Trichoptera: RhyacophilidaeIntroduction to the Rhyacophila Pictet 1834Updated 24 March 2026
TSN 115097 
Provisional Species List
Good LinksOn this website:Rhyacophila acropedes Rhyacophila tucula Rhyacophila vocala Rhyacophila vofixa Rhyacophila in checklists from all over the world Other Websites: Photos, Map, Taxon Identifier Numbers - from the Global Biodiversity Information Facility Rhyacophilidae at GBIF Photos, Map, Museums, DNA - Barcode of Life Data System family Overview - University of Alberta Entomology Collection family page Has habitat, identification, life history and more.  ReferencesAllan,JD 1975 The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56:1040-1053. PDFWidely cited longitudinal survey of Cement Creek. Allan,JD 1978 Diet of brook trout (Salvelinus fontinalis Mitchell) and Brown Trout (Salmo trutta L.) in an alpine stream. Internationale Vereinigung für Theoretische und Angewandte Limnologie Verhandlungen 20, 2045-2050. Allan,JD 1981 Determinants of diet of brook trout (Salvelinus fontinalis) in a mountain stream. Canadian Journal of Fisheries and Aquatic Sciences 38, 184-192. PDF 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." Anderson,NH 1976 The distribution and biology of Oregon Trichoptera. Agricultural Experiment Station, Oregon State University. Technical Bulletin, 134. PDF Abstract: "More than 280 species of caddisflies (Trichoptera), representing genera in 16 families, are known to occur in Oregon. Detailed distribution records within the state are given, as well as the known geographic range for each species. Biological information, such as life cycle, habitat preference, flight period, and larval feeding habits, are presented if the data are available." Buchwalter,DB; Cain,DJ; Clements,WH and Luoma,SN 2007 Using biodynamic models to reconcile differences between laboratory toxicity tests and field biomonitoring with aquatic insects. Environmental Science and Technology 41, 4821-4828. Abstract: "Aquatic insects often dominate lotic ecosystems, yet these organisms are under-represented in trace metal toxicity databases. Furthermore, toxicity data for aquatic insects do not appear to reflect their actual sensitivities to metals in nature, because the concentrations required to elicit toxicity in the laboratory are considerably higher than those found to impact insect communities in the field. New approaches are therefore needed to better understand how and why insects are differentially susceptible to metal exposures. Biodynamic modeling is a powerful tool for understanding interspecific differences in trace metal bioaccumulation. Because bioaccumulation alone does not necessarily correlate with toxicity, we combined biokinetic parameters associated with dissolved cadmium exposures with studies of the subcellular compartmentalization of accumulated Cd. This combination of physiological traits allowed us to make predictions of susceptibility differences to dissolved Cd in three aquatic insect taxa: Ephemerella excrucians, Rhithrogena morrisoni, and Rhyacophila sp. We compared these predictions with long-term field monitoring data and toxicity tests with closely related taxa: Ephemerella infrequens, Rhithrogena hageni, and Rhyacophila brunnea. Kinetic parameters allowed us to estimate steady-state concentrations, the time required to reach steady state, and the concentrations of Cd projected to be in potentially toxic compartments for different species. Species-specific physiological traits identified using biodynamic models provided a means for better understanding why toxicity assays with insects have failed to provide meaningful estimates for metal concentrations that would be expected to be protective in nature. " Buchwalter,DB; Cain,DJ; Martin,CA; Xie,L; Luoma,SN and Garland,JT 2008 Aquatic insect ecophysiological traits reveal phylogenetically based differences in dissolved cadmium susceptibility. Proceedings of the National Academy of Sciences 105 24, 8321-8326. 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 The authors discuss abundance of aquatic insects among a number of mine-polluted and clean streams and rivers in Colorado. Quote from page 633: "Rhyacophila sp.