Ecclisomyia bilera Denning 1951

Givens, Donald R., 2018, The Nearctic Ecclisomyia species (Trichoptera: Limnephilidae), Zootaxa 4413 (2), pp. 201-259 : 204-214

Ecclisomyia bilera Denning 1951

Figs. 1a–1c ; 2a–2c ; 3a–3c ; 4a–4c ; 5a , 5b ; 6a, 6b ; 7a, 7b ; 8a–8d ; 9 ; 37 ; 41 ; 42 ; 43 a-1, 43 a-2.

Ecclisomyia bilera Denning 1951 , 161, figs. 5a, 5b, 5c, male, female. Holotype male: Shasta Co., California, Lassen National Park, King's Creek Meadows, July 23, 1950, (C.P. Alexander) (CASC); Allotype female: same data as for holotype (CASC); Erman & Erman (1990, 12, 13, 15, 24, distribution, biology); Givens (2014, 159, 170, distribution).

Male diagnosis. Tergum X is foliaceous, appearing long and rectangular and apically truncate in dorsal and ventral views, curled evenly ventrad in lateral view ( Figs. 1a, 1b, 1c ); tergum X is acute or blunt in dorsal or ventral views in the other species and thicker and less curled in lateral view. The distal ends of the parameres of the phallus (ph) are angled dorsad ( Figs. 1a , 2a ); parameres are nearly or entirely straight in the other species. The inferior appendages of E. bilera lack spines, whereas E. conspersa has a stout, basal spine arising from each inferior appendage ( Figs. 10b , 11b ); E. maculosa and E. simulata have 2 black stout spines arising from the mesobasal area of each inferior appendage, with 1–4 smaller black stout spines lateral of these 2 basal spines ( Figs. 22b , 23b , 30b , 31b ).

Male description. See Denning (1951, 161, figs. 5a, 5b). Supplementary description follows: Length 8–11 mm (N = 16). Head, thoracic nota dark brown to dark fuscous. Antennae with 45–48 segments, each segment near distal end with 2 short translucent (difficult to discern) spine-like setae, except segment 1 (scape) and segment 2 with none. Antennal segments with fine black setae; scape large, pedicel short, third antennal segment 2 X length of pedicel. Sternum VIII with anterior area brown; with long reddish setae along distal ventral margin and lateral surfaces; in lateral view quadrate. Segment IX roughly triangular in lateral view, longest mesolaterally, longitudinally short dorsally and ventrally; dorsal, ventral, and anterior margins heavily sclerotized and lateral and caudal areas hyaline ( Figs. 1a , 2a ); in dorsal view, wide sclerotized bridge (strap) connecting left and right lateral portions ( Fig. 1c ); in ventral view, wide, transverse (longitudinally short) ventral strap connecting right and left portions ( Fig. 1b ). Tergum X foliaceous, appearing long and rectangular and apically truncate in dorsal and ventral views, curled evenly ventrad in lateral view; dark brown, subequal in width but slightly wider distally ( Figs. 1c , 2c ); dorsal and lateral areas with translucent (white) setae (= “spicules” of Denning 1951), best seen in dorsal view; caudal margin with deep V-shaped incision ( Fig. 2b ). Superior appendages (sa) divergent, brown, slightly wider posteriorly in ventral and dorsal views, rounded distally ( Figs. 1b, 1c ); dorsal and ventral margins and lateral surfaces clothed with translucent (white) setae. Inferior appendage (ifa) subtriangular in lateral and ventral views ( Figs. 1a, 1b , 2a ), each inferior appendage fused to form single non-articulated appendage with remnant suture visible; long, light reddish setae arising from ventral surface and distal area. Pair of parameres arising dorsally from anterior base of phallus; each with single distal hook angled dorsad ( Figs. 1a , 2a ). Phallus short, phallocrypt narrowed mesally, wider both anteriorly and caudally in lateral view ( Figs.1a , 2a ). Intermediate appendages dorsal of phallus, extending caudad; viewed ventrally ( Fig.1b ), distal portion separating into two convex extensions; dorsal apical areas appearing punctate.

