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Plecoptera

Stoneflies

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Plecoptera wings
© John Pickering, 2006-2009
Plecoptera wings
Plecoptera
© John Pickering, 2006-2009
Plecoptera

Plecoptera
© John Pickering, 2006-2009
Plecoptera
Kinds

Overview
Click here for Stonefly checklist in the Great Smoky Mountains.

Plecoptera (stoneflies) are a small order of insects of about 2000 species worldwide, with their long but fragmented fossil record dating back to the early Permian. The living suborders, Arctoperlaria and Antarctoperlaria, easily contain the earliest fossils. The more modern fossils are easily indentifiable from Miocene (38 - 54 MYA) fossils. -- (University of Texas)


Identification
Stoneflies can be easily recognized by a few simple characters. They have three segmented tarsi but their hind legs are not modified for jumping to the extent of Orthoptera (crickets and grasshoppers). Their filiform antennae are at least half the length of the body. The cerci are generally long as well, especially in the aquatic nymphs. The wings are almost always present but are sometimes very short. They are folded horizontally back over the body. These characters help distinguish them from Dermaptera and Embioptera which they superfically resemble and to which they are probably closely related. The immatures are variously called nymphs or naiads, but are most frequently referred to as nymphs. All nymphs are aquatic, and resemble the adults in many respects. They also have three-segmented tarsi. The nymphs always have long cerci and never a third central tail or median caudal filament. Gills, if they have them, can occur on various parts of the thorax and abdomen and are composed only of filiments, not plates.-- (University of Texas).

  • Immatures
    • Antennae long, filiform
    • Body flattened, legs widely separated
    • Tracheal gills present as "tufts" behind the head, at base of legs, or around the anus
    • Each segment of thorax is covered by a large dorsal sclerite
    • Cerci long, multi-segmented
  • Adults
    • Antennae long, filiform
    • Front wings long and narrow; M-Cu crossveins form distinctive boxes near center of front wing
    • Hind wings shorter than front wings; basal area of hind wing enlarged and pleated
    • Cerci long, multi-segmented-- (N.C. State University Entomology Dept.)

Phylogeny
Taxonomic Category Scientific Name Common Name
Phylum Arthropoda Arthropods
Class Insecta Insects
Order Plecoptera Stonefly

    The name Plecoptera, refering to the pleated hind wings which fold under the front wings when the insect is at rest, is derived from the Greek "pleco" meaning folded and "ptera" meaning wing. -- (N.C. State University Entomology Dept.)


Suborder HOLOGNATHA
  • Capniidae - small winter stoneflies
  • Leuctridae - rolled-winged stoneflies
  • Nemouridae - spring stoneflies
  • Peltoperlidae - roachlike stoneflies
  • Pteronarcidae - giant stoneflies
  • Taeniopterygidae - winter stoneflies
Suborder SYSTELLOGNATHA
  • Chloroperlidae - green stoneflies
  • Isoperlidae - green-winged stoneflies
  • Perlidae - common stoneflies
  • Perlodidae - perlodid stoneflies


Geographic distribution
Common in and around fast-moving streams in temperate and boreal climates.

North America Worldwide
Number of Families 10 15
Number of Species 465 2000


Natural history
Stoneflies probably represent an evolutionary "dead end" that diverged well over 300 million years ago and are regarded as the earliest group of Neoptera. Immature stoneflies are aquatic nymphs (naiads) that usually live under stones in fast-moving, well-aerated water. Oxygen diffuses through the exoskeleton or into tracheal gills located on the thorax, behind the head, or around the anus. Most species are herbivorous, feeding on algae and other submerged vegetation, but two families (Perlidae and Chloroperlidae) are predators of mayfly nymphs (Ephemeroptera) and other small aquatic insects. Adult stoneflies emerge from their streams and can generally be found on the banks next to their previous habitat. They are not active fliers and usually remain near the ground where they feed on algae or lichens. In many species, the adults are short-lived and do not have functional mouthparts. Stoneflies are most abundant in cool, temperate climates. -- (University of Texas)

    Economic Importance:

    Stoneflies require cool, well-oxygenated water for their nymphal development and are therefore very susceptible to human abuse of water courses. Farm drainage, land clearing, impoundment of water courses all of which cause changes in temperature and substrate content can eliminate stoneflies from a habitat. -- (University of Texas). Because of stoneflies' impact susceptibility, they are used by ecologists as indicators of water purity. Stoneflies are also an important source of food for game fish (e.g., trout and bass) in cold mountain streams.-- (N.C. State University Entomology Dept.)


