & A. zelmae (Arnaud & Boussy)), which make it difficult to assess intraspecific variation. Alternatively, species such as A. gillespieae (Arnaud & Boussy) have a very wide distribution amongst varying habitats, which may indicate a species complex. Despite all of the work done on the Nearctic Thaumaleidae, they remain a very under collected and understudied group with much to be learned about diversity and relationships at all levels of classification. Objectives Given that little has been done in regard to exploring the evolutionary relationships within the Thaumaleidae, the purpose of this thesis is to revise the Nearctic species of Androprosopa and propose a hypothesis on relationships between them. All Nearctic species known prior to this work and new species recognized herein are described, with the male terminalia and distributions illustrated. Molecular sequence data and male terminalia are used to test and expand existing species concepts. Molecular data is also used to associate sexes and larvae wherever possible. A revised key to males is provided to identify all known Nearctic species using new information on species limits and useful characters. Molecular data is used to provide a hypothesis on relationships between the eastern and western fauna, as well as to propose species groups therein. Dissertation Organization This dissertation is organized into four chapters. Chapter 1 summarizes the knowledge of the family Thaumaleidae and the genus Androprosopa, particularly within the Nearctic Region. Chapter 2 is a revision of the Nearctic Androprosopa, and includes descriptions of six new species, redescriptions of the remaining species, distribution maps and a revised key to the males of the Nearctic Thaumaleidae. Chapter 3 provides information on the phylogenetics of Nearctic Androprosopa. Chapter 4 is a general conclusion and includes a discussion of avenues for future research dealing with thaumaleids. As per Article 8.2 of the International Code of Zoological Nomenclature (1999), this document is not issued for the permanent scientific record or for the purposes of zoological nomenclature. Consequently, any species names contained herein should not be considered as published (sensu ICZN). LITERATURE CITED International Code of Zoological Nomenclature. (1999) Fourth edition. International Trust for Zoological Nomenclature, Natural History Museum, London. Arnaud, P.H., Jr., & Boussy, I.A. (1994) The adult Thaumaleidae (Diptera: Culicomorpha) of western North America. Myia, 5, 41-152. Bertone, M.A., Courtney, G.W. & Wiegmann, B.M. (2008) Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes. Systematic Entomology, 33, 668-687. Bezzi, M. (1913) Taumaleidi (Orfnefilidi) italiani con descrizione di nuove specie. Bollettino del Laboratorio de Zoologia Generale e Agraria della R. Scuola Superiore d’Agricoltura in Portici, 7, 227-266. Borkent, A. (2012) The Pupae of Culicomorpha – morphology and a new phylogenetic tree. Zootaxa, 3396, 1-98. Boussy, I.A., Gillespie, J.M. & Arnaud, P.H., Jr. (1994) External structure of larval Thaumalea buckae Arnaud and Boussy (Diptera: Thaumaleidae). Myia, 195-201. Brothers, D.R. (1972) A new species of Thaumalea from California (Diptera: Thaumaleidae). The Pan-Pacific Entomologist, 48 (2), 121-122. Brown, B.V. (2005) Malaise trap catches and the crisis in Neotropical Dipterology. American Entomologist, 51, 180–183. Brown, B.V. (2009) Introduction. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera, Volume 1, NRC Research Press, Ottawa, Ontario, Canada, pp. 1–7. Crampton, G.C. (1926) A phylogenetic study of the thoracic sclerites of the psychodoid Diptera, with remarks on the interrelationships of the Nematocera. Entomological News, 37, 65-70. Courtney, G.W., Pape, T., Skevington, J.H., & Sinclair, B.J. (2009) Biodiversity of Diptera. In: R.G. Foottit & P.H. Adler (Eds) Insect Biodiversity: Science and Society, Blackwell Publishing, Oxford, UK, pp.185-222. Curran, C.H. (1965) The Families and Genera of North American Diptera. Second Edition. Henry Tripp, Woodhaven, New York, pp. 12-14. Dyar, H.G. & Shannon, R.C. (1924) The American species of Thaumaleidae (Orphnephilidae) (Diptera). Journal of the Washington Academy of Sciences, 14 (18), 432-434. Edwards, F.W. (1929) A revision of the Thaumaleidae (Dipt.). Zoologischer Anzeiger, 82, 121-142. -------- (1930) Bibionidae, Scatopsidae, Cecidomyiidae, Culicidae, Thaumaleidae (Orphnephilidae), Anisopodidae (Rhyphidae). In: Diptera of Patagonia and South Chile. British Museum (Natural History), London. Part II, 3, 77-119, pls 9-11. Enderlein, G. (1936) 22. Ordnung: Zweiflügler, Diptera. Abteilung 16. In: Brohmer, Ehrmann & Ulmer. Tierwelt Mitteleuropas 6, Insekten, III Teil, Leipzig, 259 pp. Gillespie, J.M., Barr, W.F. & Elliot, S.T. (1994) Taxonomy and biology of the immature stages of species of Thaumalea occurring in Idaho and California (Diptera: Thaumaleidae). Myia, 5, 153-193. Garrett, C.B.D. (1925) Seventy new Diptera – Key to the Pseudoleria Helomyzidae. Tipulidae, Chironomidae, Dixinae, Orphnephilidae, Mycetophilidae. Privately published, Cranbrook, British Columbia, 16 pp. Hall, R.D. & Gerhardt, R.R. (2009) Flies (Diptera). In: Mullen, G.R. and Durden, L.A. (Eds.), Medical and Veterinary Entomology, Second Edition, Academic Press, New York, New York, U.S.A., pp. 137-152. Hammond, P. (1992) Species inventory. In: Groombridge, B. (Ed.), Global Biodiveristy: Status of the Earth’s Living Resources, Chapman & Hall, London, pp. 17–39. Hennig, W. (1973) Diptera. Handbuch der Zoologie 4 (2), 2, 31, 1-337. Hinton, H.E. (1958) On the nature and metamorphosis of the colour pattern of Thaumalea (Diptera, Thaumaleidae). Journal of Insect Physiology, 2, 249-260. Kovalev, V.G. (1989) [Geological history and the systematic position of the family Thaumaleidae (Diptera).] Entomologicheskoe Obozrenie, Revue d’Entomologie PURSS, 68 (4), 798-808. Mackie, G.L. (2004) Applied Aquatic Ecosystem Concepts: Second Edition. Kendall Hunt Publishing, Dubuque, Iowa, 374 pp. Marshall, S.A. (2012) Flies: the natural history and diversity of Diptera. Firefly Books Ltd., Richmond Hill, Ontario, 616 pp. Martinovský, J. & Rozkošný, R. (1976) Four new species of Thaumaleidae (Diptera) from Europe with taxonomic and distribution notes on other species. Acta entomologica bohemoslovaca, 73, 187-205. McAlpine, J.F. (1981) Key to Families - Adults. [Chapter] 4. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M., coords, Manual of Nearctic Diptera, Volume 1. Agriculture Canada Monograph, 27, pp. 89-124. McLellan, I.D. (1988) A revision of New Zealand Thaumaleidae (Diptera: Nematocera) with descriptions of new species and a new genus. New Zealand Journal of Zoology, 15, 563-575. Mik, J. (1888) Vorlaufige Diagnose einer neuen Dipteren-Art. Wiener Entomologische Zeitung, 7 (7), 243. --------(1898) Dipterologische miscellen. (2. Serie). X. Wiener Entomologische Zeitung, 2, 60-66. Moulton, J.K. (2000) Molecular sequence data resolved basal divergences within Simuliidae (Diptera). Systematic Entomology, 25, 95-113. Oosterbroek, P. & Courtney, G. (1995) Phylogeny of the nematocerous families of Diptera (Insecta). Zoological Journal of the Linnean Society, 115 (3), 267-311. Pape, T., Blagoderov, V. & Mostovski, M.B. (2011). Order DIPTERA Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148, 222-229. Pawlowski, J., Szadziewski, R., Kmieciak D., Fahrni, J. & Bittar, G. (1996) Phylogeny of the infraorder Culicomorpha (Diptera: Nematocera) based on 28S RNA gene sequences. Systematic Entomology, 21, 167-178. Ruthe, J.F. (1831) Einige Bemerkungen und Nachträge zu Meigen’s “Systematischer Beschreibung der europäischen zweiflügeligen Insecten.” Isis (Oken’s), pp. 1203-1222. Saunders, L.G. (1923) LXXIII. On the Larva, Pupa, and systematic position of Orphnephila testacea, Macq. (Diptera Nematocera). Annals and Magazine of Natural History, 9 (11), 631-640. Schmid, F. (1970) Quelque Thaumaleides nouvelles ou peu connues (Diptera: Thaumaleidae). Le Naturaliste Canadien, 97 (5), 499-510. Shimabukuro, E.M., Pepinelli, M., Perbiche-Neves, G. & Trivinho-Strixino, S. (2015) A new trap for collecting aquatic and semi-aquatic insects from madicolous habitats. Insect Conservation and Diversity, 1-6. Sinclair, B.J. (1990) The madicolous fauna of southern Ontario, with emphasis on the Niagara Escarpment. In: Conserving Carolinian Canada; conservation biology in the deciduous forest region, pp. 281-288. ---------(1996) A review of the Thaumaleidae (Diptera: Culicomorpha) of eastern North America, including a redefinition of the genus Androprosopa Mik. Entomologica scandinavica, 27, 361-376. ---------(2000) Immature stages of Australian Austrothaumalea Tonnoir and Niphta Theischinger (Diptera: Thaumaleidae). Australian Journal of Entomology, 39, 171-176. ---------(2008a) A new species of Austrothaumalea Tonnoir from New Caledonia (Diptera: Thaumaleidae). In: Grandcolas, P. (ed.), Zoologia Neocaledonica 6. Biodiversity studies in New Caledonia. Mémoires du Muséum national d’Histoire naturelle, 196, 291-295. ---------(2008b) New species of Austrothaumalea Tonnoir from Australia (Diptera: Thaumaleidae). Records of the Australian Museum, 60 (1), 1-12. ---------(2015) A new species of Afrothaumalea Stuckenberg, 1960 (Diptera: Thaumaleidae) from