Ecclisomyia simulata Banks 1920

Givens, Donald R., 2018, The Nearctic Ecclisomyia species (Trichoptera: Limnephilidae), Zootaxa 4413 (2), pp. 201-259 : 241-254

Ecclisomyia simulata Banks 1920

Figs. 30a–30c ; 31a–31c; 32a–32c; 33a, 33b; 34a, 34b; 35a, 35b; 36a–36d; 40; 41; 42; 43d.

Ecclisomyia simulata Banks, 1920 , 346, figs. 101, 106. Holotype male, M.C.Z. 10,876, Reno, Nevada (H.K. Morrison); Ross (1950, 425, fig. 15, 15a, b, male genitalia, distribution); Denning (1951, 161, 162, distribution); Schmid (1955, 59, E. simulata as synonym of E. maculosa ); Newell & Minshall (1977, 255, distribution); Ruiter et al. (2014, 27, E. simulata as valid species, distribution); Givens (2014, 160, 170, distribution).

Male diagnosis. Inferior appendages of E. simulata and E. maculosa each have 2 stout spines arising from the mesobasal area, with 1–4 smaller black stout spines lateral of these 2 basal spines. The inferior appendages of E. conspersa each have a black, stout spine ( Figs. 10a , 11b ); the inferior appendages of E. bilera are without spines ( Figs. 1a, 1b ). Ecclisomyia simulata has the distal area of the fused parameres of the phallus with a deep U- to Vshaped notch ( Figs. 30b , 31b ); that of E. maculosa has a shallow U- to V-shaped notch ( Figs. 22b , 23b ). Inferior appendages of E. simulata are longitudinally short, versus long in E. maculosa .

Male description. See Banks (1920); Ross (1950). A supplementary description follows: Length 11–14 mm (N = 15). Head, thorax, legs brown. Wings uniformly brown; hyaline; patterned with whitish spots. Head with 3 ocelli, lateral ocelli closer to eyes than to midline. Antennae each with about 52 segments; except for segments 1 (scape) and 2 each segment with 4–7 stout black to translucent spines positioned midway; spines sometimes closer to distal area of antennal segment; dorsal surfaces clothed with numerous fine black setae. Tergum VIII brown with numerous long setae ( Figs. 30a , 31c ); sternum VIII brown, with numerous short setae ( Figs. 30a, 30b ). Pleura well developed, membranous. Superior appendages broader at anterior base, narrowing distally, rounded apically, bearing hair-like setae ( Figs. 30a, 30b ; 31a, 31b). Tergum IX nearly semicircular in lateral view longest mesolaterally, longitudinally short dorsally and ventrally; dorsal, ventral, and anterior margins heavily sclerotized ( Figs 30a , 31a ); left and right lateral portions connected dorsally and ventrally with transverse sclerotized bridges (straps) ( Figs. 30b, 30c , 31c ). Tergum X elongate, extending caudad, with many small spicules ( Fig. 30a ). Intermediate appendage ventral of tergum X sclerotized; distal margin with several elongate setae ( Figs. 30a , 31a ). Apices of parameres fused, with deep U-shaped notch apically forming 2 prong-like caudal extensions ( Figs. 30b, 30c ; 31b, 31c). Phallus with broad phallocrypt; narrowing posteriorly, curving dorsad. Inferior appendages triangular in lateral aspect ( Fig. 30a ); long hair-like setae on lateral and distal dorsal and ventral surfaces; each with 2 large stout black to reddish-brown spines arising from base and extending caudad ( Figs. 30b , 31b ). Lateral of these two black spines 2–4 shorter, dark brown to black spines ( Figs. 30b , 31b ).

Female diagnosis. The distal margin of tergum X has a U-shaped incision extending about 3/4 the length of the segment ( Figs 32a, 32b ); the distal margin of tergum X of E. maculosa has a V-shaped incision extending about 1/2–3/4 the length of the segment ( Figs. 24b , 25b ). The external part of gonopod VIII is large in both species, but in E. simulata it is longer than wide ( Figs. 32a, 32b ) whereas in E. maculosa it is nearly as wide as long ( Figs. 24c , 25b, 25c ) and in E. bilera it is short and narrow ( Figs. 3b , 4a ). In all females of E. simulata examined, the caudal margin of the external part of gonopod VIII is concave. The external part of gonopod IX has a mesal V-shaped incision, whereas in E. bilera it is concave in ventral aspect. The external gonopod of segment IX sclerotized, wide, not as deeply incised as in E. maculosa , only extending posterad 1 X its width ( Figs. 32a, 32b ); in E. maculosa the external gonopod of segment IX is narrow caudal margin with deep V-shaped declivity, extending posterad about 2 X its width beyond caudal margin of the external part of segment VIII ( Figs. 24c , 25b, 25b ). The spermathecal sclerite is quadrate in E. simulata ( Figs. 32a, 32b ) and not as long as in E. conspersa ( Fig. 13a ) nor as rounded as in E. bilera ( Fig. 3b ).

Female description. Length 11–14 mm (N = 15). Tergum X fused with tergum IX; viewed from dorsal aspect incised into 2 lobes, incision about 1/2–to 3/4 length of tergum. External gonopod of segment VIII large; caudal margin concave; lateral margins straight ( Figs. 32a, 32b ). External gonopod of segment IX sclerotized, divided mesally with V-shaped incision ( Figs. 32a, 32b ); fused anteriorly, forming dorsal surface of vaginal opening; extending beyond vaginal opening and 1 X its width posterad of external gonopod of segment VIII; lobes flared laterally; flanged in lateral aspect. Spermathecal sclerite compact, extending anterad about 1/2 length of abdominal segment VIII; about as wide as long, but sometimes slightly wider than long ( Figs. 32a, 32b ).

