Isoperla, OF

Sandberg, John B. & Kondratieff, Boris C., 2013, The Isoperla Of California (Plecoptera: Perlodidae); Updated Male Descriptions And Adult Keys For 18 Western Nearctic Species, Illiesia 9 (5), pp. 34-64 : 37-39

Keys to the Isoperla of California (including I. rainiera , I. tilasqua and I. umpqua )

Males

1 Pronotum checkered black on yellow; aedeagus without sclerotized process; paraprocts abruptly tapering to blunt apical tips, usually secondarily barbed ventrally; vesicle a broadly rounded lobe, wider at base then apex ( Figs. 11 a-d) ..…... I. pinta

1’ Pronotum without checkered pigment pattern, aedeagus, paraprocts and vesicle variable …..... 2

2 Aedeagus without posterior sclerotized process ( Figs. 1 , 3 , 9-12 , 15 , 17 ) ………..……....…………. 3

2’ Aedeagus with variable posterior sclerotized process ( Figs. 2 , 4-8 , 13-14 , 16 , 18 ) ………...……. 9

3(2) Vesicle absent or weakly produced ( Figs. 9d , 15d ).………………...….………….……………… 4

3’ Vesicle present and variably shaped ( Figs. 1d , 3d , 10d , 13d , 18d ) …………..……..………………….. 5

4(3) Aedeagal posterior margin with a median patch of spinulae ( Figs. 15 a-c); apex usually with two small, pointed lobes ( Figs. 15 a-c) …………………………………………….. I. sobria

4’ Aedeagal posterobasal margin with a median patch of spinulae concentrated into two distinct longitudinal bands ( Figs. 9 a-b); and a second posteroapical patch of spinulae above posterior lobe ( Figs. 9a, c ) …………………….….. I. miwok

5(3) Ninth tergum with short stout spinulae (sometimes arranged in a bipartite patch) or ninth and tenth terga with distinct patches of long stout setae near posterior margins ( Figs. 1c , 10c , 12c ) …………………………….…………………....….. 6

5’ Ninth and tenth terga without spinulae, long setae usually scattered ……………...………..….. 8

6(5) Vesicle rounded apically, base broader than apex, occasionally appearing constricted at middle; paraproct length if straightened, subequal to combined length of first two cercal segments, tapering gradually to blunt apices; posterolateral margins of ninth and tenth terga with patches of long stout setae, fewer and smaller on tergum ten ( Figs. 12 a-d) ……………...…... I. quinquepunctata

6’ Vesicle truncated or rectangular apically; paraproct length and shape variable; ninth and tenth terga with scattered fine setae, ninth tergum with mesal patches of short stout spinulae ( Figs. 1 , 10 ) ………………………………………...…...... 7

7(6) Vesicle rectangular, length half as long as wide ( Fig. 1d ); apical 1/3 of aedeagus tube-like, directed ventrally, covered with minute spinulae and a posterior pair of finer spinule patches divided or joined apically, apex a complex nipplelike tip ( Figs. 1 a-c); paraproct length if straightened greater than combined length of first and second cercal segments, tapering gradually to sharp apices ( Fig. 1c ) …………………..… I. acula

7’ Vesicle trapezoid shaped, about as long as wide ( Fig. 10d ); apical 1/2 of aedeagus tube-like, directed posteriorly, covered with minute spinulae and concentrated somewhat above bilobed tip ( Figs. 10 a-c); paraproct length if straightened subequal to combined length of first two cercal segments, tapering gradually to blunt apices ( Fig. 10c ) ..................................... I. mormona

8(5) Abdomen yellow; when everted live and fixed in hot water, aedeagus with one large patch of spinulae concentrated below posterobasal lobe ( Fig. 3b ), a long thin patch along posteroapical margin ( Fig. 3c ), apex bulbous, deeply inverted, and without a pair of short apicolateral lobes; when cleared in KOH, apex with a single inverted tube-like process ( Fig. 20a ), CA-Domingo Spring …………………………...……...…...... I. baumanni

8’ Abdomen dark; when everted live and fixed in hot water, aedeagus with one large patch of spinulae concentrated below posterobasal lobe ( Fig. 17b ), a long thin patch along posteroapical margin ( Fig. 17c ), apex bulbous, deeply inverted, and usually with one pair of short sharp apicolateral lobes near apex (similar to I. sobria ); when cleared in KOH, apex with two inverted tube-like processes ( Fig. 20f ), OR, WA …………………………………………... I. tilasqua

9(2) Vesicle a broadly rounded lobe, wider at base then apex ( Figs. 6d , 8d , 14d , 16d ) …………...…. 10