Female diagnosis. The distal margin of tergum X ( Figs. 3c , 4c ) with V-shaped incision extending about 1/2–3/ 4 the length of the segment, unlike the shallow mesal notch in segment X of E. conspersa female ( Figs. 12 , 13a ). The external part of gonopod VIII short as in the E. conspersa female, but the distal (posterior) margin with a Vshaped indentation; contrasting with that of E. conspersa with a concave posterior margin that is sometimes convex; the external part of gonopod VIII is large in E. maculosa and E. simulata ( Figs. 24c , 32a ). The posterior margin of the external part of gonopod IX in E. bilera broadly concave; in E. conspersa with a U-shaped mesal indentation. The spermathecal sclerite round and broader posteriorly than anteriorly, whereas in E. conspersa , the spermathecal sclerite is narrower posteriorly than anteriorly ( Figs. 12 , 13a ) or subquadrate in E. maculosa and E. simulata .

Female description. Length 9–11 mm (N = 8). Tergum VIII brown, broad, extending ventrad beyond longitudinal midline ( Fig. 3a ); sternum VIII brown, short, entire, not divided; posterior dorsolateral angle triangular; pleura (pl) well developed ( Fig. 3a ); ventral caudal margin broadly and deeply concave ( Figs. 3b , 4a ). Tergum IX well developed, brown, extending laterally to pleural region; sternum IX reduced, without sclerotization, telescoped within segment VIII. Abdominal segment X conical, ventral and lateral surfaces lightly sclerotized, with numerous short black to translucent setae ( Fig. 3a ); lateral margins sinuate ( Figs. 3b , 4a ); concave elongate anal opening ventrally and anteriorly ( Figs. 3b , 4a ); in dorsal view distal margin of tergum X with deep Vshaped mesal incision for about 1/2–3/4 length of tergum. External part of fused gonopods VIII narrow, rectangular, semimembranosus or lightly sclerotized, with lateral sides straight in ventral view and curved dorsad to form small troughs, posterior margin forming V-shaped mesal indentation, not extending posterad beyond caudal margin of external part of fused gonopods IX. External part of fused gonopods IX short, concave dorsally, convex ventrally; serving as dorsal sheath for vaginal opening and extending beyond it; obscure, not heavily sclerotized; distal margin with broad U-shaped mesal indentation. Spermathecal sclerite rounded anteriorly, lateral margins heavily sclerotized, ribbed.

Pupal diagnosis. The Ecclisomyia bilera pupa may be separated from the pupa of E. conspersa by the absence of lateral line gills; with gills present on abdominal segments IIp–IVa of E. conspersa ; from E. simulata by the absence of subdorsal gills on the anterior margins of abdominal segment II ( Fig. 42 ); with gills present subdorsally on IIa of E. simulata ( Fig. 42 ). The Ecclisomyia bilera pupa has gills (dorsally and subventrally on VIIa; the E. maculosa pupa lacks such gills ( Fig. 42 ). The apical processes of E. bilera , E. maculosa , and E. simulata each have 2 stout, long, black apical setae ( Figs. 5a , 6a , 26a , 27a , 28a, 33a , 34a , 35a), whereas those of E. conspersa each have 3 long, black stout setae ( Figs. 15a , 16a, 17a, 18a).

Pupal description. Length 8–10 mm (N = 10). Head: Dark brown to dark reddish brown. Male antennae extending to abdominal apical processes; antennae in females extending to abdominal segment V. Labrum semioval, posterior margin convex, with 10 long black stout setae on dorsal surface, setae directed dorsad, distal ends of setae hooked ( Fig. 8a ). Mandibles dark brown; bases of mandibles broad, each tapering into single triangular apical tooth; ventrolateral surface with 2 long black stout setae ( Fig. 8b ). Mesal margins of mandibles finely serrated in both male and female pupae; serrations difficult to discern under light microscope.

