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Trichoptera of Gunnison County, Colorado

Introduction to the caddisfly family Hydropsychidae

Common Net Spinner Caddis, Cinammon Caddis, Light wing medium brown body caddis #16-20, Light wing tan body caddis #18-20, Medium wing tan body caddis #16-22, Spotted Sedge

Curtis, 1835

Updated Twosday 2/22/2022 :-)

This Hydropsychidae retreat was under a rock in a tributary of the East River on 18 July 2007. Notice the net they use for gathering food. The top of the Hydropsychidae's head is visible right of the net. Notice the abdomen of a Cinygmula larvae under the upper right of the retreat.

Provisional Species List

Description

Larvae are large in their last instars. They may be the biggest animals in your pan of bugs from a kick sample. Hydropsychid larvae live under rocks in most streams and rivers in Gunnison County, building a retreat (or house) and spinning a net to capture detritus and small invertebrates.
Adults are diverse in size, shape and taxonomic characteristics. Both sexes have a 5-segmented maxillary palp. All genera lack scutal warts, ocelli and preapical spurs on their front tibia (Ross, 1944).

Life History

Adults females lay eggs underwater in strings attached to submerged rocks or other objects. The eggs are in a cement-like matrix (Ross, 1944)

Good Links


Other Websites:
Photos, Map, Taxon Identifier Numbers - from the Global Biodiversity Information Facility Hydropsychidae at GBIF

Photos, Map, Museums, DNA - Barcode of Life Data System

Photos and description from The Chironomid Research Group at The University of Minnesota

References

Allan,JD 1987 Macroinvertebrate drift in a Rocky Mountain stream. Hydrobiologia 144, 261-268.
     Working in Cement Creek, Allan looked at aquatic insects drifting in the water column for 24 hour time periods during the summers of 1976 and 1977. On page 263, he briefly says "Trichoptera (mostly several species of Rhyacophila, but occasional Hydropsychidae and Brachycentrus americanus Banks) always exhibited very low drift densities."

Alstad,DN 1980 Comparative biology of the common Utah Hydropsychidae (Trichoptera). American Midland Naturalist 103, 167-174.

Balistrieri,LS; Mebane,CA and Schmidt,TS 2020 Time-dependent accumulation of Cd, Co, Cu, Ni, and Zn in mayfly and caddisfly larvae in experimental streams: Metal sensitivity, uptake pathways, and mixture toxicity. Science of the Total Environment, 732. html
     Abstract: "Conceptual and quantitative models were developed to assess time-dependent processes in four sequential experimental stream studies that determined abundances of natural communities of mayfly and caddisfly larvae dosed with single metals (Cd, Co, Cu, Ni, Zn) or multiple metals (Cd + Zn, Co + Cu, Cu + Ni, Cu + Zn, Ni + Zn, Cd + Cu + Zn, Co + Cu + Ni, Cu + Ni + Zn). Metal mixtures contained environmentally relevant metal ratios found in mine drainage. Free metal ion concentrations, accumulation of metals by periphyton, and metal uptake by four families of aquatic insect larvae were either measured (Brachycentridae) or predicted (Ephemerellidae, Heptageniidae, Hydropsychidae) using equilibrium and biodynamic models. Toxicity functions, which included metal accumulations by larvae and metal potencies, were linked to abundances of the insect families. Model results indicated that mayflies accumulated more metal than caddisflies and the relative importance of metal uptake by larvae via dissolved or dietary pathways highly depended on metal uptake rate constants for each insect family and concentrations of metals in food and water. For solution compositions in the experimental streams, accumulations of Cd, Cu, and Zn in larvae occurred primarily through dietary uptake, whereas uptake of dissolved metal was more important for Co and Ni accumulations. Cd, Cu, and Ni were major contributors to toxicity in metal mixtures and for metal ratios examined. Our conceptual approach and quantitative results should aid in designing laboratory experiments and field studies that evaluate metal uptake pathways and metal mixture toxicity to aquatic biota."

Banks,N 1905 Descriptions of new neuropteroid insects. Transactions of American Entomological Society 32, 1-20.
     Has descriptions of several of the local Hydropsychidae species.