Pupal diagnosis. The E. simulata pupa may be separated from pupae of the other three Nearctic Ecclisomyia species by the presence of dorsal gills on abdominal segment IIa, absent in the others. The E. s imulata pupa may also be separated from the pupa of E. conspersa by the absence of lateral line gills that are present on abdominal segments IIp–IVa of the E. conspersa pupa ( Fig. 42 ). Each apical process of E. simulata has 2 stout, long, black setae at the apex ( Figs. 33a , 35a ), whereas those of E. conspersa each have 3 stout, long, black setae ( Figs. 15a , 17a ).

Pupal description. Length 10–12mm (N = 11). Head: Brown to light brown. Male antennae extending to apices of apical processes; female antennae extending to anterior margin of abdominal segment VI. Labrum light brown; semicircular posterior margin sinuate, rounded anteriorly, with 10 long black setae on dorsal surface, setae directed dorsad, distal end of each seta hooked ( Fig. 36a ). Mandibles ( Fig. 36b ) reddish-brown to reddish-black; bases of mandibles broad, each curving mesad, rapidly tapering to sharp single sickle-shaped tooth; mesal margin of each mandible with minute serrations difficult to discern with light microscope.

Thorax: Pro-, meso-, and metanota brown to yellowish-brown; pro-, meso-, and metathoracic legs light brown. Procoxae each with 1 long black setae near lateral margin and 1 short black seta on ventral surface. Mesocoxae with 1 black seta near lateral surface and infrequently with 1 short black seta on ventral surface. Pro- and metafemora each with single black spine on distal margin. Profemora each also with single short black spine on ventral surface near distal margin. Mesotarsal segments 1–4 each with pair of lateral fringes of long, fine, brownblackish to reddish to opaque fine silk-like hairs, directed posterad.

Abdomen: Dorsal abdominal segments light brown to reddish brown. Lateral fringe of long fine black hairs on each side directed dorsad; originating from posterior margin of abdominal segment V; line of fringe arching ventrad, extending to posterior margin of abdominal segment VIII. Dorsal and ventral abdominal gills each singlefilament; present dorsoposteriorly and ventroposteriorly on segments II–VII; present dorsoanteriorly on abdominal segments II–VII; present ventroanteriorly on abdominal segments III–VII ( Fig. 42 ). Dorsal process of abdominal segment I quadrate ( Fig. 36d ); shape of dorsal process varying with maturity of pupa; minutely wrinkled, with 2 sclerotized lobes lateral to longitudinal axis, each posterior lobe with 7–15 reddish black spicules along posterior margin. Pairs of hook plates present anteriorly on abdominal segments III–VII ( Figs. 36c , 40 , 41 ); posteriorly on segment V. Hook plates IIIa each with 3–5 hooks; IVa with 3–4; Va with 2–6; Vp with 15–21; VIa with 3–5, and VIIa with 4–6; reddish brown, hooks reddish black, directed posterad on anterior hook plates of segments III–VII and anterad on elliptical hook plates Vp. Dorsal abdominal segments I–VIII each with 1 pair of black sa 2 setae ( Fig. 36c ); segments IV–VII sometimes with 1 pair short, black sa 1 setae; segment VI sometimes with 2 or 3 setae along lateral margins; 1 or 2 setae lateral of sa 2 setae; segment VII with 2–4 black setae lateral of sa 2 setae; segment VIII with 3–5 black setae lateral of meson on distal margin. Venter of abdominal segments II–VIII with 1 pair of short, black sa 1 setae and 1 pair of black sa 3 setae; posterior area of segment VIII with black ss setae. Ventral abdominal setae variable; number and presence of setae not consistent. Distal area formed into 2 caudal lobes (apical processes); lobes of female curved apicomesad, mesal margins slanted mesad, caudal areas of lobes angulate, dorsal surfaces of lobes wrinkled basally; lateral margins of apical processes sinuate ( Figs. 35a, 35b ). Apical processes of males straight, directed caudad ( Figs. 33a , 34a ). Dorsal surfaces of apical processes with minute reddish to translucent spicules ( Fig.33a ). Caudal lobes each with 2 long, black, stout setae on distal margin ( Figs. 33a, 33b ; 34a, 34b, 35a, 35b), with lateral reddish spur on distal margin, and with 3 short, fine setae along mesal surface. Anterodorsal area of female apical processes with transverse row of long black setae ( Fig. 35a ). Dorsomesal area of male apical processes with 3–5 long, black setae lateral of meson ( Fig. 33a ).

Pupal case. Length 11–13 mm (N = 9), width at anterior end 4 mm; width at posterior end 3 mm (N = 12). Case compact, without open interstitial spaces; constructed of large pebbles and small grains of sand ( Fig. 43d ). Both ends of case rounded, less compact, with small open areas. Cases attached to rocks such that case orientation permitting flow of water through case.