(Fig.5j) was the only caddisfly that showed a significant response to metal level and was lower at medium-metal sites." Clements,WH and Kiffney,PM 1995 The influence of elevation on benthic community responses to heavy metals in Rocky Mountain streams. Canadian Journal of Fisheries and Aquatic Sciences, 52(9), pp.1966-1977. Abstract: "We examined the influence of elevation on benthic macroinvertebrate community responses to heavy metals at 33 sites in six Colorado streams where zinc (Zn) concentration ranged from 2 to 691 µg/L and elevation ranged from 2200 to 3500 m above sea level. We assigned the 33 sites to one of four categories, on the basis of the total Zn concentration and the hardness-based criterion for Zn. Number of taxa, abundance, and species richness of mayflies (Ephemeroptera), and abundance of most mayfly and stonefly (Plecoptera) taxa were significantly reduced at sites where Zn concentration exceeded the hardness-based criterion. Although most measures of benthic community structure were not affected at sites where the Zn concentration was less than the criterion, the species richness of mayflies and the abundance of Heptageniidae, Drunella, and Rhyacophila were significantly reduced. Results of stepwise multiple regression analysis and canonical discriminant analysis showed that the effects of Zn on benthic community structure were influenced by elevation. These natural changes in benthic communities confound results of biomonitoring studies and must be considered when assessing heavy metal impacts. Total abundance and species richness of mayflies, and abundance of Heptageniidae were the most reliable indicators of heavy metal impacts in our southern Rocky Mountain streams." 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." Dallai,R; Lupetti,P and Afzeliust,BA 1995 Sperm structure of Trichoptera. IV. Rhyacophilidae and Glossosomatidae. International Journal of Insect Morphology and Embryology, 24(2)185-193. Abstract: "The families Rhyacophilidae and Glossosomatidae (Trichoptera) are considered to be the most primitive ones within the order. We examined the spermatozoa of members of these families to see whether their ultrastructure is consistent with an ancestral position. Axonemal structures, after fixation with a tannic acid-containing fixative, have been shown to be particularly useful as taxonomical indicators. It was found that 4 members of Rhyacophilidae, representing 3 subgenera (Rhyacophila, Pararhyacophila, and Hyporhyacophila) all have motile spermatozoa, with a 9 + 9 + 2 axoneme in which inner (but no outer) dynein arms are present. The accessory tubules have a wall consisting of 17 protofilaments, decreasing to 16 near the distal end, whereas the examined member of Glossosomatidae, Catagapetus nigrans, has accessory tubules with 18 protofilaments and a 9 + 9 + 2 axoneme with inner dynein arms and with motility similar to the Rhyacophilidae. Sperm motility is consistent with the inclusion of these 2 families within the suborder Integripalpia, and the axonemal pattern 9 + 9 + 2 indicates that the families indeed occupy a primitive position within Trichoptera." Denning,DG 1948 A review of the Rhyacophilidae. Canadian Entomologist 80:97-117. DeWalt,RE; Stewart,KW; Moulton,SR and Kennedy,JH 1994 Summer emergence of mayflies, stoneflies, and caddisflies from a Colorado mountain stream. Southwestern Naturalist 39 (3) 249-256. Djernæs,M 2010 Morphology, function and evolution of the sternum V glands in Amphiesmenoptera. PhD thesis, University of Alberta. 368 pages. Djernæs,M 2011 Structure and phylogenetic significance of the sternum V glands in Trichoptera Zootaxa 2884: 1-60. Abstract: "I investigated the sternum V gland in 38 families of Trichoptera, and found it to be present in 25 of these. I found that the gland is generally present in Annulipalpia, except Dipseudopsidae, and in Spicipalpia. It is widespread in Plenitentoria, while it is often absent in Brevitentoria, especially in males. The opening is slit-like and U or crescent-shaped. There is significant variation in the cuticular structures associated with the opening ranging from no apparent modification, over scaly patches to elaborate protuberances. Gland opening muscles are associated with the gland in all families except Psychomyiidae, and are divided into 2 distinct types: One originating on the front edge of sternum VI found in Philopotamidae, Rhyacophilidae, Glossosomatidae and Hydroptilidae; and 1 originating on the cuticle of sternum V found in all other trichopterans. The shape of the gland reservoir is variable, from round periform to reniform, elongate or compartmentalised. Muscle fibres are often associated with the reservoir, but are notably absent in Limnephilidae. I mapped characters based on gland structures on a phylogeny of Trichoptera, and discuss the results. The sternum V gland provides potentially important characters from the superorder to the species leve l. I discuss 2 cases where characters from the sternum V gland may solve existing phylogenetic and taxonomic puzzles: Delimitation of Dipseudopsidae versus Polycentropodidae and the relationships among the hydropsychid subfamilies. " 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 Ellsworth,SD 2000 Influence of substrate size, Cladophora, and caddisfly pupal cases on colonization of macroinvertebrates in Sagehen Creek, California. Western North American Naturalist, pp.311-319. PDF Abstract: "Colonization of stream macroinvertebrates onto artificial substrates was examined in Sagehen Creek California, during 1993. Substrate size and presence or absence of pupal cases of the caddisfly