Thorax: Pro-, meso-, and metanota dark brown to reddish brown; Pro-, meso-, and metathoracic legs brown. In mature pupa tarsal and tibial segments with both single and paired reddish-black stout spines; spines absent in immature pupa. Procoxae of both mature and immature pupae each with 1 long black seta near lateral margin and 2–5 shorter black setae on ventral surface. Mesocoxae each with 1–2 black setae on ventral surface. Profemora each with single black to brownish seta positioned near distal margin on ventral surface. Mesotarsal segments 1–4 each with pair of lateral fringes of long reddish to white hairs directed posterad.

Abdomen: Lateral fringe on each side originating from posterior area of abdominal segment V, extending to posterior margin of segment VIII; consisting of long, fine, hair-like black setae directed dorsad; line of fringe arching ventrad to segment VIII. Abdominal gills present; dorsal and ventral abdominal gills each single filament; present subdorsally on segments IIp–Va and dorsally on segments Vp–VIIp; present subventrally on abdominal segments IIp–VIIp ( Fig. 42 ). Dorsal process of segment I quadrate, rugulose; with 2 long black sa 2 setae each set laterad to meson; posterior margin sinuate, brown, forming pair of denticulate submesal lobes, each with 6–8 small, short spicules ( Fig. 8d ); shape of dorsal process varying with maturity of pupa. Hook plates on abdominal segments III–VII ( Figs. 8c , 37 , 41 ). Pairs of hook plates present anteriorly on segments III–VII, posteriorly on segment V. Hook plate IIIa with 2–5 hooks; IVa with 3–5; Va with 3–6; Vp with 13–26; VIa with 3–6; and VIIa with 3–7. Hook plates brown, hooks dark brown-black, directed posterad on anterior hook plates of segments III–VII and anterad on elliptical hook plates Vp. Dorsal abdominal segments each with 1 pair of black sa 2 setae on each of segments I–IV, 2–3 pairs on abdominal segments V–VII, and 3–4 pairs on segment VIII ( Fig. 8c ). Venter of abdominal segments III–VIII each with pair of short, black sa 2 setae and segments V–VIII each with pair of short black sa 3 setae. Apical processes distal area formed into 2 caudal lobes, brown, lightly sclerotized, curved apicomesad. Caudal lobes each with 2 long, black, stout setae on distal margin of lobe and with 2 shorter, black, stout setae along mesal surface ( Figs. 5a, 5b , 6a, 6b , 7a, 7b); several short black setae also on dorsal surface and shorter black setae on ventral surface ( Figs. 6b , 7b). Apical processes of female and male pupae with distinct lateral reddish spur set on distal lateral corner of each apical lobe. Dorsal surfaces of apical processes of male and female pupae appearing punctate due to covering of numerous spicules, barely discernable under light microscope ( Figs. 5a , 7a ).

Pupal case. Length 10–13 mm (N = 8), width 5 mm (N = 8); i.e., case approximately 2–3 X width ( Fig. 43 a-1). An inner silk-covered pupa encased within layer of large sand grains and coarse pebbles, interstitial spaces filled with much smaller grains of sand between larger pebbles and sand grains ( Fig. 43 a-2). Pupal case rounded; slightly wider anteriorly, but generally uniform in diameter. Pebbles at both ends loosely attached, leaving gaps between pebbles. In one site, cases were not attached to rocks, but lying in pools of a shallow creek without any apparent orientation to water flow; those pupal cases shorter, less streamlined ( Fig. 43 a-2).

Larval diagnosis. Of the four Nearctic species of Ecclisomyia , only the larva of E. bilera lacks mesal stripes on the head and thorax, diagnostic for this species. The posterior area of the frontoclypeus has indistinct muscle scars as occurs in E. simulata . Ecclisomyia conspersa and E. maculosa are without muscle scars on the frontoclypeus or the scars are indistinct. Each metathoracic sclerite sa 1 has 9–15 long black setae, whereas this sclerite on the other three Nearctic Ecclisomyia species each has 4–8 long black setae. The E. bilera larva does not have lateral line gills, whereas the E. conspersa larva has lateral line gills on abdominal segments II–IV ( Fig. 42 ). The larva of E. maculosa , unlike that of E. bilera , lacks anterior dorsal and ventral gills on abdominal segment VII ( Fig. 42 ). The larva of E. simulata , unlike other Nearctic Ecclisomyia species, has anterior gills on the dorsum of abdominal segment II ( Fig. 42 ).

