Nemourinae, Billberg, 1820

Grubbs, Scott A. & Baumann, Richard W., 2023, The Nemourinae (Insecta, Nemouridae) of the eastern Nearctic, Zootaxa 5306 (1), pp. 1-53 : 5

I. Adult key to eastern Nearctic Nemourinae genera

(modified from Baumann 1975 and Stewart & Stark 2008)

1 Males .............................................................................................. 2

- Females............................................................................................. 9

2 Cervical gills present, either unbranched or 4-branched ( Figs. 159 , 163 ); wing venation typical for family ( Fig. 8 ).... Zapada

- Cervical gills absent; wing venation variable ( Figs. 1, 3, 7 ).................................................... 3

3 Veins A 1 and A 2 of forewing joined near posterior margin ( Fig. 7 ); epiproct either slightly ( Fig. 105 ) or markedly bilaterally asymmetric ( Fig. 115 )............................................................................ Soyedina

- Veins A 1 and A 2 of forewing not joined ( Figs. 1‒6 ); epiproct bilaterally symmetric ( Figs. 31 , 44 , 61 )................... 4

4 Vesicle absent from sternum 9; terminal costal crossvein of forewing joins Sc vein before junction with R vein, typical “X” pattern at cord absent ( Fig. 3 ).................................................................. Paranemoura

- Vesicle present on sternum 9; terminal costal crossvein of forewing joins R vein, typical “X” pattern at cord present ( Figs. 1‒2, 4‒6 )................................................................................................ 5

5 Cerci sclerotized and developed, either with lateral spines ( Figs. 10‒12 ) or elongated and recurved ( Figs. 37‒38 )......... 6

- Cerci simple and unmodified ( Figs. 51 , 86 )................................................................. 7

6 Cerci robust with paired appressed spines (= as; Figs. 9‒13 ) and one outer spine (= os), the latter may be apically forked ( Fig. 13 ) or not ( Figs. 12 )............................................................................ Nemoura

- Cerci elongated and medially arcuate, apically pointed or bifurcated ( Figs. 19 , 25 , 31 , 37 ).................... Ostrocerca

7 10 th tergum enlarged at lateral proximal corners into long, sclerotized spiny lobes ( Figs. 95‒96 )................... Shipsa

- 10 th tergum unmodified at lateral proximal corners ( Figs. 45 , 161 )............................................... 8

8 Epiproct long and narrow throughout length, comprised mainly of the ventral sclerite, ( Figs. 61 , 69 , 77 , 85 ); dorsal sclerite reduced to lateral arms ( Figs. 67 , 82 , 88 ) or essentially absent ( Fig. 74 )..................................... Prostoia

- Epiproct short and robust, ventral sclerite T-shaped and expanded laterally ( Figs. 56‒57 ); dorsal sclerite prominent and terminates as bifurcated lateral arms ( Figs. 57‒58 ).................................................... Podmosta

9 Cervical gills present, either unbranched or 4-branched ( Figs. 155 , 159 ) wing venation typical for family ( Fig. 8 ).... Zapada

- Cervical gills absent; wing venation variable ( Figs. 1, 3, 7 ) ... 10

10. Veins A 1 and A 2 of forewing joined near posterior margin ( Fig. 7 )......................................... Soyedina

- Veins A 1 and A 2 of forewing not joined ( Figs. 1‒6 ).......................................................... 11

11 7 th sternum covering at least ¾ of 8 th sternum ( Figs. 17‒18 )............................................. Nemoura

- 7 th sternum less produced, covering <½ of 8 th sternum ( Figs. 23 , 75 , 98 )......................................... 12

12 Terminal costal crossvein of forewing joins Sc vein before junction with R vein, typical “X” pattern at cord absent ( Fig. 3 )......................................................................................... .. Paranemoura

- Terminal costal crossvein of forewing joins R vein, typical “X” pattern at cord present ( Fig. 1‒2, 4‒6 )................. 13

13 8 th sternum enlarged and modified as subgenital plate; 7 th sternum with a medial, nipple-like projection ( Figs. 24 , 30 , 35 ) (except O. truncata , Fig. 42 ).................................................................... Ostrocerca