Anagapetus (Glossosmatidae) were examined as possible influences on macroinvertebrate colonization. Macroinvertebrates were collected from substrates after 60 d of incubation and identified to the lowest feasible taxonomie unit. Dry weight of the filametous green alga Cladophora also was determined for each substrate. A general linear model indicated a significant positive relationship between both substrate size and presence pupal cases on richness (P < 0.001). There was no significant effect of cases on the E5 evenness index, which supports the passive sampling model as a mechanism giving rise to the effect of cases on richness. A multivariate general linear model indicated a significant positive effect of Anagapetus cases on Micrasema bactro, Epeorus, and Rhyacophila sp and of substrate size on Hydropsyche occidentalis and Epeorus. Neither cases nor substrate size had a significant effect on Cladophora density (P > 0.05). There were significant negative correlations between Cladophora density and both Shannon H' and J' indices. There were significant positive relationships between Cladophora density with Baetis, Simulium, and Chironomidae density. Combined main effects of substrate size, Anagapetus pupal cases, and Cladophora have dynamic influences on this macroinvertebrate fauna. This study demonstrates the complex interplay of biotic abiotic factors that determine macroinvertebrate distribution and abundance in this system." Etnier,DE; Parker,CR and Stocks,IC 2004 A new species of Rhyacophila Pictet (Trichoptera: Rhyacophilidae) from Great Smoky Mountains National Park, with illustrations of females of R. appalachia Morse and Ross and R. mycta Ross. Proceedings of the Entomological Society of Washington. 106(2): 396- 406. Friedrich,F; Schulz,J; Kubiak,M; Beckmann,F and Wilde,F 2015 The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera. Journal of Morphology, 276(12), pp.1505-1524. Abstract: "The external and internal features of the larval head of Rhyacophila fasciata (Trichoptera: Rhyacophilidae) were described in detail. Anatomical examinations were carried out using a multimethod approach including histology, scanning electron microscopy, confocal laser-scanning microscopy, microcomputed tomography, and computer-based three-dimensional reconstructions. Additionally, the information on the larval head of Limnephilus flavicornis (Limnephilidae) and Hydropsyche angustipennis (Hydropsychidae) available in the literature were reinvestigated. These anatomical data were used to address major questions of homology and terminology, that is, the ventral closure of the head capsule, the sclerites, and appendages of labium and maxilla and their muscles. These topics were discussed by summarizing the main hypotheses present in the literature and a critical inclusion of new findings. Consequently, the inner lobe of the maxilla very likely represents the galea. The distal maxillary sclerite (palpifer) is an anatomical composite formation at least including dististipes and lacinia. Based on these homology hypotheses several potential groundplan features of the larval head of Trichoptera were reconstructed. The head of Rhyacophila shows several presumably plesiomorphic features as for instance the prognath orientation of the mouthparts, the well-developed hypocranial bridge, the triangular submentum and eyes composed of seven stemmata. Derived features of Rhyacophila are the reduced antennae, the anterior directing of three stemmata and the shift of the tentorio-stipital muscle to the mentum." Giersch,JJ 2002 Revision and phylogenetic anaysis of the verrula and alberta species groups of Rhyacophila Pictet 1834 with description of a new species (Trichoptera: Rhyacophilidae). Master of Science Thesis. Montana State University PDF Abstract: "Adult, larval, and pupal characters, and data from newly associated life stages supported the monophyly of the Rhyacophila verrula- and alberta-species groups. The verrula-group is comprised of R. haddocki Denning, R. leechi Denning, R. potteri Denning in Denning and Schmid, R. rickeri Ross, R. singularis Botosaneanu, and R. verrula Milne, with a new inclusion of R. chandleri Denning. Rhyacophila autumnalis Nimmo was found to be a junior synonym of R. potteri. The monophyly of the verrula-group is based on synapomorphies that include the expanded dorsal appendage of the phallic apparatus, closed m-cell in the forewing and morphology of the larvae. The morphology of the known larvae of this group supports the hypothesis of phytophagous feeding habits, which served as an opportunity for the evolution and speciation within the group. Rhyacophila chandleri was found to be a basal member of the verrula-group, although autapomorphies suggest an early isolation from the rest of the verrula-group. Rhyacophila rickeri is basal to the rickeri-subgroup, with R. singularis and R. potteri as sister species. Rhyacophila