How to encounter
With a few exceptions in the southern hemisphere living on damp soil, stonefly nymphs dwell in aquatic habitats. The preferred habitat is rocky streams with at least a noticeable current. Some species can live in damp sandy areas. Although we know much less about lakes as habitats than rivers and streams, lakes can provide suitable habitat in the north and at high latitudes.The usual habitat in running water contains rocky, stony, or gravel substrata with more diversity in cooler, swifter water. Some studies have shown a correlation between certain species and certain habitats. For instance, Perlidae and Perlodidae are usually encountered under large stones, while Chloroperlidae tend to occur in gravel and Pteronarcyidae are frequently found in leaf packs.-- (University of Texas).


Links to other sites

Acknowledgements
This page written by Michael Howell, Ecology major at the University of Georgia, Athens.
Thanks to Sabina Gupta, Denise Lim, and Dr. John Pickering for technical and web support in developing this page.

1. http://www.cals.ncsu.edu/course/ent425/compendium/stonef~1.html
2. http://www.sbs.utexas.edu/jcabbott/courses/bio321web/labs/plecoptera/Plecoptera.pdf



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Order - PLECOPTERA
(Greek, plekein = to fold; pteron = wing)
Common Name: stoneflies
Distribution: Cosmopolitan

Description
Stoneflies look a bit like skinny cockroaches, but they do not have the head covered by the pronotum, and they are found near streams and rivers. Their membranous wings are held either flat over the body or rolled around the body like a cigar.

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A stonefly photographed near a stream in Tasmania

Order - PLECOPTERA
(Greek, plekein = to fold; pteron = wing)
Common Name: stoneflies
Distribution: Cosmopolitan

Description
Stoneflies are soft bodied insects that generally have two unequal pairs of membranous, intricately veined wings. However, there are some wingless species. The forewings are smaller than the hind wings and are often longer than the abdomen of the insect. Adult insects have large compound eyes with two or three simple eyes (ocelli). The antennae are long and composed of short, straight segments (filiform). The adult stage has mandibles but these are often reduced. In some species they may be functionless as the adult insect has not been observed to feed. Generally, there are two appendages at the end of the abdomen (cerci).


Naiad
The juveniles emerge from the eggs as miniature replicas of the adults (nymphs) but without wings. Nymphs mature through a series of moults, sometimes as many as 33. The nymphs are aquatic and possess gills as a fringe along the abdomen. The nymphs crawl about over the beds of fast flowing streams, although they can swim if necessary. Stoneflies are very quickly affected by pollution as they are unable to tolerate even minor levels of contaminants.


Members
Stoneflies.


Food
The nymphs may feed on plants, animals and detritus, or be restricted solely to plants. Some nymphs are predators on small aquatic life. Adults feed on algae, lichens, plants or even rotten wood.


Importance
No known importance apart from their presence in the food-chains of freshwater communities. The stoneflies may be useful components in fish food-chains.

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American Stonefly Web Page

American Stonefly Web Page

Stoneflies form the insect order Plecoptera. Adults are typified by a pair of tail-like cerci, two pairs of soft folded wings over the abdomen, well developed antennae, three tarsal segments, two tarsal claws, and an elongated anal region on the hindwings. (fig.1) Nymphs (fig. 2) are usually found in relatively clean streams, particularly those in mountainous reigions. More than 600 species occur in North America, a current North American Stonefly List is available on this website.

Nine families of Stoneflies occur in North America.

Capniidae


Capniidae are winter-emerging Stoneflies . Usually, they are small (less than 10 mm in length), black, and short-winged individuals are often seen. Adults have multisegmented cerci and small wedge-shaped mid-tarsal segments. They are often collected walking on snow near streams or they may be found on bridges, rocks, or streamside vegetation. Allocapnia (fig. 3) is common in eastern North America and Capnia (fig. 4) is common in the west. Dr. Riley Nelson , University of Texas-Austin , Department of Zoology, is an authority on the systematics of the Capniidae.