- 8 th sternum not enlarged, only as medial subgenital plate or medial sclerotized pattern; 7 th sternum without medial, nipple-like projection ( Figs. 60 , 84 , 98 )............................................................................ 14

14 Subgenital plate apex broad, encompassing at least 50% of the total width of the 8 th sternum, varying from scarcely ( Figs. 68 , 84 ) to moderately ( Figs. 75 , 93 ) emarginate medially................................................... Prostoia

- Subgenital plate apex comparatively smaller, encompassing much less than 50% of the total width of the 8 th sternum ( Figs. 60 , 98 )................................................................................................ 15

15 Subgenital plate with a distinct medial notch ( Fig. 59‒60 ).............................................. Podmosta

- Subgenital plate lacking a distinct medial notch, rounded distally ( Fig. 98 ).................................... Shipsa

Figures

FIGURES 157‒160. Zapada chila. 157, USA, Tennessee, Little Pigeon River, male terminalia, dorsal view; 158, USA, Tennessee, Little Pigeon River, epiproct, lateral view; 159, USA, Tennessee, Little Pigeon River, cervical gills, ventral view; 160, USA, Tennessee, Little Pigeon River, subgenital plate, ventral view. c = cercus, ds = dorsal sclerite, lg = lateral gill, ma = membranous area, mg = medial gill, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 161‒164. Zapada fumosa. 161, USA, Virginia, Lewis Fork, male terminalia, dorsal view; 162, USA, Virginia, Lewis Fork, epiproct, lateral view; 163, USA, Virginia, Lewis Fork, cervical gills, ventral view; 164, USA, Virginia, Lewis Fork, subgenital plate, ventral view. c = cercus, ds = dorsal sclerite, imb = inner medial branch, lb = lateral branch, ma = membranous area, mmb = medial medial branch, omb = outer medial branch, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 1‒8. Forewing images. 1, Nemoura arctica, USA, Michigan, Silver Creek; 2, Ostrocerca albidipennis, USA, West Virginia, Middle Fork Indian Run; 3, Paranemoura perfecta, USA, West Virginia, headwaters Sugar Creek; 4, Podmosta macdunnoughi, USA, New York, Grampus Brook, dark line at tips of arrows represent the termini of veins A1 and A2; 5, Prostoia completa, USA, Kentucky, Middle Fork Drakes Creek; 6, Shipsa rotunda, USA, Michigan, Montreal River; 7, Soyedina calcarea, USA, Kentucky, tributary Green River; 8, Zapada katahdin, Canada, Labrador, tributary Leg Lake.

FIGURES 105‒108. Soyedina amicalola. 105, USA, Georgia, Frogtown Creek. epiproct, ventral view; 106, USA, Georgia, Frogtown Creek, epiproct, lateral view; 107, USA, Georgia, Frogtown Creek, epiproct, caudal view; 108, USA, Georgia, Frogtown Creek, outer paraproct lobe. ds = dorsal sclerite, vs = ventral sclerite, im = inner member, opl = outer paraproct lobe.

FIGURES 115‒122. Soyedina carolinensis. 115, USA, West Virginia, Marsh Fork of Buckhannon River, epiproct, ventral view; 116, USA, North Carolina, Rattlesnake Springs, epiproct, ventral view; 117, USA, North Carolina, Robin Branch, epiproct, lateral view; 118, USA,North Carolina, Robin Branch, epiproct, caudal view; 119, USA, North Carolina, Rattlesnake Springs, outer paraproct lobe, lateral view; 120, USA, Maryland, Fishing Creek, outer paraproct lobe, lateral view; 121, USA, Virginia, seep into Lewis Fork, outer paraproct lobe, lateral view; 122, USA, North Carolina, tributary Connelly Creek, female, abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, im = inner member. vs = ventral sclerite.

