Stenophylax antennatus Banks, 1900, new combination (type species); Philocasca antennata (Banks, 1900), new combination; Philocasca alba Nimmo, 1977, new combination; Philocasca thor Nimmo, 1971, new combination

Adult (Fig. 1). Head to wingtip length: Montiphylax antennatus male - 19–21 mm (n = 5); antennatus female - 19–21 mm (n = 5). Montiphylax albus male - 18–19 mm (n = 8); albus female - 18–20 mm (n = 3). Montiphylax thor male - 18–21mm (n = 3). Eyes large with numerous fine setae located between ommatidia. Head rectangular in dorsal view; ocelli large, posterior ocelli set midlength. Several small setae scattered on head surface, primarily between and behind posterior ocelli. Two pair of warts evenly spread between anterior and posterior ocelli. Posterior warts small, linear with acute apices. Malar projection as long as length of 1^st labial palp segment. Antennae shorter than wing length, stout, basal segment shorter than eye width.

Pronotal warts oval, wide, nearly touching mesally. Mesonotal macrosetae in linear row, basal warts located midlength, slightly merged but not fused into a single smooth oval wart; mesocutellar macro setae scattered linearly, less fused than mesonotal wart; most dorsal head and thoracic setae pale.

Forewings (in alcohol) reddish brown, finely irrorate; third radial vein and discoidal cell with a fairly long common boundary approximately equal to length of the first transverse vein; discoidal cell longer than its pedicel; all apical cells without pedicels. Anterior and posterior anastomosis nearly perpendicular to wing length, clearly located distad from posterior anastomosis. Hindwings brown, without irrorations; discoidal cell longer than its pedicel; all apical cells without pedicels. Anterior anastomosis perpendicular to wing length, clearly located distad from posterior anastomosis. Posterior anastomosis strongly oblique to wing margin. Most setae on wing membrane recumbent.

Legs long and thin, tarsi long with basal segment more than 0.5 length of mesotibia, Apical tarsal segments without ventral spines. Spines on legs black, spurs golden, spur count 1-2-4.

Male genitalia (Figs 6, 8, 9). Eighth segment without adornment, slightly swollen on dorsal apical margin. In lateral view 9^th segment annular, narrowed dorsally and broadest at base of short, tall inferior appendages; deeply invaginated dorsally in caudal view. Tenth segment cupped anteriorly, appearing as two, slightly fused hemispheres extending broadly anteriad within the 9^th. Superior appendages of 10^th fused to intermediate appendages; in lateral view, variously broad and projecting, diagnostic by species. In dorsal view intermediate appendages of 10^th extended mesally as two long, tapered, downward directed, thin parallel projections. Inferior appendages of 10^th short, broad ventrally; nearly fused mesally. Phallus with heavily sclerotized, dorsomesal, conical projection at base of dorsolateral parameres. Parameres heavily sclerotized, projecting fingerlike, parallel to body of phallus; shape diagnostic by species. Apical portion of phallus membranous, extensile, with ejaculatory duct ending in an apical, upward directed, sclerotized cup.

Female genitalia (Figs 7, 10), female of M. thor unknown. Ninth tergite and sternite fused laterally; 10^th segment fused to 9^th, broadly projecting distad to blunt, rounded apex in lateral view. In dorsal view 10^th apex cleft to approximately midlength mesally. In ventral view, 9^th sternal lateral projections fused to mesal supragenital plate. Vulvar scale stout, broadly merged to 8^th segment laterally, with wide, smoothly conical, medial lobe slightly shorter than lateral lobes.

Larva (Figs 2–4), (n = 3 antennatus and 4 albus; larva of M. thor unknown). Head nearly circular in dorsal view, widest mid-length, rugulose, specimens in alcohol rust colored. Muscle scars indistinct. Typical limnephiloid setation, without accessory setae. Anterior ventral apotome small, equal, or shorter than ventral ecdysial suture. Submental sclerites distinctly separated from stipes and mesally separated.

