insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the most primitive groups of
Neoptera
Neoptera (Ancient Greek ''néos'' (“new”) + ''pterón'' (“wing”)) is a classification group that includes most orders of the winged insects, specifically those that can flex their wings over their abdomens. This is in contrast with the mo ...
, with close relatives identified from the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
and Lower
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
geological periods, while true stoneflies are known from fossils only a bit younger. Their modern diversity, however, apparently is of
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
origin.
Plecoptera are found in both the
Southern
Southern may refer to:
Businesses
* China Southern Airlines, airline based in Guangzhou, China
* Southern Airways, defunct US airline
* Southern Air, air cargo transportation company based in Norwalk, Connecticut, US
* Southern Airways Express, M ...
and
Northern Hemisphere
The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
s, and the populations are quite distinct, although the
evolution
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary evidence suggests species may have crossed the
equator
The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can als ...
on a number of occasions before once again becoming geographically isolated.
All species of Plecoptera are intolerant of
water pollution
Water pollution (or aquatic pollution) is the contamination of water bodies, usually as a result of human activities, so that it negatively affects its uses. Water bodies include lakes, rivers, oceans, aquifers, reservoirs and groundwater. Water ...
, and their presence in a stream or still water is usually an indicator of good or excellent water quality.
Description and ecology
Stoneflies have a generalized anatomy, with few specialized features compared to other insects. They have simple mouthparts with chewing
mandibles
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
compound eye
A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s, and two or three
ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. The legs are robust, with each ending in two claws. The
abdomen
The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
is relatively soft, and may include remnants of the nymphal gills even in the adult. Both
nymph
A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
s and adults have long, paired cerci projecting from the tip of their abdomens.
The name "Plecoptera" literally means "
braid
A braid (also referred to as a plait) is a complex structure or pattern formed by interlacing two or more strands of flexible material such as textile yarns, wire, or hair.
The simplest and most common version is a flat, solid, three-strande ...
ed-wings", from the
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''plekein'' (πλέκειν, "to braid") and ''pteryx'' (πτέρυξ, "wing"). This refers to the complex venation of their two pairs of
wings
A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expresse ...
, which are
membranous
A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. ...
and fold flat over their backs. Stoneflies are generally not strong fliers, and some species are entirely wingless.
A few wingless species, such as the Lake Tahoe benthic stonefly ('' "Capnia" lacustra'') or '' Baikaloperla'', are the only known insects, perhaps with the exception of ''
Halobates
''Halobates'' or sea skaters are a genus with over 40 species of water striders. Most ''Halobates'' species are coastal and typically found in sheltered marine habitats (a habitat where a few other genera of water striders also live), but five ...
'', that are exclusively aquatic from birth to death. Some
true water bug
Nepomorpha is an infraorder of insects in the "true bug" order (Hemiptera). They belong to the "typical" bugs of the suborder Heteroptera. Due to their aquatic habits, these animals are known as true water bugs. They occur all over the world o ...
s (Nepomorpha) may also be fully aquatic for their entire lives, but can leave the water to travel.
The nymphs (technically, "naiads") are aquatic and live in the
benthic zone
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...
of well-oxygenated lakes and streams. A few species found in New Zealand and nearby islands have terrestrial nymphs, but even these inhabit only very moist environments. The nymphs physically resemble wingless adults, but often have external gills, which may be present on almost any part of the body. Nymphs can acquire oxygen via diffusing through the exoskeleton, or through gills located on behind the head, on the thorax, or around the anus. Due to their nymph's requirement for well oxygenated water, the species is very sensitive to water pollution. This makes them important indicators for water quality. Most species are herbivorous as nymphs, feeding on submerged leaves and benthic algae, but many are hunters of other aquatic arthropods.
Life cycle
The female can lay up to one thousand eggs. It will fly over the water and drop the eggs in the water. It also may hang on a rock or branch. Eggs are covered in a sticky coating which allows them to adhere to rocks without being swept away by swift currents. The eggs typically take two to three weeks to hatch, but some species undergo
diapause
In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, with the eggs remaining dormant throughout a dry season, and hatching only when conditions are suitable.
The insects remain in the nymphal form for one to four years, depending on species, and undergo from 12 to 36 molts before emerging and becoming terrestrial as adults. Before becoming adults, nymphs will leave the water, attach to a fixed surface and molt one last time.