9’ Vesicle pedunculate, width subequal to length, constricted near base with curved lateral margins, and apex variable ( Figs. 2d , 4d , 5d , 7d , 13d , 18d ) ……………..………………………………..….… 13

10(9) Aedeagus with at least one patch of stout spinules above sclerotized process ( Figs. 6a , 8b , 15a ); apex of sclerotized process rounded in lateral view ( Figs. 6a , 8a , 15a ), distal margin either thin or expanded in posterior view ( Figs. 6b , 8b , 15b ) ………………………………..………...…..… 11

10’ Aedeagus without a spinule patch above sclerotized process ( Fig. 16a ); apex of sclerotized process expanded, flattened and projecting dorsally in lateral view ( Fig. 16b ), bifurcate subapically with two ventrally directed rounded lobes in posterior view ( Fig. 16b inset), width of lobes 0.18-0.20 mm …………………..... I. sordida

11(10) A single patch above sclerotized process composed of fine pale spinules; apex of sclerotized process rounded in lateral view ( Figs. 8 a-b), distal margin thin and blade-like in posterior view ( Fig. 8b ) ….…………. I. marmorata

11’ Spinule patch either single or triple and usually dark ( Figs. 6c , 14c ); apex of expanded sclerotized process rounded in lateral view ( Figs. 6a , 14a ), distal margin expanded slightly in posterior view ( Figs. 6c , 14c ) …………………………………..... 12

12(11) A single spinule patch above sclerotized process ( Figs. 6a & c ) ..…………………… I. fulva

12’ One bi-hemispherical shaped spinule patch above sclerotized process and a pair of smaller rounded patches laterally ( Figs. 14 a-c) ……… I. roguensis

13(9) Aedeagal sclerotized process bifurcate in posterior view ( Figs. 4-5 ) …………...…………. 14

13’ Aedeagal sclerotized process not bifurcate in posterior view ( Figs. 2 , 7 , 13 , 18 ) ……………….. 15

14(13) Sclerotized process with long apically pointed bifurcate arms; length of projecting process greater than 0.5 mm ( Fig. 4a ); abdomen mostly brown ……………………..…………... I. bifurcata

14’ Sclerotized process with short rounded bifurcate lobes; length of projecting process less than 0.5 mm ( Figs. 5 a-b); width of bifurcate apicolateral lobes 0.09-0.10 mm; abdomen mostly yellow ………………………………………....... I. denningi

15(13) Posterior margin of aedeagus with elliptical shaped patch of long stout spinulae separated from and above sclerotized process; sclerotized process digitate, with nearly parallel dorsal and ventral margins in lateral view, apex narrowly rounded ( Figs. 7 a-c) ..……………….….. I. laucki

15’ Posterior margin of aedeagus without a patch of long stout spinulae; sclerotized process otherwise ( Figs. 2 , 13 , 18 ) ………….………………...…….... 16

16(15) Sclerotized process crescent shaped in dorsal and ventral views ( Figs. 13 b-c); paraprocts gradually tapering to sharp apices ( Fig. 13c ) OR, WA.………………………….………...…. I. rainiera

16’ Sclerotized process clavate; paraprocts tapering gradually to blunt apices; OR, CA ...….……… 17

17(16) Wings hyaline; aedeagus with one distinct lobe ( Fig. 18a ); sclerotized process length approximately 0.50 mm ( Figs. 18 a-c); mesoposterior area of 8th and 9th terga with bipartite patches of stout spinulae and long fine scattered setae ( Fig. 18c ), OR ...………... I. umpqua

17’ Wings tinted; aedeagus with four distinct lobes in lateral view ( Fig. 2b ); sclerotized process length approximately 0.25 mm ( Fig. 2b ), spinule patches absent on terga 8-9, CA ….…………….. I. adunca

Females

1 Subgenital plate usually with invagination along apical margin ( Figs. 19 b-c, 19l-m, 19q-s) .........…. 2

1’ Subgenital plate usually not invaginated ( Figs. 19a, 19 d-k, 19n-p) ..…………………………….… 9

2(1) Subgenital plate with deep median invagination, the apex 3–4 times wider than depth ( Figs. 19b, 19r ) ………………………………........ 3

2’ Subgenital plate with shallow median invagination, the apex 8–10 times wider than depth ( Figs. 19c, 19 l-m, 19q, 19s) ..…………….... 4

3(2) Subgenital plate length less than half the width at base ( Fig. 19b ) …………...…………..… I. adunca

3’ Subgenital plate length approximately half the width at base ( Fig. 19r ) ………..…….… I. tilasqua

