Great Basin Naturalist

Volume 36 Number 3 Article 6

9-30-1976

The stoneflies () of Louisiana

Kenneth W. Stewart North Texas State University, Denton, Texas

Bill P. Stark North Texas State University, Denton, Texas

Thomas G. Higgins Central Hudson Gas and Electric Corp., Poughkeepsie, New York

Follow this and additional works at: https://scholarsarchive.byu.edu/gbn

Recommended Citation Stewart, Kenneth W.; Stark, Bill P.; and Higgins, Thomas G. (1976) "The stoneflies (Plecoptera) of Louisiana," Great Basin Naturalist: Vol. 36 : No. 3 , Article 6. Available at: https://scholarsarchive.byu.edu/gbn/vol36/iss3/6

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. THE STONEFLIES (PLECOPTERA) OF LOUISIANA^

Kenneth W. Stewart,^ Bill P. Stark,-* and Thomas G. Higgins*

Abstract. — A list of species, illustrated keys to nymphs and adults, distribution, and biological notes are presented for 24 stonefly species occurring in Louisiana. Leuctra moha?, Pteronarcys dor- sata, Helopicus subvarians, kansensis, Paragnetina immarginata, Phasganophora capitata, and evoluta are recorded only from the florida parishes east of the Mississippi River. Isoperla coushatta. Isoperla mohri, clymene, an undescribed Neoperla (Sp. A), Paragnetina fumosa, , Acroneuria arenosa, drymo, and placida are found both east and west of the Mississippi; and AUocapnia granulata, Allocapnia malverna, Amphinemoura nigritta, and three species of have been recorded only west of the Mississippi. Habitats are primarily sand-bottomed streams of pine-hardwood rolling hills, constituting a portion of the western Gulf Coastal Plain. Few stoneflies occur in streams of the Mississippi Alluvial Plain, the Alluvial Atchafalaya Basin, or the lowlands along the Red River.

Distribution and biology of the eco- flies along the lower Mississippi, and the logically important stoneflies are poorly fact that Louisiana was the only state that known for the Gulf Coastal Plain Physio- he had not visited. graphic Province of North America. Be- The potential habitats for stoneflies are cause this area was included in Ricker's generally limited in Louisiana to mostly (1964) "Southeastern" Glacial Refuge, sand-bottomed streams of: (1) the north its stonefly fauna has generally been con- central and western upland rolling hills, sidered similar to that of the Central and with their oak-hickory and shortleaf- Southern Appalachian and Ozark- Qua- loblolly pine forests, (2) the southwestern chita provinces. Recent discovery of three and central longleaf-slash pine-deciduous new Leuctra species and an Isoperla forest, and (3) mixed forests of the florida (James 1974) from Alabama, the en- parish region. The 50-100-mile-wide Mis- demicity of Allocapnia malverna Ross to sissippi Alluvial Plain of eastern Louisi- the Coastal Plain Province in Texas, ana, the alluvial Atchafalaya Basin along Oklahoma, and Arkansas (Stewart et al. the lower Mississippi, and the lowlands 1974), and discovery of two new Isoperla, along the Red River, cutting from the coushatta and sagittata, from East Texas northwestern corner through the central (Szczytko and Stewart 1976) suggest oc- portion of the state, are flat, characterized currence of considerable post-pleistocene by sluggish, highly silted streams, and are speciation on the coastal plains. generally devoid of stoneflies. These phys- This paper constitutes a review of Ple- iographic and vegetational features ren- coptera known from the West Gulf Coastal der Louisiana of special interest to fur- Plain and is the first comprehensive thering knowledge of present distribution treatment of the stoneflies of Louisiana. and possible past dispersals of Nearctic Previous collection effort is indicated by Plecoptera. the fact that only three species, Acroneu- ria evoluta Klapalek, Methods (Newman), and Paragnetina immarginata (Say), were listed from the state by We began collecting in Louisiana in lilies (1966). Another species, Isoperla 1970. Our effort in the florida parishes guerinii (Pictet), was considered by was concentrated during the late winter lilies a nomen dubium (et oblitum). Ross and s})ring months, Feb. -June, 1971-1973, and Ricker (1971) plotted four localities and in December 1975. Northern and cen- in central Louisiana for Allocapnia mal- tral sections were extensively collected in verna and one for Allocapnia granulata December 1970, January and December (Claassen). Thus, a total of four valid 1971, January, February, and May 1973, species was attributed to Louisiana in the and March 1974. Limited collecting at literature through 1971. Ricker (1972) selected points throughout the state was mentioned the probable scarcity of stone- done over the five-year period.

'Study supported by the Faculty Research Fund of North Texas Stale University. '•'Professor and Research Associate, respectively. North Texas State University, Deiiton, Texas 7fi20? ^Present address: (;oiitral ITikIsdii Gas atid Klectrir Corp., Piiuphkeepsie, NY. 12602

366 .

September 1976 STEWART ET AL: STONEFLIES OF LOUISIANA 367

Collecting expeditions often traversed trast. Photographs of ova were taken from over 1,000 miles. Streams were collected sf)ecimens collected in the indicated Coast- by a combination of methods, including al Plains state. sweeping streamside vegetation, using kick nets in riffles, actively searching List of Louisiana Stoneflies through leaf debris, removing and scruti- Systematic Treatment according to nizing submerged logs, actively searching Zwick (1973) among rocks and debris along stream Suborder margins, tapping emergent vegetation with Group Euholognatha a stick (good method for adult Taeniop- Family Leuctridae teryx), searching surfaces and cracks 1. Leuctra mohal Ricker^. around bridges and in some instances Family Taeniopterygidae searching around nearby lights at night 2. Taeriiopteryx burksi Ricker and Ross*. or operating a fluorescent light device at 3. Taeniopteryx lita Frison"*. streamside. 4. Taeniopteryx lonicera Ricker and Ross*. Distributional data and /or specimens Family Nemouridae 5. Amphinemura nigritta (Provancher)*. were obtained from Louisiana State Family Capniidae University at Batan Rouge, Smithsonian 6. AUncapnia granulata (Claassen). Institution, Harvard Museum of Compar- 7. AUncapnia malverna Ross. ative 7oolog\% University of Utah, Uni- Group Systellognatha versity of Kansas, and the Illinois Natural Family Pteronarcyidae History'' Surv^ey. 8. Pteronarcys dorsata (Say)*. aedeagi of The males were extruded Family Perlodidae by squeezing at time of collection or with 9. Helopicus subvarians (Banks)*. forceps in preserved specimens after 10. hoperla coushatta Szczytko and clearing in a 5 percent solution of KOH. Stewart*. 11. Isoperla mohri Frison*. Specimens of hoperla were compared to 12. hoperla sp. A. the t\'j)es of hoperla mohri Prison, ho- Family pprla namata Prison, hoperla richardsoni 13. (Newman)*. Prison, and hoperla coushatta Szcz^^ko 14. Neoperla sp. A. and Stewart. TTie recent revisions by 15. Paragnetina fumosa (Banks)*. 16. Paragnetina immarginata (Say). Stark and Gaufin f 1976a, b) were fol- 17. Paragnetina kansensis (Banks)*. lowed in determinations of Perlidae. Ova 18. Phasganophora capitata (Pictet)*. were dissected from gravid females, 19. Acroneuria abnormis (Newman)*. mounted according to the techniques of 20. Acroneuria arenosa (Pictet)*. 21. Klapalek. Koss (1968), and examined using a Zeiss 22. Perlesta placida (Hagen)*. photomicroscope equipped with phase 23. (Newman)*. contrast and Nomarski interference con- 24. Perlinella ephyre (Newman)

