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An Introduction to the Aquatic Insects of North America Third Edition

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An Introduction to the Aquatic Insects of North America Third Edition An Introduction to the Aquatic Insects of North America Third Edition Edited by Richard W. Merritt Department of Entomology Michigan State University and Kenneth W. Cummins Ecosystem Research and Implementation Department South Florida Water Management District KENDALL/HUNT PUBLISHING COMPANY 4050 Westmark Drive Dubuque, Iowa 52002 PLECOPTERA 263 Table 148. Continued Amphincmurinnc (24) ilmp/JillltJ/lUm (13) Lolic-erosionnl und Sprnwlers-clingers Shredders-netritivores. Widesprend 679, 1574,2414,2435. depositionul (in delri- facultative collectors 890,2436,3629,4578. Ill.';) -gatherers 1104,1587,4242, 2025.23·10,3841 MnleJlka (1I) West 935,2316,3334,4134 Nemourinnc (41) Leal/ill (1) Mootnnn 205,3B41 Nell/DUm (5) Latic-erosionul Spmwlers-clingcrn Shredders·dctritivores West. North, Northeast 243,247. 1574,3629, detritus) (herbivores [mucrunl· 2341,4341,3B36, gael ) 3B41,4435, 1103, 2343 Oslmrl.!tr:JJ (6) Enst, Northwest 1576, 1587,2414, 3B41 Pllra1tt!/I101lnl (I) Northen.~t 3B41 Podmann (5) West. Northell.5t 15BB, 2305, 3841 prosroia (4) Lolic-erosion111 und Shredders-dctritivorcs: Northwest, Northeust 679, 1574, 1905, 3629. depositional (fucultntive st:r.Jpm) ll04, 1587, 1622,2025, 3B41.4234 Sllipra(l) Lotic---erosionlll nnd Sllreddcrs·dctritivores, North, Northcn.~l 193,1574,3629,3841 depositional (scrnpers) SOJcdillU (7) Spring outflows Widespread 1587,2079,2414,4578, 3B41, 3924. 3926, 3919. 3841 Visoko (I) West Znpodn (10) Lotic-ermionul (de- Sprnwlcrs-clingcrs Shredders-delritiyores West, EIl~1 5BI, 1613.2079.3241.3647, mIllS) (leaf IiUer)·herbivores 2B25. 3334, 3650, 3B41. (mos.~) 3335, 3836, 42.14 Leuctridnc (53) Genemlly lotic cm- GCrlcnuly spmwleffi- Gellerally ~hredders- 1482. 1574.2021,3493, SiOTHll (course sedi- clingers derrith'llres 1310 ments, debris jams, leaf packs) and c.lepo- silionnl Megnleuc:trinne (5) Mellulellclrn (6) Springs, seeps Northwest, 3841 Appnlnchiuns Leuclrimte ('17) De,spa:do (l) Spring outOows West 3335.3B41 leI/elm (26) Louc---erosional and Shredder~-detritjvores East, North Centrol 243, 1574, 3629, 4343. 2343, depositional 3795.1104, 1576.2025,4630, 2339.2342.3841,4488,4651, 4652, 1665,3841.1057,1877 Mo,selin (I) PlIl:ific Northwest 3B41 Paraleuctm (9) Widesprcnd 3841,4234 Per/omyiu (2) West 3B41 Z!alltul'rra (8) Smnllstrcnms (snme East. Centml, Texa~ 3731,3B41 intermittent) Cnpniidac (151) Cenernlly lolk----em- Generally sprowlcrs- Generally shredders- 1307.131O,15B3 sional (course sedi- clingers dctritivorcs mellIs, wood, leaf pucks) nnd depositional AI/ocapnia (41) Clingers Shrcdder:s-detritlvores Enst 246,679. 1185, 1186, IISS, 1189,1583,2'114, ]222 Bolshecapilia (6) Small strenms, alpine lukes West tc!.l) ~ Emplll1!iw vn lmphk relulivn~hips. r h 4 PLECOPTERA K. W. Stewart P. P. Harper University of North Texas, Denton Universite de Montreal, Canada INTRODUCTION 1977; Stewart and Maketon 1991), which is a tapping upon the substrate with the tip of the abdomen. Male drummers contin­ Plecoptera (stoneflies) are primarily associated with clean, ue this activity throughout their adult life, but only virgin cool running waters, although a number of species are adapted females respond to males. Males initiate the communkation, to life in large oligotrophic, alpine, and boreal lakes. Seveml and females answer either during or immediately after the male species inhabit streams that warm or dry up in summer or that drumming call. Male and female signals are species specific, are organically enriched. Eggs and larvae of all North Ameri­ and diulect.s have been detected in European spp. (Rup­ can species are aquatic, and, with the exception of un Utacap­ Di/lra precht 1972) and North American Pteronarcella spp. (Stewart nia species living in the depths of Lake Tahoe, adult!> are ter­ et al. 1982). restrial. The following account of the biology and ecology of stoneflies can be supplemented with the comprehensive works Little courtship has been observed in stonet1ies prior to of Hitchcock (1974), Hynes (1976) and Stewart aDd Stark muting, although some Peltoperlidae do a dance involving con­ (1988). siderable wing lifting during drumming. During the intersexu­ Stonefly larvae tend to have specific water temperature, al duetting, males actively search with triangulation or other substrate type, and stream size requirements reflected in their patterns (Abbott and Stewart 1993) while females remain sta­ distribution and succession along the course of streams and tionary, until contact is made. Males mount females, curve the rivers. In addition, a distinct microdistribution of species and abdomen to the left or right side, and, using structures such as of size classes of individual species is