Stoneflies - Plecoptera: Chloroperlidae of Gunnison County, ColoradoParaperla frontalis
Hyporheic Sallfly(Banks, 1902)
Updated 26 02 2026
TSN 103234
Description
Large for a chloroperlid. Pale and thin with eyes set very far forward. The illustration on the right is from Needham and Claassen's classic stonefly study published in 1925.
Habitat
Creeks, rivers and cold lakes. Nymphs live in the hyporheic zone of rivers and floodplains. The hyporheic zone is the water between the bottom of the stream and the groundwater. This habitat can extend horizontally underneath the floodplain for some distance. Hidden from the surface for most of the year, nymphs are collected near emergence in surface water and from wells at other times of the year.
Life History
Dewalt and Stewart (1995) collected nymphs infrequently in winter months among marginal substrates of streams. Less than 10 adults were collected in early June during the 3 year study on the Rio Conejos river. Baumann et al (1977) note that adults can be collected from April to early August.
Locations Collected
Knight and Gaufin (1966) collected this species between 7200 and 9600 feet elevation. Kondratieff and Baumann note that this relatively common species is rarely collected unless you use a beat sheet or examine debris along streams.
Notes
The genus Paraperla was previously included in Perlinella. Older publications may refer to this species as Perlinella frontalis.
Good Links
Photos, Map, Taxon Identifier Numbers - from the Global Biodiversity Information Facility Paraperla frontalis at Gbif
Photos, Map, Museum specimens, DNA - Barcode of Life Data System
Plecoptera Species File Paraperla frontalis
References
Banks,N 1902 Notes and descriptions of Perlidae. Canadian Entomologist 34:123-125.
Described as Perlinella frontalis.
Baumann,RW 1979 Nearctic stonefly genera as indicators of ecological parameters (Plecoptera: Insecta). The Great Basin Naturalist, 39(3) 241-244. PDF
Quote from page 244: "Chloroperlidae are generally very sensitive to environmental eutrophication and occur only under very favorable conditions. They are generally only found in cold lotic systems. Paraperla frontalis (Banks) has, however, been collected at or near the mouth of several lakes in Glacier National Park and also in the Canadian Rockies."
Baumann,RW, Gaufin,AR and Surdick,RF 1977 The stoneflies (Plecoptera) of the Rocky Mountains. Memoirs of the American Entomological Society 31, 1-208. PDF
Quoted from page 188: "This species is found in creeks, rivers and cold lakes. The adults emerge from April to early August."
DelVecchia,AG; Stanford,JA and Xu,X 2016 Ancient and methane-derived carbon subsidizes contemporary food webs. Nature communications, 7 PDF
DeWalt,RE; Hopkins,H; Neu-Becker,U and Stueber,G 2024 Paraperla frontalis (Banks, 1902). Plecoptera Species File. Retrieved on 2024-01-13 at https://plecoptera.speciesfile.org/otus/925708/overview
I read one or another of the stonefly webpages from Plecoptera Species File regularly and sometimes add papers or other information I learned to the website you're reading https://www.gunnisoninsects.org/ :-)
DeWalt,RE and Stewart,KW 1995 Life histories of stoneflies (Plecoptera) in the Rio Conejos of southern Colorado. Great Basin Naturalist 55, 1-18. PDF
Jordan,S; DelVecchia,A; Hand,BK; Hayssen,L; Nissley,C; Luikart,G and Stanford,J 2016 Genomic data reveal similar genetic differentiation between invertebrates living under and on a riverine floodplain. bioRxiv, p.044073. PDF
Abstract: "Little is known about the life histories, population connectivity, or dispersal mechanisms of shallow groundwater organisms. Here we used RAD-seq to analyze population structure in two aquifer species: Paraperla frontalis, a stonefly with groundwater larvae and aerial adults, and Stygobromus sp., a groundwater-obligate amphipod. We found similar levels of connectivity in each species between floodplains separated by ~70 river km in the Flathead River basin of NW Montana, USA. Given that Stygobromous lacks the aboveground life stage of P. frontalis, our findings suggest that aquifer-obligate species might have previously unrecognized dispersal capacity."
Knight,AW and Gaufin,AR 1966 Altitudinal distribution of stoneflies (Plecoptera) in a Rocky Mountain drainage system. Journal of the Kansas Entomological Society 39 4, 668-675.
Kondratieff,BC and Baumann,RW 2002 A review of the stoneflies of Colorado with description of a new species of Capnia (Plecoptera: Capniidae). Transactions of American Entomological Society 128 (3) 385-401.