verrula, R. leechi, and R. haddocki form the verrula-subgroup. The wide distribution of R. verrula suggests a wider ecological tolerance than other species of the group. The alberta-group is comprised of two subgroups, R. alberta, R. tucula, and R. n. sp. comprise the alberta subgroup, and R. kincaidi and R. glaciera comprise the glaciera-subgroup. Phylogenetic reconstructions of the alberta- and verrula-groups suggest the evolution of these species groups was driven by post-Pleistocene isolation of populations in headwater streams." Holzenthal,RW; Blahnik,RJ; Prather,AL and Kjer,KM 2007 Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. PDF Abstract: "Rhyacophilidae: Rhyacophilidae is a relatively large family, originally established by Stephens (1836). At one time the family included also Glossosomatidae and Hydrobiosidae and other taxa, but its definition has progressively become more restricted. Evolutionary relationships of the family were discussed by Ross (1956) and the family was the subject of a large revision by Schmid (1970). The family is predominantly north temperate and is found in North America, Europe, and Asia, but also extends into India and the tropical areas of southeastern Asia. Currently most of the diversity is included in a single genus, Rhyacophila Pictet, the largest genus in Trichoptera, with over 700 species and additional ones regularly being described. In addition to the landmark works of Ross and Schmid on Rhyacophila, Prather and Morse (2001) studied the phylogeny of the R. invaria group from eastern North America and Mey (1999b) investigated the biogeography of Southeast Asian members of the genus. Other genera include Himalopsyche Banks (ca. 50 species, predominantly in the eastern Palearctic and Oriental regions, but with 1 species from western North America), Philocrena Lepneva (1 species from Georgia, western Palearctic), and Fansipangana Mey (a single species recently described from Vietnam). The family is 1 of 2 (the other being Hydrobiosidae) that includes species that are free-living and predaceous as larvae, constructing a domed pupal chamber of rocks at maturity. As the etymology of the family name indicates, the larvae frequent cool, fast flowing rivers and streams. Larvae in the genus Himalopsyche, and some in the genus Rhyacophila, possess abdominal and thoracic gills, quite different from those in Integripalpia or Hydropsychidae." Kiffney,PM; Clements,WH 1993 Bioaccumulation of heavy metals by benthic invertebrates at the Arkansas River, Colorado. Environmental Toxicology and Chemistry 12, 1507-1517. Quote from page 1512: "Variation among taxa: Metal concentrations in organisms collected from station AR-5 [impacted by heavy metal pollution from California Gulch] (fall, spring, summer) varied significantly among taxa (Fig 7). The highest concentrations were generally found in the mayfly Baetis spp., the stonefly Pteronarcella badia, and the caddisfly Arctopsyche grandis, whereas the lowest levels were measured in the two predators, Skwala americana, and Rhyacophila spp." Löfstedt,C; Hansson,BS; Petersson, E; Valeur,P and Richards,A 1994 Pheromonal secretions from glands on the 5th abdominal sternite of hydropsychid and rhyacophilid caddisflies (Trichoptera). Journal of Chemical Ecology (20)153-170. Martin,ID and Mackay,RJ 1982 Interpreting the diet of Rhyacophila larvae (Trichoptera) from gut analyses: an evaluation of techniques. Canadian Journal of Zoology, 60(5), 783-789. Abstract: " Three commonly used gut analysis techniques were compared for reliability in determining the food habits of fifth-instar Rhyacophila fuscula (Walker) larvae. The predaceous Rhyacophila were fed a known number of each of two prey taxa (Ephemeroptera; Ephemerellidae, and Diptera: Chironomidae) in an artificial stream under controlled conditions in the laboratory. Rhyacophila attacked equal numbers of both prey types during the experiment. Two gut analysis techniques, (1) frequency of occurrence of particular prey in the guts, and (2) the relative abundance of each prey in the guts, gave results similar to those expected. The third technique, a frequently used quantitative method which uses relative areas of the diet items, gave a significantly biased interpretation of prey selection by R. fuscula. The indiscriminate application of any of these methods may lead to errors in interpreting the diet of individual predator taxa, and consequently, incorrect conclusions in studies of comparative feeding ecology. " McLaughlin,JE; Frandsen,PB; Mey,W and Pauls,SU 2019 A preliminary phylogeny of Rhyacophilidae with peference to Fansipangana and the monophyly of Rhyacophila. Zoosymposia, 14, pp.189-192. PDF Abstract: "The phylogeny of Rhyacophilidae was explored with 28S ribosomal RNA (rRNA) and Cytochrome Oxidase Subunit I (COI) mitochondrial DNA (mtDNA). Eighty one rhyacophilids were included in the analysis. We found that although Rhyacophilidae was recovered as monophyletic, intrafamilial relationships are not well-resolved using this dataset. Bootstrap support was poor for intrageneric relationships and additional data will be required to present a more robust hypothesis. The recovered phylogeny places Fansipangana as the sister taxon of the rest of Rhyacophilidae. We found that Himalopsyche was nested inside the genus Rhyacophila with the verrula group sister to Himalopsyche and remaining Rhyacophila. These results and possible relationships should be tested with a more extensive data set." Milne,LJ 1936 Studies in North American Trichoptera. Part 3. Cambridge, Mass. : Author's publication. 