Larval description: Length (mature instar) 8–10 mm (N = 14). Probable fourth instar length 5–6 mm (N = 3). Head: Concolorous, reddish brown to dark brown, rounded posteriorly, narrowing anteriorly ( Fig. 9 ). Parietals and genae reddish brown; muscle scars distinct on parietals, posterior to eyes. Muscle scars either distinct or indistinct on posterior half of frontoclypeus. Mandibles black, each with 4 blunt apical teeth ( Fig. 9 ). Mandibular teeth of early instars not as blunt as those of mature instars and prepupae; one to two short translucent setae arise from posterolateral corner of each mandible. Mesal margins of mandibles each with tuft of translucent feather-like setae ( Fig. 19a ). Ventral apotome triangular, 1.57–2.10 X longer than ventral ecdysial line. Primary setae in positions 7, and 9–17 ( Fig. 9 ). Primary setae in position 7 black, either short or long; primary setae in position 9 long, black; primary setae in position 10 short, appressed, black to translucent; primary setae in position 11, short, difficult to discern; primary setae in position 12 black, either short or long; primary setae in position 13 black to translucent, short to long, semi-appressed, difficult to discern; primary setae in position 14 black and long; primary setae in position 15 black, short or long; primary setae in position 16 short, black to translucent, semi-appressed; primary setae in position 17 black and long. Venter of head with primary setae in positions 8 and 18 ( Fig. 19b ) short, translucent with setal sockets prominent, but may be absent or difficult to discern. Frontoclypeus wide anteriorly, narrow posteriorly, triangular; glabrous, except for primary setae in positions 1–6 ( Fig. 9 ). Primary setae in position 1 long, reddish to translucent, sometimes absent; primary setae in positions 2 long, black, may be reddish to translucent; primary setae in position 3 short, may be long, black to translucent; primary setae in position 4 short or long, appressed, black to translucent; primary setae in positions 5 and 6 short or long, black generally, but sometimes translucent. Frontoclypeal primary setae often difficult to discern, often short, black basally, translucent apically. Labrum reddish brown to dark brown, glabrous except for primary setae in positions 1–6. Primary setae in position 1 short, blunt and translucent, sometimes absent; primary setae in position 2 translucent, absent or indiscernible; primary setae in position 3 short, black to translucent; primary setae in position 4 short, translucent, difficult to discern, best seen in lateral view; primary setae in positions 5 and 6 short, those in position 6 somewhat longer, black to translucent ( Fig. 9 ). Light, short translucent setae present at anterolateral corners of labrum, forming sparse setal tuft. Cardo sclerites rectangular. Submental sclerite with 2 short black submesal setae, their setal sockets prominent.