Prothorax rust colored (in alcohol) like head; setae scattered, without distinct setal areas; in lateral view, slight transverse indentation approximately midlength; posterior margin thick and dark, with deep, preceding furrow; ventrally with a single, wide, short, prosternite between legs. Mesothorax paler than prothorax; setal areas nearly merged; posterior black margin extending along lateral margin nearly to anterior margin; ventral area with a linear row of small, dark, mesosclerites along each side of posterior margin. Dorsal metanotal setal areas on distinct sclerites, without additional setae on metonotal membrane. Ventrally, mesosternite with very pale linear row of small sclerites along each side of posterior margin, may be difficult to see, appearing as a row of indentations. Legs typical limnephiloid form with short, stout proleg and hind leg the longest. Meso and metafemur with two, long, dark setae ventrally and a row of very small, fine spinelike setae along entire ventral setal margin. Anterior face of meso and metafemur with numerous accessory setae; posterior face usually with one or two accessory setae.

First abdominal segment with numerous long setae anterior of dorsal spacing hump; lateral spacing hump with several small sclerites on dorsal margin and one larger sclerite along posterior margin; ventrally with numerous long setae and a pair of posterior setal warts on mature larvae. Chloride epithelia present ventromesally, much wider than long (albus 2–7 [n = 3]; antennatus 3–7 [n = 4]). Lateral fringe on segments 2–8 (antennatus), 3–8 (albus); segment 2 portion is very short. Forked lamellae absent. Gill clusters consist of single filaments dorsally on segment 2 through 5 or 6, ventrally on 2 through 5, lateral line gills absent.

Ninth tergite with 3–4 pairs of long setae and several additional shorter setae. Lateral sclerite of anal proleg without short, stout, pale setae. Anal claw with single dorsal accessory hook.

Pupal case of final instar slightly curved, non-tapered, wood and mineral, or all mineral construction. The limited larval material available indicates the earlier instars may build vegetation cases.

Pupa (Fig. 5), (n = 4 albus). Labral setae long, with apices spirally twisted, not hooked. Mandibles triangular with apical half strongly tapered and flattened into an acute blade; mesal edge of blade slightly serrate. Antennal scape with a ventrolateral setal tuft and a single seta at the dorsal margin. Second antennal segment with a dorsal tuft. Antennae shorter than abdominal apex. Spined ridge of 1^st abdominal tergite weak, linear with small scattered spines. Anterior hook plates present on abdominal tergites 3–7, with strong hooks directed posteriad. Small, oval posterior hook plates present on tergum 5. Abdominal lateral fringe present on segments 5–8. Anal processes elongate, slender, tubular, with patch of minute, dorsal spines at apex. Four long setae on each anal process; one located basally, and three apically.

Montiphylax from the Latin montis (mountain) and the Greek phylax (guard), referring to the mountainous habitat of this genus.

The male of M. antennatus is distinguished from M. albus and M. thor by the long, narrow superior appendages in lateral view and the blunt paramere apex. The M. thor superior appendage has a wide base, appearing more triangular in lateral view. The superior appendage of M. albus is shorter than either M. antennatus or M. thor although wide at the base like M. thor. In lateral view, the paramere apex of M. albus and M. thor is acute; in M. albus the paramere apex is downturned while it is upturned in M. antennatus and M. thor.

The female of M. antennatus is separated from M. albus by the round anterior margin of the vaginal apparatus that is more quadrate in M. albus in ventral view. The female of M. thor is unknown. The pupa of only M. albus is known.

The lateral setal fringe of M. antennatus larvae originates on the posterior margin of the 2^rd abdominal segment while the lateral fringe of M. albus starts at the anterior margin of the 3^rd segment. The larva of M. thor is unknown.

Head to wingtip length: male 19–21 mm (n = 5); female 19–21 mm (n = 5).

Male genitalia (Fig. 6): 8^th tergite without dorsal modified spines or projections. Ninth segment annular with tergite narrow and strap-like, directed distad dorsally resulting in slightly sigmoid anterior margin of 9^th in lateral view; remainder of 9^th broad in lateral view, directed downward at approximately a 45 degree angle from the narrow tergite; inferior appendages appearing separated from 9^th. In caudal view, inferior appendages slightly cupped around phallic apparatus and slightly separated mesally. Tenth segment cupped anteriorly, appearing as two, slightly fused hemispheres extending broadly anteriad within the 9^th. Superior appendage nearly twice as long as tall in lateral view. Intermediate appendages arise ventrally, strongly sclerotized and extending as two long, tapered, parallel projections, curving downward at apex. In caudal view, below the intermediate appendages are paired, curved, narrow projections nearly surrounding the anus. Phallus large, with strongly sclerotized phallicata; membranous endophallus with dorsal, strongly sclerotized band, projected upward and distad apically; strong, thick parameres originate at the base of the dorsal band, extend distally; tapering to blunt, rounded apex in lateral view and slight curved upward throughout. Parameres originate dorsolaterally and extend posteriorly over 3 times as long as wide; thick throughout, ending in blunt, rounded apex in dorsal view. The aedeagus apex a sclerotized tube within an extensile sheath; ending in the bottom of a posteriorly directed sclerotized cup.