the Western Cape (South Africa) and first description of the pupa of this genus. African Invertebrates, 56 (1), 161-166. Sinclair, B.J., Borkent, A. & Wood, D.M. (2007) The male genital tract and aedeagal components of the Diptera with a discussion of their phylogenetic significance. Zoological Journal of the Linnean Society, 150, 711-742. Sinclair, B.J. & Huerta, H. (2010) A new species of Androprosopa from Mexico (Diptera: Thaumaleidae). The Canadian Entomologist, 142, 443-447. Sinclair, B.J. & Marshall, S.A. (1987) The madicolous fauna in southern Ontario. Proceedings of the Entomological Society of Ontario, (1986), 117, 9-14. Sinclair, B.J. & Saigusa, T. (2002) A new species of the seepage midge genus Trichothaumalea Edwards from Japan. Insect Systematics & Evolution, 33, 175-184. Sinclair, B.J. & Stuckenberg, B.R. (1995) Review of the Thaumaleidae (Diptera) of South Africa. Annals of the Natal Museum, 36, 209-214. Stuckenberg, B.R. (1960) A new genus and species of Thaumaleidae from South Africa (Diptera). Proceedings of the Royal Entomological Society of London (B), 29, 107-109. Theischinger, G. (1986) Australian Thaumaleidae. Records of the Australian Museum, 38, 291-317. ---------(2004) 50: Insecta: Diptera, Thaumaleidae. In: Yule, C.M. & Yong, H.S., eds, Freshwater Invertebrates of the Malaysian Region, Academy Sciences of Malaysia, pp. 669-672 Vaillant, F. (1956) Recherches sur la faune madicole (hygropétriques s.l.) de France, de Corse et d’Afrique du Nord. Mémoires du Muséum National d’Histoire Naturelle, nouvelle série A, Zoologie, 11, 258 pp. ---------(1959) The Thaumaleidae (Diptera) of the Appalachian Mountains. Journal of the New York Entomological Society, 7, 31-37. --------- (1977) Les Diptères Thaumaleidae d’Europe. Annales de la Société entomologique de France, 13 (4), 695-710. Wagner, R. (1995) Thaumaleiden (Diptera, Thaumaleidae) aus Österreich, Slowenien und Kroatien und ein Gattungssynonym. Studia dipterologica, 2, 65-68. Wiegmann, B.M., Trautwein, M.D., Winkler, I.S., Barr, N.B., Kim, J.W., Lambkin, C., Bertone, M.A., Cassel, B.K., Bayless, K.M., Heimberg, A.M., Wheeler, B.M., Peterson, K.J., Pape, T., Sinclair, B.J., Skevington, J.H., Blagoderov, V., Caravas, J., Kutty, S.J., Schmidt-Ott, U., Kampmeier, G.E., Thompson, F.C., Grimaldi, D.A., Beckenbach, A.T., Courtney, G.W., Friedrich, M., Meier, R. & Yeates, D.K. (2011) Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences, 108, 5690–5695. Wood, D.M. & Borkent, A. (1989) Phylogeny and classification of the Nematocera. [Chapter] 114. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. coords, Manual of Nearctic Diptera, Volume 3. Agriculture Canada Monograph, 32, pp. 1333-1581. Woodley, N.E., Borkent, A. & Wheeler, T.A. (2009) Phylogeny of the Diptera. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera, Volume 1, NRC Research Press, Ottawa, Ontario, Canada, pp. 79–94. Yeates, D.K., Wiegmann, B.M., Courtney, G.W., Meier, R., Lambkin, C. & Pape, T. (2007) Phylogeny and systematics of Diptera: Two decades of progress and prospects. Zootaxa, 1668, 565–590. Chapter 2: REVISION OF THE NEARCTIC ANDROPROSOPA MIK (DIPTERA: THAUMALEIDAE) AND SIX NEW SPECIES DESCRIPTIONS Abstract The Nearctic fauna of Androprosopa Mik is revised to include thirty-two species, six of which are described as new to science. Descriptions of the adult males of A. apache, A. arnaudi, A. boussi, A. magnipelvim, A. rainierensis and A. sierra. are presented. Redescriptions of all remaining species are provided, as well as genitalic illustrations and updated distribution maps for each species. A diagnostic key to all Nearctic Thaumaleidae is also provided. Introduction The Thaumaleidae, or madicolous midges, are a family of aquatic Diptera that is found worldwide, except in Antarctica. Thaumaleids consist of roughly 185 described species within 7 genera. As the common name suggests, thaumaleids are found in and around madicolous habitats. These habitats are characterized by thin films of flowing water over rock surfaces and are also known as hygropetric zones (Mackie 2004). The larvae of thaumaleids are restricted to these madicolous habitats, which may include seepages (either natural or man-made), cascading streams, as well as splash zones near waterfalls and boulders in fast moving streams (Sinclair 2000). The larvae are recognizable by the presence of an unpaired prothoracic proleg and an unpaired anal proleg, as well a pair of short respiratory tubes on the prothorax (Theischinger 2004). The escape behaviour is also useful for identifying a larval thaumaleid, as they use a quick, sidewinding motion, seemingly gliding along the water surface to evade predation. The larvae feed on diatoms on the rock surface (Sinclair 1990). When ready to pupate, the larvae leave the madicolous habitat and seek out damp moss, debris or mud (Vaillant 1959). The adults are poor fliers and may be swept from riparian vegetation or aspirated directly off the rock surface or any moss growing on it. A combination of sweeping and aspirating is the most effective way of collecting thaumaleids, while trapping has very limited use. They are not attracted to pan traps and are only collected in malaise traps when placed directly in the appropriate habitat. Due to this habitat specificity and flying ineptitude, the Thaumaleidae are a rarely collected and understudied family. There is one described extinct species, Mesothaumalea fossilis Kovalev, that was discovered in fossil deposits in Transbaicalia, east of Lake Baikal in Siberia, Russia (Kovalev 1989). These deposits contain fossils from the Upper Jurassic/Early Cretaceous, which date back to 145mya. There is a simuliid fossil from the Jurassic period (176mya), which likely indicates that thaumaleids were present as well (Borkent 2012). Where the Thaumaleidae originated from and how they radiated is still unknown. Their phylogenetic position is as the sister family to the Simuliidae (Borkent 2012; Moulton 2000; Wiegmann et al. 2011). Ruthe first described the Thaumaleidae in 1831, when he described Thaumalea testacea in what is now Germany (Arnaud & Boussy 1994). The family was known as Orphnephilidae until Bezzi proposed the name Thaumaleidae in 1913. The first Nearctic species, Thaumalea americana Bezzi, was also described in 1913 from Ithaca, New York. There are 3 Nearctic genera of Thaumaleidae: Androprosopa Mik, Trichothaumalea Edwards and Thaumalea Ruthe. Thaumalea verralli Edwards is the sole North American representative of the genus and a likely introduction from Europe (Sinclair 1996). T. verralli has only been recorded from St. John’s, Newfoundland, at one of the oldest seaports in North America (Sinclair 1996). There are 3 known Nearctic species of Trichothaumalea, 2 of which are found in the West and 1 in the East. Androprosopa is the largest genus in the Nearctic with 25 known species: 22 from the West and 3 from the East (Arnaud & Boussy 1994; Sinclair 1996). Androprosopa was first described by Mik in 1898, with the type-species Orphnephila larvata Mik. In 1995, Wagner proposed that Orphnephilina Enderlein be recognized as a junior synonym of Androprosopa. Sinclair officially recognized this change in 1996, while also redefining the genus. This redefinition was proposed on the basis that the genus Androprosopa, along with many species of Thaumalea, including all the Nearctic species assigned to Thaumalea, shared the synapomorphy of an oblique gonostylar action (Sinclair 1996). Therefore, all of the species with this synapomorphy are now included in Androprosopa, thus increasing the known species total for the genus to 52. This paper will focus on the western Nearctic fauna of Androprosopa as Vaillant (1959) and Sinclair (1996) have studied the eastern fauna in great detail. The first description of a western Nearctic Androprosopa species was A. elnora (Dyar & Shannon) in 1924. This was followed by the description of A. fusca (Garrett) 1925, A. anolo (Schmid) 1970 and A. santaclaraensis (Brothers) 1972. In 1994, Arnaud and Boussy treated the Thaumaleidae of Western North America. In the largest study of western thaumaleids to date, the authors recognized 21 species of Androprosopa (at the time the genus was still treated as Thaumalea), 17 of which were new to science. They also included a key to species using genitalia. The most recent new species description was by Sinclair & Huerta (2010), where they described A. zempoala from Mexico. Herein, 5 new species of Androprosopa are described, range expansions are noted and an up to date identification key is provided. Table 2.1 Location and species names for Nearctic Thaumaleidae. Location Western North America Eastern North America Genus Androprosopa Androprosopa Species anolo (Schmid 1994) americana (Bezzi 1913) apache sp. nov. thornburghae (Vaillant 1959) arnaudi sp. nov. vaillantiana Sinclair 1996 becca (Arnaud & Boussy 1994) brothersi (Arnaud & Boussy 1994) buckae (Arnaud & Boussy 1994) chandlerorum (Arnaud & Boussy 1994) coloradensis (Arnaud & Boussy 1994) confracta (Arnaud & Boussy 1994) elnora (Dyar & Shannon 1924) ericfisheri (Arnaud & Boussy 1994) falciformis (Arnaud & Boussy 1994) fusca (Garrett 1925) gillespieae (Arnaud & Boussy 1994) idahoensis (Arnaud & Boussy 1994) lindsayorum (Arnaud & Boussy 