Thorax: Pro- and mesonota brown to reddish brown; metanotal sclerites brown. Lateral processes of mesonota triangular, fuscous to black. Anterior margins of pronota each with 10–17 long black setae set equidistant from each other along margin; short, translucent setae set between these setae along anterior margin. Each half of pronotum with 3–9 black setae posterior to mid-dorsal transverse groove. Posterior marginal area of pronotum black, heavily sclerotized. Mesonotum with black setae in 3 distinct groups roughly corresponding to sa 1, sa 2, and sa 3 positions; muscle scars distinct or indistinct. Metanota sa 1 sclerites elliptical, each with 8–11 black ls setae; metanota sa 2 sclerites without discrete form, variable, each with 7–10 black ls setae and 2–3 short slender (ss) black setae; metanotal sa 3 sclerites parenthesis-shaped, each with 10–15 black ls setae on anterior half of sclerite. Prosternal horn short, light in color, sclerotized. Prosternal sclerite lightly sclerotized, nearly membranous, sometimes absent. Legs brown, yellowish brown or reddish brown. Foretrochantins brown to dark brown, forming tent-like structures over procoxae. Procoxae with primary setae in positons 1 and 2. Ventral edge of each procoxa with 8–12 black ls setae. Ventral edge of each profemur with 2 long sl translucent setae and row of short translucent sl setae along ventral margin ( Fig. 21a ). Dorsal edge of each profemur with 4 ls setae. Protibiae and protarsi each with primary black seta in position 3 ( Figs. 21a, 21b ). Dorsodistal margin of each protibia with 2 long black to translucent setae. Protibiae each with 2 apical spurs. Mesocoxae each with 7 to 8 black ls setae along distal posterior margin and 8–11 black ls setae anteriorly. Meso- and metacoxae each with primary seta in position 2 ( Figs. 21b, 21c ). Metacoxae each with 5–6 black ls setae along distal posterior margin and 18–20 black ls setae on anterior surface, of which 4–5 on anterodistal margin. Trochanters of all legs with black primary seta in positions 2, 3, and 5 ( Figs. 21b, 21c ); primary seta 5 sometimes absent or difficult to discern. Ventral margins of pro- and mesotrochanters with distal trochanteral brush ( Wiggins 1996) ( Figs. 21a, 21b ). Metatrochanters each with 7 ls setae arising from anterior margin. Pro- and mesofemora with distinct sl reddish to translucent setae along ventral margins; these sl setae sometimes occurring along ventral margins of metafemora, but indistinct if present. Mesofemora each with 2 long black ls setae on ventral margin ( Fig. 21b ); with black primary setae in positions 2, 3, and 4, but primary setae in positions 3 and 4 sometimes absent. Metafemora with primary setae in positions 2, 3, and 4, but primary setae in position 3 sometimes absent ( Fig. 21c ). Meta-epimeron extending ventrally beneath metathoracic legs, each bearing 12 black ls setae. Meso- and metathoracic epimera each with 4 to 5 black ls setae on lateral surface. Meso- and metathoracic episterna each with 5–9 black ls setae arising from anterior margins; 2–5 black setae present on lateral surface.

Abdomen: Segment I lateral to dorsal hump with 14–19 black setae; setal sockets prominent; penultimate instars (5–6 mm), possibly with only 4 long, black seta lateral to dorsal hump. Dorsal hump without sclerites. Lateral humps without sclerites, each with 2 long black setae dorsally, sometimes with sclerotized bases, and 9–13 fine black ls setae, several with basal sclerotization; early instars sometimes with only 2 black ls setae; ventral base of each lateral hump with 3–12 black ls seta; small ellipsoid sternite sometimes present below each lateral hump, bearing 3–6 black ls setae. Venter of abdominal segment I with 9–30 black ls setae, most without basal sclerotization; pair of triangular sclerites or lightly sclerotized areas, each with 2–5 black setae, sometimes present lateral to midline, but some larvae without sclerotization except about bases of setae. Abdominal gills present; dorsal and ventral abdominal gills each with single filament; present dorsoposteriorly and ventroposteriorly on segment II; present dorsoanteriorly, dorsoposteriorly, ventroanteriorly, and ventroposteriorly on segments III–VII ( Fig. 42 ). Abdominal segment VII sometimes missing 1 or 2 dorsal anterior gills. Dorsum of abdomen with sa 2 setae on abdominal segments II–VIII; those on segments VI–VIII long, black; those on segments II–V ss black setae, sometimes absent or difficult to discern. Abdominal segments II and V each with 1–2 pairs of short black sa 1 setae. Segment VII sometimes with black sa 2 setae, 3 on each side. Segment VIII sometimes with 1–2 short black setae adjacent to each long black sa 2 seta. Venter of abdominal segment II with pair of short slender sa 1 setae. Abdominal tergum IX with brown, elliptical tergite bearing 7–9 black setae along caudal margin; mesal surface with 2–6 ss black seta. Lateral fringe of hl setae extending on each side from anterior of segment III to anterior of segment VIII or half way along segment VIII; lateral fringe less distinct in early instars. Chloride epithelia present on venters of abdominal segments III–VII. Anal prolegs with lateral sclerites brown, each with 1 black ls seta near posteroventral margin; caudal half with 4–10 black setae on dorsal surface and with basal tuft of 4 ls black setae, of which inner 2 stout, longer than shorter fine outer setae; narrower ventrally and curving beneath caudal lobes, appearing as narrow sclerotized bands. Sole plates brown, each with dorsal margin thick, black, bearing single short black seta near distal-ventral margin. Caudal lobes glabrous, except for minute sl reddish spicules on either side of anal opening. Anal hooks sharply recurved anteriorly, each bearing small accessory hook (denticule).