Allotype female genitalia (Fig. 7): 9^th segment fused laterally and incomplete ventrally, separated ventrally by broad supragenital plate. Ninth fused with 10^th; 9^th tergite ca. half as long as 10^th in lateral view; in lateral view 10^th with dorsal and ventral margins slightly tapered throughout to rounded apex. In dorsal view, 10^th cleft less than half the distance to base. In lateral view, ventrolateral corners of 9^th extend distally beyond remainder of 9^th; acute at apex and directed inward in ventral view. The medial lobe of the vulval scale short, shorter than lateral lobes, longer than wide. The vaginal apparatus, in lateral view, rectangular, ca. twice as long as tall; in ventral view, egg-shaped with smoothly rounded anterior margin.

WA: Whatcom County, stream near Heather Meadow Information Center on Artist Point Road, D.W. Blinn, 28 July 2009, blacklight trap, 2M; stream on Fire and Ice Trail, near top of Mt. Baker Highway, 48.85586 –121.68795, D.W. Blinn, 13 August 2012, 2L, 1M, 1F; spring stream, ca. 150 feet southeast of outhouse at Mt. Baker Information Center parking lot, near top of Highway 542, 48.85378 –121.68543, D.E. Ruiter, 5 August 2012, 1L; meltwater stream on Fire and Ice trail, in Heather Meadows, Mt. Baker, D.W. Blinn, 31 July 2009, 13M, 3F (2M to National Museum of Natural History; 2M, 1F to S. Chuluunbat; 1M, 1F to Canadian National Collection, remainder in D.E. Ruiter personal collection); Terminal Lake on Mt. Baker Highway, D. W. Blinn, black light trap, 28 July 2009, 1M.

Nimmo (1971) described M. thor from a single specimen collected within Jasper National Park, Alberta. The Barcode of Life project has collected additional males from the Willmore Wilderness in Alberta, ca. 150 km northwest of the M. thor type locality. This Barcode of Life material was compared to the M. thor holotype and found to be the same. To date, females or larvae of M. thor have not been located.

Wiggins and Anderson (1968) mentioned the dark ventral surface of the M. antennatus scape described by Banks (1900) was absent on the Idaho M. antennatus specimen that Banks (1943) described, implying it was present on the M. antennatus type. The darker ventral surface of the antennal scape is present on the Mt. Baker M. antennatus males and females examined here. However, it is absent on our M. albus material, that have a concolorous scape. Montiphylax thor males have a darkened scape with the ventral surface pale - the exact opposite coloration pattern from M. antennatus. The Banks (1943) Idaho specimen was examined and it has the light ventral surface of the scape and upturned, acute parameres of M. thor. Neither Banks (1943) nor Wiggins and Anderson (1968) recognized this Idaho specimen as a new species. Our genetic comparisons of M. antennatus and M. thor also support Nimmo’s conclusion that M. thor is a valid species. Our determination that the Banks (1943) Wallace, Idaho, record represents M. thor extends the M. thor range another 600km farther south along the Rocky Mountains. This also results in M. thor occurring both further north and south than M. albus along the Rockies. Wallace, Idaho, is a historic silver mining city in the South Fork of the Coeur d’Alene River Valley. It is likely the M. thor specimen was collected at higher elevations in the surrounding area.

Male genitalia (Fig. 8): The dark, conical, ventral sclerite of the phallic apparatus illustrated by Nimmo (1971) is absent from the holotype and the other M. thor specimens examined in this study. It is unknown what the Nimmo (1971) dark ventral sclerite represents. Eighth tergite without dorsal modified spines or projections. Ninth segment annular with tergite narrow and strap-like; remainder of 9^th broad in lateral view, directed posteroventrad at ca. a 45 degree angle from the narrow tergite, with tall, narrow inferior appendages nearly fused to 9^th. In caudal view, inferior appendages slightly cupped around phallic apparatus and slightly separated mesally. Tenth segment cupped anteriorly, appearing as two hemispheres extending broadly anteriad within the 9^th. Tenth segment appears fused mesally along anterior margin. Superior appendages approximately as long as tall, triangular in lateral view directed upward. Intermediate appendages arise ventrally, strongly sclerotized and extending as two long, tapered, parallel projections, curving downward at apex. In posterior view, below the intermediate appendages are paired, curved, narrow projections nearly surrounding the anus. Phallus large, with strongly sclerotized phallicata; membranous endophallus with dorsal, strongly sclerotized band, projected upward and distad apically; strong, thick parameres originate at the base of the dorsal band, extend distally; tapered to acute apex in lateral view and slightly curved upward throughout. Parameres originate dorsolaterally and extend posteriorly ca. 3 times as long as wide; tapering evenly throughout to acute apex in dorsal view. Aedeagal apex a sclerotized tube within an extensile sheath; ending in the bottom of an upward directed sclerotized cup.