Larval diagnosis. The E. simulata larva has single-filament gills dorsoanteriorly on dorsal abdominal segment II, whereas E. conspersa , E. bilera , and E. maculosa larvae typically lack these gills. The lack of lateral-line gills will also separate the E. simulata larva from the E. conspersa larva. The E. simulata larva has dorsoanterior and ventroanterior gills on dorsal abdominal segment VII, lacking in the E. maculosa larva ( Fig. 42 ). Some mature larvae (11 mm in length) lack dorsoanterior gills on abdominal segment II, however the gill arrangement for abdominal segment VII fits the pattern for the E. simulata larva. The E. simulata larva has a prosternal sclerite lacking in E. maculosa .

Larval description. Length (mature instar) 9–12 mm (N = 15). Head: Brown to dark brown to a blackish brown; mesal area along coronal suture with yellowish ellipsoid; muscle scars distinct lateral of coronal suture and posterior of primary setae 17. Parietals brown to dark brown, without secondary setae; muscle scars distinct posterior and lateral of eyes. Muscle scars distinct lateral of ventral ecdysical line. Mandibles black, each with 4 small blunt apical teeth. Mature larva sometimes with only 3 blunt teeth as mesal tooth sometimes worn down. One to two short translucent setae arising from posterolateral area near posterior margin of each mandible, difficult to discern; mesal margin of each mandible with minute serrations, difficult to discern, and with tuft of feather-like setae ( Fig. 19a ). Ventral apodome brown, triangular; in mature larvae 1.08–1.60 X longer than ecdysial line; in one Nevada specimen ecdysial line 0.87 X longer than ventral apotome. Primary setae in positions 7, 9–17 ( Fig. 19a ). Primary setae in position 7 reddish, long; primary setae in position 9 long, stout, reddish to black; primary setae in positon 10 short, translucent; primary setae in position 11 fine, short, reddish brown to translucent, difficult to discern, sometimes missing; primary setae in position 12 reddish, short; primary setae in position 13 short, translucent, semi-appressed, difficult to discern and best observed from lateral view; primary setae in position 14 reddish to black, long, stout; primary setae in position 15 reddish, long; primary setae in position 16 short, translucent, fine, semi-appressed, difficult to discern; primary setae in position 17 reddish to black, long. Venter of head with primary setae in positions 8 and 18 ( Fig. 19b ); setae short, translucent. Frontoclypeus smooth, posterior area without apparent muscle scars or muscle scars indistinct; without texture, glabrous, except for primary setae in positions 1–6 ( Fig. 19a ). Primary setae in positions 1–3 fine, long, translucent; primary setae in positions 4 and 5 long, translucent, appressed; primary setae in position 6 long, fine, translucent. Labrum brown, with primary setae in positions 1–6. Primary setae in positions 1 and 2 short, translucent, difficult to discern, best observed from ventral aspect; primary setae in positions 3 and 4 short, translucent; primary setae in positions 5 and 6 long, translucent. Fringes of hair-like setae present at anterolateral corners of labrum each forming sparse setal tuft. Cardo sclerites sub-rectangular. Submental sclerites each with 2 short black setae set lateral of meson.

Thorax: Pro- and mesonota brown to yellowish brown, sometimes blackish brown; nota with mesal yellow stripe; metanotal sclerites brown. Anterior margins of pronota each with 11–16 long, black slender setae set equidistant from each other along anterior margin; short translucent setae set between these setae along anterior margin. Each half of pronotum with transverse line of 8–14 black, long slender setae posterior of mid-dorsal transverse groove. Pronotum with black primary setae in sa 2 and sa 3 positions. Muscle scars on mesonotum distinct. Mesonotum with black setae in 3 distinct groups roughly corresponding to sa 1, sa 2, and sa 3 positions. Posterior marginal area of mesonotum black, heavily sclerotized forming narrow blackish brown band curving anterad on each side to lateral process. Lateral processes triangular, black. Metanotal sa 1 sclerites brown to light brown, triangular, each with 6–8 long black setae; metanotal sa 2 sclerites brown to light brown each with 6–10 black ls setae; metanotal sa 3 sclerites parenthesis-shaped, brown to light brown, each with 9–15 black setae on anterior third of sclerite. Venter of prothorax with small prosternal sclerite; sometimes only small sclerotized area present. Legs fuscus. Foretrochantins fuscus forming tent-like structures over procoxae. Pro-, meso-, and metacoxae with 6–8 black ls setae on ventral margin and 16–18 black ls setae on dorsal margin. Procoxae with primary setae in positions 1 and 2; meso- and metacoxae with primary setae in position 2. Meso- and metatrochanters with primary setae in positons 2, 3, and 5 ( Figs. 21b, 21c ). Ventral margins of pro-, meso-, and metatrochanters and pro-, meso-, metafemora with trochanteral brush, setae feather-like, translucent, may be difficult to discern ( Figs 21a, 21b, 21c ). Profemora with 2 long stout sl setae along ventral margin ( Fig. 21a ). Pro- and mesofemora with distinct short sl reddish to translucent setae along ventral margin; these setae sometimes also occurring along ventral margins of metafemora, but indistinct if present ( Figs. 21a, 21b, 21c ). Profemora with primary setae in position 2; meso- and metafemora with primary setae in positions 2, 3, and 4; setae sometimes absent in position 3. Mesofemora sometimes with 3–4 black ls setae on ventral margin. Pro- and mesopleura and pro-and mesoepimera brown.