Canton,SP and Ward,JV 1981 The aquatic insects, with emphasis on Trichoptera, of a Colorado stream affected by coal strip-mine drainage. Southwestern Naturalist 25(4) 453-460.
     They studied Trout Creek where it runs through the Edna Coal Mine in northwestern Colorado. The mine spoils were 30 meters from the edge of the creek (approximately a 100 foot buffer zone). They found the aquatic insect density (numbers per square meter) and biomass (weight in grams per square meter) did not change above and below the mine. The Shannon-Weaver Diversity index also showed no difference between sites. However the community structure (which species were present and proportions) did change. Since there were irrigation water and cattle influences at their downstream site, their results may reflect these additional water uses. They note the biggest visible change at this mine is the loss of willow and alder trees downstream of the mine. The caddisfly population changed the most between sites, shifting from a mix of families above the mine to dominance by Hydropsychidae and Glossosomatidae below the mine.
Quote from page 457: "The Hydropsychidae (Arctopsyche inermis, Hydropsyche cockerelli, and H. oslari) were unimportant at C2 (reference site), comprising only 3% of trichopteran numbers, while at C4 (mine affected), with increased abundance of Hydropsyche spp., they accounted for 16% of the density."


Cardinale,BJ; Gelmann,ER and Palmer,MA 2004 Net spinning Caddisflies as stream ecosystem engineers: the influence of Hydropsyche on benthic substrate stability. Funct. Ecol. 18: 381-387. PDF
     Abstract: "1. Organisms that physically modify or create habitat (ecosystem engineers) can have a profound influence on community and ecosystem dynamics. 2. Here evidence is presented that one of the most abundant and widely distributed lotic insects could act as an ecosystem engineer in streams by increasing the stability of benthic substrates during flooding. 3. Natural densities of larval net spinning caddisflies (Hydropsychidae) were established in stream channels that had standardized physical properties. The mobility of three particle sizes were measured during simulated flooding and the fraction of particles eroded compared with that of control streams. 4. Larvae increased the initial velocity required to erode sediments by 10-30%. At velocities sufficient to scour 87% of particles from control channels, 57-100% remained stable in channels colonized by larvae. 5. Assuming larvae have similar effects in natural streams, caddisflies could be expected to increase the recurrence interval of a substrate scouring flood from 1.67 year to 2.41 year, corresponding to a 17% decrease in the probability of bed scour per year. 6. Our study suggests these insects could play an important role in generating the spatial 'refuges' that moderate the resistance of lotic communities to flooding. It is argued that, as has occurred in marine systems, a better understanding of how freshwater organisms engineer their physical environment has much potential to complement our historical focus on the abiotic forces that constrain populations and communities."

Clements,WH; Carlisle,DN; Lazorchak,JM and Johnson,PC 2000 Heavy metals structure benthic communities in Colorado mountain streams. Ecological Applications 10(2)626-638. Abstract

Clements,WH; Cherry,DS and Cairns,J 1988 Impact of heavy metals on insect communities in streams: a comparision of observational and experimental results. Canadian Journal of Fisheries and Aquatic Sciences 45 11, 2017-2025.
     Working in the Clinch River of Russell County, Virginia and outdoor experimental streams, they measured population responses of macroinvertebrates to natural conditions and 12 µg of copper and zinc in the artifical streams. They used 6 replicates of substrate-filled trays everywhere and counted all the animals (no subsampling). Both stream mesocosm experiments and Clinch river sites showed similar results. They found abundance or total numbers of aquatic insects declined at all high effluent sites associated with the Clinch River coal-fired power plant, recovering 3- 4 kilometers downstream. Low levels of copper and zinc reduced species richness (number of different taxa) and total numbers as well as caused a shift in the species composition of dominant taxa. Metal contamination caused macroinvertebrate populations to shift from control (clean) sites dominated by Mayflies and Tanytarsini Midges to polluted sites dominated by Hydropsychidae caddisflies and Orthocladiinae midges.

Coutant,CC 1982 Evidence for upstream dispersal of adult caddisflies (Trichoptera: Hydropsychidae) in the Colombia River. Aquatic Insects 4:61-66
     Abstract: "The radioisotope 65Zn, introduced to the Columbia River in discharges of the U.S. Atomic Energy Commission's Hanford reactors, was used to test the hypothesis that adult caddis flies migrate upstream after emerging from the aquatic environment. The larval stages living downstream of the effluents are known to accumulate appreciable levels of 65Zn. The radioisotope was found in levels above background in shoreline swarms of adult caddis flies as far as 16 km above the uppermost reactor effluent. Whether movement was only upstream and the precise distances flown remain unclear. The upstream movement is important for understanding the biology of riverine aquatic insects and for evaluating the upstream dispersal of radioactive contaminants."