128 pages Nimmo,A 1971 The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta and eastern British Columbia and their post glacial origin. Quaestiones Entomologicae 73: 3-234. Obernborfer,RY; McArthur,JV; Barnes,JR and Dixon,J 1984 The effect of invertebrate predators on leaf litter processing in an alpine stream. Ecology, 65(4), pp.1325-1331. Abstract: " The effect of the predators Megarcys signata (Plecoptera: Perlodidae) and Rhyacophila sp. (Trichoptera: Rhyacophilidae) on the abundance of macroinvertebrates and on the rates of leaf processing in artificial leaf packs was studied by using manipulative field experiments. Predators were confined within artificially constructed leaf packs in an alpine stream in Utah. Both predators significantly reduced the rate of breakdown of leaves in the fall, but had no effect in late winter, when the most important shredder, Zapada cinctipes, emerges, Predation on shredders caused a reduction in breakdown rates and an increase in the residence time of the leaf litter in the stream. In experimental treatments where predators significantly reduced the numbers of shredders and decreased the rate of leaf processing, leaf pack half-life increased an average of 10.3 d. These experiments demonstrate that invertebrate shredders can contribute substantially to the rate of leaf processing in streams and suggest that one of the factors limiting the abundance of natural shredder population is predation." Pauls,SU; Graf,W; Hjalmarsson,AE; Lemmon,A; Lemmon,EM; Petersen,M; Vitecek,S and Frandsen,PB 2023 Gill structure linked to ecological and species diversification in a clade of caddisflies. Arthropod Systematics & Phylogeny, 81, pp.917-929. PDF Abstract: "Streams represent a special case of directional environmental gradients where ecological opportunity for diversification may be associated with upstream and downstream dispersal into habitats that differ in selective pressures. Temperature, current velocity and variability, sediment erosion dynamics and oxygen saturation are key environmental parameters that change in predictable ways from springs to river mouth. Many aquatic insects occupy specific longitudinal regions along these gradients, indicating a high degree of adaptation to these specific environmental conditions. In caddisflies, the evolution of tracheal gills in larval and pupal stages may be a major driver in oxygen uptake efficiency and ecological diversification. Here we study the evolution of larval gill structure in the Rhyacophila vulgaris species group using phylogenomic methods. Based on anchored hybrid enrichment, we sequenced 97 kbp of data representing 159 independent nuclear protein coding gene regions to infer the phylogeny of the R. vulgaris species group, whose species exhibit both high diversity of gill types and varied longitudinal preferences. We find that the different gill types evolved independently as derived characters in the genus and that gill structure is linked to the longitudinal habitat preference, thereby serving as a possible ecological key innovation in the R. vulgaris group."  Peck,DL and Smith,SD 1978 A revision of the Rhyacophila coloradensis complex (Trichoptera: Rhyacophilidae). Melanderia 27, 1-24. Pictet,FJ 1834 Recherches pour servir l'histoire et l'anatomie des Phryganides. A. Cherbuliez, Geneva. Prather,AL and Morse,JC 2001 Eastern Nearctic Rhyacophila species, with revision of the Rhyacophila invaria group (Trichoptera: Rhyacophilidae). Transactions of American Entomological Society 127 1, 85-166. ERRATUM http://entweb.clemson.edu/research/rhyacophila.htm Has updated keys to Eastern United States (US) Rhyacophila Females, Males, Larvae. To find these keys, look at the bottom of the webpage. Rader,RB and Ward,JV 1988 Influence of regulation on environmental conditions and the macroinvertebrate community in the upper Colorado River. Regulated Rivers: Research and Management 2:597-618. PDF Quote from page 611 and 612: "The reference site was represented by twelve species of caddisflies, including relatively abundant populations of Arctopsyche grandis and Rhyacophila acropedes. Trichopterans at the regulated site, however, were represented by nine rare and three slightly more abundant