2 – 5 3 – 5 3 – 6 13 – 26 3 – 6 3 – 7

I II III IV V VI VII VIII E. bilera Denning [based on 10 pupae examined]

3 – 4 2 – 5 3 – 5 10 – 22 3 – 5 2 – 5

I II III IV V VI VII VIII E. conspersa Banks [based on 7 pupae examined]

2 – 7 2 – 8 3 – 8 9 – 17 2 – 6 3 – 7

I II III IV V VI VII VIII E. maculosa Banks [based on 7 pupae examined]

3 – 5 3 – 4 2 – 6 15 – 21 3 – 5 4 – 6

I II III IV V VI VII VIII E. simulata Banks [based on 3 pupae examined]

41

Abdomen: Dorsal and lateral humps of abdominal segment I without any sclerites. Abdominal segment I with 16–28 black ls setae anterior to, and lateral of dorsal hump. Dorsal hump without sclerites; with 3–10 black ls setae. Lateral humps with 3–6 short black setae; these setae sometimes on 2 small sclerotized areas or setae with only basal sclerotization. Ventral base of each lateral hump with 6–13 short black setae, setal sockets prominent; sometimes setal bases sclerotized. Venter of abdominal segment I with pair of brown sclerites lateral of meson, each with 2–4 black ls setae; area anterior and lateral of each of these sclerites with 4–17 black ls setae. Dorsal and ventral abdominal gills each single-filament; present in mature larvae dorsoanteriorly on segments II–VII; ventroanteriorly on segments III–VII; dorsoposteriorly and ventroposteriorly on segments II–VII ( Fig. 42 ). Gill arrangement variable, depending on instar stage; some specimens 5 mm in length with only anterior gills on abdominal segments II–VII, and only few posterior gills; many segments without posterior gills. All larvae 3 mm in length possessing gills, but lacking full complement of gills observed in mature larvae. Dorsum of abdomen with black primary sa 2 setae on abdominal segments II–VIII; sa 2 setae on segments II–V sometimes difficult to discern; sa 2 on segments VI–VIII long and slender. Dorsum of abdominal segment VIII sometimes with additional short, black seta adjacent to each sa 2 setae. Abdominal segment IX with elliptical sclerite bearing 5–8 stout reddish-black ls setae spaced equidistant from each other along posterior margin; setae immediately adjacent to meson longer and stouter than other setae along posterior margin; surface of sclerite with transverse row of 6–8 fine black setae; areas lateral of sclerites each with 2–3 short, fine, black setae. Sternum glabrous, except for 2–4 long reddish-black setae on venter of segment IX. Lateral fringes of hl setae extending on each side from anterior of abdominal segment III to posterior of segment VII. Chloride epithelia present on venter of abdominal segments III–VII. Anal prolegs each with lateral sclerite brown; narrowing ventrally and forming thin strap, extending 3/4 distance to meson of abdominal segment IX, ending at first black seta on venter of segment IX; with basal tuft of 4 long reddish black setae, mesal 2 longer and thicker than lateral 2 setae; with 3–4 black ls setae on dorsoposterior margin; and 7–8 black setae on dorsal surface. Sole plates each bearing only single black seta on distoventral margin. Anal hooks each bearing smaller accessory hook (denticule); caudal margin with short, black setae and ventral margin with 1 short, black to translucent seta. Caudal lobes glabrous, except for minute sl tan to reddish spicules on either side of anal opening.

Distribution. Ecclisomyia simulata has been recorded in the Cascade Mountain range of Oregon and Washington ( Ross 1950) and the Sierra-Nevadas of California and Nevada ( Banks 1920; Ross 1950; Denning 1951; Givens 2014). I have collected and examined E. simulata from California, Idaho, Nevada, Oregon, and Washington.

Bionomics. Ecclisomyia simulata is a higher altitude species, with altitudes ranging from 1223–2225 m. Emergence begins in late April extending into mid-August. The creeks and streams in which larvae occur tend to be narrow, 0.6–3.05 m, however E. simulata larvae may also be found in larger and deeper streams, up to 4.6 m wide and 0.6 m deep. Typically, stream depth ranges from 5.08–30.5 cm. These streams are generally rapid, fast flowing, frequently turbulent, with cobble, gravel, and sand substrate. In common with the other Nearctic Ecclisomyia species, much instream woody debris often occurs. Ecclisomyia simulata has been reported sympatric with E. bilera and E. conspersa ( Givens 2014) .