Crichton,MI 1957 The structure and function of the mouth parts of adult caddis flies (Trichoptera). Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 241(677), pp.45-91. PDF
     Abstract: "The paper gives a detailed account of the structure and function of the mouth parts of Phryganea striata L., followed by a comparative study of these structures throughout the order Trichoptera. Observations on the feeding of caddis flies are reviewed. Consideration is given to homologies and phylogeny. In Phryganea the head is produced ventrally into a proboscis to which all parts of the mouth complex contribute. A detailed account is given of external and internal structure, musculature, and nervous system of the head and mouth parts. The central area of the anterior surface of the head capsule is interpreted as a frontoclypeus because of the origin of muscles to the foregut. The elongate labrum covers a sclerotized groove or sitophore. Mandibles are reduced to small lobes. The cardines and stipites of the maxillae contribute to the base of the proboscis. The single maxillary lobe is interpreted as a lacinia on grounds of musculature. The distinctive protrusible haustellum is regarded as derived from the hypopharynx. It is traversed by a common salivary duct, provided with a muscular valve. The anterior surface of the haustellum is covered with a system of channels which converge to the sitophore. These channels are formed by cuticular outgrowths arranged in lines and drawn out into filaments which roof the channels thus formed. These outgrowths, which are named pectinate hairs, differ in form according to their position on the haustellum. The labium forms part of the base of the proboscis. There is no ligula. Extension of the proboscis is brought about both by muscle action on sclerites and increased blood pressure affecting the flexible areas of cuticle. Relaxation results from reduction in blood pressure, and contraction of retractor muscles. The haustellum functions as an organ for taking up liquids. A direct drinking and a lapping attitude are described. The comparative study includes observations on fifty-three species, which are representative of each of the thirteen families found in Britain. All species examined have a protrusible haustellum, and are capable of drinking. The most highly developed condition is seen in the Phryganeidae and Limnephilidae. A channelled haustellum is also found in the Sericostomatidae, Beraeidae, Molannidae, Odontoceridae, Leptoceridae and Polycentropidae. A simple granulose haustellar surface, devoid of channels, is present in the Hydropsychidae, Psychomyidae, Philopotamidae, Rhyacophilidae and Hydroptilidae. The mandibles are of doubtful function. They are largest in the Hydropsychidae and Rhyacophilidae, and most reduced in Limnephilidae. Small lobes, which are thought to represent the ligula of the labium, are seen in the Philopotamidae, Hydropsychidae, Psychomyidae and Polycentropidae. These differing conditions of the mouth parts are shown to accord with views on the phylogeny of the Trichoptera, which are derived from other data. An account is given of published descriptions of modified mouth parts in some exotic species. The nature of these modifications is discussed. Published observations on the feeding of caddis flies are reviewed. It is concluded that using the haustellum to drink nectar and water is a normal activity of caddis flies."

Curtis,J 1835 British Entomology being Illustrations and Descriptions of the Genera of Insects found in Great Britain and Ireland Containing Coloured Figure from Nature of the Most Rare and Beautiful Species, and in Many Instances of the Plants Upon Which They are Found. Richard Taylor, London. vol. XII, 530-577.



Deutsch,WG 1985 Swimming modifications of adult female Hydropsychidae compared with other Trichoptera. Freshwater Invertebrate Biology, 4(1) 35-40.
     Abstract: "Mesothoracic legs of 10 of 12 species of adult female Hydropsychidae (not Parapsyche almota or P. elsis) are modified for swimming; at least four of the ten species are known to swim to stream substrates to oviposit. Modified tibiae and tarsi of females differed from those of males in being widened, concave, and often, with a fringe of longer hair on the posterior edge. Modified mesothoracic legs were also found on some adult female Glossosomatidae, Polycentropodidae, and Psychomyiidae. Differences in swimming modifications found among hydropsychids and glossosomatids suggest a diversity of ovipositing behavior within these families."

Dodds,GS and Hisaw,FL 1925 Ecological studies on aquatic insects. IV. Altitudinal range and zonation of mayflies, stoneflies and caddisflies in the Colorado Rockies. Ecology 6(4)380-390. Abstract PDF

Elliot,AG; Hubert,WA and Anderson,SH 1997 Habitat associations and effects of urbanization on macroinvertebrates of a small, high-plains stream. Journal of Freshwater Ecology 12 (1) 61-73.

Fairchild,MP and Holomuzki,JR 2005 Multiple predator effects on microdistributions, survival, and drift of stream hydropsychid Caddisflies. J. NABS 24: 101-112.