4(2) Subgenital plate apex truncated, with nearly parallel lateral margins ( Fig. 19c ); deflected ventrally in lateral view, CA-Domingo Spring ……………………………………...…. I. baumanni

4’ Subgenital plate apex rounded and not deflected ventrally in lateral view ( Figs. 19 l-m, 19q, 19s) ………………………………………………........... 5

5(4) Subgenital plate width at base approximately half the width of segment 8 and nearly twice that of the subgenital plate length ( Fig. 19m ) ………………………………...... I. quinquepunctata

5’ Subgenital plate width at base greater than half the width of segment 8 ( Figs. 19l, 19q, 19s ) …… 6

6(5) Subgenital plate lateral margins broadly rounded ( Fig. 19q ) ………….…….…… I. sordida

6’ Subgenital plate with short, nearly parallel lateral margins ( Figs. 19l, 19s ) ………………….…..……. 7

7(6) Pronotum with checkerboard pattern ( Fig. 12d inset) …………………………...……………. I. pinta

7’ Pronotum without checkerboard pattern …….... 8

8(7) Apical margin of subgenital plate sometimes invaginated (broadly rounded in Fig. 19j ), apex reaches approximately mid-length of ninth sternum (CA) ………………… I. miwok (in part)

8’ Apical margin of subgenital plate typically invaginated ( Fig. 19s ), apex not reaching midlength of ninth sternum (OR) ………... I. umpqua

9(1) Subgenital plate with lateral margins nearly parallel ( Figs. 19j, 19p ) ........................................ 10

9’ Subgenital plate with lateral margins not nearly parallel ( Figs. 19a, 19 d-i, 19k, 19n-o) ................... 11

10(9) Subgenital plate width nearly as wide as segment 8 ( Fig. 19j ) …………...…………. I. miwok

10’ Subgenital plate width distinctly less than width of segment 8 ( Fig. 19p ) ……..……...…… I. sobria

11(9) Subgenital plate triangular ( Figs. 19g, 19o ) …..………………….. I. fulva , I. roguensis (in part)

11’ Subgenital plate rounded ( Figs. 19a, 19 d-g, 19h-i, 19k, 19n) ………….…………………..……….…. 12

12(11) Subgenital plate width at base approximately 4-5 times that of the length ( Figs. 19 d-f, 19i, 19n) ………………………………………….……….... 13

12’ Subgenital plate width at base approximately 2-3 times that of the length ( Figs. 19a, 19h, 19k ) …………………………………………………...... 16

13(12) Subgenital plate completely suffused with dark pigment ( Fig. 19n ) ………………. I. rainiera

13’ Subgenital plate partially patterned with pigment or completely light ( Figs. 19 d-f, 19i) ……….…. 14

14(13) Subgenital plate partially patterned with pigment ( Figs. 19d, 19 f-g).…. I. bifurcata , I. fulva

14’ Subgenital plate completely light ( Figs. 19e, 19i ) …………………………………………………...... 15

15(14) Subgenital plate width at base approximately 3-4 times the length ( Fig. 19e ) ………… I. denningi

15’ Subgenital plate width at base 6-7 times the length ( Fig. 19i ) …………………….……… I. marmorata

16(12) Subgenital plate width at base approximately 3 times as wide than length; apex slightly truncated or broadly rounded ( Fig. 19k ) ...…… I. mormona

16’ Subgenital plate width at base approximately twice that of the length ( Figs. 19a, 19h ) …..…. 17

17(16) Subgenital plate darkly pigmented ( Fig. 19h ) ……………………………………..……………. I. laucki

17’ Subgenital plate not darkly pigmented ( Fig. 19a ) ……………………………………………... I. acula

Figures

Figs. 11-12. Adult male characters of Isoperla pinta, Butte Cr., Butte Co., CA and I. quinquepunctata, Yuba R., Yuba Co., CA & Nehalem R., Clatsop Co., OR, 32-50x. Fig. 11a. I. pinta aedeagus, lateral. Lobes A: posterobasal, B: posteroapical lobe, C: anterolateral pair. Fig. 11b. Aedeagus, ventral. Fig. 11c. Paraprocts (arrow). Fig.11d. Vesicle; prothoracic pigment pattern (inset-dorsal view). Fig. 12a. I. quinquepunctata aedeagus, lateral. Lobes A: posterobasal, B: posteroapical, C: dorsal, D: anterior. Fig. 12b. Aedeagus, ventral. Fig. 12c. Paraprocts (black arrow); abdominal spinules (white arrows). Fig. 12d. Vesicle.