1994) magnipelvim sp. nov. melanderi (Arnaud & Boussy 1994) palouse (Arnaud & Boussy 1994) rainierensis sp. nov. santaclaraensis (Brothers 1972) schmidiana (Arnaud & Boussy 1994) sierra sp. nov. sonorensis (Arnaud & Boussy 1994) waha (Arnaud & Boussy 1994) zelmae (Arnaud & Boussy 1994) zempoala Sinclair & Huerta 2010 Genus Thaumalea Thaumalea Species verralli Edwards 1929 Genus Trichothaumalea Trichothaumalea Species pilosa (Garrett 1925) elakalensis Sinclair 1992 pluvialis (Dyar & Shannon 1924) Materials and Methods Terms used for adult structures follow those of McAlpine (1981), except wing venation where the interpretations of Sinclair (in press) and Saigusa (2006) are accepted. Homology of the male terminalia follows that of Sinclair (1992). For best results, thaumaleids should be collected into 95% ethanol. They may later be critical-point-dried or dried using the hexamethyldisilazane (HMDS) method, as outlined by Brown (1993). The latter method was used for the preparation of the newly described species. Males were identified using genitalic characters and were cleared using a hot, 85% lactic acid bath. Females and larvae were identified by association, when possible, as well as through molecular methods as outlined in (should I reference my molecular paper?). Label data for primary type specimens are presented exactly as they appear. Data are listed from the top downward on the staging pin, with data from each label enclosed in quotation marks; lines are delimited by a forward slash mark. The repository is given in parentheses. #’s of specimens studied, from where Taxonomy Androprosopa Mik Androprosopa Mik, 1898: 65. Type-species: Orphnephila larvata Mik (original designation). Orphnephilina Enderlein, 1936: 96. Type-species: Orphenphila nigra Loew (monotypy). Psychothaumalea Vaillant, 1970: 166. Type-species: Orphnephila nigra Loew (original designation). Protothaumalea Vaillant, 1978: 698. Type-species: Thaumalea tarda Loew (original designation). Diagnosis. Males are distinguished from Thaumalea by obliquely to dorsoventrally directed gonostyli. Androprosopa is distinguished readily from other thaumaleid genera, except Thaumalea, by a complete subcostal vein. Description. Male. Scutum with rows of undifferentiated setulae; scutellum with row of marginal setae. Supralar region flat, not produced into prominent ridge. Wing membrane lacking macrotrichia; Sc complete; basal section of R proximal to origin of Rs straight, not weakened; R+R1+R1+2 with macrotrichia along entire length; R2 situated close to base of R1; CuA1 branching close to CuA2; CuA2 lacking angular base and basal appendage. Abdominal sternite 8 rectangular with stout anterior margin projecting onto preceding segment. Terminalia: Hypandrium reduced to narrow sclerite. Epandrium often with posterolateral processes. Gonocoxites short, width subequal to length; gonocoxal plate normally broad, thinly sclerotized. Gonostyli articulate obliquely to dorsoventrally; usually apical spines present. Parameres divided into pair of long blades; dorsal parameral apodeme memranously connected to the posterolateral margin of the epandrium; membranous connection often sclerotized, forming pair of venterolateral epandrial plates. Female. Similar to male except as follows: hypogynial valves generally truncate apically, often width of notch narrow. Tergite 9 lacking lateral projections. Distribution. Known from the Nearctic, Palearctic and Oriental Regions. Key to adult males of North American Thaumaleidae 1. Sc not reaching C; macrotrichia on membrane of wing as well as veins………….. ……………………………………………..…….…...Trichothaumalea Edwards (3 spp.)…2 -Sc reaching C; macrotrichia confined to veins…………………………………………....4 2. Pair of sickle-shaped gonocoxal blades arising from base of gonocoxite; eastern species………………………………………….…….…..Trich. elakalensis Sinclair -Lacking gonocoxal blades; western species…………………………………………….....3 3. Gonstyli tapering to a narrow tip with three subapical spines in a row……… ……………………………………………………………………………..…..Trich. pilosa (Garrett) -Gonstyli with broad apex and scattered spines…………………………………………. ……………………………………………………….…...Trich. pluvialis (Dyar and Shannon) 4. Gonocoxal plate divided into pair of slender, pointed rods; parameres fused medially……………………………………………….……...…Thaumalea verralli Edwards -Gonocoxal plate undivided; parameres divided into pair of rods …………………………………………………………….….…Androprosopa Mik (30 spp.)…5 5. Gonostyli with 4 or more stout, apical spines; eastern species……………………..6 -Gonostyli with 4 or less apical spines, if 4 present then spines elongate; western species…………………………………………………………………………………………8 6. Epandrium with posterolateral process……………...A. thornburghae (Vaillant) Epandrium without posterolateral process………………………………………………...7 7. Gonostyli cylindrical, not tapering; bearing row of 8 apical spines………………… ……………………………………………………………………………..…A. vaillantiana Sinclair -Gonostyli tapering; bearing crown of 5-6 apical spines………………………………... …………………………………………………………………………….....….A. americana (Bezzi) 8. Body disctinctly yellow in colour…………………………………..……………………………9 Body colour brown, black or dark grey, at most brownish orange……………...10 9. Gonocoxite elongate with gonostyli extending beyond hind margin of epandrium; parameres distinct, with subapical projection……………………………. ………………………………………………………………..……A. elnora (Dyar and Shannon) -Gonocoxite short, quadrate with gonostyli not reaching hind margin of epandrium; paramers reduced…………….….A. ericfisheri (Arnaud and Boussy) 10. Gonostyli raptorial in appearance, outer margin bearing 2 setae, basal lobe setose, apex with a stout spine and longer, sickle-shaped process; parameres with small flange before apex……………………..….A. santaclaraensis (Brothers) -Gonostyli not as above…………………………………………………………………………....11 11. Gonostyli with 3-4 elongated, claw-like spines at apex…………………………..…..12 -Gonostyli with less than 3 apical spines…………………………………………………...13 12. Gonocoxite with lindsayorum - sonorensis compare types 13. Gonostyli with distinct lateral projection or process near midpoint, nearly devoid of setae………………………………………………………………………………………...14 -Gonostyli without lateral projection or process…………………………….....………15 14. Gonostyli strongly curved, with a large, thickened seta on medial inner margin; parameres without distinct bend…………A. rainierensis new species -Gonocoxite with posterolateral process short, blunt, not extending laterad of of gonostyli; gonostyli elbowed at apical third with lateroventral process at elbow; parameres with strong bend apically……………………………………………... ……………………………………………………………..A. falciformis (Arnaud and Boussy) 15. Gonocoxite with posterolateral process long, sharp, extending at least midway down laterad of gonostyli; parameres strongly curved…………………16 -Gonocoxite without posterolateral process, at most bluntly lobed; parameres never strongly curved……………………………………………………………………….……..19 16. Gonostyli with a distinct tip bearing a single seta at apex……………..………..…..17 -Gonostyli either expanding apically or with subapical seta……………………….18 17. Gonocoxal process extending to about middle of gonostyli; distal half of gonostyli expanded………………………………...……………………..…A. anolo (Schmid) -Gonocoxal process extending length of gonostyli; distal half of gonostyli narrowed……………………………………………………………..………..…A. fusca (Garrett) 18. Parameres bent apically at right angle, undivided at apex…………………………... ……………………………………………………………….......A. zelmae (Arnaud and Boussy) -Parameres bifurcate apically……………………....…A. waha (Arnaud and Boussy) 19. Lateral epandrial process present…………....…………………...………………………….20 -Lateral epandrial process absent…………………………………………. 20. pal/schmid/gil/buck/Idaho/zemp/AZ/NV/178/Col Species Diagnosis and Descriptions Androprosopa americana (Bezzi) Orphnephila testacea (Ruthe); authors (misident.) Thaumalea americana Bezzi, 1913: 250. Thaumalea johannis Dyar and Shannon, 1924: 434. Androprosopa anolo (Schmid) Thaumalea anolo Schmid, 1970: 501. Androprosopa anolo (Schmid); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ pinned with a minuten on a piece of card labeled, “U.S.A., Ore./Marion Co./French Creek/18-VI-1965/F. Schmid”; “HOLOTYPE/CNC No. 11.306/Thaumalea/anolo Schmid.” (CNC) The abdomen and right wing are mounted on separate microscope slides and labeled with the same label data as the holotype. Recognition The species is recognized by its strongly recurved parameres and is distinguished from the closely related A. falciformis by lacking elbowed gonostyli. Redescription Wing length: 2.64-2.91mm. Colouration: Head blackish-brown. Mesonotum and pleura dark brown; abdomen blackish-brown, dull; halter and legs yellow-brown, tarsi darker; terminalia brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin, then slightly divergent just before margin. Abdomen: Male abdominal sternites 2-7 rectangular with basal half well sclerotized, lightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 with few setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium nearly cylindrical, slightly longer than wide; posterior margin not indented; each posteroventral corner bearing a pair of double-dentate apical epandrial processes. Hypandrium convex medially, slightly arching into sternite 8. Gonocoxite longer than wide, with posterolateral process about half the length