Key to the Adult Stoneflies of Louisiana

1. Paraglossae and glossae equal in length (Pig. 1) 2 Paraglossae longer than glossae (Pig. 2) 6 2(1). Gill remnants present on thorax and first two abdominal sterna;

body length greater than 20 mm .. Pteronarcyidae; Pteronarcys dorsata^. Gill remnants absent from abdominal sterna; body length less than 15 mm 3 3(2). Pirst and second tarsal segments approximately equal in length; coxae with small round membranous area ventrally (Pig. 5) Taeniopterygidae; Taeniopteryx 19 Second tarsal segment shorter than first (Pig. 6); coxae without membranous area ventrally 4 4(3). Cerci multisegmented. Capniidae; Allocapnia 21 Cerci wdth one segment 5

5(4). Apical marginal space of forewing with oblique crossvein (Pig. 10); cervical gills present Nemouridae; Amphinemura nigritta

'Louisiana records represented by nvmphs only. 368 GREAT BASIN NATURALIST Vol. 36, No. 3

Apical marginal space of forewing without oblique crossvein; cervical gills absent Leuctridae; Leuctra moha?^.

6(1). Gill remnants usually present on sides and venter of thorax; arms of mesosternal Y-ridge poorly developed, but reaching to, or near, anterior corners of furcal pits (Fig. 4). Perlidae 7 Gill remnants absent from sides and venter of thorax; arms of mesosternal Y-ridge well developed and reaching posterior cor- ners of furcal pits (Fig. 3). Perlodidae 18

7(6). Apex of abdomen with a pair of dorsal sclerotized genital hooks (Figs. 11, 12) 8 Apex of abdomen without dorsal hooklike structures; posterior margin of eighth sternum variously modified 13

8(7). Genital hooks developed from hind margin of tergum 10 anterior to cerci (Fig. 11); abdominal sternum without a hammer 9 Genital hooks developed from paraprocts, originating below cer-

eal bases (Fig. 12) ; abdominal sternum 9 with or without ham- mer 11

9(8). Two ocelli Neoperla clymene' Three ocelli 10

10(9). Genital hooks projecting to anterior margin of tergum

- PJiasganophora capitata Genital hooks projecting to middle of tergum 9. Paragnetina 22

11(8). Sternum 9 without a hammer Perlesta placida Sternum 9 with a hammer (Fig. 13) 12

12(11). Forewing with at least one crossvein beyond the anal cell between

A^ and A- (Fig. 9) Perlinella ! 24 Forewing without crossveins beyond anal cell between A^ and A-; patches of spinules on abdominal terga 9 and 10 (Fig. 12) Acroneuria 25

13(7). Forewing with at least one crossvein beyond the anal cell between A^ and A^ (Fig. 9). Perlinella 24 Forewing without crossveins beyond anal cell between A^ and A- 14

14(13). Two ocelli Neoperla clymene'' Three ocelli 15

15(14). Subgenital plate with median notch on posterior margin 16

16(15). Forewing length less than 15 mm Perlesta placida Forewing length more than 1 7 mm Paragnetina 22 17(15). Thoracic sterna with transverse dark bands of pigmentation Phasganophora capitata Thoracic sterna without dark bands of pigmentation. Acroneuria 25

18(6). Submental gills present; male tergum 10 cleft; wings and most of body black flelopicus suhrarians^ Submental gills absent; male tergum 10 entire; wings and body color variable, pale yellow to brown. Isoperla 27

19(3). Vesicle usually j)resent on male steniiun 9; paraprocts narrow and pointed at tip; notch of female subgenital plate margined by dark V-shaped bands

"Louisiana record consists of one female from Washington Parish, 'The genus Neoperla is preeently imder revision; some specimens will he n sorond species (sec \c

Vesicle absent from male stenium 9; paraiirocts broad at base and truncate or broadly rounded at tips; notch of female subgenital plate margined by dark U-shaj)ed band 20

20(19). Basal width of paraprocts equal to or greater than length Taeniopteryx lita Basal width of paraprocts approximately half the length Taeniopteryx lonicera

21(4). Process of male tergimi 8 cleft; female subgenital plate [)ointed mesally Allocapnia granulata Process of male tergum 8 entire; female subgenital plate with apical margin convex Allocapnia malverna

22(10,16). Body color pale yellow^; male tergum 5 unproduced Paragnetina immarginata Body color dark browTi to black; male tergum 5 posteriorly pro- duced and mesally notched 23

23(22). Male genital hooks pointed at apex; ovum with collar at least half as wide as greatest diameter Paragnetina immarginata Male genital hooks bluntly rounded at apex; ovum with collar less than one-third greatest diameter Paragnetina jumosa

24(12,13). Pronotum with median dark band of pigmentation Perlinella drymo Pronotum without distinctive color pattern Perlinella ephyre

25(12,17). Male paraprocts broadly triangular and flattened; female sub- genital plate not produced Acroneuria abnormis Male paraprocts slender, fingerlike; female subgenital plate pro- duced over at least one-third of sternum 9 26

26(25). Male paraprocts notched apically; female subgenital plate trun- cate Acroneuria arenosa Male paraprocts evenly tapered to a point; female subgenital plate rounded, lateral margins constricted Acroneuria evoluta^

27(18). Abdominal terga with longitudinal rows of dots; male paraprocts not produced over tergum 10; female subgenital plate rounded Isoperla mohri Abdominal terga without rows of dots; male paraprocts produced

over tergum 10; female subgenital plate triangular in outline .... Isoperla coushatta

Key to Nymphal Stoneflies of Louisiana

1. Paraglossae and glossae equal in length (Fig. 1) 2 Paraglossae longer than glossae (Fig. 2) 6

2(1). Branched gills present on thorax and first two abdominal sterna Pteronarcyidae; Pteronarcys dorsata Gills absent from abdominal sterna 3

3(2). Unbranched. fingerlike gills present on each coxa; first and sec- ond tarsal segments approximately equal Taeniopterygidae; Taeniopteryx Coxae without gills 4

4(3). Branched cervical gills present; hind wingpads widely divergent Nemouridae; Amphinemura nigritta Cervical gills absent; hind wingpads parallel to axis 5

5(4). Abdominal terga with distinct posterior setal fringe; segments 1-9 370 GREAT BASIN NATURALIST Vol. 36, No. 3

divided laterally by membranous fold (Species indistinguish- able at this time) Capniidae; Allocapnia Abdominal terga without postrior setal fringe; segments 1-6, at most, divided by lateral membranous fold Leuctridae; Leuctra

6(1). Branched gills present on sides and ventor of thorax. Perlidae 7 Branched gills absent from thorax. Perlodidae 12

7 {6). Occiput with a straight, evenly spaced, complete row of spinules (Fig. 7) 8 Occiput with or without spinules, if present, the row is sinuate, unevenly spaced or incomplete (Fig. 8) 10 8(7). Two ocelli Neoperla clymene Three ocelli 9 9(8). Abdominal terga 1-9 with numerous scattered spinules in ad- dition to posterior fringe. Paragnetina 13 Abdominal terga 1-9 with 5 or less scattered spinules Phasganophora capitata 10(7). Occiput with a sinuate, unevenly spaced row of spinules; abdomi- nal terga freckled Perlesta placida Occiput with, at most, a few spinules near postocular fringe; ab- dominal terga without freckles 11