Figs. 1-2. Adult male characters of Isoperla acula, Deadman Cr., El Dorado Co., CA and I. adunca, Campbell Cr. & Trib., Butte Co., CA, 32-50x. Fig. 1a. I. acula aedeagus, lateral; fine spinule patch (arrow). Fig. 1b. Aedeagus, ventral; fine spinule patches (arrow & inset-posterior view). Fig. 1c. Paraprocts, (black arrow); abdominal spinules (white arrow). Fig. 1d. Vesicle; brush setae (inset). Fig. 2a. I. adunca aedeagus, lateral; sclerotized process (arrow & inset-lateral view). Lobes A: posterobasal, B: posteromedian, C: anterior, D: dorsomedian pair. Fig. 2b. Aedeagus, ventral; sclerotized process (arrow & inset-posterior view). Fig. 2c. Paraprocts (arrow). Fig. 2d. Vesicle.

Figs. 3-4. Adult male characters of Isoperla baumanni, Domingo Springs, Plumas Co., CA and I.bifurcata, Mosquito Spring, Plumas Co., CA, 25-50x. Fig. 3a. I. baumanni aedeagus, lateral; large posterobasal spinule patch & thin apical patch (arrows). Lobes A: posterobasal, B: posteromedian. Fig. 3b. Aedeagus, posterior; large basal spinule patch (arrow).Fig. 3c. Paraproct (white arrow); apical thin spinule patch (black arrow). Fig. 3d. Vesicle. Fig. 4a. I. bifurcata aedeagus, lateral. Lobes A: posterobasal, B: posteromedian, C: dorsal, D: anterior lobe.Sclerotized process characters 1: base, 2: median arm, 3: bifurcate arms, 4: subapical barb & apical tine. Fig. 4b. Aedeagus, ventral. Fig. 4c. Paraproct (arrow). Fig. 4d. Vesicle

Figs. 9-10. Adult male characters of Isoperla miwok, Campbell Cr., Butte Co., CA and I. mormona, Big Chico Cr., Butte Co., CA, 40-50x. Fig. 9a. I. miwok aedeagus, lateral; posteroventral pair and posteroapical spinule patches (arrows). Lobes A: posterior, B: dorsal. Fig. 9b. Aedeagus, ventral; posteroventral pair of spinule patches (black arrow), small pointed apicolateral lobe (white arrow). Fig. 9c. Paraprocts (white arrow); posteroapical spinule patch (black arrow). Fig. 9d. Vesicle. Fig. 10a. I. mormona aedeagus, lateral; paraproct (black arrow); fine dorsoapical spinulae (white arrow). Lobe Adorsoanterior. Fig. 10b. Aedeagus, ventral; paraproct base (arrow). Fig. 10c. Paraprocts (black arrow); fine dorsoapical spinulae (left white arrow); abdominal spinulae (right white arrow). Fig. 10d. Vesicle.

Figs. 15-16. Adult male characters of Isoperla sobria, Tyee Spring, Skamania Co., WA and I. sordida, Cold Water Cr., Mono Co., CA, 25-40x. Fig. 15a. I. sobria aedeagus, lateral (Tyee Sp., in copula); spinule patch (arrow). Lobes A: anterior, B: dorsal pair, C: posteroapical (aberrant). Fig. 15b. Aedeagus, posterior; spinule patch (arrow). Fig. 15c. Paraprocts (black arrow), Lk. Davis-Corral Sp.; spinulae patch, lateral (white arrow); dorsal pair of lobes in typical position. Fig. 15d. Vesicle. Fig. 16a. I. sordida aedeagus, lateral; sclerotized process (white arrow); paraproct (black arrow). Lobes A: posteroventral, B: posteromedian, C: dorsal, D: anterior. Fig. 16b. Aedeagus ventral; sclerotized process (arrow & inset-posterior view).Fig. 16c. Paraprocts (black arrow); sclerotized process (white arrow). Fig. 16d. Vesicle.

Figs. 17-18. Adult male characters of Isoperla tilasqua, Meacham Cr., Umatilla Co., OR and I. umpqua, Muir Cr., Douglas Co., OR, 25-40x. Fig. 17a. I. tilasqua aedeagus, lateral; large basal spinule patch & thin apical patch (arrows). Lobes A-B: posterior. Fig. 17b. Aedeagus, posterior; large basal spinule patch (arrow). Fig. 17c. Paraprocts (white arrow); apical thin spinule patch (right arrow); large basal patch (left arrow). Fig. 17d. Vesicle. Fig. 18a. I. umpqua aedeagus, lateral; sclerotized process (arrow). Lobe A: dorsoanterior. Fig. 18b. Aedeagus, ventral; sclerotized process (arrow & insetposterior view). Fig. 18c. Paraproct (left white arrow); sclerotized process (right white arrow); abdominal tergite 9 spinulae (black arrow). Fig. 18d. Vesicle.