Larval case. Larval case of penultimate instar larva 5–8 mm (N = 10). Case constructed of small grains of sand and plant material, slightly cornucopia-shaped, slender with slight posterior curvature. Stone portion of case consisting of tightly meshed sand pebbles, anterior fourth of case composed primarily of plant material. Plant material sometimes attached to opposing sides of case; plant pieces also sometimes attached anteriorly, angling outward from case.

Distribution. Specimens of E. bilera during this study were from Shasta and Tehama counties, California, and Washoe County, Nevada. Ecclisomyia bilera was previously recorded from Shasta and Tehama counties, California ( Givens 2014). This species has also been recorded from Nevada and Sierra counties in California ( Erman & Erman 1990), and Mono County, California ( Maciolek & Tunzi 1968). Ecclisomyia bilera appears to be restricted to the Sierra Nevada Mountains of California and Nevada. Ecclisomyia bilera is sympatric with both E. conspersa and E. simulata ( Givens 2014) .

Bionomics: Ecclisomyia bilera is a high-altitude species, occurring at elevations of 1737–2455 m. Emergence commences in late June, and extends into the middle of August. This species has been collected from larger creeks with widths up to 3.05 m and depths up to 0.91 m, but this species seems to occur primarily in narrow (0.30–1.8 m), shallow (7.6–45.7 cm), rapidly flowing creeks with cobble, gravel, and sand substratum. These streams usually have mixed plant debris, frequently with wood and bark debris.