11(10). Postocular spinule row reduced to a single seta, or absent; pro- notum without well-developed setal fringe. Perlinella 15 Postocular spinule row with several stout setae; pronotum with front and hind margins fringed with spinules. Acroneuria 16

12(6). Submental gills present (Fig. 2) Helopicus subvarians Submental gills absent. Isoperla 18

13(9). Abdominal terga patterned in yellow and brown; a longitudinal mesal fringe of long setae extending from head to apex of ab- domen Paragnetina immarginata Yellow pigmentation absent or restricted to a few abdominal terga; body without a mesal longitudinal fringe of setae 14

14(13). Brown pigmentation extending completely across frons forward of median ocellus Paragnetina kansensis Brown pigmentation interrupted on frons by a longitudinal stripe of yellow forward of median ocellus Paragnetina fumosa

15(11). Two ocelli Perlinella ephyre Three ocelli Perlinella drymo

16(11). Anal gills absent Acroneuria abnormis Anal gills present 17

17(16). Abdominal terga with alternating transverse bands of brown and yellow; dorsum of head with broad yellow W-mark Acroneuria evoluta Abdominal terga without distinct transverse yellow bands; head pattern variable Acroneuria arenosa and mela

18(12). Maxillary lacinia with a single tooth and setae present along its entire inner margin, continuing around the outer margin; man- dibles deeply cleft with serrations on inner margin of outer tooth Isoperla mohri Maxillary lacinia with two apical teeth and complete inner row of setae; mandibles shallowly cleft, with no serrations on outer tooth Isoperla coushatta .

September 1976 stkwart et al: stoneflies of Louisiana 371

,^r\>\fP ^^^'^

^por

Fig. 1-8. Louisiana Plecoptera: 1, Pteronarcys, nymphal labium (p = paraglossae. g = glossae); 2, Hydroperla crosbyi. nymphal labium (sg = submental gill); 3, H. crosbyi, nymphal mosostemal grooves; 4. Paragnetina fumosa. nymphal mososternal grooves (a = arm of Y-ridge, fp = furcal pit); 5, Taeniopteryx lita, nymphal tarsi; 6. Allocapnia granulata, nymphal tarsi; 7, Paragnetina fumosa. nymphal head (or = occipital spinule row. por = post occular spinule row); 9. Perlesta placida. nymphal head (oc = anal crossvein) 372 GREAT BASIN NATURALIST Vol. 36, No. 3

Figs. 9-n. Louisiiiriii Plornptera: 9, Perlinella cpliyre. forowing (ac anal i rossvoin) ; 10, Amphinemura nigritta, forewing (amc = apical marginal crossvoiii); 11. Parnprirlina futnosa, male terminalia (c = cercus, gh = genital hook); 12, Acroncuria ahnonuis, male torminalia (c = cer- cus, pp = paraproct, sp = spinule patch); \i, Aroneuria arenosa, male sternum 9 (h = hammer). September 1976 STEWART ET AL: STONEFLIES OI-^ LOl'ISIANA 373

Leuctra tnoiia'} Ricker Hanson. Harley P. Brown recently sent us a Leuctra nymph from Tishomingo Leuctra rnoha Ricki'i. 19:52: 171. County, Mississip])i. Distribution in Louisiana: Washington, Small Crt'ek Jet. Hwy 10-62 near Sheridan (Fig. 14).

Only one female was collected at the Taenioptcryx hurksi Ricker and Ross locality R. above by K. W. Stewart, W. Taeniopteryx burksi Ricker and Ross, 1968: 1425. Baiimann, and B. P. Stark, December 1, Distribution in Louisiana: Bienville. Black 1975. It fits the description and illustra- Lake Bayou; Saline R. (plus other parislies tion of a supposed Leuctra tnoha fe- from which only presentlj" indistinguishable Taeniopteryx have been collected. male (Ricker 1952, Fig. 123). The col- nymphs Fig. 15). lector, Dr. P. W. Fattig, had taken two types of females along with the holotypo; We collected a large series of adults the one illustrated w'as more accordant in and only two nymphs at the above two size to the holotype, but Ricker noted that localities in Bienville Parish on 4 Feb- correlation was not certain. All previously ruary 1972, indicating a January-Feb- reported records of this species have been ruary emergence. This widely distributed from Georgia (Ricker 1952), during the species emerges as late as April in Quebec month of October. No Leuctra species (Harper and Magnin 1969). In Canada have been reported from immediately ad- eggs hatch a few weeks after deposition, jacent states; how^ever, ap})roximately 12 nymphs undergo a long summer dia- species occur from the Carolinas westward pause until fall, and then njTuphs grow (lilies 1966). James (1974) reported three very rapidly (Harper and Hynes 1970; new Leuctra from the Gulf Coastal Plain Harper and Magnin 1969). Nymphs occur state of Alabama, in addition to Leuctra in debris in small sand-bottom streams biloba Claassen and Leuctra alexanderi and slow-floNving bayous ; good results in

3 Leuctra moha'i' • Allocapnia granulata A. malverna

Allocapnia sp. nymphs Annphinennura sp Annphinennura nigntta Pteronarcys dorsata

Fig. 14. Distribution of Capniidae, Nemouridae, and Pteronarcyidae in Louisiana. 374 GREAT BASIN NATURALIST Vol. 36, No. 3

. Taeniopteryx burksi

I T. lita

I T. lonicera

I Taeniopteryx sp. nymphs

I

Fig. 15. Distribution of Taeniopterygidae in Louisiana. collecting adults can be obtained by tap- lustrations of the adult genitalia and ping emergent vegetation sharply with a wings were given by Frison (1952). Our stick or net handle, then netting the adults examination of the holotype from the from the water's surface. Nymphs have INHS indicated that the dorsal processes been figured by Harper and Hynes of the cerci are much stouter than the

( 1971 ) , but since lonicera and lita nymphs "small and finger-like" description and are undescribed, Louisiana specimens can- illustration given by Frison (1942). Hitch- not be separated with confidence. cock (1974) and Stewart and Szczytko (1974) illustrated the male paraprocts, Taeniopteryx lita Prison and Stewart and Szczytko (1974) com- pared the central plate (subgenital plate) Taeniopteryx lita Prison. 1942: 249. of the female abdominal sternum 8 of 7. Distribution in Louisiana: Beauregard, Whis- lita, T. lonicera, and T. starki. The nymph key Chitto Cr. Franklin, Big Cr. Grant, 1 mi SE Montgomery; near Pollock (Fig. 15). of T. lita is undescribed. This species and lonicera can be distin- guished from burksi by absence of a vesi- Taeniopteryx lonicera Ricker and Ross cle on sternite'' and by the broadly round- Taeniopteryx lonicera Ricker and Ross. 1968: 1427. ed or truncate tips of their paraprocts. Distribution in Louisiana: Beauhkc;ahd. Whis- Adults appear in December and January key Chitto Cr. (Fig. 15). in Louisiana. In their revision of Taeniop- teryx, Ricker and Ross (1968) reported Two males were collected with a large this species from adjacent Mississippi. iiiiinl)(>r of T. lita on 26 December 1970, It also occurs in the east Texas pine hills in Whiskey Chitto Creek. They exhibit in sluggish streams and bayous (Stewart the distinct slender, twisted cerci, with et al. 1974; Szczytko and Stewart 1977). inner surfaces deeply excavated for the A closely related species, 7'. starki, occurs distal 0.7 of its length, and paraprocts as in the sluggish Leon River in central illustrated by Ricker and Ross (1968). Texas (Stewart and Szczytko 1974). Il- This species has been reported from adja- ;