Ge,X; Zang,H; Ye,X; Peng,L; Wang,B; Lian,G and Sun,C 2022 Comparative mitogenomic analyses of Hydropsychidae revealing the novel rearrangement of protein-coding gene and tRNA (Trichoptera: Annulipalpia). Insects, 13(9), p.759. PDF
     Abstract: " Gene rearrangement of the mitochondrial genome of insects, especially the rearrangement of protein-coding genes, has long been a hot topic for entomologists. Although mitochondrial gene rearrangement is common within Annulipalpia, protein-coding gene rearrangement is relatively rare. As the largest family in Annulipalpia, the available mitogenomes from Hydropsychidae Curtis, 1835 are scarce, and thus restrict our interpretation of the mitogenome characteristic. In this study, we obtained 19 novel mitogenomes of Hydropsychidae, of which the mitogenomes of the genus Arctopsyche are published for the first time. Coupled with published hydropsychid mitogenome, we analyzed the nucleotide composition evolutionary rates and gene rearrangements of the mitogenomes among subfamilies. As a result, we found two novel gene rearrangement patterns within Hydropsychidae, including rearrangement of protein-coding genes. Meanwhile, our results consider that the protein-coding gene arrangement of Potamyia can be interpreted by the tandem duplication/random loss (TDRL) model. In addition, the phylogenetic relationships within Hydropsychidae constructed by two strategies (Bayesian inference and maximum likelihood) strongly support the monophyly of Arctopscychinae, Diplectroninae, Hydropsychinae, and Macronematinae. Our study provides new insights into the mechanisms and patterns of mitogenome rearrangements in Hydropsychidae."

Gray,LJ and Ward,JV 1979 Food habits of stream benthos at sites of differing food availability. American Midland Naturalist 102 1, 157-167. PDF

Holzenthal,RW; Blahnik,RJ; Prather,AL and Kjer,KM 2007 Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. PDF
     "Hydropsychidae: Hydropsychidae, with about 1,500 described species, is the 3rd largest family in Trichoptera and the most diverse of the net-spinning annulipalpians. It was established by Curtis (1835) and, like most of the families described early in the history of the order, its composition has changed substantially over the years, achieving its modern definition with the work of Ulmer (1907b). Five subfamilies are currently recognized: Arctopsychinae, Macronematinae, Hydropsychinae, Diplectroninae, and Smicrideinae. The monophyly and phylogenetic relationships among these subfamilies has been the subject of recent studies (Geraci et al. 2005, Schefter 1996, 2005), but a stable generic classification has yet to be achieved. Arctopsychinae, often recognized as a separate family (e.g., Fischer 1963, Schmid 1968), is widespread in the Holarctic and Oriental regions. Three genera are recognized, with Arctopsyche McLachlan and Parapyche Betten the largest, each with about 2 dozen species, and Maesaipsyche Malicky and Chantaramongkol the smallest, with 2 Southeast Asian species. Macronematinae, with 17 genera, is found in all biogeographical regions. The largest genera include Leptonema Guerin-Meneville with more than 125 species, largely in the Neotropics, but with a significant fauna in Africa and Madagascar and some extending into the southwestern USA, and the cosmopolitan Macrostemum Kolenati, with about 100 species. Other genera, with about 20-30 species each, include Macronema Pictet (Neotropical), Amphipsyche McLachlan (Afrotropical, Oriental, Palearctic), and Polymorphanisus Walker (Afrotropical, Oriental). The other genera each contain 10 or fewer species and include: Aethaloptera Brauer (India, Africa, Russian far east), Baliomorpha Neboiss (Australasian), Blepharopus Kolenati (Neotropical), Centromacronema Ulmer (Neotropical), Leptopsyche McLachlan (Australasian), Oestropsyche Brauer (Sri Lanka), Plectromacronema Ulmer (Neotropical), Protomacronema Ulmer (Afrotropical), Pseudoleptonema Mosely (Oriental), Pseudomacronema Ulmer (Neotropical), Synoestropsis Ulmer (Neotropical), and Trichomacronema Schmid (Oriental). Hydropsychinae currently contains 19 genera from all biogeographic regions, although it barely reaches the northern Neotropical region, and is absent from South America, where it is replaced by species of Smicrideinae and Macronematinae. The largest genera include Hydropsyche Pictet (ca. 275 species), Ceratopsche Ross and Unzicker, often considered a subgenus of Hydropsyche (ca. 100 species), and Cheumatopsyche Wallengren (ca. 260 species). Other, medium-sized genera (10-50 species) include Potamyia Banks (mostly eastern Palearctic and Oriental, but with 1 Nearctic species), Hydromanicus Brauer (Afrotopical, Oriental), and Calosopsyche Ross and Unzicker (Central America, Greater Antilles). The other genera contain less than 10 species each and include: Abacaria Mosely, Aoteapsyche McFarlane, Austropsyche Banks, Caledopsyche Kimmins, and Orthospsyche McFarlane (Australasian), Herbertorossia Ulmer (Australasian, Oriental), Hydatomanicus Ulmer (Oriental), Hydatopsyche Ulmer and Hydronema Martynov (Oriental, Palearctic), Symphitopsyche Ulmer (Afrotopical), Mexipsyche Ross and Unzicker and Plectropsyche Ross (Neotropical), and Streptopsyche Ross and Unzicker (Greater Antilles). The subfamily Smicrideinae was relatively recently established by Schefter (1996). The species had been previously placed in Hydropsychinae. Smicrideinae contains only 3 genera: Smicridea McLachlan, with 2 large subgenera, Smicridea and Rhyacophylax Müller (175 species total, mostly Neotropical in distribution, but extending into the southwestern United States), Smicrophylax Neboiss (5 Australasian species), and Asmicridea Mosely (2 Australasian species). Diplectroninae is a small subfamily with most of its diversity in a single genus, Diplectrona (ca. 120 species from all biogeographic regions, but concentrated in the Oriental and Australasian regions and absent from South America). Schefter (1996) provided evidence that neither the genus nor the subfamily may be monophyletic. Other genera included in Diplectroninae, none with more than a dozen species, include Homoplectra Ross (North America), Oropsyche Ross (eastern North America), Sciadorus Barnard (South Africa), and Diemeniluma Neboiss (Tasmania). In addition to the generic phylogenetic studies listed above, Mey (1996, 1999c, 2003) has provided important phylogenetic and biogeographic analyses of southeast Asian Hydropsyche. The nets of hydropsychids are typically attached to rocks in flowing waters and are used to capture detritus or microorganisms from the stream, but some species, particularly those in the subfamily Arctopsychinae capture and consume live prey. Net structure and function varies considerably. In the genus Macrostemum, a subdivided subterranean structure is constructed of rocks to function like a siphon, drawing the current over a specially constructed net, which is tended by the larva. Larvae of hydropsychids are often very abundant and are important in the nutrient dynamics of the rivers and streams they inhabit."