of gonostylus. Gonostylus shorter than gonocoxite; sudden thickening at midpoint, bearing single apical claw-like setae; with few setae. Parameres paired; broad basally, narrowed sharply medially and distal portion strongly recurved laterally. Gonocoxal plate attaining length of posterolateral gonocoxal processes. Additional Material Examined U.S.A.: IDAHO: Shoshone Co.: Marble Ck Rd, FR 321, St. Joe NF, 20.vi.2014 47°11.557N116°04.549W ex. roadside str., 890m, B.J. Sinclair (1♂); OREGON: Linn Co.: Santiam SF, Monument Peak Rd, 10.vii.2013, 44°42'34"N122°22'16"W, 820m, B.J. Sinclair, (1♂); Marion Co.: Willamette NF, French CK Rd, 10.vii.2013, 44°46'58"N122°11'12"W, trib, 660m, B.J. Sinclair (2♂); WASHINGTON: Mt. Rainier NP, E Christine Falls, 16.vii.2012, 46°46'47"N121°46'21W, roadside seeps, 1125m, B.J. Sinclair (1♂); Mt Rainier NP, below Comet Falls, 3.vii.2013, 46°47'31"N 121°46'59"W, seeps, 1425m, B.J. Sinclair (4 larvae); Pierce Co.: Mt. Rainier N.P., WA-123, 1.vii.2014, 46°49'33.4"N 121°32'36.1"W, ex. roadcut seeps/waterfalls, J.K. Moulton & R.J. Pivar (11♂); Skamania Co.: Gifford Pinchot NF, FR 23 mi 22, 7.vii.2013, 46°19'54"N121°43'12"W, 730m, roadside streams, B.J. Sinclair (1♀). Distribution Found in the Cascade Mountain Range (Washington & Oregon) as well as the Columbia Plateau in Idaho. Androprosopa becca (Arnaud & Boussy) Thaumalea becca Arnaud & Boussy, 1994: 60. Androprosopa becca (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ mounted on microscope slide under two cover glasses labeled, “British Columbia/14mi. S. Valemont,/Hwy. 5 15 Aug., 1969/E.M. & J.L. Fisher/♂” (CAS Entomology Type #17000). Allotype: ♀ with same collection data also deposited in CAS. Paratypes: CANADA: BRITISH COLUMBIA: 14 miles south Valemont, Highway 5, 15.viii.1969, E.M. & J.L. Fisher (14♂, 4♀; CAS, CDFA, USNM); USA: IDAHO: Idaho Co.: 49.5miles northeast Lowell, Warm Springs Creek, 23-V-1972, J.M. Gillespie (6♂; CAS, UIC); 55miles northeast of Lowell, 15-16.viii.1971, J.M. Gillespie (3♂, 1♀; CAS); 69miles northeast Lowell, 16.vii.1971, J.M. Gillespie (5♂, 3♀; CAS); WASHINGTON: Skagit Co., Marblemount, 31.v.1965, F. Schmid (4♂, 3♀; CAS, CNC). Recognition This species is recognized by gonostyli armed with a long terminal spine (nearly ½ the length of gonostyle) and parameres with a subapical, lateral beak. Redescription Wing length: 2.86-3.51mm. Colouration: Head brown, dull. Mesonotum and pleura brown, shiny; abdomen dark brown to blackish-brown, dull; halter and legs light brown, tarsi darker; terminalia light brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with basal half well sclerotized, not dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 with few setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium nearly cylindrical, slightly longer than wide; posterior margin not indented; without apical epandrial processes. Hypandrium strongly arching into sternite 8, distinct. Gonocoxite rectangular, longer than wide. Gonostylus shorter than gonocoxite; arched; apex bearing long, thin apical spine, nearly half the length of gonostylus. Parameres paired, subequal in length compared to gonostyli; widening medially, rounded at apex with subapical on lateral margin. Gonocoxal plate shorter than posterior margin of gonocoxites; with thin medial line. Additional Material Examined CANADA: ALBERTA: Canmore, Grassi Lakes Trail, 13.vii.2015, 51°04.87"N115°23.64"W, 1420m, ex. streams, B.J. Sinclair (17♂, 8♀); Hwy. 40, Lougheed Pk., 20.viii.1987, 50°39'10"N115°10'49"W, J. Troubridge (1♂; DEBU); BRITISH COLUMBIA: 13km W. Revelstoke, 20.vii.2012, 50°57'45"N 118°22’30”W, roadcut seep, 560m, B.J. Sinclair (1♂); East Kootenay Moyie R., Hwy 3/95, 19.vi.2014, 49°10.279’N116°00.487’W, 900m, ex. roadcut seeps, B.J. Sinclair (1♀, 7 larvae, 1 pupa); East Kootenay, Hwy 3/95, 22.vi.2014, 49°10'18.4"N116°00'31.9"W, ex. roadcut seeps, J.K. Moulton & R.J. Pivar (19♂, 2♀, 47 larvae); Moyie River, Hwy 3, 12.vii.2012, 49°10'18" -116°00'31", roadcut seep, 915m, B.J. Sinclair (11 larvae)put pupae reared info; nr. Kimberley, Meachen Ck. FR, 17.vii.2015, 49°36.21"N 116°13.45"W, 1027m, ex. roadside falls, B.J. Sinclair (1♂); nr. Kimberley, Meachen Ck. FR, 17.vii.2015, 49°36.22"N116°13.33"W, 1021m, ex. roadside falls, B.J. Sinclair (14♂); W. Creston-Hwy 3, 18.vii.2015, 49°06.24"N116°53.07"W, 1130m, ex. roadcut seep, B.J. Sinclair (2♂); U.S.A: MONTANA: Lincoln Co.: Hwy. 37, 24.vi.2014, 48°41'24.6”N115°18'49.2"W, ex. roadcut seeps, J.K. Moulton & R.J. Pivar (13♂, 1♀, 7 larvae); WASHINGTON: Pierce Co.: Mt. Rainier N.P. Christine Falls, 2.vii.2014, 46°46'51.2"N121°46'45.5"W, ex. small falls E of Chris. Falls next to road, J.K. Moulton & R.J. Pivar (2♂). Distribution Found from Mt. Rainier in the Cascade Mountains, East to the Columbia Plateau in North Central Idaho and North to Columbia Mountains of northwest Montana and southeast British Columbia. Androprosopa brothersi (Arnaud & Boussy) Thaumalea brothersi Arnaud & Boussy, 1994: 63. Androprosopa brothersi (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ point-mounted with apex of abdomen removed is labeled: “Uvas Canyon/Santa Clara Co./Cali. XI-5-72”; “Swanson Creek”; “D.R. Brothers/Acc. No. 9758”; “Arnaud and Boussy/ Dissection No./23.04.93”; “HOLOTYPE Male/Thaumalea/brothersi/Arnaud and Boussy/1994; CAS Ent. Type No./17001.” Apical portion of the abdomen of the holotype is mounted on a microscope slide with equivalent label information (CAS Entomology Type #17001). Allotype: ♀ mounted on two microscope slides under three cover glasses is labeled: “ALLOTYPE Female/Thaumalea/brothersi/Arnaud and Boussy/1994”; “Swanson Creek/Uvas Canyon/County Park/Santa Clara Co., Calif., III-9-73/reared from/immature/D.R. Brothers/Acc. No. 10020”; Arnaud and Boussy/Dissection No./17-IV-1973”; “Arnaud & Boussy/wing print, 1993” (CAS). Paratypes: U.S.A: CALIFORNIA: Santa Clara Co., 2♂ with same label data as holotype but dated “III-9-73” and “Acc. No. 10018” and “Acc. No. 10030”; 1♂ with same label data as holotype but dated “IX-21-72” and “Acc. No. 9748” (CAS, DRB). 2♀ with similar collection data as allotype but with accession numbers “10021” and “10022” (CAS, DRB). Recognition Distinguished from the closely related A. melanderi by the hourglass shaped gonocoxal plate, and from A. confracta by having only one apical seta on the gonostyli. Redescription Wing length: 2.24-2.96mm. Colouration: Head dull, dark brown. Mesonotum and pleura dark brown and dull; scutellum slightly darker; abdomen blackish-brown; halter and legs pale brown, tarsi darker; terminalia brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 weak; R3 and R4+5 converging toward wing margin, then R4+5 diverging weakly at wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized and slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium a truncated cone, slightly wider than long; posterior margin with shallow indentation; apical epandrial processes not produced. Hypandrium very slender, faint; slighty convex. Gonocoxite ovoid, slightly longer than wide. Gonostylus stout; shorter than gonocoxites; base broad, tapering towards tip; apex bearing single, strong claw-like spine. Parameres paired, longer than gonostyli; lateral margin with irregular serration; pointed lateral projection at apex. Gonocoxal plate broad, extending past posterior margin of gonocoxites; produced distomedially into hourglass-like shape, ending near the middle of gonocoxite. Additional Material Examined U.S.A.: CALIFORNIA: Nevada Co.: culvert along Sagehen Crk., 11.vii.2012, 39°26'04.4"N120°16'52.2"W, J.M. Cumming (1♂, 1♀); Tahoe N.F., River Rd. Hwy. 89, 9.vii.2014, 39°15’56.3”N120°12’37.1”W, ex. small roadside waterfall , J.K. Moulton & R.J. Pivar (1♂,3♀). Distribution Found in the Santa Cruz Mountains and the Sierra Nevada’s in California. Androprosopa buckae (Arnaud & Boussy) Thaumalea sp., Arnaud 1968: 83 (First record of the family in central California, at Marin County.) Thaumalea buckae Arnaud & Boussy, 1994: 66. Androprosopa buckae (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ mounted on a microscope slide under two cover glasses is labeled: “HOLOTYPE/Thaumalea/buckae/Arnaud & Boussy/1994/♂ KOH/USA: CALIFORNIA: Marin Co./Mill Valley, Lee Street, 360’ el./ex dripping cliff 22 III 1969/ coll. I. A. Boussy.” (CAS Entomology Type #17002). Allotype: ♀ mounted on a microscope slide under two cover glasses with same collection data as the holotype except the date and collector reads “17 May 1967 by Paul H. Arnaud, Jr.” There has been an incorrect wing association with the allotype (CAS). Paratypes: U.S.A.: CALIFORNIA: Marin Co.: Mill Valley, Lee Street, 360 feet: 13.v.1967, P.H. Arnaud, Jr. (2♂, 1♀; CAS); 15.v.1967, P.H. Arnaud, Jr. (4♂, 2♀; CAS); 17.v.1967, P.H. Arnaud, Jr. (6♂, 1♀; CAS); 21.v.1967, P.H. Arnaud, Jr. (2♂; CAS); 22.v.1967, PH. Arnaud, Jr. (12♂, 2♀; CAS); 24.ii.1969, I.A. Boussy (1♂; CAS); 2.iii.1969, I.A. Boussy (3♂; CAS); 22.iii.1969, I.A. Boussy (12♂, 3♀; CAS); 28.iii.1971, I.A. Boussy (1♂, 1♀; CAS); 8.xi.1971, I.A. Boussy (2♂; CAS); Sausalito, 27.v.1969, P.H. Arnaud, Jr. (1♂; CAS); Stinson Beach State Park, 2nd inlet S main beach, 16.iii.1969, overcast day, I.A. Boussy (2♂; CAS); Santa Clara Co.: Black Rock Falls, Uvas Canyon County Park, 27.v.1972, Accession #9026, D.R. Brothers (1♂; DRB); Uvas Creek, Uvas Canyon County Park, 27.v.1972, Accession #9029, D.R. Brothers (1♂; DRB); same locality, 1.viii.1972, Accession #9045, D.R. Brothers (1♂; DRB); same locality, 7.x.1972, Accession #9802, D.R. Brothers (1♂; DRB); Seep along Uvas Creek, Uvas Canyon County Park, 27.v.1972, Accession #9031, D.R. Brothers (1♂; DRB); Swanson Creek, Uvas Canyon County Park, 21.ix.1972, Accession #9748, D.R. Brothers (1♂; DRB); Santa Cruz Co.: Big Basin Redwoods State Park, Opal Creek, 5.viii.1972, Accession #9123, D.R. Brothers (1♂; DRB); Paratype males have also been depositied in the following collections: AMNH, BJS, BMNH, CDFA, CNC, FVC, JFM, LACM, SEM, SJSU, UCB, UCD, UCR, UIC, USNM, USU and WSU. Recognition Similar to A. idahoensis, but distinguished by the paramere widening prior to narrowed apex and lacking sub-apical spines on gonostyli. Redescription Wing length: 2.42-3.12mm. Colouration: Head dark brown. Mesonotum and pleura dark brown to blackish-brown and somewhat shiny; abdomen dark brown to blackish-brown, dull; halter pale yellow; legs brown, tarsi darker; terminalia brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 with few setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium quadrate, wider than long; posterior margin indented; apical epandrial processes triangular, shorter than long; posterolateral epandrial processes hook-like, longer than wide, with serrate inner margin. Hypandrium faint, thin. Gonocoxite quadrate, slightly longer than wide. Gonostylus slightly shorter than gonocoxite; gently tapering toward apex; apex bearing a terminal claw-like spine. Parameres paired, longer than gonostyli; widening medially with slight arching, then narrowing toward apex. Gonocoxal plate extending past posterior margin of gonocoxites; distal margin concave creating two rounded lobes. Additional Material Examined U.S.A: CALIFORNIA: Contra Costa Co.: Tilden Regional Park, Wildcat Crk @Grizzly Picnic area, 9.viii.2013, 37°52'56.03"N122°13'35.79”W, 457m, R.I. Madriz & K. Lindsay (9 larvae); Tilden Regional Park @ Wildcat Crk. near Grizzly Picnic Area, 7.vii.2014, N37°53'00.8" W122°13'40.7"W, ex. small seep (creek almost dried up), J.K. Moulton & R.J. Pivar (1♀, 3 larvae); Del Norte Co.: Gasquet, 18.ix.1934, A.L. Melander (♂; USNM); Humboldt Co.: Trinidad State Beach, 4.vi.2009, 41°03'25.9"N 124°09'00.0"W, sea cliff face, B.J. Sinclair (4♂); Shasta Co.: McArthur-Burney Falls Memorial State Park, base of falls, 1.viii.1970, P.H. Arnaud, Jr. (1♂, 1♀; CAS); OREGON: Wallowa Co., Imnaha, waterfall, 1.vii.1969, K. Goeden (4♂; CAS, OSDA). Distribution Known primarily from the Coastal Range in California, but also collected in the northern Sierra Nevada’s and the Columbia Plateau. Androprosopa chandlerorum (Arnaud & Boussy) Thaumalea chandlerorum Arnaud & Boussy, 1994: 77. Androprosopa chandlerorum (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ is point mounted with abdomen and right wing removed, labeled: “NE Carpenter Ridge/Sagehen Crk/Nevada Co. 8000’ el./CAL 23 VII 1970”; “I.A. Boussy/Collector”; “Arnaud & Boussy/Diss. No. 4VIII70.1”; “HOLOTYPE Male/Thaumalea chandlerorum/Arnaud and Boussy/1994.” The right wing and abdomen of the holotype are mounted on a microscope slide under two cover glasses with label data equivalent to that cited above (CAS, Entomology Type #17003). Allotype: ♀ mounted under three cover slips on two microscope slides, same data as holotype (CAS). Paratypes: U.S.A.: CALIFORNIA: Nevada Co.: northeast Carpenter Ridge, Sagehen Creek, 8,000 feet, 22.vii.1970, D.S. Chandler (1♀; CAS); same locality but collected on 23-V-1970, I.A. Boussy (1♂, 3♀; CAS). Recognition This species is recognized by the parameres with a short, forked process dorsobasally, the distal third arcuate laterally and the apex produced dorsolaterally. Redescription Wing length: 3.04-mm. Colouration: Head dull, ferruginous to dark brown. Mesonotum and pleura shining ferruginous to dark brown; abdomen dark brown and dull; halter and legs pale yellow to brown; terminalia light brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium spherical, wider than long; posterior margin with broad indentation; apical epandrial processes produced into pointed, finger-like projections that may be simple or bifurcate. Hypandrium very slender, faint; slightly concave. Gonocoxite quadrate, slightly longer than wide; inner margin with strong setae. Gonostylus densely setose; four strong spines in a row on dorsal surface of the apex, the most apical one being the longest. Parameres paired, longer than gonostyli; with short forked process dorsobasally and distal third arcuate laterally and apex produced dorsolaterally. Gonocoxal plate extending past posterior margin of gonocoxites; distal half produced medially into tear-drop shaped keel. Additional Material Examined U.S.A.: CALIFORNIA: Nevada Co.: Carpenter Ridge, Cirque Lake and stream, 12.vii.2012, 39°25'01.4"N120°18'33.2"W, S.E. Brooks (1♀); Carpenter Ridge, Cirque Lake and stream, 13.vii.2012, 39°25'01.4"N120°18'33.2"W, J.M. Cumming (1♂, 1♀). Distribution Known only from Carpenter Ridge on the eastern slope of the central Sierra Nevada of California. Androprosopa coloradensis (Arnaud & Boussy) Thaumalea coloradensis Arnaud & Boussy, 1994: 79. Androprosopa coloradensis (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ is mounted on two microscope slides under three cover glasses, labeled: “HOLOTYPE Male/Thaumalea/coloradensis/Arnaud & Boussy/1994/CAS Ent. Type No./17004”; “U.S.A.: COLORADO:/San Juan County./Red Mountain Pass,/north side/3292m 14-VIII-1973/ Paul H. Arnaud. Jr.”; “Arnaud/Dissection No. 27-VIII-1973.” (CAS Entomology Type #17004). Recognition This species is recognized by the apically broadened parameres, narrow gonostyli and posteroventral corners of epandrium heavily sclerotized and claw-like. Redescription Wing length: 3.82-3.89mm. Colouration: Head blackish-brown. Mesonotum and pleura brown; abdomen dark brown, dull; halter light brown; legs pale brown, tarsi darker; terminalia brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, not dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium spherical, wider than long; posterior margin with broad indentation; apical epandrial processes heavily sclerotized, claw-like. Hypandrium very slender, faint. Gonocoxite rectangular, slightly longer than wide; inner margin concave. Gonostylus strongly narrowed beyond basal fifth; ending in a terminal claw-like spine. Parameres paired, shorter than gonostyli; basal two-thirds widened; notched before apex, which is rounded. Gonocoxal plate extending past posterior margin of gonocoxites; produced medially into long, thin rod. Additional Material Examined U.S.A.: COLORADO: San Juan Co.: San Juan N.F., Hwy. 550, 10.vii.2014 37°42'29.5N107°46'17.8W, ex. Coal Crk. waterfalls and seeps on W side of road, J.K. Moulton and R.J. Pivar (2♂, 6 larvae). Distribution Known only from the San Juan Mountains of southwestern Colorado. Androprosopa confracta (Arnaud & Boussy) Thaumalea confracta Arnaud & Boussy, 1994: 82. Androprosopa confracta (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ pinned with a minuten pin and abdomen removed, labeled: “U. S.A., Ore./Linn Co./Lost Prairie/20-VI-1965/F. Schmid”; “CNC 121”; HOLOTYPE Male/Thaumalea/confracta/Arnaud and Boussy/1994.” The abdomen of the holotype is mounted on a microscope slide under a cover glass (CNC Type #21928). Allotype: ♀ pinned with a minute and has its abdomen removed, labeled: “U.S.A., Wash./Yakima Co./American River/26-VI-1965/F. Schmid”; “Arnaud/Dissection/18-IV-1973”; “ALLOTYPE Female/Thaumalea/confracta/Arnaud and Boussy/1994.” The abdomen of the allotype is mounted ona microscope slide without cover glass (CNC). Paratype: U.S.A.: WASHINGTON: Yakima Co.: American River, 26.vi.1965, F. Schmid (1♂; CAS) Recognition This species can be distinguished from its close relatives A. melanderi and A. brothersi by the gonocoxal plate and the presence of a subapical spine on the right gonostylus. Redescription Wing length: 2.93-3.45mm. Colouration: Head dull, dark brown. Mesonotum, pleura and abdomen dark brown and dull; halter light brown; legs brown, tarsi darker; terminalia brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 weak; R3 and R4+5 nearly parallel toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized and slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 with few setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium a truncated cone, wider than long; posterior margin with wide, very shallow indentation; apical epandrial processes not produced. Hypandrium very slender, faint; very slighty convex. Gonocoxite rectangular, slightly longer than wide. Gonostylus stout; shorter than gonocoxites; base broad, tapers slighty midway; apex of right gonostylus bearing a terminal claw-like seta and one subapical spine, subequal in length to the latter; left gonostylus bearing a single apical claw-like seta. Parameres paired, longer than gonostyli; thickened along median margin; lateral margin with irregular serration. Gonocoxal plate broad, extending past posterior margin of gonocoxites; produced distomedially into triangle, ending near the middle of gonocoxite. Additional Material Examined U.S.A.: OREGON: Marion Co.: Willamette NF, French CK Rd, French CK, 10.vii.2013, 44°46'25"N122°12'40"W, 840m, B.J. Sinclair (1♂); Willamette NF, French CK Rd, 10.vii.2013, 44°46'58"N122°11'12"W, trib, 660m, B.J. Sinclair (4♂); WASHINGTON: Lewis Co.: Gifford Pinchot NF, Skate Crk Rd mile 14, 7.vii.2013, 46°40'00" -121°46'24"W, 660m, roadside streams, B.J. Sinclair (3♂); Santiam SF, Monument Peak Rd, 10.vii.2013, 44°42'34"N122°22'16"W, 820m, B.J. Sinclair (3♂); Pierce Co.: Mt. Rainier N.P., @ Longmire Meadow, 1.vii.2014, 46°45’13.9”N121°48’46.7”W, ex. stream, J.K. Moulton (1♂); Mt. Rainier NP, Narada Falls to Reflection Lk tr., 6.vii.2013, 46°46'15" -121°44'44"W, small stream, 1415m, B.J. Sinclair (1♀, 1 larva); Mt. Rainier NP, S of Nisqually River, 30.viii.2011, 46°46'55.2"N121°45'58.9W, J.K. Moulton (1♂); Mt. Rainier N.P., WA-123, 1.vii.2014, 46°49'33.4"N121°32'36.1"W, ex. small waterfall, J.K. Moulton & R.J. Pivar (1♂); Skagit Co.: Marblemount, Cascade R. Rd.: Mt. Baker-Snoqualmie NF, 18/19.vii.2012, 48°32'10"N121°17'47"W, 410m, roadside seep, B.J. Sinclair (2♀). Distribution This species is found throughout the Cascade Range in Washington and Oregon. Androprosopa elnora (Dyar & Shannon) Thaumalea elnora Dyar & Shannon, 1924: 434. Androprosopa elnora (Dyar & Shannon); Sinclair 1996: 376 (comb.n.). Type Material Lectotype: ♂ mounted on a microscope slide, labeled: “Thaumalea/elnora/Type D. & S./Moscow Mt., Ida/July 25. 