frequently observed the hooks of the subanal lobes or the epiproct, gain apposition within a particular stream reach. Microhabitats include boulder with the subgenital plate (Chap. 2). The aedeagus is everted surfaces, cobble and gravel interstices, debris accumulations from beneath the ninth sternum and curves upward between the and leaf packs, as well us the hyporheal (Isocapllia sp., Pamp­ male cerci into the female genital opening. Sperm are either erla spp., some European [metra spp., and early instars of conveyed internally directly from, the aedeagus (Stewart et al. many other species). Stonefly adults live on riparian vegetation 1969) or through a specially grooved epiproct (Brinck 1956), or or among rocks or debris. introduced externally into a pocket just outside the genital Food ingested by larvae may vary depending on species, opening, after which the female aspirates the speml into the developmental stage, or time of day (Stewart and Stark 1988). bursa (Stewart and Sturk 1977). The gravid female releases Some species are, for example, shredders or predators through­ eggs over the stream surface or deposits them in the water. out development; however, others may change their feeding Eggs are attached to the substrate by either a stick)' gelatinous habit as development proceeds. Gut content analyses indicate covering or specialized anchoring devices. that shifts from herbivory-detritivory in the earliest instars to In most species, embryonic development proceeds direct­ omnivory-carnivory in later periods are common. Predators ly and is complete within 3-4 weeks; hatchlings then undergo usually ingest their prey whole. Stoneflies may either be gradual development requiring some 12-24 months, depending opportunistic or select for particular food items in low abun­ on the species, sex, and environmental conditions. In most dance. Details on individual families are given in Table 14B. cases, growth requires 10-11 months (annual life cycle, univol­ Adults of Taeniopterygidae, Nemouridae, Leuctridae, Ci.ipni­ tine), although growth rates are often low during winter idae, Chloroperlidae, and some Perlodidae feed on epiphytic months. Semivoltine (2-3 year) cycles are known or suspected algae or young leaves and buds of riparian vegetation. in many Pteronarcyidae and Perlidae. as well as in a few Plecoptera species emerge in clear succession throughout species of Nemouridae, Leuctridae. and Perlodidae (Table the year, except at latitudes or altitudes experiencing seasons of 14A). Such cycles are ternled slow seasonal in Hynes' (1961) drought or frost. In general, species with predaceous larvae classification, have an emergence period restricted to the spring and summer In other species, particularly those inhabiting intermittent months (Table 14A). Adults live about 1-4 weeks, although aquatic habitats or streams subjected to extremes in temperature, winter species often have greater longevity. embryonic development may be arrested for 3-6 months and In many stonefly species. male and female adults are hatching delayed until environmental conditions are more favor­ attracted by drumming (Rupprecht 1967; Zeigler and Stewart able (Table 14A). Hatchlings must then develop rapidly in order 217 218 PLECOPTERA to complete a univoltine cycle in the remaining 6~8 months. A 14.57). Gills may persist in adult stages, but tend to shrivel aod variation of this strategy is found in some Taeniopterygidae, sometimes leave scars (Fig. 14.124) or stubs (Figs. 14.114 Capniidae, and Leuctridae in which diapause occurs in the early 14.122, 14.137). ' larval instars (Harper and Hynes 1970). This type of cycle is Cerci: As a rule, cerci of both adults and larvae are loon called fast seasonal. Under drought conditions, species of whiplike appendages divided into numerous segments (Fig; Zealellctra are able to extend the embryonic diapause for an 14.1-14.2); however, in a few families, cerci ofthe adults ofone extra year (Snellen and Stewart 1979a). These observations and or both sexes are reduced to the basal segment (Figs. 14.138_ Hynes and Hynes' (1975) studies on Australian stonctlies sug­ 14.139, 14.145-14.150, 14.174-14.177), and, in some cases, are gest considcmble flexibility in stonefly life cycles. Patterns of modified as accessory male copulatory organs (Figs. 14.146 egg development, life histories and life cycles, and larval growth 14.174-14.175). ' are reviewed by Stewart and Stark (1988). WIngs: The wings appear in half-grown larvae as tiny bUds on the thorax, They grow with each successive instar and Wing pads reach their full development in the tenninal instar (Figs. EXTERNAL MORPHOLOGY 14.1, 14.36, 14.38). Most species are winged; however, some may be apterous or brachypterous (shortened wings; Fig. Larvae and Adults 14.37), often depending on sex. Venation is simple and veins are easily recognized (Fig. 14.2).
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