Ivanov,VD; Melnitsky,SI and Perkovsky,EE 2016 Caddisflies from Cenozoic resins of Europe. Paleontological Journal, 50(5), pp.485-493.
     Abstract: "Analysis of the available data on the findings and taxonomical structure of caddisflies (Insecta, Trichoptera) in the Cenozoic fossil resins of Europe shows that there are four European amber regions (Baltic, Rovno, Saxonian, and Danish) are characterized by a relatively abundant trichopteran fauna, comprising 27 families, 72 genera, and 256 species. These faunas show the dominance of Psychomyioidea (families Polycentropodidae, Psychomyiidae, and Ecnomidae) with Polycentropodidae comprising up to 75% of all records. The amber faunas are second in the dominance of Polycentropodidae only to the terminal Eocene of Florissant (84% of Polycentropodidae). No modern caddisfly species have been found. The amber regions are significantly different in the species composition of Trichoptera although the generic and family structures are similar. Comparison with the modern faunas of Europe shows the absence of advanced Limnephilidae, which are characteristic of the Holocene faunas of Europe, and the rarity of recently abundant Hydropsychidae and Hydroptilidae. The overall composition of amber Trichoptera suggests that it is structurally related to the faunas of Caucasus and Southeastern Asia and might be evidence of seasonally low-contrast (equable) climate in the Late Eocene of Europe."

Kiffney,PM 1996 Main and interactive effects of invertebrate density, predation and metals on a Rocky Mountain stream macroinvertebrate community. Canadian Journal of Fisheries and Aquatic Sciences 53 (7) 1595-1601. PDF

Metcalfe,AN; Kennedy,TA; Marks,JC; Smith,AD and Muehlbauer,JD 2020 Spatial population structure of a widespread aquatic insect in the Colorado River Basin: Evidence for a Hydropsyche oslari species complex. Freshwater Science, 39(2), pp.309-320. PDF

Nimmo,AP 1987 The adult Arctopsyche and Hydropsyche (Trichoptera) of Canada and adjacent United States. Questiones Entomologicae 23:1-189.