1920/R.C. Shannon”; “LECTOTYPE/Thaumalea/elnora ♂/Dyar & Shannon/Designated by/Arnaud and Boussy, 1994” (USNM). Lectoallotype: ♀ pinned, labeled: “Mts. Moscow/Ida. 7.25.20/RC Shannon”; “LECTOALLOTYPE/Thaumalea elnora Dyar/ & Shannon ♀/Designated by Arnaud and Boussy, 1994” (USNM). Recognition Thorax yellow with abdomen yellow-brown. Distinguished from A. ericfisheri by distinct, broad parameres with subapical beak. Redescription Wing length: 3.06-3.38mm. Colouration: Head yellow with golden setae; mouthparts and antennae yellow-brown. Mesonotum and pleura yellow with humerus and sclerites below base of halter darkened; halter and legs concolourous with thorax; abdomen yellow-brown, some brownish-black colouration intermixed; terminalia yellow-brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized and slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium quadrate, wider than long; posterior margin strongly indented; apical epandrial processes triangular, somewhat produced. Hypandrium very slender, faint. Gonocoxite rectangular, wider than long; many setae on inner margin; with short posterolateral processes. Gonostylus with many setae; shorter than gonocoxites; base broad, tapering midway to narrow apex; apex bearing a terminal claw-like seta and one smaller, dorsoapical spine. Parameres paired, blunt, very broad, distal third bearing subapical beak-like process. Gonocoxal plate broad, extending to posterior margin of gonocoxites; produced medially into nearly straight line. Additional Material Examined CANADA: BRITISH COLUMBIA: Perry Ck Falls tr., 11.vii.2012, 49°32'58"N 115°59'50"W, cascade stream, 1095m, B.J. Sinclair (1♂, 4♀); U.S.A.: IDAHO: Latah Co.: Moscow Mountain: 4.vii.1915, A.L. Melander (1♀; USNM); 7.vii.1918, A.L. Melander (18♂, 5♀; USNM); 5.vii.1919, A.L. Melander (1♀; USNM); 10.viii.1924, A.L. Melander, (1♀; USNM); WASHINGTON: Pierce Co.: Mt. Rainier NP, below Comet Falls, 16.vii.2012, 46°47'29"N121°46'55"W, streams/seeps, 1350m, B.J. Sinclair (7 larvae); Spokane Co.: Bald Knob Campground, Mount Spokane SP, 4800-5200 feet, 24.vii.1978, W.J. Turner, sweeping (16♂, 8♀; WSU, CAS); Mt. Spokane SP, Deadman Crk., 12.vii.2012, 47°54'20"N117°06'35"W, 1325m, B.J. Sinclair (16♂); WYOMING: Park Co.: Yellowstone National Park, Dunraven Pass, 28.viii.1934, A.L. Melander (2♂; USNM). Distribution A northwestern species ranging from northwest Wyoming, to southern British Columbia and west to Mt. Rainier in Washington. Androprosopa ericfisheri (Arnaud & Boussy) Thaumalea ericfisheri Arnaud & Boussy, 1994: 87. Androprosopa ericfisheri (Arnaud & Boussy); Sinclair 1996:376 (comb.n.). Type Material Holotype: ♂ is mounted on a microscope slide under two cover glasses, labeled: “HOLOTYPE Male/Thaumalea/ericfisheri/Arnaud and Boussy/1994”; “Alberta: Rocky Mtns./Grizzly Crk. at/Kananaskis Rd./18 mi. S. Hwy 1 5600’/17 Aug. 1969 E.M. Fisher” (CAS Entomology Type #17005). Allotype: ♀ pinned with left wing and abdomen removed, labeled: “CANADA: BRITISH COLUMBIA/Mount Revelstoke National/Park, small cascading creek 6.3 road mi below summit Mount Revelstoke/8 August 1986/Paul H. Arnaud, Jr./Calif. Acad. Sci. Coll.”; “Swept from pools and/cascades small creek,/sunny day, creek shaded/by mountain, 1630-1700/hours.”; “Arnaud and Boussy/Dissection No./042093”; “ALLOTYPE Female/Thaumalea/ericfisheri/Arnaud & Boussy/1994” (CAS). Paratypes: CANADA: ALBERTA: same collection information as holotype (7♂; CAS, CDFA, CNC, USNM); BRITISH COLUMBIA: same collection information as allotype (2♂, 3♀; CAS, CNC). Recognition Thorax light yellow-brown with abdomen blackish brown. Distinguished from A. elnora by having very short, pointed parameres with curved apex. Redescription Wing length: 3.90-3.95mm. Colouration: Head brownish black. Mesonotum yellow with tip of scutellum, sclerites below base of halter darkened; pleural area with some dark spots; halter and legs concolourous with thorax, tarsi slighty darker; abdomen blackish-brown; terminalia brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized and slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium quadrate, wider than long; posterior margin without indentation; apical epandrial processes slighty produced. Hypandrium very slender, faint. Gonocoxite quadrate, slightly longer than wide; many setae on inner margin; with short posterolateral processes. Gonostylus with many setae; shorter than gonocoxites; base broad, tapers dorsally and slightly ventrally to blunt apex; apex bearing three, short spines. Parameres paired, short, ending before middle of gonostylus; pointed and sinuous; curved ventrally. Gonocoxal plate weak, extending nearly to posterior margin of gonocoxites; apical margin trapezoidal. Additional Material Examined CANADA: BRITISH COLUMBIA: Golden, Quartz Creek, Small Tributary, 10.ix.2011 51°27'55.2"N117°21'03”W, J.K. Moulton (2♂, 1♀). Distribution Only known from the southern most portion of the Canadian Rockies. Androprosopa falciformis (Arnaud & Boussy) Thaumalea falciformis Arnaud & Boussy, 1994: 91 Androprosopa falciformis (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ mounted on a microscope slide under two cover glasses, labeled: “HOLOTYPE Male/Thaumalea/falciformis/Arnaud and Boussy,/1994/USA: IDAHO, Idaho Co., 4.6 mi E Lowell/Larva: 5.IV.1970/Adult: 20.IV.1970/S.T. Elliott, J.M. Gillespie/F.W. Spray/CODE AD-1.” (CAS Entomology Type #17006). Allotype: ♀ mounted on two microscope slides under three cover glasses, similar collection data to holotype, but adult dated 18-IV-1970 with collection number KK and specimen # 4 (CAS). Paratypes: CANADA: BRITISH COLUMBIA: Terrace, Mount Thornhill, 26.vii.1960, C.H. Mann (1♂; CNC). U.S.A.: IDAHO: Clearwater Co.: 5.5 miles west of Orofino [similar stream to population 2], larva 5.iv.1970, adult 16.iv.1970, S.T. Elliott, J.M. Gillespie, F.W. Spray; Code G, specimen #3 (1♀; CAS); same locality but larva 5.iv.1970, adult 19.iv.1970, code AC, specimen #4 (1♂; CAS); 17.6 miles west of Orofino [small permanent cascading stream, 80 angle], elevation ca 1,100 feet, larva 5.iv.1970, adult 14.iv.1970, S.T Elliott, J.M. Gillespie, F.W. Spray, Code Q, specimen #1 (1♂; CAS); same locality but larva 5.iv.1970, adult 17.iv.1970, code F specimen #3 (1♂; CAS); same locality but larva 5.iv.1970, adult 19.iv.1970, code AC specimen #4 (1♂; CAS); same locality but larva 5.iv.1970, adult 19.iv.1970, code ZZ, specimen #5 (1♂; CAS). Idaho Co.: 2.3 miles west of Lowell, larva 5.iv.1970, adult 14.iv.1970, S.T. Elliott, J.M. Gillespie, F.W. Spray, code U, specimen #13 (1♀; CAS); 4.6 miles E of Lowell [small cascading stream otherwise similar to stations 2 and 3], larva 5.iv.1970, adult 14.iv.1970, S.T. Elliott, J.M. Gillespie, F.W. Spray, code GG specimen #1 (1♂; CAS); same locality but larva 5.iv.1970, adult 18.iv.1970, code JJ specimen #7 and code SS specimen #8 (2♀; CAS); same locality but larva 5.iv.1970, adult 20.iv.1970, code AD-2 specimen #-) (1♂;CAS); same locality but larva 5.iv.1970, adult 17.iv.1970, code C specimen #2 (1♂; CAS); Latah Co.: Kendrick, [no date or name of collector; with identification label by A. Stone as Thaumalea fusca (Garrett)] (1♂; CU); 3.9mi east of Laird Park, larva 19.iv.1970 [no date for adult], S.T. Elliott, J.M. Gillespie, F.W. Spray, code AS (1♂;CAS). Recognition Distinguished by the strongly recurved parameres (similar to A. anolo), gonostyli elbowed bearing a distinct blunt, lateroventral process. Redescription Wing length: 2.60-2.86mm. Colouration: Head brownish-black. Mesonotum and pleura light brown, scutellum brown; abdomen dark brown, dull; halter pale yellow-brown; legs brown, tarsi darker; terminalia brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin, then slightly divergent just before margin. Abdomen: Male abdominal sternites 2-6 rectangular with basal half well sclerotized, not dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 7 & 8 with few