Taxonomy Notes
Sexual dimorphism and morphology
Notes on Plecoptera
Plecoptera Taxa List for North America Stonefly Emergence Patterns
NRE 516 Taxonomic List for Plecoptera

Principal Taxonomic Literature:

Claassen, P. W. 1934. Plecoptera Nymphs of America north of Mexico. Entomological Society of America. Thomas Say Foundation 3.
Frison, T. H. 1935. The stoneflies, or Plecoptera, of Illinois. Bulletin of the Illinois Natural History Survey 20:281-471. Also see his two updates: Frison, T. H. 1937. Descriptions of Plecoptera, with special reference to the Illinois species. Bulletin of the Illinois Natural History Survey 21(3):78-99, and Frison, T. H. 1942. Studies of North American Plecoptera, with special reference to the fauna of Illinois. Bulletin of the Illinois Natural History Survey 22:235-355.
Hitchcock, S. W. 1974. Guide to the Insects of Connecticut. Part VII. The Plecoptera or Stoneflies of Connecticut. State Geological and Natural History Survey of Connecticut, Bulletin No. 107. 262pp.
Illies, J. 1965. Phylogeny and zoogeography of the Plecoptera. Annual Review of Entomology 10: 117-140.
Stewart, K. W., and B. P. Stark. 1988. Nymphs of North American Stoneflies. Entomological Society of America. Thomas Say Foundation 12.
Zwick, P. 1973. Insecta: Plecoptera. Phylogenetisches System und Katalog. Das Tierreich 94. Walter de Gruyter, Berlin.
Zwick, P. 2000. Phylogenetic system and zoogeography of the Plecoptera. Annual Review of Entomology 45:709-746.

Major taxonomic classification is taken from Illies (1965) as presented in Stewart and Stark (1988):

Order Plecoptera
Group Euholognatha (Suborder Filipalpia sensu strictu)
    Family Capniidae
    Family Leuctridae
    Family Nemouridae
    Family Taeniopterygidae
Group Systellognatha (Suborder Filipalpia sensu latu)
    Family Peltoperlidae
    Family Pteronarcyidae
Group Systellognatha (Suborder Setipalpia sensu strictu)
    Family Perlidae
    Family Perlodidae
    Family Chloroperlidae

Zwick (1973) recognizes two suborders, the Arctoperlaria (northern hemisphere stoneflies) and Antarctoperlaria (southern hemisphere stoneflies), the former divided into two groups, Euholognatha and the Systellognatha.

Some recent synonymies for taxa found in and near Michigan:

• Agnetina (Perlidae). Considered valid for species previously assigned to Phasganophora or Neophasganophora (Zwick 1984)
• Capnura = Capnia (part)
• Chloroperla (Chloroperlidae). Name applied by Banks to a variety of species currently placed in several general, including Haploperla, which is found in Michigan
• Clioperla = Isoperla (part)
• Haploperla = Hastaperla
• Helopicus (Perlodidae). Valid name for four eastern Nearctic species formerly placed in Isogenus and Hydroperla.
• Isogenus (Perlodidae). Formerly used to include North American species now placed in Cultus and Isogenoides.

Refer to Stewart and Stark (1988: 430-436) for additional information about generic-level nomenclatural changes for the North American taxa .

Sexual dimorphism and morphology in adult stoneflies

Males are separated from females by the presence of hooks or projections of the genitalia apparatus, which are generally visible dorsally. The female dorsum is unornamented. Ventrally, some males have a lob or knob on a posterior abdominal segment, whereas the female commonly has one abdominal sternite (usually the 8th) expanded.

Wing venation follows the basic insect pattern. Most species have four membranous wings: the front wings are elongate and rather narrow and usually have a series of cross veins between M and Cu 1 and between Cu 1 and Cu 2 ; the hind wings are slightly shorter than the front wings and usually have a well-developed anal lobe that is folded fan-wise when the wings are at rest. A few species of stoneflies have the wings reduced or absent in the male. Stoneflies at rest hold the wings flat over the abdomen.