September 197(1 STEWART ET AE: STONEFLIES OF LOUISIANA 375 cent Mississippi (Ricker and Ross 1968) fall. Prison (1929) discussed the life cycle and Texas (Stewart et al. 1974; Szczytko and habits in Illinois. Excellent accounts and Stewart 1977). The n>anphs are un- and illustrations of the morphology of described and the life cycle unreported. adults, Nearctic distribution, and evolu- tion are given by Ross and Ricker (1971). Nymphs were figured by Frison Amphinemura nigritta (Provancher) (1929), Harden and Mickel (1952), and Harper Nemoura nigritta Provancher, 1876: 214. and Hynes (1971). Nemoura venosa Banks, 1897: 21. Nemoura striata Banks. 1920: 324. Nemoura nigritta, venosa. Claasson. 1940: 60. 66. Allocapnia malverna Ross Nemoura {Amphinemura) nigritta. Ricker. Allocapnia malverna Ross, 1964: 170. 19'52: 2-5. Distribution in Louisiana: Beauregard, Whis- Nemoura nigritta. venosa. Gaufin. 19*^6: 322. key Chitto Cr. Claiborne, 2 mi E Lisbon; DisTRinuTiON IN Louisi.\NA: C.\T.\Houi,A, Bircls DESOTO. 2 and 4 mi E Mansfield; Grant. 6 mi Cr.; DESOTO. Four Mile Bayou; Jackson. S Georgetown, 1 mi SE Montgomery. J.\ckson, stream; Lasai.le. Cr. ; unnamed Trout Natchi- unnamed stream. Natchitoches. Kisatchie Cr.

toches, Kisatchie Cr.. ; Sabine, Winn Cr. Winn Cr. Rapides, Hemphill Cr. Vernon, Blackwell Cr. Calcasieu R.; Comrade Cr. Webster, 6 mi E Minden. Lasalle, 15 mi SW Jena; Earl Cr.; This species is distributed primarily in Whitewall (Ross and Ricker 1971). central and western I^ouisiana (Fig. 14). Ricker (1952) illustrated variations in the A. malverna appears to be restricted to male paraprocts; the present name nigritta the western Gulf Coastal Plain of Louisi- probably includes a complex of species. ana (Fig. 14) and adjacent states of Ar- The paraprocts of Louisiana males varv^ kansas (Ross and Ricker 1971). Oklahoma from those figured by Ricker (1952) and (Stark and Stewart 1973), and Texas Hitchcock (1974) and may be an un- (Stewart et al. 1974; Szczytko and Stewart described species. Specimens have been 1977). Males are figured by Ross and sent to R. W. Baumann for further study. Ricker (1971); females cannot at present Adults have been collected in March- be distinguished from niohri and mystica April, and n^^nphs as late as 14 April. (both figured by Ross and Ricker 1971). Hitchcock (1974) reported a May to We have collected adults from 26 De- mid-.Tune emergence in Connecticut and cember to 5 February in Louisiana. indicated that adults came to blacklights Nymphs are un described, and the life traps. Nymphs have been figured by cycle is unknown. Claassen (1931) and Harper and Hynes (1971). Harper (1973b) described the life Pteronarcys dorsata (Say) cycle. Louisiana nymphs have four Sialis dorsata Say. 1823: 164. branched cers'ical gills, appearing as PlfTonarr^s rrgalis Newman. 1838: 176. prominent gill tufts in the neck region. Kollaria insignis Pictet. 1841: 123. Pteronarcys nobilis Jia^en. 1861: 15. Pteronarcys frigida Gersiaec\ier. 1873: 65. Allocapnia granulata (Claassen) Pteronarcys rectus Provancher, 1876: 191. Pteronarcys flavicarnis ProYancher. 1876: 191. Capnella granulata Claassen. 1924: 44. Pteronarcys lahradoriensis Samal. 1933: 95. Distribution in Louisiana; Tasam.e. Hairs Cr. Petronarcys shelfordi Frison. 1934: 25. Grant. Small stream 1 mi N Pollock; central Distribution in I-ouisiana: E. Feliciana? (label Louisiana ("one plot. Ross and Ricker 1971). -^Fla? Parish). Living.ston, Tickfaw R. .5 mi This widely distributed eastern species Tangipahoa, Tchefuncta R.; E Kent- wood. is apparently much less common than A. malvcrna and appears to be restricted to Only n>Tnphs of this Pteronarcys have the pine hills near the Kisatchi National been collected from 29 January to 27 Forest in central Louisiana (Fig. 14). May in Louisiana. The backward projec- Stark and Stewart (1973) and Stewart tion of sternum 9 in male nymphs is et al. (1974) reviewed its distribution in wider and has the sides more parallel as adjacent states of Oklahoma and Texas. described and illustrated bv Harden and Adults appear in December-January and Mickel (1952, Fig. 7 of Plate IX). Male probably undergo a n\'mphal diapause iiATTiphs of pictctii exhibit a subtriangular in summer (Harper and Hynes 1970) projection (Harden and Mickel 1952, Fig. with rapid nymphal development in the 8 of Plate IX). Female nymphs collected 376 GREAT BASIN NATURALIST Vol. 56, No. 3

in May 1972 and 1973 are over 40 mm in Helopicus subvarians (Banks)

length. Harden and Mickel (1952) stated Prrld subvarians Banks. 1920: 317. that female Minnesota nymphs of both Pcrla postica Needham and Claassen, 1925: 82. dorsata (over 40 mm) and pictetii (less lincta Needham and Claassen, 1925: 89. Perla Claassen, 1936: than 39 mm) could be consistently dif- tinctata 623. Hydroperla subvarians. Prison. 1942: 292. size. are ferentiated by their Nymphs Isogenus (Helopicus) subvarians. Ricker. found clinging to logs in larger rivers in 1952: 103. the florida parishes (Fig. 14), and emer- Distribution in Louisi.xna: Tangipahoa, Terry gence probably occurs in May and June. Cr. Washington. Tchefuncta R. Haj's Cr. 3 mi S Jet. 25-38 near Clifton. The range of dorsata is perhaps the widest of any Nearctic stonefly, being transcon- Only the distinctive nymphs with a tinental from Alaska to Labrador in the black band across the head (Ricker 1952) north, extending down the Cordillera to have been collected east of the Mississippi Wyoming in the west, and previously to River (Fig. 16) in Louisiana. Little is Georgia in the east. Louisiana is the known of the biology of this species. farthest southwestward that the species Hitchcock (1974) based its presence in has been reported, so collection and study Connecticut also on nymphal collections. of the morphology and biology of adults is needed. Nebeker (1971) reported the ef- hoperla coushatta Szczytko and Stewart fects of different laboratory temperatures on development, feeding, emergence, egg Isoperla coushatta Szczytko and Stewart, 1976: 99. production, and adult longevity. Nelson Distribution in Louisiana: Catahoula, tribu- tary Bird Cr. Lasalle, Hemphill Cr. ; Trout geni- and Hanson (1971) figured adult Cr. Lincoln, Bayou D'Arbonne. Natchi- talia, and Needham and Claassen (1925) toches. Kisatchie Cr. ; Middle Cr. R'^pides, and Knight et al. (1965) figured the egg. Castor Cr. Tangipahoa, Terry Cr. Washing- Hoke (1924) illustrated the head and ton, Hays Cr. mouthparts of a nymph that might have This species is distributed in small, been dorsata. sand-bottomed creeks throughout central