FIGURES 31‒36. Ostrocerca prolongata. 31, male abdominal terminalia, dorsal view; 32, Canada, Quebec, tributary to Lac Croche, epiproct, dorsal view; 33, Canada, Quebec, tributary to Lac Croche, epiproct, lateral view; 34, Canada, Quebec, tributary to Lac Croche, epiproct, ventral view; 35, USA, West Virginia, Seneca Creek, female abdominal terminalia, ventral view; 36, USA, West Virginia, Gandy Creek, female abdominal terminalia, ventral view. bc = basal cushion, c = cercus, ds = dorsal sclerite, hy = hypoproct, ipl = inner paraproct lobe, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 43‒48. Paranemoura claasseni. 43, male abdominal terminalia, lateral view; 44, USA, Maine, Crystal Brook, epiproct, dorsolateral view; 45, male abdominal terminalia, dorsal view; 46, Canada, New Brunswick, Williams Brook, epiproct, dorsal view; 47, female abdominal terminalia, ventral view; 48, Canada, New Brunswick, Williams Brook, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, ep = epihook, vs = ventral sclerite.

FIGURES 61‒68. Prostoia completa. 61, USA, Wisconsin, Plover River, epiproct, dorsal view; 62, male abdominal terminalia, lateral view; 63, USA, Wisconsin, Plover River, epiproct, lateral view; 64, USA, Wisconsin, Plover River, epiproct apex, dorsolateral view; 65, USA, Wisconsin, Plover River, epiproct apex, dorsal view; 66, epiproct base, dorsal view, USA, Virginia, Roanoke River; 67, Canada, Newfoundland, Walsh River, epiproct base, lateral view; 68, USA, Kentucky, Trammel Fork, female, abdominal terminalia, ventral view. c = cercus, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 9‒16. Nemoura arctica. 9, USA, Wyoming, West Glacier Lake, male abdominal terminalia, lateral view; 10, USA, Wyoming, West Glacier Lake, male abdominal terminalia, dorsal view; 11, USA, Iowa, Twin Springs, epiproct, dorsal view; 12, Canada, New Brunswick, Coombs Brook, male cercus, distal view; 13, USA, Iowa, Twin Springs, male cercus, distal view; 14, Canada, New Brunswick, Coombs Brook, epiproct, lateral view; 15, USA, Iowa, Twin Springs, epiproct apex; 16, Canada, New Brunswick, Coombs Brook, epiproct apex. ap = apical prong, as = appressed spine, bc = basal cushion, c = cercus, ds = dorsal sclerite, la = lateral area, os = outer spine, vs = ventral sclerite.

FIGURES 37‒42. Ostrocerca truncata. 37, male abdominal terminalia, dorsal view; 38, Canada, Ontario, Clare River, male abdominal terminalia, dorsal view; 39, Canada, Ontario, Clare River, epiproct, lateral view; 40, male abdominal terminalia, ventral view; 41, USA, New York, Cascadilla Creek, female abdominal terminalia, ventral view; 42, USA, West Virginia, headwaters Davis Creek, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, hy = hypoproct, ipl = inner paraproct lobe, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 49‒54. Paranemoura perfecta. 49, male abdominal terminalia, lateral view; 50, Canada, Nova Scotia, Silvey Brook, epiproct, dorsolateral view; 51, male abdominal terminalia, lateral view; 52, Canada, Nova Scotia, Silvey Brook, epiproct, dorsal view; 53, female abdominal terminalia, ventral view; 54, USA, West Virginia, Glady Fork, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, ep = epihook, vs = ventral sclerite.

FIGURES 85‒92. Prostoia similis. 85, USA, Virginia, McClure River, epiproct, dorsal view; 86, male abdominal terminalia, dorsal view; 87, male abdominal terminalia, lateral view; 88, USA, Virginia, McClure River, epiproct, lateral view; 89, USA, Virginia, McClure River, epiproct apex, lateral view; 90, USA, Virginia, McClure River, epiproct apex, dorsal view; 91, USA, Virginia, McClure River, epiproct base, caudal view; 92, USA, Virginia, Big Run, epiproct base, dorsal view. c = cercus, la = lateral arm, vs = ventral sclerite.