Leuctridae


Leuctridae are small dark stoneflies, generally similar to the Capniids. They differ in having 1 segmented cerci and the wings are rolled around the sides of the body. Emergence is somewhat variable, but most species are spring or summer emergers; a smaller number of primarily southern and midwestern species emerge in the fall or winter. Leuctra (fig.5 ) is the most common genus in eastern North America. Megaleuctra (fig.6 ) is an uncommon group of large, spring seep-inhabitating, Leuctrids found in the Appalacians, northern Rocky Mountains, Cascades, Sierras, and northern Coast ranges.

Nemouridae


Nemouridae are small brown stoneflies with 1-segmented cerci. The Wings are held flat over the abdomen and most species have a distinctive "X" pattern of venation near the wing apex. Amphinemura is a common North American genus, and a few other Nemouridae genera such as Zapada (fig. 7 ) , have ventral gills in the neck region. Nemouridae are typically spring emerging stoneflies. Dr. Richard Baumann (mail), Brigham Young University , Department of Zoology, is an authority on Neumorid systematics.

Taeniopterygidae


Taeniopterygidae is a group of medium sized, dark bodied stoneflies. Emergence times range from winter through spring for most species. Males have 1-segmented cerci but females have short multi-segmented cerci. Adults are usually recognized by their relatively large mid-tarsal segment which is about the same size as the basal segment. Taeniopteryx (fig. 8) is a common eastern and northern genus and Taenionema (fig. 9) is the common western group.

Chloroperlidae


Chloroperlidae is a group of small to medium size, mostly pale yellow or green stoneflies. Most species are spring or summer emerging and adults are often collected while sweeping streamside vegetation during their emergence season. Adults have a reduced anal area to the hindwings. Triznaka (fig. 10 ) and Sweltsa (fig.11 ) are common genera. Dr. Rebecca Surdick, Laboratory of Entomology, Winchester, Virginia is an authority on this group. Mary Leigh Lyon and Mark Lyon are currently studying the epiproct complex for several Alloperla (fi g. 12 ) species found in western North America and southeastern North America respectively. Kevin Alexander is currently working with Dr. Ken Stewart, University of North Texas , Depar tment of Biology on Suwallia

Peltoperlidae


Peltoperlidae is a small group of medium sized stoneflies found primarily in the mountains of eastern North America and the west. These stoneflies are somewhat roachlike in appearance and they emerge in spring and summer. Ad ults have only two ocelli and the labial palps are short and are usually not visable from above. Tallaperla (fig. 13 ) is a common peltoperlid found in streams of the southern Appalachians. Dr. Bill Stark , Mississippi College , Biology Department is an authority on this group.

Perlidae


Perlidae is a family of mostly medium or large summer-emerging stoneflies. Nymphs have branched gills on their thoracic segments and the adults often retain these as vestiges around the leg bases. The front femora of adult perlid s bears an anterovenural row of long bristles. Acroneuria (fig. 14 ) is a common large perlid of the east while Hesperoperla (fig.15 ) is common in the west. Dr. Bill Stark , Mississippi College , Biology Department is an authority on this group.

Perlodidae


Perlodidae includes stoneflies that are similar to perlids in general apperance but most lack gill vestiges and long bristles are absent from the front femora. Perlodids usually emerge in spring. Isoperla (fig.1 6 ) is a common North American Perlodid genus and Frisonia (fig.17 ) is found in the pacific northwest. Dr. Boris Kondratieff , Colorado State University , Department of Entomology and Dr. Stan Szczytko , Universit y of Wisconsin-Stevens Point , Colllege of Natural Resources are authorities on this group.

Pteronarcyidae


Pteronarcyidae include the giant salmonflies famous among flyfishers. Adults are usually recognized by the presence of two or more cross vein rows in the forewing anal area. Pteronarcys (fig. 18 ) is fou nd in both eastern and western North America. Dr. Charlie Nelson(mail) , University of Tennessee-Chattanooga , Department of Biology, is an authority on this group.

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Representatives of Stonefly Families (Plecoptera)


Tips for identification - Stoneflies look similar to certain flattened mayflies and many beginners have trouble distinguishing the groups. The relative length of the antennae may be used to tell flattened mayflies from stoneflies. Stoneflies have antennae that are many times, four to six times, longer than the width of their head, while those of flattened mayflies are only about twice the width of their head.

Here are some other tips.