Material Examined. CALIFORNIA: Butte Co., Humboldt Rd. above Colby Creek, N40.186 W121.825, 25- v-2014, 1 M (BCK and CJV) (CSUC). Fresno Co., Billy Creek, at Huntington Lake, 26-vi-2011, 1 M P, 1 F P, 1 P, 1 PP (D.J. Burdick) (CASC). Plumas Co., Benner Creek, 10.8 km NW Chester (Chester-Juniper Lake Rd.), Benner Creek Cmpgd., 25-vii-2012, 3 M, 1 F (DRG) (DRGPC). Shasta Co., Hat Lake, 02-vii-1947, 2 F P, 1 P (H.P. Chandler) (CASC); E. Fork Hat Creek, 18.3 km S of NW entrance to Lassen Volcanic Nat'l. Park, Dersch Meadows, (Lassen Park Rd.), 12-vii-2013, 1 M P, 1 L (DRG) (CSUC); E. Fork Hat Creek, 20.3 km S of NW entrance to Lassen Volcanic Nat'l. Park, (Lassen Park Rd.), 13-vii-2013, 1 L (DRG) (CSUC); W. Fork Hat Creek, 15 km S of NW entrance to Lassen Volcanic Nat'l. Park, (Lassen Park Rd.), 20-vii-2012, 3 P, 2 PP (DRG) (CSUC); 03-viii-2012, 6 M, 10 F (DRG) (CSUC); W. Fork Hat Creek, Paradise Meadows, Lassen Volcanic Nat'l. Park, 13- vii-2013, 1 M, 1 F (DRG) (CSUC); small creek, Paradise Meadows, Lassen Volcanic Nat'l. Park, 13-vii-2013, 1 PP (DRG) (CSUC); small creek, 1.2 km up trail to Paradise Meadows, Lassen Volcanic Nat'l. Park, (trailhead 15 km NW entrance to Lassen Volcanic Nat'l. Park), 13-vii-2013, 1 F P (DRG) (CSUC); small creek, 1.6 km up trail to Paradise Meadows, Lassen Volcanic Nat'l. Park, (trailhead 15 km NW entrance to Lassen Volcanic Nat'l. Park), 17- vii-2013, 1 F P (DRG) (CSUC); King's Creek, 26.5 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 20-vii-2012, 1 M (DRG) (CSUC); 01-viii-2012, 1 M (DRG) (CSUC); 13-viii-2012, 1 P (DRG) CSUC); 01-vii-2013, 2 M P, 1 P, 1 PP (DRG) (CSUC); King's Creek, King's Creek Picnic Site 28 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 04-viii-2012, 5 M, 4 F (DRG) (CSUC); 10-viii-2012, 1 M (DRG) (CSUC); 13-viii-2012, 1 M, 1 L (DRG) (CSUC); 28-vi-2013, 1 L (DRG) (CSUC); 19-vii-2013, 2 PP (DRG) (CSUC); King's Creek, 28.1 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 04-viii-2012, 1 M, 4 F (DRG) (CSUC); King's Creek, 26.8 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), King's Creek Upper Meadow, 28-vi-2013, 4 M P, 2 F P, 6 P, 1 PP (DRG) (CSUC); King's Creek, 27.5 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), King's Creek Upper Meadow, 11-vii-2013, 1 M P, 5 P, 6 L (DRG) (CSUC); Lost Creek, 11.9 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 26-vii-2012, 1 F (DRG) (CSUC); 03-viii-2012, 6 M (DRG) (CSUC); 14-vii-2013, 2 M (DRG) (CSUC); Manzanita Creek, 2.2 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 02-viii-2012, 4 M, 2 F (DRG) (CSUC). Sierra Co., Big Canyon Creek, 2.4 km E Yuba Pass, 26-vi-1973, 4 M (DRG and SDS) (DRGPC); 1.6 km E of Bassetts (Hwy. 49), 24-vi-2008, 1 F (PAO) (CSUC); 27-vi-2016, 1 M (PAO) (CSUC); 3.2 km E of Bassetts (Hwy. 49), 19-vi-2006, 4 M (PAO) (CSUC); 12-vii-2012, 3 M (PAO) (CSUC); 14-vii-2012, 11 M, 7 F (PAO) (CSUC); 09-vii-2012, 4 M (PAO) (CSUC); 3.2 km E of Bassetts, San Francisco State University, (SNFC), (Hwy. 49), 28-vi-1999, 2 M (PAO) (CSUC); 20-v-2000, 2 M, 1 F (PAO) (CSUC); 24-vi-2000, 12 M, 8 F (PAO) (CSUC); 26-vi-2000, 22 M, 20 F (PAO) (CSUC); 26-v-2006, 10 M, 2 F (PAO) (CSUC); 30-v-2006, 1 M (PAO) (CSUC); 14- vii-2012, 3 F (PAO) CSUC); N. Fork Yuba River, 1.6 km E of Bassetts (Hwy. 49), 23-vi-2002, 2 M (PAO) (CSUC); 17-vi-2003, 1 M, 1 F (PAO) (CSUC); 27-v-2006, 1 M, 1 F (PAO) (CSUC); 15-vii-2009, 1 M (PAO) (CSUC). Siskiyou Co., Castle Lake, 24-vi-2000, 2 M (J. Powell) (EMEC). Tehama Co., small creek 1.1 km S of SW entrance to Lassen Volcanic Nat'l. Park, 28-vi-2013, 8 M, 10 F (DRG) (DRGPC). Tuolumne Co. West side of Sonora Pass, 07-viii-1963, 3 M, 2 F (C.W. O’Brian) (EMEC). IDAHO: Cassia Co., Fourth Fork Rock Creek, Lower Penstemon Cmpgd., 01-viii-1990, 6 M, 3 F (R.W. Baumann) (BYUC); Franklin Co., Cub River Canyon S of Thomas Spring, 07-viii-1985, (G.W. Knowlton) (BYUC). NEVADA: Elko Co., Weeks Creek, 4.8 km NE on Weeks Creek Access Rd., (off State Rd. 232), Humboldt-Toiyabe Nat’l. Forest, N40.9433 W115.0793, 02-vii- 2016, 1 M (DRG) (DRGPC); 03-vii-2016, 1 M P (DRG) (DRGPC); Washoe Co., Third Creek to Ophir Creek diversion, ca. 0.8 km from Incline Lake along Hwy. 431, N39.3011 W119.92064, 14-vi-2013, 1 PP, 8 L (DER) (CSUC). White Pine Co., Baker Creek, Great Basin Nat'l. Park, 28-vii-2002, 1 M, 1 F (CSUC); 15-v-2015, 1 L (BCK) (CSUC); Lehman Creek, 11.3 km W Baker, 07-viii-1978, 1 M, 3 F (J. Powell) (EMEC); Lower Lehman Cmpgd., Great Basin Nat'l. Park, 14-vii-2014, 5 M (PAO) (CSUC); Upper Lehman Water Supply and Stream, Great Basin Nat'l. Park, 16-v-2015, 1 L (DER) (DRGPC); S. Fork Snake Creek, 16-v-2015, 12 L (DER) (DRGPC); Timber Creek, 6.8 km on Timber Creek Rd., (off State Hwy. 486), Humboldt-Toiyabe Nat’l. Forest, N39.461 W114.652, 27-vi-2016, 1 M P, 2 F P, 2 P, 5 PP, 16 L (DRG) (DRGPC). OREGON: