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Plecoptera of Gunnison County, Colorado

Introduction to the Nemouridae

Winter Stoneflies, Tiny Early Winter Black Stonefly, Tiny Late Winter Black Stonefly, Spring Stoneflies, Nemourid Stonefly

Updated 10 January 2026
TSN 102517
This shows the classic "Neumoridae X" on the top part of the wing.

Provisional Species List

Amphinemura banksi
Malenka californica
Malenka coloradensis
Malenka flexura
Podmosta decepta
Podmosta delicatula
Prostoia besametsa
Zapada cinctipes
Zapada frigida
Zapada haysi
Zapada oregonensis

Good Links


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

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

Nymph Key, Photos and more from Troutnut

References

Allan,JD 1978 Trout predation and the size composition of stream drift. Limnology and Oceanography 23 (6) 1231-1237. PDF

Allan,JD 1987 Macroinvertebrate drift in a Rocky Mountain stream. Hydrobiologia 144, 261-268.

Baumann,RW 1975 Revision of the stonefly family Nemouridae (Plecoptera): A study of the world fauna at the generic level. Smithsonian Contributions Zoology 211, 1-74. PDF
     Abstract: "The Northern Hemisphere family Nemouridae is revised and divided into two subfamilies: Amphinemurinae, new subfamily, and Nemourinae. Of the 17 recognized genera in the Nemouridae, 14 are described and figured and 3 are described as new: Illiesonemoura, Indonemoura, and Mesonemoura; 373 species are assigned to genera or listed as incertae sedis. Keys to subfamilies and genera are given for males, females, and nymphs. Structures of male and female terminalia are described and the terminology defined. A phylogeny is constructed showing the relationships of the genera. Distributions are given for all species, the zoogeography of the family is discussed, and a composite map of the distribution of all genera in Nemouridae is provided."

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
     Quote from page 17: "The members of this family are the most common stoneflies in most habitats in the Rocky Mountains. The nymphs are small and stout bodied with numerous spines and hairs on the dorsal surface and appendages (fig. 63). The adults are easily recognized by the distinctive nemourid "X" in the forewings at the cord (figs 140-143).
This family is very important from an ecological standpoint because they are often the dominant primary consumers in flowing water ecosystems. They are detritovores and often act as shredders of heterotrophic material, such as leaves, that enter the ecosystem from outside."


Caruso,C and Wichard,W 2010 Overview and descriptions of fossil stoneflies (Plecoptera) in Baltic Amber. Entomologie Heute, (22) 85-97. PDF
     Abstract: "Three new fossil species of stoneflies (Plecoptera: Nemouridae and Leuctridae) from Eocene Baltic amber are being described: Zealeuctra cornuta n. sp., Lednia zilli n. sp., and Podmosta attenuata n. sp.. Extant species of these three genera are found in Eastern Asia and in the Nearctic region. It is very probably that the genera must have been widely spread across the northern hemisphere in the Cretaceous period, before Europe was an archipelago in Eocene. The current state of knowledge about the seventeen Plecoptera species of Baltic amber is shortly presented. Due to discovered homonymies, the following nomenclatural corrections are proposed: Leuctra fusca Pictet, 1856 in Leuctra electrofusca Caruso & Wichard, 2010 and Nemoura affinis Berendt, 1856 in Nemoura electroaffinis Caruso & Wichard, 2010."