setulae, anterior margin strongly dipped into preceding segment. Male terminalia: epandrium a truncated sphere, slightly longer than wide; posterior margin not indented; each posteroventral corner bearing single, small tooth-like apical epandrial processes. Hypandrium convex medially, arching into sternite 8. Gonocoxite longer than wide, with posterolateral process bluntly produced. Gonostylus slightly shorter than gonocoxite; bent laterally and apically elbowed dorsally with stout claw-like setae at apex, bearing a blunt lateroventral process at elbow. Parameres paired; broad basally, tapering medially then sharply produced laterally and curved anteriorly toward apex; protruding slightly beyond posterior epandrial margin. Gonocoxal plate exceeding length of gonocoxites. Additional Material Examined U.S.A: IDAHO: Boundary Co.: 2673 Moyie River Rd. next to Feist Creek Falls Resort, 23.vi.2014, 48°54'43.0"N116°10'23.2", ex. small rock trickle at end of drive, J.K. Moulton & R.J. Pivar (3♂); same locality info as above except, ex. headwaters of small rock trickle at end of drive, J.K. Moulton & R.J. Pivar (8♂); same locality info as above except, ex. small waterfall passed base of driveway, J.K. Moulton (1♂); same locality info as above except, 24.vi.2014, ex. drain pipe under road, J.K. Moulton & R.J. Pivar (7 larvae); Kootenai Co.: St. Joe NF, Hwy 97, FR 438, 19.vi.2014, 47°36.392’N116°40.099’W, 660m, ex. tribs of Beauty Ck, B.J. Sinclair (2♂); Latah Co.: Moscow Mt., 7.vii. 1918, A.L. Melander (1♂; USNM); Moscow Mt., Hatter Ck. Rd., 13.vii.2012, 46°49'16"N116°49'28"W, 1075m, B.J. Sinclair (1♀); Moscow Mt., Moscow Mtn Rd., 22.vi.2014, 46°47.721’N 116°54.093’W, 1000m, ex. cascading str., B.J. Sinclair (3♂); Shoshone Co.: St. Joe NF, Rd. 50, E. Calder, 20.vii.2014, 47°15.820’N116°08.803’W, ex. roadside streams/seeps, 680m, B.J. Sinclair (4♂); WASHINGTON: Mt. Spokane SP, Deadman Crk, 12.vii.2012, 47°54'20"N 117°06'35"W, 1325m, B.J. Sinclair (3♂). Distribution Known from northern Idaho and from one male in mid-western British Columbia. Androprosopa fusca (Garrett) Orphnephila fusca Garrett, 1925: 10. Thaumalea fusca (Garrett); Arnaud & Boussy 1994: 94 (comb.n.). Androprosopa fusca (Garrett); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ is glued to a paper point with most of abdomen and left wing removed, labeled: “MARYSVILLE/B.C. 12-VIII/C GARRETT”; “HOLO/TYPE/ORPHNEPHILA/FUSCA ♂/CGarrett./C.B, D. GARRETT”; “Arnaud & Boussy/Dissection No./69III24-1.” The left wing and abdomen are mounted under separate cover glasses on a microscope slide with similar label data (CNC Entomology Type #7912). Recognition Readily distinguished with combination of gonocoxite with finger-like posterolateral process as long as gonostylus and paramere very broad basally and narrowed sharply with distal portion recurved laterally. Redescription Wing length: 2.99mm. Colouration: Head brown. Mesonotum and pleura brown; apex of scutellum blackened; abdomen brown; legs with coxae and trochanter brown, femora and tibia light yellow-brown, tarsi darker; halter pale yellow knob with brown base; terminalia brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin, then R4+5 slightly diverging slightly at margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, 2-5 not dipped into preceding segment, 6-7 slightly dipped into preceding segment; sternites 2-7 bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium spherical, slightly longer than wide; posterior margin without indentation; apical epandrial processes produced into short, double dentate processes; posterolateral epandrial processes absent. Hypandrium thin, convex and dipping into sternite 8. Gonocoxite broadly conical, longer than wide; bearing glabrous, finger-like, posterolateral process that is equal in length to gonostylus. Gonostylus shorter than gonocoxite; basal half wider, then tapering medially, apex bearing a single apical claw-like spine; few setae on gonostylus. Parameres paired, longer than gonostyli; very broad basally, narrowed sharply at apical third with distal portion strongly recurved laterally. Gonocoxal plate extending to length of gonocoxal plate, including posterolateral process; produced into medial keel. Additional Material Examined Only the holotype male is known. Distribution Known from the single locality of Marysville in the Canadian Rockies of southeastern British Columbia. Marysville is no longer recognized on maps, but previously shown as located between Cranbrook and Kimberley. Androprosopa gillespieae (Arnaud & Boussy) Thaumalea gillespieae Arnaud & Boussy, 1994: 97. Androprosopa gillespieae (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Type Material Holotype: ♂ male is mounted on a microscope slide under two glasses, labeled: “HOLOTYPE/Thaumalea gillespieae/Arnaud and Boussy, 1994/ Ore.: Linn Co./Monument Peak Guard/Sta. s.21, T105, R4E/4000’ E.M. Fisher/♂ July 28, 1969” (CAS Entomology Type #17007). Allotype: ♀ mounted on a microscope slide under two cover glasses with same label data as holotype (CAS). Paratypes: CANADA: BRITISH COLUMBIA: ca. 11km east Lytton, waterfall, 12.viii.1974, P.H. Arnaud, Jr. (4♂; CAS); Cultus Lake, 24.v.1965, F. Schmid (1♂; CNC); Kinnaird, 21.v.1965, F. Schmid (1♂; CNC); Kitchener, 5.v.1965, F. Schmid (1♂; CNC); U.S.A.: IDAHO: Bonner Co.: Priest Lake, 1.viii.1916, A.L. Melander (2♂, 2♀; USNM); Priest Lake, Cavanaugh Bay, 18.viii.1919, A.L. Melander (2♂; USNM); Idaho Co.: Apgar Creek, U.S. Highway 12, 15.vii.1971, collection #24 specimen #1, J.M. Gillespie (1♂; CAS); Holly Creek, U.S. Highway 12, 33 miles NE Lowell, 2,600 feet, 19.viii.1969, E.M. Fisher (14♂, 2♀; CAS); Tumble Creek, Highway 12, 16 miles northeast of Lowell, 2,000 feet, 19.viii.1969, E.M. Fisher (1♂; CAS); Tumble Creek, U.S. Highway 12, 15.viii.1971, J.M. Gillespie (2♂, 3♀; CAS); 2.3 miles west of Lowell, larvae collected 5.iv.1970, adults emerged from 12 to 18.iv.1970, J.M. Gillespie, F.W. Spray, S.T. Elliott (8♂, 1♀; CAS, UIC); OREGON: Clackamas Co.: Eagle Creek, Forest “Res.”, 1.viii.1917, A.L. Melander (1♂; USNM); Lane Co.: w. side Cascade Mtns., Mack Creek, 20.vii.1982, 800-900m (1♂, 2♀; USNM); w. side Cascade Mtns., Mack Creek, 11.viii.1982, 800-900m (1♂, 1♀; USNM); Linn Co.: same locality information as holotype except 26.vii.1969 (3♀; CAS); same locality information as holotype (4♂, 5♀; CAS); same locality as holotype except 2.viii.1969 (1♂, 1♀; CAS); WASHINGTON: Pierce Co.: Mount Rainier, Hansen Camp, 31.vii.1922, A.L. Melander (1♂; USNM); Skagit Co.: Baker Lake, 27.v.1965, F. Schmid (1♂; CNC); Marblemount, 31.v.1965, F. Schmid (2♂; CNC). Recognition This species is recognized by the large, lateral triangular shaped flange on the apical third of the paramere, with apex narrowed and blunt. Redescription Wing length: 2.60-3.4mm. Colouration: Head dark brown to blackish-brown. Mesonotum and pleura dark brown to blackish-brown and somewhat shiny, however some specimens intensely black and others reddish-brown; abdomen blackish-brown, dull; halter pale yellow-brown; legs light brown, tarsi darker; terminalia lighter brown than abdomen. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 lacking setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium quadrate, wider than long; posterior margin strongly indented; apical epandrial processes triangular, rounded; posterolateral epandrial processes produced as flattened flanges, longer than wide, with serrate inner margin. Hypandrium thin, slighty concave. Gonocoxite conical, slightly longer than wide. Gonostylus slightly shorter than gonocoxite; apical third tapers to apex; apex bearing a strong terminal claw-like spine. Parameres paired, longer than gonostyli; very strong lateral flange on apical third, apex narrow and blunt. Gonocoxal plate extending past posterior margin of gonocoxites; distal margin three lobed. Additional Material Examined CANADA: BRITISH COLUMBIA: Cathedral Prov Pk, 19.vii.2015, 49°08.52"N120°01.66"W, 725m, ex. stream, B.J. Sinclair (1♂); Hwy 23, 13km N Revelstoke, 21.vii.2012, 51°04'03"N118°10'31"W, 600m, B.J. Sinclair (1 larva); Manning Prov. Pk., Three Falls Tr., below Derek Falls, 20.vii.2015, 49°03.78"N120°56.34"W, 1220m, B.J. Sinclair(1♂,1♀); nr. Kimberley, Meachen Ck. FR, 17.vii.2015, 49°36.22"N116°13.33"W, roadside falls, 1021m, B.J. Sinclair (7♂); nr Kimberley, Meachen Ck. FR, roadside falls, 17.vii.2015, 49°36.21"N116°13.45"W, 1027m, B.J. Sinclair (1♂); Perry Ck falls, 11.vii.2012, 49°32'50"N115°59'55"W, spray zone, 1105m, B.J. Sinclair (2♂); Perry Ck Falls Tr., 49°32'58"N115°59'50"W, 1095m, cascad. str., 11.vii.2012, B.J. Sinclair (4♂, 2♀); W. Creston-Hwy 3, Topaz Ck., 18.vii.2015, 49°08.76"N116°45.65"W, 825m, B.J. Sinclair (1♂); U.S.A.: CALIFORNIA: Monterey Co.: Los Padres National Forest, Mill Crk @ Mill Crk trail, 6.viii.2013, 35°59'30.68"N121°28'28.06"W, 165m, R.I. Madriz & K. Lindsay (1♂); San Bernardino Co.: Crestline, 4.vii.1942, A.L. Melander (1♂; USNM); IDAHO: Boundary Co.: NFR-2517, 23.vi.2014, 48°58'46.3"N116°09'26.5"W, ex. Brass Creek, R.J. Pivar (1♂, 2♀); Idaho Co.: Boise NF, 10 Mile Cpgd, Hwy 21, 11.vi.2014, 43°53.840’N115°42.669’W, 1470m, ex. 10 Mile Ck, B.J. Sinclair (6♂, 5♀); Boise NF, East of Lowman, Chapman Ck, 12.vi.2014, 44°06.162’N115°18.753’W, 1450m, B.J. Sinclair (9♂, 2♀); Hwy. 95, 