Clackamas Co., Crater Creek (tributary to Clackamas River), 01-vi-2000, 170 L (RWW) (RWWPC); headwaters tributaries of Still Creek and West Fork Salmon River, (Old Timberline Rd.), Mt. Hood Zigzag Ranger District, Timberline Lodge Ski Area, 09- vii-2010, 1 M, 1 L (RWW) (RWWPC); tributary to Clear Fork of Sand River, (Jct. of Forest Service Rds. 10 and 18), 22-v-2014, 1 F P (RWW and DER) (DERPC). Deschutes Co., Squaw Creek nr. Falls, N44.167 W121.667, 09- viii-1986, 1 M (GWC) (RWWPC). Douglas Co., Diamond Lake, N43.183 W122.166, 25-vi-1987, 1 M (R. Roberts) (RWWPC); 02-vii-1987, 1 M (R. Roberts) (RWWPC). Harney Co., headwaters Big Indian Creek, N42.667 W118.583, 30-vii-1987, 3 M P, 4 F P, 24 P, 1 PP (RWW) (RWWPC). Hood River Co., Meadows Creek, Mt. Hood Meadows Ski Area, 14-vii-1998, 2 M (RWW) (RWWPC). Jackson Co., Mt. Ashland Bull Cmpgd., 26- vi-2000, 1 M (D. Rubinoff) (EMEC); Lower Gap Creek, Rogue River Nat’l. Forest, Ashland Ranger District, 40S 5W Sec. 34, 19-vi-2000, 2 M (RWW) (RWWPC); Upper McDonald Creek, Rogue River Nat’l. Forest, Ashland Ranger District, 10-vii-2000, 1 M, 1 F (RWW) (RWWPC); E. F. Ashland Creek, Rogue River Nat’l. Forest, Ashland Ranger District, 09-vii-2001, 17 M, 1 F (RWW) (RWWPC). Jefferson Co., Jack Springs, N44.47664 W121.72327, 12-v-2014, 2 M (CDK) (DERPC). Lane Co., Roaring River, N43.95 W122.083, 09-vii-1986, 1 M (GWC) (RWWPC); 22-viii-1986, 1 M (GWC) (RWWPC); Titbits Creek, (at first bridge upstream from Blue Lake), N44.24595 W122.36639, 24-vii-2012, 3 M, 1 F (CDK) (DERPC); Titbits Creek, ca. 4.8 km upstream from Blue River Lake, N44.24421 W122.30082, 24-vii-2012, 1 F (CDK) (DERPC). Linn Co., Lost Lake Creek, (Hwy. 20), 05-viii-1998, 1 M (S. Fitzgerald) (CSUC); Mann Creek, (County Rd. 15), N44.30371 W122.16827, 16-vii-2012, 1 M (CDK) (DERPC). Wallowa Co., small creek 35.2 km SE Jct. County Rd. 39 and Hwy. 350, (which is 12.8 km NE Joseph), Wallowa-Whitman Nat'l. Forest (Hells Canyon Rec. Area), 09-vii-2014, 1 M (DRG) (DRGPC). WASHINGTON: Columbia Co., N. Fork Touchet River, 30.6 km SW Dayton, (Touchet Rd.) (Jct. of USFS Rd. 64 and Touchet Corral Trail), Umatilla Nat’l. Forest, N46.093 W117.847, 29-v-2015, 28 F (DRG) DRGPC); N. Fork Touchet River, 29 km SW Dayton, (Touchet Rd). (Jct. of State Forest Rd. 9115 and USFS Rd. 65), Umatilla Nat’l. Forest, N46.118 W117.823, 30-v-2015, 7 F (DRG) DRGPC). Clallam Co., Moose Lake outlet, side pool, Olympic Nat’l. Park, Grand Valley, 13-vii-1987, 2 M (RWW) (RWWPC). Lewis Co., Ohamopecosh River, Ohamopecosh River Cmpgd., Mt. Rainier Nat’l. Park, 13-viii-1986, 1 M (Furnish) (RWWPC); small stream, nr. Narada Falls, 23- vii-2001, 1 M (RAL) (CSUC); Clear Fork Cowlitz River 26.5 km E of Packwood, (Hwy. 12), Gifford-Pinchot Nat’l. Forest, N46.629 W121.425, 20-iv-2015, 11 L (DRG) (DRGPC). Pierce Co., Paradise River, Paradise Lodge, Mt. Rainier Nat’l. Park, 13-viii-1986, 1 M (RWW) (RWWPC); Longmire, Mt. Rainier Nat'l. Park, 10-vii-1998, 1 M (RAL) (CSUC); tributary to Tahoma Creek, (Jct. of Westside Rd. and Hwy. 706), Mt. Rainier Nat'l. Park, 08- viii-1998, 5 M, 3 F (BCK) (CSUC); Paradise River, at fourth crossing, Mt. Rainier Nat'l. Park, 16-viii-1999, 1 M (BCK) (CSUC); boggy stream, tributary to Tahoma Creek, (Westside Rd.), Mt. Rainier Nat'l. Park, 17-viii-1999, 2 M, 6 F (BCK) (CSUC); tributary to Tahoma Creek, (Jct. of Hwy. 70 and Westside Rd.), Mt. Rainier Nat'l. Park, 08- vii-1999, 6 M, 3 F (BCK) (CSUC); tributary to Tahoma Creek, 3 km from Hwy. 706 (Westside Rd.), Mt. Rainier Nat'l. Park, 21-vi-2003, 4 M, 3 F (BCK, REZ and J. Schmidt) (CSUC); Fish Creek, at Westside Rd., Mt. Rainier Nat'l. Park, 14-vii-2000, 2 M (RAL) (CSUC); stream, 2.9 km N of Paradise Rd. on Westside Rd., Mt. Rainier Nat'l. Park, 01-vi-2001, 1 M (RAL) (CSUC); 17-vii-2001, 1 M (RAL) (CSUC); 2.7 km N of Paradise Rd., Mt. Rainier Nat'l. Park, 17-vi-2001, 1 M (RAL) (CSUC); 3 km N of Paradise Rd., Mt. Rainier Nat'l. Park, 09-vii-2001, 1 M (RAL) (CSUC); Saint Andrews Creek, (Westside Rd.), Mt Rainier Nat'l. Park, 14-vii-2002, 3 M (RAL) (CSUC); Wetland North 0.5 km E Tahoma (Hwy. 706), Mt. Rainier Nat'l. Park, 19-vi-2003, 1 F (BCK) (CSUC); Ghost Lake, 1 km N Jct. Hwys. 123 and 410, Mt. Rainier Nat'l. Park, 12-vi-2004, 1 M (M. Grove) (CSUC); 06-vii-2004, 1 M (M. Grove) (CSUC). Skagit Co., Bacon Creek, N48.65 W121.417, 30-vii-1986, 1 M (GWC) (RWWPC). Skamania Co., Clearwater Creek, Mt. Saint Helens, 01-viii-1986, 6 M, 1 F (Meyerhoff) (RWWPC); creek, nr. June Lake, Mt. Saint Helens, 12-vii-1997, 1 M (S. Fitzgerald and A. Foley) (CSUC); Snohomish Co., Snohomish River, N48.183 W121.183, 31-vii-1986, 1 M (GWC) (RWWPC). Yakima Co., American River, Pleasant Valley Cmpgd., 28-vii-2000, 1 M (D.L. Strenge) (UMSC); American River (Lodgepole Cmpgd.), 24-vii-2003, 21 M, 2 F (D. Strenge) (DERPC).