Quoted from pages 394-395: "This is a relatively common species, but adults, especially males, are rarely collected unless by using a beating sheet or examining streamside debris piles along Mountain and Plateau streams and rivers. Nymphs are hyporheic until near emergence."
Malison,RL; DelVecchia,AG; Woods,HA; Hand,BK; Luikart,G and Stanford,JA 2020 Tolerance of aquifer stoneflies to repeated hypoxia exposure and oxygen dynamics in an alluvial aquifer. Journal of Experimental Biology, 223(16). PDF
Abstract: "Aquatic insects cope with hypoxia and anoxia using a variety of behavioral and physiological responses. Most stoneflies (Plecoptera) occur in highly oxygenated surface waters, but some species live underground in alluvial aquifers containing heterogeneous oxygen concentrations. Aquifer stoneflies appear to be supported by methane- derived food resources, which they may exploit using anoxia-resistant behaviors. We documented dissolved oxygen dynamics and collected stoneflies over 5 years in floodplain wells of the Flathead River, Montana. Hypoxia regularly occurred in two wells, and nymphs of Paraperla frontalis were collected during hypoxic periods. We measured mass-specific metabolic rates (MSMRs) at different oxygen concentrations (12, 8, 6, 4, 2, 0.5 mg l-1 , and during recovery) for 111 stonefly nymphs to determine whether aquifer and benthic taxa differed in hypoxia tolerance. Metabolic rates of aquifer taxa were similar across oxygen concentrations spanning 2 to 12 mg l-1 (P>0.437), but the MSMRs of benthic taxa dropped significantly with declining oxygen (P<0.0001; 2.9-times lower at 2 vs. 12 mg l-1 ). Aquifer taxa tolerated short-term repeated exposure to extreme hypoxia surprisingly well (100% survival), but repeated longer-term (>12 h) exposures resulted in lower survival (38-91%) and lower MSMRs during recovery. Our work suggests that aquifer stoneflies have evolved a remarkable set of behavioral and physiological adaptations that allow them to exploit the unique food resources available in hypoxic zones. These adaptations help to explain how large-bodied consumers might thrive in the underground aquifers of diverse and productive river floodplains."
Malison,RL; Ellis,BK; DelVecchia,AG; Jacobson,H; Hand,BK; Luikart,G; Woods,HA; Gamboa,M; Watanabe,K and Stanford,JA 2020 Remarkable anoxia tolerance by stoneflies from a floodplain aquifer. Ecology, 101(10), p.e03127. PDF
Needham,JG and Claassen,PW 1925 A Monograph of the Plecoptera of North America. Entomological Society of America, Lafayette, Indiana. 397 pages. PDF
Illustration at top of this webpage is from page 129.
Nelson,CH 2009 Surface ultrastructure and evolution of tarsal attachment structures in Plecoptera (Arthropoda: Hexapoda). Aquatic Insects, (31)523-545. Html
The author used scanning electron microscopy (SEM) to image the plantar surfaces of the stonefly tarsomeres and pretarsus of Paraperla frontalis and a number of other species.
Quote from page 534 "The tarsus of this species shown in Figure 35 is very similar to those of Styloperlidae and Peltoperlidae in that tarsomere 1 is short, approximately two-fifths the length of
tarsomere 3. The euplantulae of tarsomeres 1 and 2 resemble those of the pteronarcyid Pteronarcys californica shown in Figures 25 and 26 in that their surface profile is smooth. The plantar region of tarsomere 3 is identical to those of Pteronarcella badia (Pteronarcyidae) shown in Figure 17, Yoraperla brevis (Peltoperlidae) shown in Figure 29, Megarcys signata (Perlodidae) shown in Figure 31, and Claassenia sabulosa shown in Figure 32 in exhibiting a specialised attachment surface consisting of a hairy median longitudinal unscelerotised band. The claws of P. frontalis lack setae. However, there are two setae on the arolium plantar surface and as many as four setae on the orbicula."
Newell,RL; Baumann,RW and Stanford,JA 2008 Stoneflies of Glacier National Park and Flathead River basin, Montana. International Advances in the ecology, zoogeography, and systematics of mayflies and stoneflies. University of California Publications in Entomology, Berkeley and Los Angeles, pp.173-186. PDF
Quote from page 176: "Stoneflies recorded from hyporheic habitats (pumped wells) included: Alloperla severa, Capnia confusa, Claassenia sabulosa, Diura knowltoni, Hesperoperla pacifica, Isocapnia crinita, I. grandis, I. integra, I. vedderensis, Isoperla fulva, Kathroperla, Paraperla frontalis, and P. wilsoni"
Radford,DS and Hartland-Rowe,R 1971 Emergence patterns of some Plecoptera in two mountain streams in Alberta. Canadian Journal of Zoology, 49(5), 657-662.