caddisflies (Hydroptila sp., Brachycentrus americanus, and Hesperophylax designatus). The abundance of net-spinning caddisflies was significantly reduced in the regulated site compared to both reference and recovery locations (p=0.05), as has been reported by several workers (Armitage and Capper, 1976; Müller, 1962; Ward, 1987). The thirteen species of Trichoptera in the recovery site included four of the most abundant species of macroinvertebrates at that site: Glossosoma ventrale, Brachycentrus americanus, Lepidostoma ormeum, Oligophlebodes minutes. Ward (1987) summarized the effects of regulation on Trichoptera in Rocky Mountain streams and concluded that Rhyacophila and Hydropsyche were the only genera commonly occurring at both regulated and reference locations, whereas Brachycentrus, Glossosoma, Arctopsyche, and Lepidostoma were often reduced or absent in regulated segments. With the exception of Brachycentrus americanus, which was significantly more abundant in the regulated and recovery sites, compared to the reference site (p=0.05), data from this study concur with previous conclusions concerning the influence of reguation on Trichoptera (Ward, 1987)." Ross,HH 1950 New species of nearctic Rhyacophila (Trichoptera, Rhyacophilidae). Journal of the Washington Academy of Sciences 40 8, 260-265. Ross,HH 1956 Evolution and classification of the mountain caddisflies. University of Illinois Press, Urbana, 213 pages. Has keys, illustrations, discusses phylogeny and biogeography of Rhyacophila among other primative caddisflies. Schmid,F 1970 Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Memoires de la Societe Entomologique du Canada 66:1-230. Résumé: "Le genre Rhyacophila présente un intérèt exceptionel parmi les Trichoptères, car il est le plus primitif de l'ľordre et ľun des plus importants. Peuplant presque tout le réseau lotique de la région holarctique, il contient 465 especes connues a ce jour. Dans ce travail, les caractères imaginaux sont décrits en détail, leur primitivité évaluéc et le genre situé dans ľascendance presque directe des autres families. La musculature des genitalia du ♂ est décrite et commentće, de même que les mouvements probables des pièces génitales et luer position à l'accouplement. Les Rhyacophilides sont aujourd'hui réduits aux seuls genres Rhyacophila et Himalopsyche dont les relations sont définies ici. Pour la première fois chez les Trichoptères de nombreux exemples de mimétisme sont signalés. Le corps de l'ouvrage est cinstitué par une étude phylétique détaillć de l'évolution et de la différenciation du genre, au cours de laquelle une approche diachronique du problème a été tentée. Seuls les caractères des génitalia du ♂ ont été utilisés, car ils se sont révélés les seuls utiles. Mais leurs variations sont si riches et si complexes qu'elles ont rendu relativement faciles l'identification des lignées et l'articulation de beucoup d'entre elles. Les effectifs du genre sont repartis en quatre branches, elles-memes divisées en rameaux dans lesquels se classent 78 groupes d'espèces. Le centre d'apparition peut ètre situé dans la région orientale et l'âge du genre suppose crétace inférieur. Durant le Tertiaire, un petit mombre de lignćes se sont dispersćes en Europe et en Amérique néarctique. Enfin, 157 especes et sous-espèces sont décrites comme nouvelles. On en trouvera la liste à la page 119." Abstract translated via Google Translate https://translate.google.com/ The genus Rhyacophila is exceptionally interesting among the caddisflies, as it is the most primitive of order and one of the most important. Populating almost the entire Holarctic region, it contains 465 species known to date. In this work, the imaginal characters are described in detail, their primitiveness evaluated and the almost direct descent from other families discussed. The muscles of the ♂ genitalia are described along with the likely movements of the genital parts and their position while mating. The only Rhyacophilides genera left are Rhyacophila and Himalopsyche , whose relations are defined here. For the first time in caddisflies many examples of mimicry are reported. The body of the book consists of a detailed phylogenetic study of the evolution and differentiation of the genus, in which a diachronic approach to the problem was attempted. Only characters of the genitalia of the ♂ have been used because they were proven useful. But their variations are so rich and so complex that they have made it relatively easy to identify lineages and relations between them. The genus is divided into four branches, themselves divided into branches in which rank 78 groups of species. The center of appearance can be located in the eastern area of their range and age implies lower Cretaceous. During the Tertiary, a small number of lineages dispersed into the European and American nearctic. Finally, 157 species and subspecies are described as new. The