Figures

FIGURES 1a–1c. Ecclisomyia bilera Denning, male genitalia. 1a, left lateral; 1b, ventral; 1c, dorsal; Abbreviations: ifa = inferior appendage (paired); ima = intermediate appendage; IX = segment IX; par = paramere (paired); phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; IXt = tergum IX; Xt = tergum X.

FIGURES 3a–3c. Ecclisomyia bilera Denning, female genitalia. 3a, left lateral; 3b, ventral; 3c, dorsal. Abbreviations: e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; IXt = tergum IX; pl = pleuron (paired); spt.sc = spermathecal sclerite;ssc.o = spermathecal sclerite opening;VIIIs = sternumVIII; VIIIt= tergum VIII;Xt = tergum X.

FIGURES 4a–4c. Ecclisomyia bilera Denning, female genitalia. 4a, ventral; 4b, left lateral; 4c, dorsal. Abbreviations: e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; spt.sc = spermathecal sclerite; ssc.o = spermathecal sclerite opening; VIIIs = sternum VIII; Xt = tergum X.

FIGURES 10a–10c. Ecclisomyia conspersa Banks, male genitalia. 10a, left lateral; 10b, ventral; 10c, dorsal. Abbreviations: ifa= inferior appendage (paired); ima = intermediate appendage; IX= segment IX; par = parameres (paired); ph = phallus; phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; Xt = tergum X.

FIGURES 15a, 15b. Ecclisomyia conspersa Banks, male pupa,apical processes.15a, dorsal; 15b, ventral. FIGURES 16a, 16b. Ecclisomyia conspersa Banks, male pupa,apical processes.16a, dorsal; 16b, ventral.

FIGURES 17a, 17b. Ecclisomyia conspersa Banks, female pupa, apical processes.17a, dorsal; 17b, ventral. FIGURES 18a, 18b. Ecclisomyia conspersa Banks, female pupa, apical process.18a, dorsal; 18b, ventral.

FIGURES 19a, 19b. Ecclisomyia conspersa Banks, larval head. 19a, dorsal, with primary setae numbered; 19b, ventral, with primary setae numbered.

FIGURES 21a–21c. Ecclisomyia conspersa Banks, larva, right legs. 21a, prothoracic leg; 21b, mesothoracic leg; 21c, metathoracic leg.Abbreviations:ls = long slender setae; sl = spike-like setae; tb = trochanteral brush.