Nimmo,AP 1995 New species of Hydropsychidae and Limnephilidae (Insecta, Trichoptera) from the far east of Russia, with description of a new genus of Limnephilidae (Limnephilini). Occasional Papers on Trichoptera Taxonomy 1, 1-15.

Oláh,J and Johanson,KA 2008 Generic review of Hydropsychinae, with description of Schmidopsyche, new genus, 3 new genus clusters, 8 new species groups, 4 new species clades, 12 new species clusters and 62 new species from the Oriental and Afrotropical regions (Trichoptera: Hydropsychidae). Zootaxa, 1802 (1), 1-248. https://doi.org/10.11646/zootaxa.1802.1.1 PDF
     Abstract: "The taxonomy of the genera of Hydropsychinae is revised. The following genera are synonymized: Hydatomanicus Ulmer, 1951 and Hydatopsyche Ulmer, 1926 are synonymized with Hydromanicus Brauer, 1865; Streptopsyche Ross and Unzicker, 1977 is synonymized with Calosopsyche Ross and Unzicker, 1977; Symphitopsyche Ulmer, 1907, Mexipsyche Ross and Unzicker, 1977, Ceratopsyche Ross and Unzicker, 1977 and Herbertorossia Ulmer, 1957 are synonymized with Hydropsyche Pictet, 1834; and Plectropsyche Ross, 1947 is synonymized with Cheumatopsyche Wallengren, 1891. A new genus, Schmidopsyche Oláh and Schefter, new genus is established and the single species in the genus, S. rossi Oláh and Schefter, new species is described and illustrated, based on material collected by Schmid in Sikkim in 1959. The additional 61 new species are described and illustrated: India: Hydromanicus krsamgin Oláh and Barnard, new species; H. sacso Oláh and Johanson, new species; H. naraik Oláh and Barnard, new species; H. palnis Oláh and Barnard, new species; H. topali Oláh and Johanson, new species; Hydropsyche januha Oláh and Barnard, new species; H. kamenga Oláh and Schefter, new species; H. mizora Oláh and Schefter, new species; H. narayana Oláh and Johanson, new species; H. khasigiri Oláh and Barnard, new species; H. igunapali Oláh and Schefter, new species; H. kangra Oláh and Barnard, new species; H. kiogupa Oláh and Schefter, new species; H. minutanga Oláh and Johanson, new species; H. tritiyaha Oláh and Barnard, new species; H. ekaropa Oláh and Schefter, new species; H. yathetima Oláh and Schefter, new species; H. keralana Oláh and Barnard, new species; H. semkala Oláh and Johanson, new species; H. shillonga Oláh and Barnard, new species; Cheumatopsyche barakambra Oláh and Johanson, new species; C. mariannae Oláh and Johanson, new species; C. sandrae Oláh and Johanson, new species; Potamyia assamana Oláh and Schefter, new species. Vietnam: Hydromanicus demden Oláh and Johanson, new species; and Hydropsyche soinha Oláh and Johanson, new species (also from Malaysia); H. tanung Oláh and Johanson, new species; H. lomdom Oláh and Johanson, new species; H. sangbung Oláh and Johanson, new species; H. keoda Oláh and Johanson, new species; H. xenga Oláh and Johanson, new species; and Cheumatopsyche simayorum Oláh and Johanson, new species. Malaysia: Hydropsyche gekilara Oláh and Barnard, new species; H. tanua Oláh and Barnard, new species; and H. namea Oláh and Barnard, new species. Philippines: Hydropsyche meyi Oláh and Johanson, new species; Cheumatopsyche anonima Oláh and Johanson, new species; C. kuranishii Oláh and Johanson, new species; and C. montapo Oláh and Johanson, new species. Papua New Guinea: Hydropsyche flintorum Oláh and Johanson, new species; and H. carolae Oláh and Johanson, new species. Tibet: Hydropsyche gyantsana Oláh and Barnard, new species. Nepal: Hydropsyche nepalarawa Oláh and Johanson, new species; and Cheumatopsyche chitawana Oláh and Johanson, new species. China: Hydropsyche chengdu Oláh and Barnard, new species; and Cheumatopsyche yangmorseorum Oláh and Johanson, new species. Iran: Hydropsyche masula Oláh and Johanson, new species. New Caledonia: Orthopsyche anulmika Oláh and Johanson, new species. Japan: Cheumatopsyche okinawana Oláh and Johanson, new species; and C. tanidai Oláh and Johanson, new species. Taiwan: Cheumatopsyche davisi Oláh and Johanson, new species; and C. tungpa Oláh and Johanson, new species. Sri Lanka: Cheumatopsyche hasalaka Oláh and Johanson, new species. South Africa: Cheumatopsyche krugerana Oláh and Johanson, new species. Madagascar: Cheumatopsyche mafana Oláh and Johanson, new species; C. nondra Oláh and Johanson, new species; C. tiokalamkita Oláh and Johanson, new species; C. fianara Oláh and Johanson, new species; C. masiposa Oláh and Johanson, new species; C. ranoma Oláh and Johanson, new species; and C. rantsoa Oláh and Johanson, new species. The following species are synonymized: Hydropsyche kawamurai Tsuda, 1940 is synonymized with H. kozhantschikovi Martynov, 1924, Hydropsyche belos Malicky and Chantaramongkol, 2000 is synonymized with H. klimai Mey, 1999, Hydropsyche taiwanensis Mey, 1997 is synonymized with H. spinata Kobayashi, 1987, and Cheumatopsyche copiosa Kimmins, 1956 is synonymized with C. columnata Martynov, 1935. Recognized genera are divided into previously established and new species groups, defined by shared diagnostic characters. All Oriental species of Hydropsyche Pictet and Hydromanicus Brauer are grouped into different levels of species groups based on similar morphology in male genitalia."