Caruso,C and Wichard,W 2011 Paleogeographic Distribution of Leuctridae and Nemouridae genera preserved in Baltic amber, with the description of Palaeopsole weiterschani n. gen., n. sp.(Plecoptera). Entomologie heute, 23, pp.69-77.
     Abstract: " The description of the new fossil species Palaeopsole weiterschani n. gen., n. sp. adds the genus Palaeopsole to the extant six genera of the families Leuctridae and Nemouridae, which have been found in Eocene Baltic amber. These Eocene fossils are the oldest representatives of their extant genera, so far. But the Eocene European archipelago is neither place nor time in which these genera originated. The origin of these seven genera is unknown, but took very probably place in the northern hemisphere before the Eocene. After Laurasia had finally broken apart at early Jurassic, probably not till then the genera developed and spread out across the warm-temperate and subtropical northern hemisphere. The extant Nearctic genera - Lednia, Megaleuctra, and Zealeuctra, including the transberingian Podmosta, and including Leuctra and Nemoura which are nearly distributed all over the Holarctic region - spread probably across the northern hemisphere via the temporary Beringian and Turgai Strait land bridges at Cretaceous. Towards the end of Eocene the temperature decreased worldwide and forced the Nearctic and Holarctic genera to adapt to temperate and boreal climates, whereas the recent relatives of genus Palaeopsole, probably genus Rhopalopsole, continued and live in the subtropical area of the Oriental region and Southeast Asia."

Chen,ZT 2025 Three new stoneflies (Insecta: Plecoptera) from Eocene Baltic amber. European journal of taxonomy, 1026, pp.107-122. PDF
     Abstract: "Three new stonefly species are described and illustrated from Eocene Baltic amber found in Lithuania: †Palaeopsole spinosa sp. nov. (family Leuctridae Klapálek, 1905), †Podmosta biloba sp. nov. (family Nemouridae Newman, 1853), and †Isoperla lituana sp. nov. (family Perlodidae Klapálek, 1909). The generic diagnosis of †Palaeopsole Caruso & Wichard, 2011 is also emended. These new fossils enhance the understanding of the extinct Plecoptera fauna in Baltic amber."

Cummins,KW; Wilzbach,MA; Gates,DM; Perry,JB and Taliaferro,WB 1989 Shredders and riparian vegetation. BioScience, 39(1), pp.24-30. PDF

Duffield,RM and Nelson,CH 1998 Stoneflies (Plecoptera) in the diet of brook trout (Salvelinus fontinalis Mitchell) in Libby Creek, Wyoming, USA. Hydrobiologia 380, 59-65.
     Quote from page 61: "The Nemouridae were represented by five species in the stomach samples. Malenka flexura, Podmosta decepta, P. delicatula, Zapada cinctipes, and Z. haysi were recovered largely as larvae. The latter species, constituting 12.3% of the total stonefly dietary component, was the second most abundant species recovered. Adult female:male ratio of this species was approximately 1.2:1."

Gamboa,M; Muranyi,D; Kanmori,S and Watanabe,K 2019 Molecular phylogeny and diversification timing of the Nemouridae family (Insecta, Plecoptera) in the Japanese Archipelago. PloS one, 14(1), p.e0210269. PDF
     Abstract: " The generation of the high species diversity of insects in Japan was profoundly influenced by the formation of the Japanese Archipelago. We explored the species diversification and biogeographical history of the Nemouridae Billberg, 1820 family in the Japanese Archipelago using mitochondrial DNA and nuclear DNA markers. We collected 49 species among four genera: Indonemoura Baumann, 1975; Protonemura Kempny, 1898; Amphinemura, Ris 1902 and Nemoura Latreille, 1796 in Japan, China, South Korea and North America. We estimated their divergence times-based on three molecular clock node calibrations-using Bayesian phylogeography approaches. Our results suggested that Japanese Archipelago formation events resulted in diversification events in the middle of the Cretaceous (<120 Ma), speciation in the Paleogene (<50 Ma) and intra-species diversification segregated into eastern and western Japan of the Fossa Magna region at late Neogene (20 Ma). The Indonemoura samples were genetically separated into two clades-that of Mainland China and that of Japan. The Japanese clade clustered with the Nemouridae species from North America, suggesting the possibility of a colonisation event prior to the formation of the Japanese Archipelago. We believe that our results enhanced the understanding both of the origin of the species and of local species distribution in the Japanese Archipelago."