FIGURES 25a–25c. Ecclisomyia maculosa Banks, female terminalia. 25a, left lateral; 25b, ventral; 25c, ventral. Abbreviations: e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; spt.sc = spermathecal sclerite; ssc.o = spermathecalsclerite opening; VIIIs = sternum VIII; IXs =sternum IX; Xt = tergum X.

FIGURES 30a–30c. Ecclisomyia simulata Banks, male genitalia. 30a, left lateral; 30b, ventral; 30c, dorsal. Abbreviations: ifa = inferior appendage (paired); ima = intermediate appendage;IX = segment IX; par = paramere (fusedpair); ph = phallus; phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; Xt = tergum X.

FIGURES 31a–31c. Ecclisomyia simulata Banks, male genitalia. 31a, left lateral; 31b, ventral; 31c, dorsal. Abbreviations:ifa = inferior appendage (paired); ima = intermediate appendage; IX = segment IX; par = parameres (fused pair); ph = phallus; phcr = phallocrypt; sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt= tergum VIII; Xt = tergum X.

FIGURES 32a–32c. Ecclisomyia simulata, female genitalia. 32a, ventral; 32b, ventral; 32c, left lateral. Abbreviations: e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; spt.sc = spermathecal sclerite; ssc.o = spermathecal sclerite opening; VIIIs = sternum VIII; IXs = sternum IX; Xt = tergum X.

FIGURES 33a, 32b. Ecclisomyia simulata Banks, male pupa, apical process. 33a, dorsal; 33b, ventral.

FIGURES 34a, 34b. Ecclisomyia simulata Banks, male pupa, apical process. 34a, dorsal; 34b, ventral.

FIGURES 35a, 35b. Ecclisomyia simulata Banks, female pupa, apical process.35a, dorsal; 35b, ventral.

FIGURE 41. Pupal schematic hook plate diagrams, dorsal, for Ecclisomyia bilera Denning, Ecclisomyia conspersa Banks, Ecclisomyia maculosa Banks, Ecclisomyia simulata Banks. Abbreviations: I–VIII = dorsal abdominal segments, with Arabic numerals representing the average number of hooks per plate.

FIGURE 42. Pupal and larval gill schematic diagrams, left lateral, for Ecclisomyia bilera Denning, Ecclisomyia conspersa Banks, Ecclisomyia maculosa Banks, Ecclisomyia simulata Banks. Abbreviations: I–VIII = dorsal abdominal segments, with Arabic numerals representing the number of gills per segment in anterior and posterior positions at dorsal, lateral or lateral line (dorsolateral and ventrolateral), and ventral levels.

FIGURES 43a–43d. Pupal cases. 43a-1 and 43a-2, Ecclisomyia bilera Denning (8–11 mm); 43b, Ecclisomyia conspersa Banks (16 mm); 43c, Ecclisomyia maculosa Banks (10.5 mm); 43d, Ecclisomyia simulata Banks (11 mm).

FIGURES11a–11c. Ecclisomyia conspersa Banks, male genitalia.11a, left lateral; 11b, ventral; 11c, dorsal. Abbreviations:ifa = inferior appendage (paired); ima = intermediate appendage; IX = segment IX; par = paramere (paired); ph = phallus; phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; Xt= tergum X.

FIGURES22a–22c. Ecclisomyiamaculosa Banks, male genitalia.22a, left lateral; 22b,ventral; 22c,dorsal. Abbreviations:ifa = inferior appendage (paired); ima = intermediate appendage; IX = segment IX; par = parameres (fused pair); phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; Xt= tergum X.

FIGURES23a–23c. Ecclisomyiamaculosa Banks, male genitalia.23a, left lateral; 23b, ventral; 23c, dorsal. Abbreviations:ifa = inferior appendage (paired); ima = intermediate appendage; IX = segment IX; par = parameres (fused pair); ph = phallus; phcr = phallocrypt; pl = pleuron (paired); sa = superior appendage (paired); VIIIs = sternum VIII; VIIIt = tergum VIII; Xt = tergum X.

FIGURES24a–24c.Ecclisomyia maculosa Banks, female terminalia.24a, leftlateral; 24b, dorsal;24c, ventral.Abbreviations: e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; IXs = sternum IX; IXt = tergum IX; pl = pleuron (paired); spt.sc = spermathecal sclerite; ssc.o = spermathecal sclerite opening; VIIIs = sternumVIII; IXs = sternum IX; IXt = tergum IX; Xt = tergum X.

FIGURES13a, 13b. EcclisomyiaconspersaBanks, femalegenitalia.13a, ventral;13b, left lateral.Abbreviations:e.gon.VIII = external part of gonopod VIII; e.gon.IX = external part of gonopod IX; spt.sc = spermathecal sclerite; ssc.o = spermathecal sclerite opening; IXs = sternum IX; Xt = tergum X.

FIGURES36a–36d. Ecclisomyia simulata Banks,pupa.36a,labrum,dorsal;36b, mandibles, ventral; 36c abdominal setaeand hook plates, dorsal; 36d, dorsal process of abdominal segment I, dorsal.

FIGURES 39, 40. Ecclisomyia spp., pupal hook plates. 39, Ecclisomyia maculosa Banks; 40, Ecclisomyia simulata Banks. Abbreviations: III–VII = abdominal terga III through VII; a = anterior; p = posterior.