23.vi.2014, 45°20.547’N116°20.999’W, cascade at Rest Stop, 650m, B.J. Sinclair (5♀); Nez Perce-Clearwater N.F., Rt. 12, Apgar Crk., 19.vi.2014, 46°12'55"N 115°32'12"W, J.K. Moulton (1♀); Selway-Bitterroot Wildnerness, North Fork Moose Creek, 23-30.vi.1989, F.W. Merickel (1♂; UIC); Latah Co.: Moscow Mt. Moscow Mtn Rd., 22.vi.2014, 46°47.721N116°54.093’W, 1000m, ex. cascading str., B.J. Sinclair (4♂); St. Joe NF, FR447, E of Laird Pk, 12.vi.2011, 46°57'37.7"N116°35'51.0"W, B.J. Sinclair (1♀); OREGON: Baker Co.: Wallowa-Whitman N.F. Bennet Crk., 29.vi.2014, 44°59'54.8"N117°24'00.8"W, R.J. Pivar (1 larva); Linn Co.: Cascadia SP, nr. Sweet Home, 9.vii.2014, 44°23’56.1”N122°28’52.3”W, 261m, ex: along Soda Crk, S.E. Brooks (21♂); Willamette NF, 1175m, fork of No. 410 & 420, small stream, Hackleman Crk area, 11.vii.2013, 44°23'24"N122°02'48"W, B.J. Sinclair (3♂); Willamette NF, Rd 2025, along Moose Crk, 11.vii.2013, 44°25'11"N122°24'59"W, 335m, streams, B.J. Sinclair (1♂, 3♀); Marion Co.: Willamette NF, French CK Rd, French CK, 10.vii.2013, 44°46'25"N122°12'40"W, 840m, B.J. Sinclair (5♂); Multnomah Co.: Wahkeena Falls & Ck, 8.vii.2013, 45°34'31"N122°07'40"W, 45m, B.J. Sinclair (24♂, 6♀, 5 larvae); WASHINGTON: Lewis Co.: Gifford Pinchot N.F., Boulder Creek @ Skate Crk. Rd., 29.viii.2011, 46°39’10.9"N121°43’17.1"W, J.K. Moulton (1♀); Gifford Pinchot N.F., Boulder Crk @ Skate Crk Rd., 28.vi.2014, 46°39’10.9”N 121°43’17.1”W, ex. boulder splash zones, J.K. Moulton & R.J. Pivar (1♀); Gifford Pinchot N.F., N.F.R.-23, 3.vii.2014, 46°16'52.1"N121°36'50.2"W, ex. creek, J.K. Moulton & R.J. Pivar (2♂); Gifford Pinchot NF, Skate Crk Rd mile 14, 7.vii.2013, 46°40'00"N121°46'24"W, 660m,roadside streams, B.J. Sinclair (1♂); Gifford Pinchot NF, Skate Crk Trib, Skate Crk Rd, 15.vi.2011, 46°40'59.1"N121°46'25.3"W, J.K. Moulton (1♀); Pierce Co.: Mt. Rainier NP, Comet Falls trail, 16.vii.2012, 46°46'43"N121°47'01"W, streams, 1100m, B.J. Sinclair (1♂, 1♀); Mt. Rainier N.P., Comet Falls trail, 2.vii.2014, 46°46'55.7"N121°46'47.1"W, ex. Van Trump Crk trib., J.K. Moulton & R.J. Pivar (4♂, 2♀); Mt. Rainier N.P. Falls Crk. falls, 1.vii.2014, 46°45'36.2"N121°33'35.6"W, ex. boulder splash zone beneath bridge, J.K. Moulton & R.J. Pivar (2♂); Mt. Rainier N.P. Falls Crk. falls, 2.vii.2014, 46°45'36.2"W 121°33'35.6"W, ex. boulder splash zone beneath bridge, J.K. Moulton & R.J. Pivar (1♂); Mt. Rainier NP, Narada Falls to Reflection Lake tr., 2.vii.2013, 46°46'15"N 121°44'44"W, small stream, 1415m, B.J. Sinclair (1♂); Mt. Rainier NP, S of Nisqually River, 30.viii.2011, 46°46'55.2"N121°45'58.9"W, J.K. Moulton (1♂); Mt. Rainier NP, Stevens Creek tr., 4.vii.2013, 46°45'20"N121°38'31"W, 860m, B.J. Sinclair (1♀); Mt. Rainier NP, Sunrise Bridge, 11.vi.2011, 46°54'27"N121°32'32.8"W, J.K. Moulton (1♀); Mt. Rainier N.P., WA-123, 1.vii.2014, 46°49'33.4"N121°32'36.1"W, ex. small waterfall, J.K. Moulton & R.J. Pivar (♂, ♀); WA: Camas Creek 47°28'37.4" -120°37'04.1"UTAH: UT: Utah Co.: Mt. Timpanogos, Stewart Ck., 8.vi.2001, 40°23'11.1"N111°36'14.7"W, 8.vi.2001, B.J. Sinclair (18♂, 10♀); Sundance, 8.vi.2001, 40°23'41.9" -111°34'56.5", B.J. Sinclair (6♂). Distribution The most widespread of the Nearctic Androprosopa, it’s range encompassing British Columbia to the north, south to California and east to Colorado. The most commonly encountered Western thaumaleid. Androprosopa idahoensis (Arnaud & Boussy) Thaumalea idahoensis Arnaud & Boussy, 1994: 101. Androprosopa idahoensis (Arnaud & Boussy); Sinclair, 1996: 376 (comb.n.). Type Material Holotype: ♂ is mounted on a microscope slide under two cover glasses, labeled: “HOLOTYPE Male/Thaumalea/idahoensis/Arnaud and Boussy, 1994/U.S.A.: IDAHO, Nez Perce Co./Juliaetta Falls/larva: --/adult: 5.IV.1970/S.T. Elliott, J.M. Gillespie, F.W. Spray/ Code Y” (CAS Entomology Type #17008). Allotype: ♀ mounted on a microscope slide under two cover glasses, collection data the same as the holotype, except larva collected 5 April 1970, adult 18 April 1970, code MM specimen #12 (CAS). Paratypes: IDAHO: Latah Co.; Juliaetta [no date/collector given], A.L. Melander Collection, 1961 (3♂, 1♀; USNM); Nez Perce Co.: Juliaetta Falls, ca 900 feet, larva 15.iii.1970, adult 29.iii.1970, J.M. Gillespie (1♂; CAS); same locality, larva 15.iii.1970, adult 5.iv.1970, Code BB (1♀; CAS); same locality, larva 5.iv.1970, adult 16.iv.1970, code E (1♂; CAS); same locality, larva 5.iv.1970, adult 17.iv.1970, code L specimen #3 (1♀; CAS); same locality, larva 5.iv.1970, adult 18.iv.1970, code MM specimen #12, code NN specimen #4, code PP specimen #6, code RR specimen #10, code TT specimen #11 (2♂, 3♀; CAS, UIC); same locality, larva 5.iv.1970, adult 19-IV-1970, code WW, specimen #14 (1♀; CAS); same locality, larva 5.iv.1970, adult 21-IV-1970, code AI specimen #24 (1♂; CAS); same locality, larva 5.iv.1970, [no adult date], code II specimen #8 (1♂; CAS). Recognition Recognized by the shape of the apex of the paramere, which is twisted nearly 180° towards the laterad, as well as the strongly lobed base. Redescription Wing length: 2.68-3.38mm. Colouration: Head dark brown to blackish-brown, shiny near apical margin of eye. Mesonotum and pleura brown to blackish-brown and shiny; abdomen blackish-brown, dull; halter and legs light brown, tarsi darker; terminalia dark brown. Wing: lightly infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 strongly convergent toward wing margin. Abdomen: Male abdominal sternites 2-7 rectangular with anterior margin well sclerotized, slightly dipped into preceding segment, bearing microtrichia and numerous setulae; sternite 8 with few setulae, anterior margin strongly dipped into segment 7. Male terminalia: epandrium quadrate, wider than long; posterior margin strongly indented; apical epandrial processes triangular, shorter than long; posterolateral epandrial processes hook-like, partially arched, longer than wide, with serrate inner margin. Hypandrium thin, convex toward sternite 8. Gonocoxite broadly conical, slightly wider than long. Gonostylus nearly subequal in length to gonocoxite; gradual taper to apex; apex bearing one large spine at apex and two smaller ones subapically on dorsal surface. Parameres paired, longer than gonostyli; with strongly developed basal lobe; apex expanded slightly and twisted toward laterad. Gonocoxal plate extending past posterior margin of gonocoxites; distal margin with two lateral lobes and a very lightly convex medially. Additional Material Examined U.S.A.: CALIFORNIA: Sierra Co.: Tahoe National Forest, waterfall 4 miles E of Downieville, 14.viii.2013, 39°33'31.97"N120°45'40.48"W, 1001m, R.I. Madriz (15 larvae); IDAHO: Boise Co.: Hwy 21, seep near hot spring, 11.vi.2011, 44°05'26.5"N116°03'09.8"W, B.J. Sinclair (4 larvae); Latah Co.: Juliaetta Falls, 14.vii.2012, 46°31'23"N116°44'47"W, 286m, ex.roadcut seeps, B.J. Sinclair (14 larvae); Juliaetta Falls Hwy 3, 15.vi.2014, 46°31.394’N116°44.798’W, 286m, ex. roadcut stream/seeps, B.J. Sinclair (3♀, 5 larvae) see excel; Nez Perce Co.; Juliaetta Falls, 12.vi.2011, 46°31'22.2"N,116°44'49.7"W, B.J. Sinclair (3♂, 2♀); Juliaetta Falls, 19.v.1993, J.B. Johnson (2♂). WASHINGTON: Lewis Co.: Gifford Pinchot NF, Boulder Crk @ Skate Crk Rd., 15.vi.2011, 46°39'10.9"N121°43'17.1”W, J.K. Moulton (2♂). Distribution Known from the Pacific Northwest in Washington (Mt. Rainier), Idaho and the northern Sierra Nevada mountain range in California. Androprosopa lindsayorum (Arnaud & Boussy) Thaumalea lindsayorum Arnaud & Boussy, 1994: 105. Androprosopa lindsayorum (Arnaud & Boussy); Sinclair 1996: 376 (comb.n.). Would I mention here the mis ident. As sonorensis? Type Material Holotype: ♂ pinned, lacking left midleg, left wing and abdomen, labeled: “Hermiston, OR/9 July 1922/A.L. Melander”, “ALMelander Collection/1961”; “Arnaud & Boussy/Dissection No./69.IX.26-3”; “HOLOTYPE Male/Thaumlea/lindsayorum/Arnaud & Boussy/1994.” The left wing and abdomen are mounted under two cover glasses on a microscope slide and labeled with similar data as holotype (USNM). Allotype: ♀ is cleared and mounted on two microscope slides, labeled: “U.S.A., Wash./Skagit Co./Baker Lake/27-V-1965/F. Schmid”; “ALLOTYPE Female/Thaumalea/lindsayorum/Arnaud & Boussy/ 1994”; “Arnaud & Boussy/Dissection No./5-V-1973” (CNC). Paratypes: U.S.A.: CALIFORNIA: Del Norte Co.: Gasquet, 4.vi.1965, F. Schmid (2♂, 1♀; CAS, CNC); OREGON: Benton Co.: Corvallis, 21.viii.1925, A.L. Melander (3♂; CAS, UCB, USNM); WASHINGTON: Skagit Co.: Baker Lake, 27.v.1965, F. Schmid (3♂, 1♀; CAS, CNC); Marblemount, 30.vi.1965, F. Schmid (1♂; CNC); Whatcom Co.: Glacier, 25.v.1965, F. Schmid (2♂, 2♀; CNC). Recognition Recognized by the three to four long, dorsomedial claw-like spines on the gonostyli. Differs from son? Redescription Wing length: 3.27-3.61mm. Colouration: Head blackish-brown. Mesonotum and pleura ranging from brown to orange-brown, shiny; scutellum light brown; abdomen blackish-brown, dull; halter and legs light brown, tarsi darker; terminalia yellow-brown. Wing: infuscate; R+R1+R1+2 with macrotrichia along entire length, remaining veins bare; R2 situated at basal third of R1+2; bend in R3 gentle; R3 and R4+5 convergent toward wing margin, but R4+5 diverging weakly at margin. Abdomen: Male abdominal sternites 2-7 rectangular with basal half well sclerotized, not dipped into