list is on page 119. Short,RA and Ward,JV 1980 Macroinvertebrates of a Colorado high mountain stream. The Southwestern Naturalist, 23-32. PDF Abstract: "The macroinvertebrate community and associated environmental parameters of a high mountain stream were studied biweekly during the open season of two consecutive years. Physicochemical parameters were similar both years indicating a cold, clear stream with soft waters, near neutral pH, and sparse algal growth. Aquatic insects comprised approximately 95% of the total macroinvertebrate numbers each year. Similar numbers of taxa were collected in 1975 (48) and 1976 (40). Although there were differences in abundance between years, the predominant organisms were generally the same. Ephemeropterans were by far the most abundant organisms both years. A high degree of sympatry was exhibited by the trichopteran Rhyacophila; six species were collected from a single riffle. A lower mean macroinvertebrate density in 1976 (774 org/m2) than in 1975 (1467 org/m2) can be attributed to heavy rainfall resulting in a 400% increase in discharge over a short time period. The differential effect of a short-term flood on the fauna also explains composition differences between years. Coarse particulate organic matter exhibited qualitative differences, lower standing crop values, and earlier seasonal maxima than a similar montane stream in the same drainage basin. Collectors and scrapers were more abundant and shredders much less abundant (and composed of different dominants) than in small eastern woodland streams." Smith,SD 1968 The Rhyacophila of the Salmon river drainage of Idaho with special reference to larvae. Annals of the Entomological Society of America 61 (3) 655-674. Abstract: "Eighteen species of Rhyacophila were found in the Salmon River drainage of Idaho in collections made during 1964, 1965, and 1966. The larvae of Rhyacophila acropedes Banks, angelita Banks, bifila Banks, coloradensis Banks, hyalinata Banks, oreta Ross, rotunda Banks, tucula Ross, vaccua Milne, vagrita Milne, vao Milne, vepulsa Milne, verrula Milne, and vocala Milne were found, and are described and figured. Distributional and bionomical data are presented for most species. Some of the Rhyacophila species encountered exhibited differences in their seasonal occurrence. Differences in adult diurnal activity periods also were noted and are recorded. An attempt was made to evaluate the preferred larval habitat of each species, but information was not sufficient to permit definite conclusions." Smith,SD 1976 A progress report on the phylogeny of Rhyacophila larvae. Pages 5-6 in Proceedings of the 1st International Symposium on Trichoptera (H. Malicky, ed.) Dr. W. Junk, The Hague. Smith,SD 1984 Larvae of Nearctic Rhyacophila, part I: acropedes group. Aquatic Insects 6:37-40. Thomas,JA; Frandsen,PB; Prendini,E; Zhou,X and Holzenthal,RW 2020 A multigene phylogeny and timeline for Trichoptera (Insecta). Systematic Entomology, 45(3), pp.670-686. PDF Abstract: "The Trichoptera, or caddisflies, are traditionally split into two taxonomic subdivisions: the ‘retreat-making’ Annulipalpia and the ‘case-making’ Integripalpia (sensu Ross). The monophyly of these groups is well documented; however, the establishment of a third subdivision, ‘Spicipalpia’, and the positions of the five ‘spicipalpian’ families is much debated. In contrast to previous molecular studies using nuclear ribosomal RNA, a recent trichopteran study (using nuclear protein-coding genes) placed one of these ‘spicipalpian’ families, the free-living predatory Rhyacophilidae, as the sister taxon to the rest of Trichoptera, a result that has significant implications for both the understanding of trichopteran evolution and its timing. This paper sets out to investigate the relationships of Trichoptera using several newly sequenced genes, together with previously published gene sequences. This dataset is the largest trichopteran dataset to date, covering six independent genes and > 10 000 nucleotides, and containing 185 species representing 49 families. With all data included, likelihood and Bayesian analyses support a monophyletic Annulipalpia and a monophyletic Integripalpia, which includes the ‘spicipalpians’ as a paraphyletic grade at the base of this clade. However, an analysis of the protein-coding data alone using similar analytical methods recovers Rhyacophilidae as the most basal taxon in Trichoptera, with low support. A reanalysis correcting for nucleotide composition bias provides support for the placement of the ‘spicipalpian’ taxa as sister to the Integripalpia, consistent with the total data analysis, suggesting that the basal position of Rhyacophilidae in the uncorrected analysis could be (or is probably) an artefact of base composition. We find it likely that ancestral trichopterans made incipient cases and retreats, and these had independent origins as precocious pupal