Figs. 13-14. Adult male characters of Isoperla rainiera, Tyee Spring, Skamania Co., WA and I. roguensis, Butte Cr., Butte Co., CA & Mad R., Humboldt Co., CA, 32-50x. Fig. 13a. I. rainiera aedeagus, lateral; sclerotized process (arrow). Lobes A: posterior, B: dorsal, C: anterior. Fig. 13b. Aedeagus, ventral; sclerotized process (arrow).Fig. 13c. Paraprocts (black arrow); sclerotized process (white arrow). Fig. 13d. Vesicle. Fig. 14a. I. roguensis aedeagus, lateral; sclerotized process (white arrow); spinule patches (black arrows). Lobes A: posterior, B: dorsal, C: anterior. Fig. 14b. Aedeagus, ventral (Mad R.); sclerotized process (arrow); inset-posterior view (Butte Cr). Fig. 14c. Paraprocts (left arrow); sclerotized process (right arrow). Fig. 14d. Vesicle.

Figs. 20a-f. The cleared aedeagus of six western Nearctic Isoperla species after live evertion; membranous opening of internal tube like process or apex (white arrows); tube like internal process (black arrows). 50x (20b, d & e, cropped and enlarged). 20a. I. baumanni, lateral view, partially everted; 20b. I. fulva, lateral view; 20c. I. marmorata, lateral view; 20d. I. miwok, dorsoapical view, tube like process openings along anteroapical margin; 20e. I. roguensis, ventral view, tube like process openings along anteroapical margin; 20f. I. tilasqua, lateral view.

Figs. 5-6. Adult male characters of Isoperla denningi, San Mateo Cr., San Diego Co., CA, and I. fulva, Sand Cr., Larimer Co., CO & Pole Cr., Juab Co., UT, 32-50x. Fig. 5a. I. denningi aedeagus, lateral; sclerotized process (arrow & inset-lateral view). Lobes A-B: posterobasal, C: posteroapical, D: dorsolateral pair. Fig. 5b. Aedeagus, ventral; sclerotized process (arrow & inset-posterior view). Fig. 5c. Paraprocts (arrow). Fig. 5d. Vesicle. Fig. 6a. I.fulva aedeagus, lateral; sclerotized process (left white arrow & inset-lateral view); spinule patch (right white arrow & inset); paraprocts (inset-black arrow). Lobes A: posterior, B: dorsal, C: anterior. Fig. 6b. Aedeagus, ventral; sclerotized process (arrow & inset-posterior view). Fig. 6c. Aedeagus, dorsal; sclerotized process (black arrow); spinule patch (white arrow). Fig. 6d. Vesicle.

Figs. 7-8. Adult male characters of Isoperla laucki, Dragsaw Spring, Humboldt Co., CA, and I. marmorata, Big Chico Cr., Tehama Co., CA, 32-40x. Fig. 7a. I. laucki aedeagus, lateral; sclerotized process (white arrow & inset-lateral view); posteroapical and posterobasal spinule patches (black arrows). Lobes A: posterior, B: dorsal, C: anterolateral pair. Fig.7b. Aedeagus, ventral; sclerotized process (white arrow & inset-dorsal view); posterobasal spinule patch (black arrow). Fig. 7c. Paraprocts (left white arrow); sclerotized process (right white arrow); posteroapical spinule patch (black arrow). Fig. 7d. Vesicle. Fig. 8a. I. marmorata aedeagus, lateral; sclerotized process (white arrow & inset-lateral view); paraprocts (black arrow). Lobes A: posterior, B: dorsal, C: anterior. Fig. 8b. Aedeagus, posterior; spinule patch & sclerotized process (black arrow). Fig. 8c. Paraprocts (arrow). Fig. 8d. Vesicle.

Figs. 19a-s. Isoperla subgenital plates, 50x. Horizontal arrows = subgenital plate width at base, vertical line (two lines when subgenital plate apex invaginated) = subgenital plate length, horizontal line = invagination width at apex. 19a. I.acula; 19b. I. adunca; 19c. I. baumanni; 19d. I. bifurcata; 19e. I. denningi; 19f. I. fulva, rounded; 19g. I. fulva, triangular; 19h. I. laucki; 19i. I. marmorata; 19j. I.miwok; 19k. I. mormona; 19l. I. pinta; 19m. I. quinquepunctata; 19n. I. rainiera; 19o. I. roguensis; 19p. I.sobria; 19q. I. sordida; 19r. I. tilasqua; 19s. I. umpqua.