Sandberg,JB and Stewart,KW 2003 Continued studies of drumming in North American Plecoptera; Evolutionary implications. In: Research Update on Ephemeroptera and Plecoptera. Ed: Gaino,E University of Perugia, Perugia, Italy, 73-81.
PDF
Has the sonogram of a three-way duet between two males and a female P. frontalis from the San Miguel River near Placerville over in San Miguel County, Colorado.
South,EJ; Skinner,RK; DeWalt,RE; Davis,MA; Johnson,KP; Teslenko,VA; Lee,JJ; Malison,RL; Hwang,JM; Bae,YJ and Myers,LW 2021 A new family of stoneflies (Insecta: Plecoptera), Kathroperlidae, fam. n., with a phylogenomic analysis of the Paraperlinae (Plecoptera: Chloroperlidae). Insect Systematics and Diversity, 5(4), p.1. PDF
Table 1 indicates a P. frontalis specimen was used in phylogenetic analyses.
Stanford,JA; DelVecchia,AG; Giersch,JJ and Malison,RL 2024 Amphibitic stoneflies (Plecoptera) are integrators of ecosystem processes in alluvial aquifers of gravel-bed river floodplains. Wiley Interdisciplinary Reviews: Water, p.e1720. https://doi.org/10.1002/wat2.1720
Abstract: "Over 50 years ago nymphs of the Plecoptera species, Paraperla frontalis Banks, 1906 (Plecoptera: Chloroperlidae), were shown to exist in a shallow floodplain aquifer of the Tobacco River, a gravel-bed river in western Montana and later they were documented throughout the main stems of the Flathead River system. Nymphs are almost never found in surface waters, until they emerge on the river shorelines. As teneral adults, they mate and subsequently deposit fertilized eggs into the river. This novel life cycle is termed "amphibitic". Over the years we and others have found P. wilsoni Ricker, 1965 (Plecoptera: Chloroperlidae), Kathroperla perdita Banks, 1920 (Plecoptera: Kathroperlidae), and five species of Isocapnia Banks, 1938 (Plecoptera: Capniidae), including long-winged and brachypterous adults and wingless dwarfs (male and female), occupying amphibitic niches in the alluvial aquifers of rivers in Washington, Idaho, Colorado, Alaska and British Columbia. These stoneflies are remarkably tolerant of hypoxia which allows them to exist as abundant consumers in aquifer food webs subsidized by ancient methane. Indeed, stonefly tissues contain carbon that is up to 7000 years old, underscoring the existence of a strong interaction involving the uptake of labile carbon derived from methanogenic and methanotrophic process in aquifers. Details of life cycles, trophic relationships, distribution and abundance have been documented by a suite of studies on the Nyack Floodplain of the Middle Flathead River, Montana. In this paper we review the ecophysiology and ecology of these unique stoneflies in the context of their functional role in gravel-bed river ecosystems."
Stanford,JA and Gaufin,AR 1974 Hyporheic communities of two Montana rivers. Science 185:700-702. PDF
The authors report P. frontalis from the hyporheic zone of the Flathead River in Montana, USA. Figure 1 is a photo of P. frontalis nymphs from the hyporheic zone of the Tobacco River. From nymph collections, they report that P. frontalis spends 2 or 3 years in the hyporheic zone growing to nearly 2cm in length before emerging as adults.
Stanford,JA and Ward,JV 1988 The hyporheic habitat of river ecosystems. Nature, 335(6185), pp.64-66. PDF
Stanford,JA and Ward,JV 1993 An ecosystem perspective of alluvial rivers: connectivity and the hyporheic corridor. Journal of the North American Benthological Society, 48-60.
Stark,BP; Baumann,RW; Kondratieff,BC and Stewart,KW 2013 Larval and egg morphology of Paraperla frontalis (Banks 1902) and P. wilsoni Ricker 1965 (Plecoptera: Chloroperlidae). Illiesia 9(08):101-108. PDF
Abstract:Eggs and larvae of Paraperla frontalis (Banks) and P. wilsoni Ricker are described from scanning electron micrographs. The eggs of both species are coarsely punctate over the entire chorion except for a narrow ring which surrounds the sessile collar, however, punctations are larger and more conspicuous on P. frontalis eggs. Larvae of P. wilsoni are similar to those of P. frontalis, but lack the lacinial pecten row and vertical cercal fringe found in the latter species.