chambers. Molecular dating analysis in beast, using the birth-death model of speciation, with a relaxed-clock model of sequence evolution informed by 37 fossil constraints, suggests that the most recent common ancestor of Trichoptera appeared in the Permian (c. 275 Ma) in line with the first appearance of Trichoptera in the fossil record, and that vicariance explains the distribution of most trichopteran taxa. A new infraordinal name, Phryganides, is introduced for the tube-case-making families of Integripalpia." Thut,RN 1969 Feeding habits of larvae of seven Rhyacophila (Trichoptera: Rhyacophilidae) species with notes on other life-history features. Annals of the Entomological Society of America, 62(4), pp.894-898. Abstract: "Of 7 species of Rhyacophila recovered from an experimental stream, 5 were carnivorous, 1 omnivorous, and 1 herbivorous. Chironomidae, Copepoda, and Acari were the most important constituents of the diets of the carnivorous forms. The 2 last-mentioned constituents were preyed upon in excess of their relative abundance on the stream bottom. Data relating to other features of the life histories are presented, including emergence periods, current preferences, and a breakdown of the duration of each of the instars and life stages. The ecological isolation of these congeneric species in a homogenous environment is discussed." The United States Geological Survey (USGS) National Water Quality Assessment Data Warehouse (NAWQA) shows this genus is present in Gunnison County. Data as of 1Sep2005 Wagner,R; Aurich,M; Reder,E and Veith,HJ 1990 Defensive secretions from the larvae of Apatania fimbriata (Pictet)(Trichoptera: Limnephilidae). Chemoecology, 1(3), pp.96-104. PDF Abstract: "When the larvae of the caddis fly Apatania fimbriata (Pictet) are threatened, drops of fluid appear on their head capsules. The secretions are produced in a gland in the dorsal part of the prothorax. The neck region contains an eversible sac with numerous single setae, groups of 3 setae, or rows of setae on the surface. The secretion is released through two paris of orifices on the lateral sides of the sac. For the most part the secretion is composed of some 30 fatty acids, with the major components having 12—14 carbon atoms and up to 4 double bonds (approx. 1—2 µg secretion per specimen). Biotests with synthetic saturated acids (C6—C12) in a stream and in the laboratory demonstrated a paralysing effect on small invertebrate predators (Rhyacophila sp.,Plectrocnemia conspersa, Hydropsyche sp., larvae, all Trichoptera). In choice experiments, Rhyacophila sp. larvae preferred larvae of Agapetus fuscipes and Drusus annulatus (Trichoptera) as food as compared with Apatania fimbriata larvae. Larger predators, such as Dinocras cephalotes (Insecta, Plecoptera) and the fish Cottus gobio, did not discriminate between Apatania fimbriata and other prey species. The use of fatty acids in defensive secretions is interpreted as an adaptation to the running water environment. They are effective repellents against Rhyacophila sp. larvae, the most important predator in the natural environment of Apatania larvae." 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 and Caspers,N 1991 Caddisflies of Baltic amber - 2. Fossil species of the genus Rhyacophila. Pages 447-451 in Proceedings of the 6th International Symposium on Trichoptera (C. Tomaszewski, ed.) Adam Mickiewicz University Press, Poznan, Poland. Wichard,W; Schmidt,HH and Wagner,R 1993 The semipermeability of the pupal cocoon of Rhyacophila (Trichoptera: Spicipalpia). Pages 25-27 in Proceedings of the 7th International Symposium on Trichoptera (C. Otto, ed.) Backhuys Publishers, Leiden, The Netherlands. Wiggins,GB 1996 Larvae of the North American Caddisfly Genera (Trichoptera). 2nd Edition. University of Toronto Press, 457 pages. Quote from page 114: "Rhyacophila is the largest genus in the Trichoptera, with close to 500 species widely distributed through the Holarctic and Oriental regions." Williams,DD 1980 Some relationships between stream benthos and substrate heterogeneity. Limnology and oceanography, 25(1), pp.166-172. PDF Abstract: "Baskets filled with stones, to give four different degrees of heterogeneity, were placed in a stream and made available for colonization by benthic invertebrates. No difference resulted in the total numbers and weights of animals colonizing; 26% of the species, however, showed clear substrate preferences, 35% showed slight preferences, and 39% showed none. The last group made up most of the benthic invertebrate biomass. A high quantity of detritus accumulated in each substrate, and this may account for the similar densities of detritivores collected on all four substrates." Wold,JL 1974 Systematics of the genus Rhyacophila (Trichoptera: Rhyacophilidae) in western North America with special reference to the immature stages. M.S. thesis, Oregon State University, Corvallis. PDF |