Peckarsky,BL 1986 Colonization of natural substrates by stream benthos. Canadian Journal of Fisheries and Aquatic Sciences 43, 700-709. PDF

Ross,HH 1944 The Caddis Flies, or Trichoptera, of Illinois. Natural History Survey of Illinois 23 Los Angeles, CA. 326 pages.
     Contains keys to genera for adult Hydropsychidae.

Rutherford,JE and MacKay,RJ 1985 The vertical distribution of hydropsychid larvae and pupa (Trichoptera: Hydropsychidae) in stream substrates. Canadian Journal of Zoology 63:1306-1315.

Schefter,PW 2005 Re-evaluation of genera in the subfamily Hydropsychinae (Trichoptera: Hydropsychidae). Aquatic Insects 27(2) 133 - 154.
     Abstract: "The problem of generic attributions in the subfamily Hydropsychinae is reviewed. Historic morphological characters are evaluated and new morphological characters are introduced, including characters from all life history stages when available. A natural classification depends on maximizing the number of specimens studied and the number of universally applicable characters used. A cladistic analysis was performed based on examination of representative species of the world hydropsychine fauna and use of characters that embody the panoply of morphological attributes expressed in the subfamily. The analysis suggests that decisions based on insufficient evidence may have allowed six presently recognized genera to be reassigned from Hydropsyche. Therefore, a conservative approach toward classification changes within the subfamily is urged. A key to males of 11 genera resulting from the analysis is presented."

Stoaks,RD and Kondratieff,BC 2014 The aquatic macroinvertebrates of a first order Colorado, USA Front Range stream: what could the biodiversity have been before irrigated agriculture?. Journal of the Kansas Entomological Society, 87(1), pp.47-65. PDF
     Abstract: "The aquatic macroinvertebrates of Sand Creek, a first order Front Range Colorado stream, were surveyed from March-September 2012 with an additional winter adult collecting trip on 2 March 2013 using both qualitative and quantitative techniques, emphasizing EPT (Ephemeroptera/Plecoptera/Trichoptera) taxa. One hundred and thirty macroinvertebrate taxa were identified from both qualitative samples and quantitative benthic samples. Of these, 53 mayfly/stonefly/caddisfly species were collected from Sand Creek, a remarkable number for such a small Front Range stream. The macroinvertebrate community of Sand Creek is dominated by collector-gatherers (50-76%), collector-filterers (9-33%) and shredders (3-24%), with the other feeding guilds comprising low overall percentages of the community. The collector-gatherer taxa were primarily mayfly taxa, chironomid midges, stratiomyid flies, the elmid genus Optioservus, and oligochaete worms; collector-filterers were primarily net spinning hydropsychid caddisflies; and shredders were primarily nemourid stoneflies. Sand Creek should be protected from cattle grazing which causes degradation of the riparian zone and stream bank integrity."