The adults generally only survive for a few weeks, and emerge only during specific times of the year when resources are optimal. Some do not feed at all, but those that do are herbivorous. Adults are not strong fliers and generally stay near the stream or lake they hatched from.
Systematics
Traditionally, the stoneflies were divided into two
suborder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
s, the "Antarctoperlaria" (or "Archiperlaria") and the
Arctoperlaria
Arctoperlaria is a suborder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the ...
. However, the former simply consists of the two most basal superfamilies of stoneflies, which do not seem to be each other's closest relatives. Thus, the "Antarctoperlaria" are not considered a natural group (despite some claims to the contrary).
The Arctoperlaria, though, have been divided into two
infraorder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
s, the
Euholognatha
Arctoperlaria is a suborder of stoneflies.Plecoptera of North America, http://plsa.inhs.uiuc.edu/plecoptera/default.aspx
External linksFilipalpia) and the Systellognatha (also called Setipalpia or Subulipalpia). This corresponds to the
phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
with one exception: the Scopuridae must be considered a basal family in the Arctoperlaria, not assignable to any of the infraorders. Alternatively, the Scopuridae were placed in an unranked
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
"Holognatha" together with the Euholognatha (meaning roughly "advanced Holognatha"), but the Scopuridae do not appear significantly closer to the Euholognatha than to the Systellognatha.
In addition, not adopting the clades Antarctoperlaria and Holognatha allows for a systematic layout of the Plecoptera that adequately reproduces phylogeny, while retaining the traditional ranked taxa.Nelson (1996b)
Basal lineages ("Antarctoperlaria")
* Superfamily Eusthenioidea
** Family Diamphipnoidae
** Family
Eustheniidae
Eustheniidae is a family of insects in the order Plecoptera, the stoneflies. They are native to Australia, New Zealand, and Chile.Zwick, P. (1979)Revision of the stonefly family Eustheniidae (Plecoptera), with emphasis on the fauna of the Austra ...
Arctoperlaria
Arctoperlaria is a suborder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the ...
Euholognatha
Arctoperlaria is a suborder of stoneflies.Plecoptera of North America, http://plsa.inhs.uiuc.edu/plecoptera/default.aspx
External linksCapniidae
The Capniidae, the small winter stoneflies, are a family of insects in the stonefly order (Plecoptera). It constitutes one of the largest stonefly families, containing some 300 species distributed throughout the holarctic. Their closest relative ...
(about 300 species) – small winter stoneflies
** Family Leuctridae (390+ species) – rolled-winged stoneflies
** Family
Nemouridae
The Nemouridae are a family of Plecoptera, stoneflies containing more than 700 described species, occurring primarily in the Holarctic region.Nelson, C. Riley. 1996. Nemouridae. Version 1 January 1996 (under construction). http://tolweb.org/Nem ...
Chloroperlidae
Chloroperlidae are a family of stoneflies, commonly known as green stoneflies, with more than 200 species and 22 genera. They appear green to yellow in colour, and are popularly used among fisherman as bait for trout fishing. Green stoneflies ...
(over 180 species) – green stoneflies
** Family
Perlidae
The Perlidae are a family of stoneflies, with more than 50 genera and 1,100 described species. The majority of the Perlidae are found in eastern North America, but they occur worldwide except for Antarctica and parts of Africa. Their lifecycle ...
(about 400 species) – common stoneflies
** Family
Perlodidae
The Perlodidae, also known as the perlodid stoneflies, stripetails, or springflies, are a family of stoneflies.
Natural history
The family Perlodidae is composed of at least 50 genera and over 350 species, with the fossil records extending at ...
(350+ species)
** Family
Peltoperlidae
The Peltoperlidae, also known as roach-like stoneflies or roachflies, are a family of stoneflies.
The family Peltoperlidae comprises 11 genera and 46 known species. Species are semivoltine, meaning their lifecycles last 1 to 2 years. Adults of ...
(about 68 species) – roachlike stoneflies
** Family
Styloperlidae
Styloperlidae is a family of stonefly, stoneflies in the order Plecoptera. There are at least 2 genera and 9 described species in Styloperlidae. The species etymology is based on the type locality near Sapa.
History
The Styloperlidae family wa ...
(about 10 species)
** Family
Pteronarcyidae
The Pteronarcyidae, also known as giant stoneflies or salmonflies, are a family of the order Plecoptera.
Natural history
Two genera of Pteronarcyidae are found; ''Pteronarcys'' is found in all of North America, while ''Pteronarcella'' is f ...