Holotype male - Canadian National Collection Type # 10,588, Alpine Meadows, east of Mt. Edith Cavell, Jasper National Park, Alberta, habitat: Mountain Tarn, A. P. Nimmo, 10–12am, 4 July 1975; Alberta, Willmore Wilderness, 53.6858, -119.419, Hilchie & MacAuley, 21 July 2007, 3 males (Bold # 10ABCAD-004, 10ABCAD-006, & 10ANCAD-007).

Nimmo (1977) described M. albus and included characters to separate M. albus from M. thor. The second author reared from associated pupae (Alberta, Middle Fork Creek, Marmot Creek Experimental watershed, 1800 meters altitude, 50.94820 –115.15151) the male and female of M. albus, and provided figures of the male, female, and pupa. These are from the same watershed as the Nimmo paratype collection. Based on this M. albus material, we add descriptions and images for the M. albus adults, pupa and larva.

Head to wingtip length: male - 18–19 mm (n = 8); female - 18–20 mm (n = 3). General description of adults, pupa, and larva contained in genus description above.

Male genitalia (Fig. 9): 8^th tergite without dorsal modified spines or projections. Ninth annular with tergite narrow and strap-like; remainder of 9^th broad in lateral view; with tall, narrow inferior appendages nearly fused to 9^th. In caudal view, inferior appendages curved around margin of 9^th base to meson of 9^th, appearing fused mesoventrally. Tenth segment cupped anteriorly, appearing as two hemispheres extending broadly anteriad within the 9^th. Tenth segment appears split mesally. Superior appendages taller than long in lateral view, directed downward and distad at ventral margin. Intermediate appendages arise ventrally, strongly sclerotized and extending as two long, tapered, parallel projections, curving downward at apex. In caudal view, ventrad to the intermediate appendages paired, curved, narrow projections nearly surround the anus. Phallus large, phallicata strongly sclerotized; endophallus with strongly sclerotized band dorsally, projected upward and distad apically; strong, thick parameres originate at the base of the dorsal band, extend distally, tapered to acute apex in lateral view and slightly curved downward throughout. In dorsal view, parameres originate dorsolaterally and extend posteriorly ca. 3 times width, tapering evenly to bluntly curved apex. Aedeagus apex a sclerotized tube within an extensile sheath, ending in the bottom of an upward directed sclerotized cup.

Allotype female genitalia (Fig. 10): 9^th segment fused laterally and incomplete ventrally. In lateral view, 9^th fused with, and nearly as long as 10^th. Tenth with dorsal and ventral margins parallel throughout most of its length until rounded apex. In dorsal view 10^th cleft to approximately midlength. In lateral view, ventrolateral corners of 9^th clearly shorter than remainder of 9^th; in ventral view, broadly rounded at apex and directed outward. Medial lobe of the vulval scale shorter than lateral lobes, approximately as wide as long. Vaginal apparatus rectangular in lateral view, ca. twice as long as tall; in ventral view, rectangular with anterolateral corners projecting outward.

Holotype male: Canadian National Collection # 151765, Rowe Brook, near Lower Rowe Lake, 6350’, Waterton National Park, Alberta, 12 June 1975, D.B. Donald [near 49.054706, -114.052434]; Alberta, Twin Creek, Marmot Creek Experimental watershed, 1800 meters elevation, R. Mutch, 1978, 4L, 50.94820 -115.15151; light trap, 1 July 1979 4M4F; 16 April 1980, 1 pupa reared to male; 16 April 1980 1 female pupa; 26 April 1980, 1 larva reared to female; 26 April 1980, 1 larva reared to male; 1 May 1980, 1 larva reared to male.