StonefliesC. Riley Nelson
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The Plecoptera (stoneflies) are a small order of exopterygote insects of about 2000 species worldwide. The order has a long, but rather fragmented, fossil record extending back to the early Permian. These Permian fossils can be rather easily contained in the living suborders, Arctoperlaria and Antarctoperlaria. The modern families are clearly identifiable among specimens from the Baltic amber, which is of Miocene age (38-54 million years ago) as well as a few other compression fossils.
The nymphs of stoneflies dwell in aquatic habitats, although there are some species in the southern hemisphere which spend much time on damp land. In general the preferred habitat is rocky streams with a noticeable current, but there are species that live in sandy places. Cold lakes and ponds are also suitable habitats in the north and at high altitudes. We know far less about lakes as habitats than we do about streams and rivers. In running water the usual nymphal habitats are rocky, stony or gravel substrata, and there are more species in cooler, swifter water. Many studies have shown that the various species are each to be found in particular situations. For instance, large Perlidae and Perlodidae are usually found on and under large stones. Chloroperlidae tend to occur in gravel while Pteronarcyidae, and Nemouridae are most frequently encountered in leaf packs.
Finally, it should be stressed that because stoneflies depend upon cool, well-oxygenated water for their nymphal development, they are very susceptible to human abuse of water courses. Any effluent that reduces the oxygen content of the water quickly extirpates them. Even quite minor pollution sources such as farm drainage can elimate stoneflies from nearby streams. Also clearing the land or impoundment or water courses, both of which raise the summer temperature of the water, can eliminate stoneflies from the habitat. Plecoptera, then, as a whole serve as indicators of healthy streams and rivers.
Stoneflies are 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 such as crickets and grasshoppers. They have long filiform antennae at least half 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 larvae, or 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 filaments, not plates.
Both Zwick (1973) and C. H. Nelson (1984) list Nemouridae and Notonemouridae as sisters. This is an interesting arrangement because the nemourids are uniquely northern hemisphere in distribution and the notonemourids are uniquely southern hemisphere in distribution. This is the only pattern of this sort at the family level (and below!) in Plecoptera. Nemouridae + Notonemouridae is then placed as sister to Capniidae + Leuctridae.
The Pteronarcyidae form a monophyletic group which is most closely related to the Peltoperlidae + Styloperlidae clade (Uchida & Isobe 1989). These three families plus the Perloidea (Perlidae + Perlodidae + Chloroperlidae) have paraglossae and glossae of approximately equal length, and have been called the Systellognatha (sensu Uchida & Isobe 1989).
Pteronarcyids, styloperlids, and peltoperlids (Pteronarcyoidea, sensu Uchida & Isobe 1989), however, are more bland in coloration than the predatory systellognaths (Perloidea sensu Uchida & Isobe 1989) as nymphs. The Pteronarcyoidea are herbivorous throughout their aquatic lives. In contrast, the three perloidean families are all carnivores, at least as mature nymphs. The predatory behavior is a synapomorphy for the Perloidea and a symplesiomorphy for the Pteronarcyoidea.
The relationship of peltoperlids with other stonefly families has been in question since their initial recognition. Zwick (1973) placed the family as the sister group to Subulipalpia (Perlidae, Perlodidae, and Chloroperlidae) with synapomorphies of: 1, body stout, head prognathous, cockroach-like nymphal body form, and 2, male cercal segments fused. Additional apomorphic features which suggest that the family is monophyletic include: 1, nymphal coxae with flap-like lobe (Claassen 1931); 2, nymphal thoracic sterna enlarged into prominent plates, and 3, molar area of nymphal mandibles with pectinate surface (Stark & Stewart 1981). Stark and Stewart (1981) proposed that the family is more closely related to Pteronarcyidae than the carnivorous Subulipalpia based on: 1, nymphal lacinia tridentate; hemispherical, dorsally flattened eggs; and 3, nymphal tergum 10 with apical spine-like process.
The chloroperlids, along with the perlids and perlodids, form a monophyletic trichotomy which has defied splitting despite numerous attempts to determine which family is the nearest sister to which other family. Zwick (1973) listed the chloroperlids as sister to the perlids and cited the synapomorphies for this scheme as having the sternal coxal rotator muscles reduced and having having attachment of muscle I ism 22 shifted from the tip of the furcal arm to near the base (his Fig. 16b). C. H. Nelson (1984) found no resolution to the trichotomy using characters derived from Zwick (1973) and additional characters he explored using numerical cladistic analysis.
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I thank Noel Hynes for graciously granting access to the manuscript from which the Introduction and References were distilled.
C. Riley Nelson
Brigham Young University, Provo, Utah, USA
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Nelson, C. Riley. 1996. Plecoptera. Stoneflies. Version 01 January 1996 (under construction). http://tolweb.org/Plecoptera/8245/1996.01.01 in The Tree of Life Web Project, http://tolweb.org/