Voelz,NJ; Poff,NL and Ward,JV 1994 Differential effects of a brief thermal disturbance on caddisflies (Trichoptera) in a regulated river. American Midland Naturalist 132 (1) 173-182.

Voelz,NJ and Ward,JV 1996 Microdistributions, food resources and feeding habits of filter-feeding Trichoptera in the Upper Colorado River. Archiv fur Hydrobiologie 137 (3) 325-348.

Warnick,SL and Bell,HL 1969 The acute toxicity of some heavy metals to different insects. Journal WPCF 41 2, 280-284.

Weaver III,JS and Morse,JC 1986 Evolution of feeding and case-making behavior in Trichoptera. Journal of the North American Benthological Society, 5(2) 150-158. PDF
     Abstract: "A phylogeny of the families of Trichoptera is reviewed to provide a basis for understanding the probable evolution of feeding tactics and case or retreat constructions by larvae. At least 48 hierarchically inclusive homologues are known, mostly from larval, pupal, and adult morphology. Their resulting phylogeny indicates that Rhyacophilidae, Hydrobiosidae, Glossosomatidae, and Hydroptilidae are more closely related to Philopotamidae, Hydropsychidae, and their allies than to Limnephilidae, Leptoceridae, and their allies. This phylogeny implies that the ancestral caddisfly larva was probably a tube-dwelling detritivore, inhabiting humus and detrital mats near the shores of lentic or lotic-depositional habitats. This ancestor evolved into a tube-case-making detritivore and scraper in the ancestor of Integripalpia and into a retreat-making collector-gatherer in the ancestor of Annulipalpia. All other larval feeding and case-making tactics evolved from these ancestral habits."

Wichard,W Kraemer,MM and Luer,C 2006 First caddisfly species from Mexican amber (Insecta: Trichoptera). Zootaxa, 1378(1), pp.37-48. PDF
     Abstract: "The first 4 new species of caddisflies (Trichoptera) are described from Mexican amber: Culoptila aguilerai n. sp. (Glossosomatidae), Plectropsyche alvarezi n. sp. (Hydropsychidae), Antillopsyche mexicana n. sp. (Dipseudopsidae), and Xiphocentron chiapasi n. sp. (Xiphocentronidae). Culoptila, Plectropsyche, and Xiphocentron are typical members of the Neotropical fauna and the fauna of Mexico; they now are also represented in Miocene Mexican amber. The genus Antillopsyche, previously known from the Greater Antilles and from Dominican amber, is now reported from Mexican amber."

Wiggins, GB 1996 Larvae of the North American Caddisfly Genera (Trichoptera). 2nd Edition. University of Toronto Press, 457 pages.
     A key to genera starts on page 128. Wiggins states that there are 145 species in 10 genera in North America.

Zuellig,RE; Kondratieff,BC and Rhodes,HA 2002 Benthos recovery after an episodic sediment release into a Colorado Rocky Mountain river. Western North American Naturalist 62 (1) 59-72.
     Abstract: "During late September 1996, approximately 7000 m3 of clay- to gravel-sized sediment was flushed from Halligan Reservoir, Larimer County, Colorado, into the North Fork Cache la Poudre River during dam inspections. Approximately 9.6 km of this river was partially or completely affected by this episodic sediment release. Pools up to 3.2 km downstream from the dam lost 50% of their volume. Hess samples taken from October 1996 to September 1997, 100 m downstream from the dam (site 1) and 3.2 km downstream (site 2), revealed effects of sediment on recovery patterns of benthic communities. A 2-way ANOVA was used to determine significant interactions using site and date as main factors. Pairwise differences were then compared using least squares means to determine significant dates within and between sites. Ten days after the sediment release, both density and taxa richness at site 1 (55 organisms per m2 , 5 taxa) were significantly lower (P < 0.05) than site 2 (1156 organisms per m2, 25 taxa). These differences remained until June when species richness and densities increased. Plecoptera and Trichoptera colonized from June to September after being eliminated at site 1 and reduced at site 2. No permanently flowing tributaries exist within the study area; therefore, passive downstream drift from such inputs apparently did not influence recovery. Increased densities of taxa such as Baetidae, Hydroptilidae, Hydropsychidae, Chironomidae, Simuliidae, and Oligochaeta occurred plausibly by rapid reproduction. Based on pre-event data, community function completely changed at site 2 from a scraper community to one dominated by collector-gatherers."

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Brown,WS 2005 Trichoptera (Caddisflies) of Gunnison County, Colorado, USA
www.gunnisoninsects.org