  • stoneflies always have two elongate tails
  • head with long slender antennae
  • each leg ends in two claws
  • length one quarter inch to two inches excluding antennae and tails

Family Image Details
Perlidae photo of plecoptera
Perlidae photo of plecoptera Head with variable color
patterns
Perlidae photo of plecoptera Finely branched gills only on sides of
thoracic segments, never extending to
abdominal segments
Perlodidae photo of plecoptera Tails equal in length or longer than
abdomen
Perlodidae photo of plecoptera No branching gills behind base of legs
Perlodidae photo of plecoptera Metathoracic wingpads strongly diverging
from body
Taeniopterygidae photo of plecoptera
Taeniopterygidae photo of plecoptera With single gill at base of each leg
or with triangular plate underneath
next to last segment of abdomen
Taeniopterygidae photo of plecoptera
Taeniopterygidae photo of plecoptera Tarsal segments 1 and 2
about same length
Nemouridae photo of plecoptera Extended hind legs reaching
approximately to end of abdomen
Nemouridae photo of plecoptera Tarsal segment 2 much
shorter than 1
Pteronarcyidae photo of plecoptera Body black
Nemouridae photo of plecoptera Some branching gills
on ventral abdominal
segments
Peltoperlidae photo of plecoptera Thorax with large dorsal plates
Peltoperlidae photo of plecoptera Thoracic sternal plates shield like,
overlapping succeeding
segment
Chloroperlidae photo of plecoptera Nota usually without pigmented pattern.
Tails 3/4 or less than length of abdomen.
Abdominal terga widest posteriorly.
Chloroperlidae photo of plecoptera
Chloroperlidae photo of plecoptera
Capniidae photo of plecoptera

Abdominal terga widest posteriorly.

(This family is very difficult to separate from the family Leuctridae )

Capniidae photo of plecoptera Head
Capniidae photo of plecoptera Paraglossa and glossa
about equal in length.
Leuctridae photo of plecoptera Abdominal segments essentially
parallel-sided

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Order Plecoptera

(Stonefly)

Soil & Water Conservation Society of Metro Halifax ( SWCSMH )

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July 26, 2006       Freshwater Benthic Ecology and Aquatic Entomology Homepage


Contents:



Linnean system of hierarchical classification ( Williams & Feltmate, 1992 ):

  • Superphylum Arthropoda
    • (jointed-legged metazoan animals [Gr, arthron = joint; pous = foot])
  • Phylum Entoma
  • Subphylum Uniramia
    • (L, unus = one; ramus = branch, referring to the unbranched nature of the appendages)
  • Superclass Hexapoda
    • (Gr, hex = six, pous = foot)
  • Class Insecta
    • (L, insectum meaning cut into sections)
  • Subclass Ptilota
  • Infraclass Neopterygota


Introduction

The Plecoptera (stoneflies), all of which are aquatic as nymphs, are considered to be the most primitive order of living Neoptera. Plecopterans number about 1718 species in 239 genera belonging to 15 families. Nymphs feed on fresh or decayed vegetable matter, but may be carnivorous in later instars.

The order Plecoptera belongs to the infraclass Neoptera because stoneflies' wings fold over their backs at rest. Wings develop in external wingpads, a characteristic that places Plecoptera in the division Exopterygota. North American stoneflies are generally divided into two groups, Euholognatha and Systellognatha, based on major differences in mouthpart morphology and, hence, feeding biology.

The taxonomy of this order, like that of the Ephemeroptera, is poorly known because the larvae of many species have not been associated with adults.

Modern plecopterans are thought to have been derived from the Protoperlaria of the Permian (360-286 million years B.P.) and the fossil record is quite respectable with more than 30 species described from the different strata of the Permian to the middle Tertiary.


Life History

The stoneflies are terrestrial as adults, but in the nymphal stages they are strictly aquatic, and most are restricted to flowing waters of relatively high oxygen concentrations. Fertile eggs, laid over or in the water, requite two to three weeks for hatching in many species, and several months among some larger forms. The nymphal instars, from 10 to over 30 moltings, occur in one to three years. Adults live from 1 to 4 weeks. Most adults are winged, although a few species are wingless (apterous) or have short wings (brachypterous). None fly well and this has prevented them from crossing even small geographical barriers. Thus, like the mayflies, stoneflies are useful tools in studies of historical biogeography.