Helopicus subvarians Isoperla coushatta

• Isoperla sp. nymphs

• I. nnohn

a Isoperla sp. A

Fig. 16. Distribution of Perlodidae in Louisiana. Septonih(>r 1076 STIAVAHT ET AI/ STONE} 'LIES OF EOUISIANA Ml and iiortlieni Louisiana and the northern the male aedeagus and genitalia, female florida parish region (Fig. 16). The male subgenital plate and terminalia, and aedeagus is entirely nienibranous, some- nyniphnl mouthparts. what (lub-shajDed when extruded, and the female subgenital ])late is triangular, pro- Isoperla sp A. duced backward over not more than one- tliird of tiie stenuun 9. The lacinia of the One fully winged male Isoperla from n>^llJ)h has two apical teeth and a com- the Tulane University collection, labeled plete inner row of setae. Nymphal mani- La, N. 0. 19-IV-51, J. LI., Lot No. 2425, bles are shallowly cleft, with no serrations apj)ears to be an undescribed species. It on the outer tooth. Szczytko and Stewart is similar to /. lonpiseta, differing primar- (1976) figured the terminalia and extrud- ily in having shorter paraprocts, lack of ed aedeagus of the male, subgenital distinct spinule patterns on tergas 9 and plate and terminalia of the female, and 10, and different shape of the lobe on the complete mouthparts of the nymphs. [posterior margin of sternum 8. We are at- Adults were taken in April in Louisiana. tempting to collect additional specimens.

Isoperla rnohri Prison Neoperla clymene (NewTuan) Isoperla mohri Prison, 1935: 455. Chloroperla clymene Newman, 1839: 87. Distribution in Louisi.'^n.'v: Bienville, Black Perla occipilalis Pictet, 1841: 254. Lake Bayou. Caddo. 6 mi N Jet. 525-169. C.xLDWELL. unnamed stream. Claiborne. Cor- Distribution in Louisiana: Beauregard. Bayou

; Chitto ney Lake Spillway. Desoto. 3 mi E Mansfield, Anococo; Bundick Cr. Whiskey Cr. 4 mi N Logansport. Grant, unnamed stream. Catahoula, Birds Cr. tributar\'. East Baton Jackson, 11 mi E Jonesboro. Lincoln. Bayou Rouge, Baton Rouge. East Feliciana, near D'Arbonne. Red River, Grand Bayou. Sabine. Magnolia. Grant, Big Cr.; Fish Cr.; Camp Hardtner; Pollock. Lasalle. Trout Cr. Liv- Blarkwell Cr.; Pliillips Cr.; 3 mi W Robeline. Tangipahoa. Wilson Branch. 5 mi E Wilmer. ingston. Little Natalbany R.; Tickfaw R.; LTnion. Little Corney Bayou. near Magnolia. Rapides. Germany Branch; Hempliill Cr.; Spring Cr. St. Helena, Amite This species is widespread in Louisiana R. Tangipahoa, Big Cr. Washington, Bogue Chitto Silver Mill west of the Mississippi (Fig. 16), east R.; Hays Cr.; Little Cr.; Cr. Texas fSzxzytko and Stewart 1976, 1977), and Oklahoma (Stark and Stewart 1973). This is one of the most frequently en- It has been previously reported only from countered species in the state (Fig. 17) Illinois and Missouri. The male aedeagus and throughout the Gulf Coastal Plains, bears a distinct posterior spine below- although population densities are seldom which is a double row of sclerotized fin- great. Stewart et al. (1974) discussed the gers (Szcz\i;ko and Stewart 1976); the Nearctic distribution and unique south- paraprocts are reduced, not curving up western relict populations of the species. over tergum 10 as in most Isoperla. The Stark and Gaufin (19763") have subse- female subgenital plate is broadly round- quently pointed out the existence of a ed, sometimes with a shallow notch, and is complex of several species which are cur- much produced, covering most of the ster- rently under revision. Louisiana materials num 9. Nymphs have distinctive mouth- examined as part of this revision reveal parts. Their mandibles are deeply cleft, the presence of a second form herein des- with serrations on the inner margin of ignated "S])ecies A." Vaught and Stewart the outer tooth. The lacinia bears a single fl974) detailed the life history and ecolo- apical tooth, with setae present along the gy of A^. clymene based on a study of entire inner margin, continuing arotmd Texas populations. Accounts and illustra- the outer margin; paraglossae bear a dis- tions of adult morphology were given by tinct apical nipple. The emergence of /. Stark and Gaufin (1976a). Nymphs were mohri is spontaneous, apparently cued figured by Frison (1935) but are not defi- to water temperature, and lasts usviallv nitely associated with this species. Photo- less than two weeks in late March to mid- graphs of eggs definitely of this species April. Frison ('1935) figured the geni- are found in Vaught and Stewart (1974) talia, nymph, and n\TTiphal mouthparts. and Stark and Gaufin (1976a). Details of Szczytko and Stew^art (1976) gave a de- the chorion and collar are shown in Fig. tailed analysis of the species and figured 20. 378 GREAT BASIN NATURALIST Vol. 36, No. 3

Neoperia clymene

Neoperia sp. A Paragnetina kansensis P immarginata P fumosa Phasganophora capitata

Fig. 17. Distribution of Perlinae in Louisiana.

Neoperia sp. A et al. (1974) discussed its distribution,

Distribution in Louisian.a: Grant, near Pollock. which extended westward to the Black- Livingston, Little Natalbany R. Vernon, Ft. land Prairie of Texas. Two })opulation Polk. centers have been located in Louisiana, This is a smaller and more darkly pig- one in the Kisatchie National Forest and mented form than typical clymene. The the other in the florida f)arishes (Fig. 17). two species broadly overlap in range over Ricker (1949) figured the achdt genitalia. the Gulf Coastal Plains and have been Nymphs are undescribed but, based on our taken together in the same light trap col- reared material, may be distinguished lection in Livingston Parish (Fig. 17). from kansensis by the color variation Eggs dissected from gravid females are noted in the key. Like kansensis, female shown in Figs. 21 and 22. The chorion is nymphs j)()ssess anal gills. No published finely and regularly punctate, which con- data are available on the life history or trasts markedly with the striate pattern behavior of this species. The egg (Fig. of typical clymene (Fig. 20). 23) is oval with a distinctly stalked collar. The lip of the collar is bent outward and Paragnetina fumosa (Banks) has irregular, sharp emarginations; chorionic reticulation is not evident. Perla fumosa Banks, 1902: 123. Perla imniarginata Ncedham & Claassen. 1925: 102. Paragnetina immarginata (Say) Distribution in Louisiana: Rapides. Castor Cr.;

