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Trichoptera: Rhyacophilidae

Introduction to the Rhyacophila

Freeliving Caddisflies, Green Rock Worm, Green Caddis, Green Sedge, Mountain Caddisflies, medium olive body caddis #16-20

Pictet 1834

Updated 17 June 17
TSN 115097

Provisional Species List

Good Links

On this website:
Rhyacophila acropedes
Rhyacophila tucula
Rhyacophila vocala
Rhyacophila vofixa
Rhyacophila in checklists from all over the world

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Allan,JD 1975 The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56:1040-1053. PDF
     Widely 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."

Buchwalter,DB; Cain,DJ; Clements,WH; 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; 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; 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. "

Denning, DG 1948 A review of the Rhyacophilidae. Canadian Entomologist 80:97-117.

DeWalt,RE; Stewart,KW; Moulton,SR; 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

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.

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 http://virgin.msu.montana.edu/Joe/

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. "

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."

Peck,DL; 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.

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.

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

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."

Wold,JL 1973 Systematics of the genus Rhyacophila (Trichoptera: Rhyacophilidae) in western North America with special reference to the immature stages (Doctoral dissertation). PDF

Brown,WS 2004 Trichoptera or Caddisflies of Gunnison County, Colorado, USA