Material Examined. CALIFORNIA: Shasta Co., East Fork Hat Creek, 20.5 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 20-vii-2011, 1 PP (DRG) (CSUC); East Fork Hat Creek, 18.3 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 22-vii-2011, 1 L (DRG) (CSUC); West Fork Hat Creek 15 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 11-ix-2013, 12 L (DRG) (CSUC); King's Creek, 26.5 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 01-vii-2011, 2 M (DRG) (CSUC); 20-vii-2011, 2 M (DRG) (CSUC); 01-vii-2013, 1 PP, 1 P (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, 1 F P, 3 P (DRG) (CSUC); King's Creek, 27.1 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), King's Creek Upper Meadow, 01-vii-2013, 2 M P (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 (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.), 14-vii-2011, 2 M, 1F (DRG) (CSUC); 26-vii-2011, 1 F, 5 M P, 47 P, 3 PP (DRG) (CSUC); 28-vii-2011, 1 M P, 2 F P (DRG) (CSUC); 14-vii-2012, 6 M, 4 F, 4 PP (DRG) (CSUC); 20-vii-2012, 24 M, 3 F, 3 M P, 2 F P, (DRG) (CSUC); 21-vii-2012, 1 M, 3 P, 2 L (DRG) (CSUC); 27-vii-2012, 11 M, 5 M P, 8 P (DRG) (CSUC); 01-viii-2012, 4 M, 1 F, 5 M P, 4 F P, 11 P (DRG) (CSUC); 04-viii-1012, 1 F, (DRG) (CSUC); 10-viii-2012, 10 M, 1 F (DRG) (CSUC); 11-viii-2012, 1 M P (DRG) (CSUC); 13-viii-2012, 3 M, 1 F P, 2 P (DRG) (CSUC); 14-viii-2012, 13 M, 2 F, 3 P (DRG) (CSUC); 27-vi-2013, 2 M, 1 M P, 1 F P, 6 P, 1 L (DRG) (CSUC); 28-vi-2013, 1 M, 1 F, 2 M P, 3 F P, 7 P, 2 PP (DRG) (CSUC); 01-vii-2013 1 M P, 1 F P (DRG) (CSUC); 19-vii-2013, 1 L (DRG) (CSUC); 09-ix-2013, 1 L (DRG) (CSUC); King's Creek, 28.2 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 27-vii-2012, 2 M, 1 F, 1 F P (DRG) (CSUC); 01-viii-2012, 1 F P, 2 P (DRG) (CSUC); 11-viii-2012, 1 F P (DRG) (CSUC); 27-viii-2012, 2 M P, 3 P (DRG) (CSUC); 19-vi-2013, 1 M P, 3 P (DRG) (CSUC); 27-vi-2013, 1 M, 4 M P, 2 F P (DRG) (CSUC); 30-vi-2013, 1 P (DRG) (CSUC); 11-vii-2013, 1 P, 1PP (DRG) (CSUC); King's Creek, 28.3 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 01-viii-2012, 1 M P (DRG) (CSUC); 04- viii-2012, 1 F (DRG) (CSUC); small creek, Paradise Meadows, Lassen Volcanic Nat'l. Park, 14-vii-2013, 1 M P, 5 P (DRG) (CSUC); small creek about 1.2 km up the trail to Paradise Meadows (trailhead 15 km NW entrance to Lassen Volcanic Nat'l. Park), 10-ix-2013, 1 L (DRG) (CSUC); small tributary to King's Creek, King's Creek Picnic Site, 28 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 11-vii-2013, 2 P (DRG) (CSUC); 12-vii-2013, 1 P, 1 PP (DRG) (CSUC); small tributary to King's Creek, King's Creek Upper Meadow 28.2 km S of NW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 28-vi-2013, 2 M, 2 P, 3 L (DRG) (CSUC); 29-vi- 2013, 4 M P, 3 F P, 7 P, 4 PP (DRG) (CSUC); 30-vi-2013, 1 M P, 1 PP (DRG) (CSUC); 01-vii-2013, 1 P, 1 PP (DRG) (CSUC); 07-vii-2013, 1 M P (DRG) (CSUC); 19-vii-2013, 2 M, 1 M P, 1 F P (DRG) (CSUC). Tehama Co., small snowmelt puddle 0.3 km N of SW entrance to Lassen Volcanic Nat'l. Park (Lassen Park Rd.), 29-vi-2013, 1 M P, 1 F P, 1 P (DRG) (CSUC). NEVADA: Washoe Co., tributary to Galena Creek, about 0.2 km NW of Mount Rose Ski Area well pad near Hwy. 431, N39.31948 W119.89963, 13-v-2015, 1 M, 2 F, 1 M P, 1 F P, 21 P (N. Kang and DER) (CSUC).

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 2a–2c. Ecclisomyia bileraDenning, male genitalia.2a, left lateral; 2b, left caudodorsal; 2c, ventral; 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.

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 5a, 5b. Ecclisomyia bilera Denning, male pupa, apical processes.5a, dorsal; 5b, ventral.

FIGURES 6a, 6b. Ecclisomyia bilera Denning, male pupa, apical processes.6a, dorsal; 6b, ventral.

FIGURES 7a, 7b. Ecclisomyia bilera Denning, female pupa, apical processes. 7a, dorsal; 7b, ventral.

FIGURES 8a–8d. Ecclisomyia bilera Denning, pupa. 8a, labrum, dorsal; 8b, mandibles, ventral; 8c, dorsal abdominal setae and hook plates, dorsal;6d, dorsal process abdominal segment I, dorsal.

FIGURE 9. Ecclisomyia bilera Denning, larval head.9, dorsal,with primary setae numbered.

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 12. Ecclisomyia conspersa Banks, female genitalia, ventral. 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 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 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 26a, 26b. Ecclisomyia maculosa Banks, male pupa, apical process.26a, dorsal; 26b, ventral.

FIGURES 27a, 27b. Ecclisomyia maculosa Banks, male pupa, apical process.27a, dorsal; 27b, ventral.

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.

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).

FIGURES 37, 38. Ecclisomyia spp., pupal hook plates. 37, Ecclisomyia bilera Denning; 38, Ecclisomyia conspersa Banks. Abbreviations: III–VII = abdominal terga III through VII; a = anterior; p = posterior.

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.

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.

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.