Hotaling,S; Shah,AA; Dillon,ME; Giersch,JJ; Tronstad,LM; Finn,DS; Woods,HA and Kelley,JL 2021 Cold tolerance of mountain stoneflies (Plecoptera: Nemouridae) from the high Rocky Mountains. Western North American Naturalist, 81(1), pp.54-62. PDF
     Abstract: "How aquatic insects cope with cold temperatures is poorly understood. This is particularly true for high-elevation species, which often experience a seasonal risk of freezing. In the Rocky Mountains, nemourid stoneflies (Plecoptera: Nemouridae) are a major component of mountain stream biodiversity and are typically found in streams fed by glaciers and snowfields, which are rapidly receding due to climate change. Predicting the effects of climate change on mountain stoneflies is difficult because their thermal physiology is largely unknown. We investigated cold tolerance of several alpine stoneflies (Lednia tumana, Lednia tetonica, and Zapada spp.) from the Rocky Mountains, USA. We measured the supercooling point (SCP) and tolerance to ice enclosure of late-instar nymphs collected from a range of thermal regimes. SCPs varied among species and populations, with the lowest SCP measured for nymphs from an alpine pond, which was much more likely to freeze solid in winter than flowing streams. We also show that L. tumana cannot survive being enclosed in ice, even for short periods of time (<3 h) at relatively mild temperatures (-0.5 °C). Our results indicate that high-elevation stoneflies at greater risk of freezing may have correspondingly lower SCPs, and despite their common association with glacial meltwater, these stoneflies appear to be living near their lower thermal limits."

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.
     Has brief comments on many of the stoneflies of Colorado.

Mihuc,TB; Mihuc,JR 1995 Trophic ecology of five shredders in a Rocky Mountain stream. Journal of Freshwater Ecology 10(3) 209-216. PDF

Stark,BP; Szczytko,SW; Baumann,RW 1986 North American stoneflies (Plecoptera): systematics, distribution, and taxonomic references. Great Basin Naturalist 46, 383-397. PDF

Stewart,KW and Alexander,NH 2008 The nymphs of three Nemouridae species (Plecoptera) from Oregon temporary headwaters streams. Transactions of the North American Entomological Society 134: 173-183. PDF

Stewart,KW and Stark,BP 2002 Nymphs of North American Stonefly Genera. 2nd edition The Caddis Press, Columbus, Ohio. 510 pages.

Wang,Y; Guo,C; Yue,X; Fan,X; Fan,Y and Cao,J 2024 Mitochondrial genomes of Nemourinae species (Plecoptera: Nemouridae) and the phylogenetic implications. Journal of Insect Science, 24(2), p.4. PDF
     Abstract: " Currently, the classification system of 2 subfamilies within Nemouridae has been widely accepted. However, monophyly of 2 subfamilies has not been well supported by molecular evidence. To date, only mitogenomes from genus Nemoura of the subfamily Nemourinae were used in previous phylogenetic studies and produced conflicting results with morphological studies. Herein, we analyzed mitogenomes of 3 Nemourinae species to reveal their mitogenomic characteristics and to examine genus-level classification among Nemouridae. In this study, the genome organization of 3 mitogenomes is highly conserved in gene order, nucleotide composition, codon usage, and amino acid composition. In 3 Nemourinae species, there is a high variation in nucleotide diversity among the 13 protein-coding genes (PCGs). The Ka/Ks values for all PCGs were far lower than 1, indicating that these genes were evolving under purifying selection. The phylogenetic analyses highly support Nemurella as the sister group to Ostrocerca. Meanwhile, Nemoura is recovered as the sister group of Malenka; they are grouped with other Amphinemurinae and emerged from a paraphyletic Nemourinae. More molecular data from different taxonomic groups are needed to understand stoneflies phylogeny and evolution."

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."

Brown,WS 2004 Plecoptera or Stoneflies of Gunnison County, Colorado
www.gunnisoninsects.org