Currently 578 species of Plecoptera in 100 genera and 9 families (Table 2) are recognized for North America north of Mexico. They represent about 27% of the known world fauna, but this percent-age should decline as Oriental and Neotropical faunas become better known. Only the Oriental realm, with 662 species, exceeds the Nearctic in numbers. Illies (1966), Zwick (1973), and Stark et al. (1986) have provided catalogue information for world species. The North American stoneflies show relatively strong biogeographic affinities with the Palearctic and Oriental realms. We estimate that another 50 North American species await description primarily in the genera Isoperla, Perlesta, Capnia, and Leuctra. California has recently been, and is anticipated to continue to be, the site of major discoveries at both the generic and species level. Unlike the mayflies, all species of stoneflies are known in the adult stage. Nearly all species are known from both sexes (3% from one sex). Some 45% of North American species are known as larvae (nymphs) (Stewart & Stark 1988), although not necessarily formally described. Considerable effort should be placed on rearing, associating, and describing larvae. Placing larvae within a generic classification will be aided by the new work of Stewart & Stark (1988). Larval keys should soon be available for Acroneuria, Neoperla, Isogenoides, and Isoperla. The eggs of 205 species are known, mostly from the Systellognatha. Careful descriptions or redescriptions of genitalia of most species are needed to document ultra-structure. Nelson & Baumann (1987), Stark & Szczytko (1988), and Stark (1989) demonstrated their value in the Capniidae, Perlodidae, and Perlidae. Keys to species are critically needed for far western US and Canada as well as eastern North America. Works by Jewett (1959, 1960) for the West and Hitchcock (1974) for the East are considerably out-of-date. Adults and known larvae of Rocky Mountain species are usually identifiable with the keys of Baumann et al. (1977). An overview of the status and need of revisionary work on the 9 North American families follows:

1) Capniidae -- C. R. Nelson is conducting research and beginning to examine world genera as an approach to revising polyphyletic genera;
2) Leuctridae -- Genera are relatively well known, but a new genus is thought to occur in California, and certain species revisions are needed;
3) Nemouridae -- Genera are well known (Baumann 1975), but species revisions are required for some, which are being studied by R. W. Baumann;
4) Taeniopterygidae -- Ricker & Ross (1968, 1975) established the foundation for understanding this group; revisions are being carried out by J. A. Stanger & R. W. Baumann, and extensive stage correlations are being conducted by K. W. Stewart & J. A. Stanger on western species and by R. F. Kirchner on eastern species; some unpublished electrophoretic work has also been conducted by D. H. Funk;
5) Chloroperlidae -- A family revision was recently completed by Surdick (1985), and species-level revisions are now needed for larger genera;
6) Peltoperlidae -- Stark & Stewart (1981) reviewed the Nearctic genera, and Stark (1983a,b) and Stark & Kondratieff (1987) have revised larger genera;
7) Perlidae -- Stark & Gaufin (1976) reviewed Nearctic genera. The greatest problems remain in the genus Perlesta. Stark (unpublished) has reviewed the types, and B. C. Poulton and K. W. Stewart are working on the Ozark-Ouachita species;
8) Perlodidae -- Stewart & Stark (1984) described known larvae, and Stark & Szczytko (1984) used egg morphology to revise trivial classification; certain western species were revised by Szczytko & Stewart (1979), and Szczytko is now revising eastern species; new genera continue to be discovered in isolated areas of California;
9) Pteronarcyidae -- Stark & Szczytko (1982) have studied the eggs, and Nelson (1988) has presented a species phylogeny; revised keys to adults and larvae are needed.

With respect to special stonefly habitat considerations, spring seeps are probably the most critical because they have been historically overlooked by many collectors. As a result, they contain a high percentage of species that are not well known or that could be considered rare. Also, the relatively small insulated habitats provided by spring seeps and their usual patchy geographic distribution provide an ideal situation for studying geographic speciation and adaptive radiation in certain groups of stoneflies (e.g., Ross & Ricker 1971). Several genera have been named from spring seeps in recent years, and several other "rare" genera are also found in this habitat. Unfortunately, spring seeps are often aesthetically pleasing, and thus are frequently enclosed by parks or private property where water use can be incompatible with stonefly survival.

The bulk of systematic research on North American stoneflies in the past 15 years has been carried out by 8 researchers in scattered localities. North Texas State University and Brigham Young University currently have the most active programs in Plecoptera systematics for Ph.D. students.

Large collections are housed at these institutions, the United States National Museum of Natural History, the California Academy of Sciences, and the Illinois Natural History Survey. A computer file of North American species and distributions is maintained by B. P. Stark, S. W. Szczytko, and R. W. Baumann. Annual bibliographies are compiled by these workers and published by the North American Benthological Society. In addition, "Perla" is a regular international newsletter of Plecoptera.

Plecoptera Taxa List for North America

Alphabetical list of families and genera of extant Plecoptera known from North America north of Mexico. Numbers of currently valid nominal species in the area, and numbers known as larvae, are indicated respectively after each genus.