FIGURES 19‒24. Ostrocerca albidipennis. 19, male abdominal terminalia, dorsal view; 20, USA, New York, Oquaga Creek, male abdominal terminalia, dorsal view; 21, USA, New York, Oquaga Creek, epiproct, dorsal view; 22, USA, New York, Oquaga Creek, epiproct, ventral view; 23, female abdominal terminalia, ventral view; 24,USA,Ohio, tributary Pike Run, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, ipl = outer paraproct lobe, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 25‒30. Ostrocerca complexa. 25, male abdominal terminalia, dorsal view; 26, USA, New York, Stoney Creek, epiproct, dorsal view; 27, USA, New York, Stoney Creek, epiproct apex, dorsolateral view; 28, USA, New York, Stoney Creek, epiproct, caudal view; 29, female abdominal terminalia, ventral view; 30, USA, West Virginia, tributary Hills Creek, female abdominal terminalia, ventral view. bc = basal cushion, c = cercus, ds = dorsal sclerite, hy = hypoproct, ipl = inner paraproct lobe, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 95‒98. Shipsa rotunda. 95, USA, Virginia, Meherrin River, male abdominal terminalia, dorsal view; 96, USA, Virginia, Meherrin River, male abdominal terminalia, ventral view; 97, Canada, Ontario, Skootamatta River, female abdominal terminalia, ventral view; 98, Canada, Quebec, Riviere du Diable, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, la = lateral arm, sgp = subgenital plate, sl 10 = spiny lobes of 10th tergum.

FIGURES 69‒76. Prostoia hallasi. 69, USA, Virginia, Washington Ditch, epiproct, dorsal view; 70, male abdominal terminalia, lateral view; 71, USA, Virginia, Washington Ditch, epiproct, lateral view; 72, USA, Illinois, tributary to Alcorn Creek, epiproct apex, lateral view; 73, USA, Virginia, Washington Ditch, epiproct apex, lateral view; 74, USA, Virginia, Washington Ditch, male abdominal terminalia, caudal view; 75, female abdominal terminalia, ventral view; 76, USA, Virginia, 1 mi SE Dunnsville, female abdominal terminalia, ventral view. c = cercus, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 77‒84. Prostoia ozarkensis. 77, USA, Arkansas River, epiproct, dorsal view; 78, USA, Oklahoma, Baron Fork, epiproct, dorsal view; 79, USA, Arkansas River, epiproct, lateral view; 80, USA, Oklahoma, Baron Fork, epiproct apex, lateral view; 81, USA, Oklahoma, Baron Fork, epiproct apex, lateral view; 82, USA, Oklahoma, Baron Fork, epiproct base, caudal view; 83, USA, Illinois, Hutchins Creek, epiproct base, caudal view; 84, USA, Illinois, Hutchins Creek,, female abdominal terminalia, ventral view. c = cercus, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 55‒60. Podmosta macdunnoughi. 55, Canada, Labrador, male abdominal terminalia, dorsal view; 56, Canada, Quebec, Jacques Cartier River, epiproct, dorsal view; 57, Canada, Quebec, Jacques Cartier River, epiproct, dorsal view; 58, Canada, Quebec, Jacques Cartier River, epiproct, lateral view; 59, Canada, Labrador, female abdominal terminalia, ventral view; 60, Canada, Nova Scotia, Roop Brook, female abdominal terminalia, ventral view. c = cercus, ds = dorsal sclerite, la = lateral arm, sgp = subgenital plate, vs = ventral sclerite.

FIGURES 151‒156. Soyedina washingtoni. 151, USA, Maine, Thorndike Brook, epiproct, ventral view; 152, USA, Maine, Thorndike Brook, epiproct apex, ventral view; 153, USA, Maryland, spring into Elk Lick Run, epiproct, lateral view; 154, USA, Pennsylvania, Fourmile Creek, epiproct, caudal view; 155, USA, Maryland, Mill Run, outer paraproct lobe, lateral view; 156, USA, Maryland, spring into Blue Lick Run, female, abdominal terminalia, ventral view. ds = dorsal sclerite, im = inner member, opl = outer paraproct lobe, vs = ventral sclerite.

FIGURES 17‒18. Nemoura arctica. 17, female abdominal terminalia, ventral view; 18, USA, New Hampshire, Lakes of the Clouds, female abdominal terminalia, ventral view. c = cercus, sgp = subgenital plate.

FIGURES 93‒94. Prostoia similis. 93, female abdominal terminalia, ventral view; 94, USA, West Virginia, Little Coal River, female abdominal terminalia, ventral view. sgp = subgenital plate.