Germany Br.; Hemphill Cr.; Spring Cr. ; St. Sidlis imrnarpinatn Say. 1823: 164. Helena, .5 mi E Chipola. Tangipahoa, Terry Prria lurida Hagen. 1861: 21. Cr. Tangipahoa-St. Tammany. Tchefuiuta Dlstuihution in Louisiana: Ori.e.\ns, New Or- R. Washington, Silver Cr.; Bogue Cliitto R.; leans (Fig. 17). Hays Cr.; 'j mi K Franklin; J( t. Hwy. 10-62; 3 mi S Jtt. 25-58 near Clifton. The record of this species in Louisiana This is the common Paragnetina spe- is based on Plagen's Perla lurida type cies of the (iulf (Coastal Plains. vStewart specimens in the MCZ and on a similar Sej)tenil)er U)7(i STKWAHT KT AL: ST()NKPLI1> ()!• LOl'ISIANA 379

specimen mentioned by Ricker (1949) Acroneuria abnormis (Newman) among Klapalek's material. No recent ma- Perla abnormis Newman. 1838: 177. terial of this Appalachian species has been Perla sonans Newport, 1851: 449. taken in the state; and. based on its pres- Acroneuria eidrnanni Samal, 1933: 96. ent distribution and habitat [ireference of Distribution in Louisiana: Grant, Big Cr. cool mountain streams, it is doubtful that Lasaele, Trout Cr. Livingston, Little Natal- any will bo forthcoming. bany R.; Tickfaw R. Rapides. Brown Cr.; Cas- tor Cr.; Hemphill Cr. T.^ngip.mio.a. Chapepeela Cr.; Tangipahoa R.; Terry Cr. Tangipahoa-St. Tammany, Tcliefuncta R. Washington, Sil- Paragnetina kansensis (Banks) ver Cr.; Tchefuncta R.

Per la kansensis Banks, 1905: 56. This is the most widely distributed Togoperla Sp. A Frison. 1935: 414. Acroneuria in North America; however, Distribution in Louisi.\na: Livingston, Tickfaw R. St. Helena, Amite R. Tangipahoa, Tangi- in Louisiana it is somewhat of a rarity. pahoa R. Washington, Rogue Chitto R.; Hays The distribution shown in Figure 18 re- Cr.; Pushpatapa Cr. flects .several specific locations but most of these collections consist of This species is generally distributed one or two over the midwestem states and south to specimens. Stark and Gaufin (1976b) the Gulf Coastal Plains. Present Louisi- presented illustrations of male and female ana records are from the florida parishes genitalia and eggs; Frison (1935) figured the Biological avail- (Fig. 17), and it has not been reported nymph. data are not west of the Mississippi in the southern able for the species. part of its range in other recent studies (Stewart et al. Stark 1974; and Stewart Acroneuria arenosa (Pictet) 197^). Adult genitalia were figured by Frison (1937) and the nymph by Frison Perla arenosa Pictet. 1841: 178. Perla pennsylvanica Rambur, 1842: 456. (1935). Female nymphs of this species Perla trifuncta Walker, 1852: 153. differ from Frison's description in haying Distribution in Louisiana: Be.\uregard, Bun- small tuft anal gills. life a of The cycle dick Cr. Grand, 3 mi NW Drv Prong, near is unknown. Pollock. Lasalle, Trout Cr. Livingston, Tickfaw R. N.-vtchitoches, Middle Cr. Rapides, Brown Cr.; Castor Cr.; Germany Branch; Phasganophora capitata (Pictet) Hemphill Cr.; Indian Cr. Tangipahoa, Big Cr.; Chapepeela Cr.; Natalbany R,; Terry Cr. Perla capitata Pictet, 1841: 214. W.-^shington, Little Silver Cr.; .5 mi E Enon; Perla Iristis Hagen, 1861: 22. Jet. Hwy. 10-62. Prrla annulipes Hagen. 1861: 22. Perla flavescens Walsh, 1862: 363. This species ranges from Pennsylvania Perla hieroglrphica Provancher, 1876: 211. down the Atlantic Coast to Florida and Perla niarginipes Provancher, 1876: 212. westward in the Gulf Coastal Plains to Perla americana Banks, 1900: 243. (Stark Gaufin 1976b). In Perla: illustris Banks. 1908: 256. Texas and Perla innota Banks, 1918: 6. Louisiana, it has been taken in a variety Harrisinla nigriscens Banks. 1948: 119. of streams ranging from small creeks to Harrisiola klapaleki Biuiks. 1948: 121. medium-sized rivers. Populations are Harrisiola rnodesta Banks. 1948: 121. in the florida parishes and in the Distribution in Louisiana: Tangipahoa, Tangi- centered Kisatchie National Forest area (Fig. 18). pahoa R.; Terrv Cr. ; 1.5 mi E Amite (Fig. 17). Illustrations of the male and female geni- talia and the egg are presented by Stark This species is widely distributed over and Gaufin (1976b); the nymph is in- the eastern United States and Canada but distinguishable from Frison's (1935) has not been found west of the Mississippi Acroneuria sp. A illustration. No data are River in the southern part of its range. available on the biology of this species. Although common in much of the Gulf I Coastal Plains area, the species is a rarity in Louisiana. Illustrations of adult geni- Acroneuria evoluta Klapalek talia, nymphs, and eggs are given by Frison (1935) and Stark and Gaufin Acroneuria evoluta Klapalek. 1909. Acroneuria arida Frison (Not Hagen), 1935: 395. (1976a). Harper (1973a) presented data Acroneuria prolonga Claassen, 1937: 42. on the emergence, oviposition, hatching, Distribution in Louisiana: Orleans. New Or- and growth of the species in Canada. leans (Fig. 18). ;

380 GREAT BASIN NATURALIST Vol. 36, No. 3

A Acroneuria abnormis A. arenosa n A. evoluta # Perlinella drymo • P ephyre

Fig. 18. Distribution of Acroneurinae (excluding Perlesta) in Louisiana.

The name evoluta has been applied by ingston, Little Natalbanie R.; Magnolia; Prison (1947) to a midwestern species Tickfaw R.; Natchitoches; Kistachie Cr.; Mid-

dle Cr.; Winn Cr. Rapides, Brown Cr. ; Castor that ranges south into the Oklahoma Cr.; Chenywinche Cr.; Germany Branch; Ozarks. It is doubtful if this midwestern Roaring Cr.; Spring Cr. 2 mi S Calcasieu. form is the same as the type specimen Red River, Grand Bayou. S.'\bine, Blackwell which came from New Orleans. Stark C; Crib Cr.; 3 mi N Many. St. Helena, Amite R. Tangipahoa, Chapepeela Cr.; Natal- and Gaufin (1976b) suggest Klapalek's banie R.; Tangipahoa R.; Tcliefuncta R.; Terry evoluta types may be synonymous with Cr.; Wilson Branch. Vernon, Comrade Cr; Prison's mela; however, this specimen Fort Polk; Whiskey Chitto Cr. Washington, needs to be examined before the matter Bogue Chitto R.; Hays Cr.; Little Silver Cr.; Mill Cr.; Silver Cr.; Tchefuncta R. Webster, can be settled. Sawsman Cr.