Family Capniidae
Allocapnia 41,10 Capnura 7,1 Mesocapnia 13,2 Paracapnia 3,2
Bolshecapnia 6,1 Eucapnopsis 1,1 Nemocapnia 1,1 Utacapnia 10,3
Capnia 55,4 Isocapnia 11,6

Family Leuctridae
Despaxia 1,1 Megaleuctra 6,1 Paraleuctra 9,2 Zealeuctra 8,1
Leuctra 25,8 Moselia 1,1 Perlomyia 2,2

Family Nemouridae
Amphinemura 10,5 Nemoura 5,2 Podmosta 5,3 Soyedina 7,2
Lednia 1,1 Ostrocerca 6,2 Prostoia 4,3 Visoka 1,1
Malenka 11,3 Paranemoura 1,1 Shipsa 1,1 Zapada 9,5

Family Taeniopterygidae
Bolotoperla 1,1 Oemopteryx 4,3 Taenionema 8,3 Taeniopteryx 12,11
Doddsia 1,1 Strophopteryx 7,2

Family Chloroperlidae
Alloperla 29,4 Kathroperla 2,1 Plumiperla 2,1 Sweltsa 21,4
Bisancora 2,1 Neaviperla 1,1 Rasvena 1,1 Triznaka 2,2
Chloroperla 1,0 Paraperla 2,1 Suwallia 5,2 Utaperla 2,2
Haploperla 4,2

Family Peltoperlidae
Peltoperla 2,1 Soliperla 6,4 Viehoperla 1,1 Yoraperla 2,2
Sierraperla 1,1 Tallaperla 6,1

Family Perlidae
Acroneuria 12,9 Beloneuria 3,2 Eccoptura 1,1 Paragnetina 5,4
Agnetina 3,3 Calineuria 1,1 Hansonoperla 1,1 Perlesta 2,2
Anacroneuria 2,1 Claassenia 1,1 Hesperoperla 2,2 Perlinella 3,3
Attaneuria 1,1 Doroneuria 2,2 Neoperla 13,1

Family Perlodidae
Arcynopteryx 1,1 Diploperla 4,3 Kogotus 2,2 Pictetiella 1,1
Baumannella 1,1 Diura 3,3 Malirekus 2,2 Remenus 1,1
Calliperla 1,1 Frisonia 1,1 Megarcys 5,1 Rickera 1,1
Cascadoperla 1,1 Helopicus 4,3 Oconoperla 1,1 Salmoperla 1,1
Chernokrilus 3,1 Hydroperla 3,3 Oroperla 1,1 Setvena 3,3
Clioperla 1,1 Isogenoides 9,8 Osobenus 1,1 Skwala 2,2
Cosumnoperla 1,1 Isoperla 57,38 Perlinoides 1,1 Yugus 2,2
Cultus 6,3

Family Pteronarcyidae
Pteronarcella 2,2 Pteronarcys 8,8

Stonefly Emergence Patterns

Some common stoneflies from the Au Sable River Drainage basin with dates of emergence.
______________________________________________________________________________________
Taxa                             Jan     Feb       Mar     April     May     June     July     Aug     Sept
______________________________________________________________________________________

Taeniopteryx maura                              xxxxxxxxxx
Allocapnia pygmaea                             xxxxx
Paracapnia opis                                   xxxxxxxxxxxxx
Nemoura rotunda                                             xxxxxxxxx
Nemoura completa                                                   xxxxxxxx
Isoperla slossonae                                                   xxxxxxxxxx
Isoperla (Hydroperla) olivacea                                    xxxxxxx
Isoperla bilineata                                                      xxxxxxx
Isoperla signata                                                        xxxxxxxxxxx
Cultus decisa                                                              xxxxxxxxxxx
Isoperla confusa                                                          xxxxxx
Nemoura venosa                                                           xxxxxxxxxxx
Isoperla lata                                                                 xxxxxxxxx
Acroneuria carolinensis                                                   xxxxxxx
Isoperla transmarina                                                       xxxxxxxxxxxxxxxx
Nemoura trispinosa                                                         xxxxxxxxxxxxxx
Isoperla truncata                                                             xxxxxxxxxxxxxxxx
Isoperla dicala                                                                 xxxxxxxxxxxxxxxxxxxxxx
Pteronarcys dorsata                                                                             xxxxx
Haploperla brevis                                                                                    xxxxxxx
Perlesta placida                                                                                              xxxxxx

NRE 516 Taxonomic List for Plecoptera

Additional References

Baumann, R. W. 1975. Revision of the stonefly family Nemouridae (Plecoptera): a study of the world fauna at the generic level. Smithsonian Contributions to Zoology 211. Smithsonian Institution Press, Washington, DC.

Dewalt, R. E., D. W. Webb, and T. N. Kompare. 2001. The Perlesta placida (Hagen) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, and an identification key. Proceedings of the Entomological Society of Washington 103: 207-216.

Nelson, C. H. 1996. Placement of Helopicus rickeri Stark in Hydroperla Frison (Plecoptera: Perlodidae) with the description  of the adult female, nymph, and egg and a cladistic  analysis of Hydroperla. Proceedings of the Entomological Society of Washington 98: 237-244.

Poulton, B. C., and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecoptera). Memoirs of the American Entomological Society 38:1-116.

Surdick, R. F. 1985. Nearctic genera of Chloroperlinae (Plecoptera: Chloroperlidae). Illinois Biological Monographs 54. University of Illinois Press, Urbana, Illinois.

Page created: January 26, 2001
Page last edited: February 06, 2003 (EB)