Temperate species that overwinter as nymphs often do not stop growing even in water temperatures close to 0°C. It seems that it is warm water temperatures rather than cold ones that punctuate stonefly life cycles. The ability to spend the summer in diapause enables some species to live in temporary streams .


Feeding

Generally, stonefly nymphs are either shredders or predators. Some groups that are predaceous as late instars have been reported to be herbivorous or detritivorous in early instars, while late instars of large detritivores may consume some prey. Predators are engulfers, that is, they swallow their prey whole or bite off and swallow parts of prey. They are active search or pursuit predators, using their long filamentous antennae to locate prey using tactile, wave disturbance, and chemical cues. Many species are opportunistic feeders, consuming prey in proportion to their relative abundance. Other species are selective for prey species or sizes. In some families adults feed, and in others they do not.


Habitat and Ecological preference

Plecopteran nymphs are restricted to cool, clean streams with high dissolved oxygen content. some species, however, may be found along the wave-swept shores of large oligotrophic lakes. When subjected to low dissolved oxygen concentration, the nymphs of many species exhibit a characteristic "push-up" behaviour that increases the rate of movement past the gills. The gills are variously placed among species on the neck, thorax and abdomen. However, some species have no gills and respiration in these is assumed to be across the cuticle surface.

The high water quality requirements of the nymphs bars all but a very few species from habitats subject to low oxygen levels, siltation, high temperatures and organic enrichment, and this has led to their effective use as biological indicators of environmental degradation.

Field surveys clearly show that the nymphs of many species are associated with particular sections of a stream bed or lake shore. The specific microhabitat occupied depends on a variety of environmental factors such as the nature of the substratum (particle size and configuration), current regime, presence of other organisms, and local variations in water chemistry and temperature. Habitat preference often changes as the nymphs develop and with season. Prior to emergence, final instar nymphs tend to migrate towards the bank where they crawl out of the water to shed their skins.

Most stoneflies are classified as clingers or sprawlers, as they are closely associated with the substrate or leaf litter. A few species have been reported from the hyporheic zone.


Some physiological and ecological tolerances and requirements ( Mackie , 2001)

Species General habitat Feeding pH Oxygen %
Acroneuria lycorias rocks, streams predator of insects <7 - >7 approx. 100
Allocapnia spp. rocks, streams shredder >7 approx. 100
Amphinemura delosa gravel, rocks, streams gatherer, shredder <7 - 7 100
Isoperla bilineata plants, rocks, streams predator of insects, gatherer >7 100
Isoperla clio plants, streams predator of insects >7 100
Isoperla fulva plants, rocks, streams predator of insects, scraper, gatherer ≥7 50-100
Nemoura trispinosa plants, rocks, streams shredder <7 - >7 100
Peltoperla maria leaf litter, streams shredder ≥7 approx. 100
Perlesta placida rocks, leaves, streams predator of insects, gatherer >7 approx. 100
Pteronarcys spp. rocks, logs, leaves, streams predator, scraper, shredder ≥7 approx. 100
Taeniopteryx maura rocks, logs, leaves, streams gatherer, shredder <7 - >7 approx. 100

References and web URLs:

  • Hutchinson, G.E. 1993. A Treatise on Limnology. Vol. IV, The Zoobenthos. Ed. Y.H. Edmondson. John Wiley & Sons, Inc. Xx, 944pp.
  • Mackie, G.L. 2001. Applied Aquatic Ecosystem Concepts. Kendall/Hunt Publishing Company. xxv, 744 pp. ISBN 0-7872-7490-9
  • Narf, R. 1997. Midges, bugs, whirligigs and others: The distribution of insects in Lake "U-Name-It". Lakeline. N. Am. Lake Manage. Soc. 16-17, 57-62.
  • Peckarsky, B.L., P.R. Fraissinet, M.A. Penton, and D.J. Conklin, Jr. 1990. Freshwater Macroinvertebrates of Northeastern North America. Cornell Univ. Press. xii, 442pp.
  • Wetzel, R.G. 1983. Limnology. 2nd ed. Saunders College Publishing. Xii, 767pp, R81, I10.
  • Williams, D.D., and Feltmate, B.W. 1992. Aquatic Insects. CAB International. ISBN: 0-85198-782-6. xiii, 358p.




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