This is the most abundant and widely Perlesta placida (Hagen) distributed Louisiana stoiicfly (Fig. 19). Perla placida Hagen, 1861: 28. Nymphs have been collected in gravel, Perla decipiens Walsh, 1862: 364. leaf litter, and debris; adults have been Perla brunnpippnnis Wnhh. 1862: 367. taken in May and June by sweeping vege- Chloroperla uirf^inica ]inuks, 1898: 199. Perlinella cinctipes Hanks, 1905: 56. tation or at light traps. Illustrations of Isoperla texana Banks, 1914: 611. male and female genitalia are jiresented Perlesta costalisYAixY^nXaV, 1921: 150. by Prison (1935), Stewart et al. (1969), Perlesta virginica inimaculata Klapalek, and Stark and Gaufin (1976a). Prison ,1921: 150. Perlesta placida nit ida 'Ranks, 1948: 115. (1935) figured the nym])h and com- Distribution in Louisiana: Beauregard, Bayou mented on its carnivorous food habits. Anococo. BiENVii.i.E, urniainod stream. Bossier, Photomicrographs of eggs were given in 9 mi R Red River. Caihx). ^ mi N .fct. 525-169. Stewart et al. (1969) and Stark and Gau- Cataiioula, Tributary Biids (>'r. Desoto, ^ mi fin (1976a). The egg (Pig. 23) is oval E Mansfield, 4 mi N Logansport. ChiANT, Fisli Cr.; Hardtner; Pollock. Lasai.i.e, Hemphill Cr. with a distinctly stalked collar. The lip of Trout Cr. Lincoln, Bayou D'Arbonne. Liv- the collar is l)ent outward and has ir- September 1976 stewaht kt al: stoneflies or Louisiana 381

A Perlesta placida

Fig. 19. Distribution of Perlesta placida in Louisiana.

re^lar sharp emar^nations. Chorionic re- tion and food habits of this species. The ticulation is not evident. No data are avail- egg (Fig. 25) is oval, lacks chorionic re- able on the growth cycle of this species; ticulation, and has a small, elegantly however, the appearance of onh' very stalked collar topped by a massive mush- small nymphs from November to January room-shaped anchor plate. suggests a summer egg diapause and very rapid spring growth in southern latitudes. Perlinella ephyre (Newman) Stewart et al. (1969) discussed the mating Chloroperla ephyre Newman. 1839: 87. behavior of Texas adults. Perla producta Walsh, 1862: 365. A toperla consors Banks, 1948: 128. Distribution in Louisiana: St. Helena, Amite Perlinelln drymo (Newman) R. Washington, Bogue Chitto R.; Harp Cr.; Pushpatapa Cr. Isogenus drymo Newman, 1839: 86. Perla elongata Walsh. 1862: 366. This species is widely distributed across Perla trivittala Banks, 1895: 313. the American midwest and (iulf Coastal Distribution in Louisi.\na: Vernon, unnamed Plains. We have collected it only in the stream. W.^shington, Bogue Chitto R.; Little florida parish portion of Louisiana (Fig. Silver Cr.; Jet. Hwy. 10-62. 18), and it has not been taken west of the This widely distributed eastern species Mississippi in the southern part of its is rare among collections from Louisiana range. Details of male and female geni- at present (Fig. 18). Zwick (1971) talia were figured by Zwick (1971); the figured male and female genitalia, and nymph was illustrated by Frison (1935). Stewart et al. (1974) reviewed the south- No biological data are available for this western distribution. Stark and Gaufin species. (1976a) figured male and female geni- Acknowledgment talia, eggs, and selected characters of the m-mph. Prison (1930) figured the whole We are greatly indebted to Dr. Joan nymph and discussed the microdistribu- B. Chapin, Louisiana State University at ..

382 GREAT BASIN NATURALIST Vol. 36, No. 3

Figs. 20-25. Photomicrographs of stonefly eggs: 20, Neoperla clymene, Etowah River. Georgia; 21, Neoperla sp. A, Blackwater River. Florida; 22, Neoperla sp. A, Blackwater River, Florida; 23. Parag- netina fumosa, Jasper, Texas; 24, Perlesta placida, Etowali River. Georgia; 25, Perlinella drymo, Blackwater River, Florida.

Baton Rouge, for the loan of specimens Nearctic neuropteroid . Trans. Amer. for study; and to Dr. Richard W. Bau- — 1902. Notes and descriptions of Perli mann, Brigham Young University, who dae. Canadian Entomol. 34: 123-125. provided specimens, collection records, — 1905. New genera and species of Perli and gave much helpful advice during dae. Psyche 12: 55-57.

. Neuropteroid insects: notes and manuscript preparation. We extend many 1908. descriptions. Trans. Amer Entomol. Soc. 34: thanks to Stanley W. Szczytko who ex- 255-267.

amined all hopcrla collections and to Dr. - -. 1914. Perlidae. In: New Tieuropteroid Roy E. Rhame for help with fioldwork. insects native and exotic. Proc. Acad. Natur. Sci. Philad. 66: 608-611.

1 Bull. . 1918. New neuropteroK insects. I^ITERATIJRK ClTED Mus. Comp. Zool. 62: 3-22. - neuropteroid — . ,1920. Perlidae. In: New Mus. Com. Zool. 64: Banks, N. 1895. New neuropteroid insects. insects. Bull. Harvard Trans Amer. Entomol. Soc. 22: 313-316. 314-325. Perlidae. Psyche 55: — . 1948. Note; . 1897. Perlidae. In: New North Anieri can neuropteroid insects. Trans. Amer. Ento- 11 3- MO. \\ l<>21'. New s|)ecies of Noiih mol. Soc. 24: 21-22. Ci.AASSKN. American Capniidae (PIccoptera"). Canadian -. 1898. Perlidae. In: Descriptions of new- 54-57. North Amer. neuropteroid insects. Trans, Entomol. 56: 43-48. of America. Amer. Entomol. Soc. 25: 199-200. . 1931. Plecoptera nvMii)lis Thomas Say Foundation. !'>'> pp. . 1900. New genera and species of September 1976 STI'.WAKT ET AL: STONRFIJES OF LOITISIATVA 383

. 10S6. No\v tuniios for stoneflios fPlo- Nf.beker. a. V. 1971. Effect of water tempera- copternV Ann, Entnninl. Soc. Amer. 29: 622- ture on nymphal feeding rate, emergence, 623. and adult longevifv of tlie stonefly Ptern-

— . 1937. New spories of sfonoflios fPlo narcYS dnrsnta. .1. Kansas Entomol. Soc. 44: coptera). J. Kniis. Enfomol. Soc. 10: 42-'5l. 21-26.