Quote from page 106: "Egg. Outline oval (Fig. 1). Length ca. 305 µm, equatorial width ca. 227 µm. Collar sessile, surrounded by a smooth circular zone ca. 43 µm wide forming a circular plaque-like structure with diameter of ca. 107 µm (Figs. 2, 3). Chorion covered throughout (except smooth collar zone) with irregularly sized pits; larger pits, ca. 6.5 µm in diameter, give a coarse aspect to chorionic surface. Micropylar row equatorial, orifices without rims (Fig. 4).
Larva. Body length pre-emergent specimens 15-17 mm. General color pale brown without distinctive pigment pattern. Body covered with thin clothing hairs and short, thick setae, usually restricted to posterior segmental fringes and lateral clusters. Posterior fringes of abdominal sterna interrupted mesally except on sternum 10. Lacinia with two teeth, 2nd much smaller than 1st; and not reaching mid length of larger tooth (Figs. 7-8); a prominent row of ca. 30 spine-like pecten comprise a comb-like structure extending from base of 1st lacinial tooth, along midline of tooth to beyond base of 2nd tooth (Figs. 9-10). Basal cercal segments with terminal whorl of setae, but mid and apical segments bear a dense vertical fringe of long, fine swimming hairs (Figs. 13-15)."
Stewart,KW and Oswood,MW 2006 The Stoneflies (Plecoptera) of Alaska and Western Canada. In The Caddis Press, Columbus, Ohio. vi. + 325.
Stewart,KW and Ricker,WE 1997 The stoneflies of the Yukon. pgs 201-222 in Danks,HV and Downes,JA (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp.
Quote about the genus Paraperla from page 213: "Western Nearctic; the 2 species occur from the Yukon to California and New Mexico. Adults are brown, 16-20 mm. They emerge March-August, depending on elevation and latitude. Nymphs occur in the hyporheal beneath and beside larger streams, and are collected in surface layers only just prior to emergence. Nymph sizes before and after emergence suggest 2-3 year, semivoltine life cycles. "
Stewart,KW and Stark,BP 2002 Nymphs of North American Stonefly Genera. 2nd edition The Caddis Press, Columbus, Ohio. 510 pages. Illustrations of nymph on pages 266-267, figures 11.11-11.12
Stewart,KW and Zeigler,DD 1984 Drumming behavior of twelve North American stonefly (Plecoptera) species: First descriptions in Peltoperlidae, Taeniopterygidae and Chloroperlidae. Aquatic Insects. 6(1) 49 - 61. Abstract
Part of abstract: "Both chloroperlids, Kathroperla perdita and Paraperla frontalis, had di-phasic male calls, with 6.1 ± 0.7 and 74 ± 10.6 beats, respectively, in the first phases, and with distinctly different beat intervals. "
Zuellig,RE; Heinold,BD; Kondratieff,BC and Ruiter,DE 2012 Diversity and Distribution of Mayflies (Ephemeroptera), Stoneflies (Plecoptera), and Caddisflies (Trichoptera) of the South Platte River Basin, Colorado, Nebraska, and Wyoming, 1873-2010. U.S. Geological Survey Data Series 606, 257 p. PDF - caution 46MB
Quote from page 42: " Larvae of this species distinctly are hyporheic and rarely are encountered until near adult emergence; adults also are rarely collected (DeWalt and Stewart, 1995)." The elevation range is 5,200-10,750 feet and the adults emerge from July-August.
Zwick,P 1973 Insecta: Plecoptera Phylogenetisches System und Katalog. Das Tierreich - Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen., 94, 465 pp.
Zwick,P 2006 New family characters of larval Plecoptera, with an analysis of the Chloroperlidae: Paraperlinae. Aquatic Insects, 28:13-22.
Abstract: "Larval structures diagnostic of several Plecoptera families, subfamilies or genera are described and illustrated. They concern the following: distinction of larval Leuctridae from Capniidae; an additional apomorphy of Nemouridae; a synapomorphy of Perlidae and Perlodidae; monophyly of each of the two subfamilies of Chloroperlidae, and the generic limits among Chloroperlidae: Paraperlinae. Paraperla lepnevae Zhiltzova is transferred to Utaperla. Some comments on Notonemouridae are included."
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