. 1940. A catalog of tho Plecoptera of Needii.am. J.. .AND P. Cr.A.AssEN. 192'i. A mono the world. Mom. Conipll Univ. Agric. Expt. graph of the Plecoptera or stoneflios of Sta. 252: 1-23^. America north of Mexico. Thomas Say Foun- Fni.<;oN. T. H. 1929. iFall and winter stoneflies. dation. 397 pp. or Plecoptera of Illinois. Illinois Natur. Hist. Nelson. C. .and J. Hanson. 1971. Contribution Surv. Bull. 18: 343-409. to the anatomy and phylogony of tho family

. 1934. Four new species of 'Stoneflios Pteronarcidae. Trans. Amer. Entomol. Soc.

from Nort.h America f Plecoptera V Canadian 97: 123-200. Entomol. 66: 2*^-30. Newman. E. 1838. Entomological notes. Ento-

. 193*1. Tlie stoneflios. or Plecoptera. of mol. Mag. '5: 168-181. 372-402. 483-'500. Illinois. Illinois Natur. Hist. Surv. Bull. 20: 1839. On the synonymy of Perlites. 277-471. together with brief characters of the old. and

— . 1942. Studies of North American Ple- a few new species. Ann. Mag. Natur. Hist. coptera. Illinois Natur. Hist. Surv. Bull. 22: 3: 32-37, 84-90. 23'i-3'5'^. Newport. G. .18'51. On the anatomy and af- G.\uFiN. A. B. 19-if). Ati annotated list of the finities of PteronarcYs regalis. Trans. Linn.

stoneflios of Ohio (Plecoptera). Ohio ,T. Sci. Soc. London. 20: 42'5-4'52.

'^6: 321-324. PiCTET. F. .1. 1841. Histoiro naturolle generale Gf.rst.\eckkr. a. 1873. tlber P/rrnnarcys et particulioro des insoctes Nevropteros. Per- Newn. und eine zweite. im Imago-stadium lides: 1-423. mit Trackoonkiemon versehene Perlarien Provancher. L. 1876. Petite faune entomolo- Gattung. In: Yau- l\Tor[)hologie dor Orthoptora giquo du Canada. I.es Nevropteres. Natur. amphihiotica. Fostschr. Ges. Natin-f. Froundo: Canad. 8: 177-191. 209-218. 60-74. Bamrttr, J. P. 1842. Tribu des Perlides. In: HvGF.N. H. A. 1861. Svnopsis of the Nouroptora Histoiro naturelle des insectos. Nevropteres: of North America. 347 pp. 449-462. H\RDEN. P.. .AND C. MiCKEi.. '1952. Tlio stono- Bicker. W. E. 1949. The North American spe- flies of Minnesota (Plecoptera ). Univ. Min- cies of Parngnrtina. Ann. Entomol. Soc. nesota Agric. Expt. Sta. Tech. Bull. 201: Amer. 42: 279-288.

1-84. . 19'52. Svstematic studies in Plecoptera. Harper. P. P. 1973a. Emergence, reproduction, Ind. Univ. Publ. Sci. Ser. 18: 1-200.

and growth of setipalpian Plecoptera in . 1964. Distribution of Canadian .stone- southern Ontario. Oikos. 24: 94-107. flies Gewasser u. Abwasser 34/3*1: '50-71.

. 1973h Eifo histories of Nemouridao . 1972. Besearch and management. Key- and I-ouctridae in southern Ontario CPle- note session 101st. Ann. Meet. Trans. Amer. copteraV Hvdrohiologia. 41: 309-3'o6. Fish. Soc. 101: 150. Harper, P. P.. .and E. Magnin. 1969. Cycles Bicker. W. E.. and H. H. Boss. 1968. North vitaux de quolques Plecopteres des Lauren- American species of Taeniopteryx (Plecop-

tides rinsectesV Candaian .1. Zool. 47: 483- tera. InsectaV J. Fish. Bes. Bd. Canad. 25: 494. 1423-1439. H\RPER. P. P.. .AND H. B. N. Hynes. 1970. Dia- Boss, H. H. 1964. New species of winter stone- pause in the nvmphs of Canadian wintei- flies of the genus Allncapnia (Plecoptera. stoneflios. Ecologv '51: 92*1-927. Capniidae). Entomol. News. 75: 169-177.

. 1971. The nvmphs of the Taenionterv- . 1968. Morphology and taxonomic use gidao of eastern Canada. Canad T. Zool. 49: of Ephemeroptora eggs. Ann. Entomol. Soc. 941-947. Amor. 61: 696-721. Hitchcock. S. W. 1974. Tho Plecoptera or Boss, H. H.. AND W. E. Bicker. 1971. The stoneflios of Connecticut. Connecticut Geol. classification, evolution, and dispersal of the and Natur. Hist. Surv. Bull. 107: 1-262. winter stonefly genus Allncapnia. Illinois Hoke. G. 1924. The anatomv of the head and Biol. Monogr. 45: 1-166. mouth-parts of Plecoptera. T. Morphol. 38: Samai.. J. 1933. Beitrage zur K(>initnis der Plo- 347-38'=;. copteronfauna Sudlabradors. Zool. Anz. 102:

Trr.TES. .1. 196f). Kataloc dof rezonton Plecoptoia. 95-100. Das Tierroirh 82: 1-632. Say. T. 1823. Descriptions of insects belonging T\MEs. A. M. 1974. Four now species of stone- to the order Nouroptora Linn, and Latr. col- flios in North America rPleconteraV Ann. En lected bv tho expedition under the command tomol. ,Soc. Amer. 67: 964-966. of Maior Long. Godman's Western Quart. Klap.at.ek. F 1900 Bevision dor Gattune Arrn- Bep. 2: 160-165. nciiria Pict. Bull. Intern. .A( ad. Sci. Bolieino Stark. B P.. and A. B. Gatjfin. 1976a. The 14: 234-247. Nearctic genera of Perlidae (Plecoptera). press.) . 1921. Plecopteres nouvoaux. Ann. Soc. Misc. Pub. Entomol. Soc. Amor. (In

Entomol. Belgique. 61: '57-67. 146-1'i0. 320. . 1976b. The Nearctic species of Acro- 327. neuria (Plecoptera: Perlidae). Tour. Kans. Kntght. a. W.. A. V Nereker. and A. B. G.au- Entomol. Soc. (In press.) FIN. 196'>. Fuither descrintions /-"f ''" ""--c Stark. B. P.. and K. W. Stewart. 1973. Distri- of Plecoptera of western T"^nitod States. Ento- bution of stoneflios (Plecoptera) in Oklahoma. mol. News. 76: 233-239. .Tour. Kansas Entomol. Soc. 46: 563-577. 384 tJREAT BASIN NATIIHALIST Vol. 36, No. 3

Stewart, K. W., G. L. Atmar, and B. M. Solon. Szczytko, S. W. and K. W. Stewart. 1976. 1969. Reproductive morphology and mating Three new sepcies of Nearctic Isoperla (Ple- behavior of Perlesta Placida (Plecoptera: coptera). Great Basin Nat. 56: 99-108. Perlidae). Ann. Entomol. Soc. Amer. 62: Walker, F. il852. Perlides. In: Catalogue of the 1433-1438. specimens of Neuropterous insects in the col- Stewart, K. W., R. W. Baumann, and B. P. lection of the British Museum. 1: H6-192. St.^rk. 1974. The distribution and past dis- Walsh, B. D. 1862. Perla. In. List of the persal of southwestern United States Ple- Pseudoneuroptera of Illinois. Proc. Acad. coptera. Trans. Amer. Entomol. Soc. 99: Natur. Sci. Philad.: 363-567. 507-546. ZwicK, p. 1971. iNotes on the genus PerlineUa Stew.\rt, K. W.. and S. W. Szczytko. 1974. A and a generic synonymy in North American new species of Taeniopteryx from Texas (Ple- Perlidae. Fla. Entomol." 54: 315-320.

coptera: Taeniopterygidae). Jour. Kansas En- . 1973. Phylogenetisches Systems und tomol. Soc. 47: 451-458. Katalog. Das Tierreich 94: 1-465.