Montiphylax, (Trichoptera, Limnephilidae), a new genus to accommodate the western North American species: Stenophylaxantennatus Banks, 1900, Philocascathor Nimmo, 1971, and Philocascaalba Nimmo, 1977

Abstract Montiphylax, a new genus within the Limnephilidae, is proposed to include Philocascaalba, P.antennata, and P.thor. Characters for the adults and larvae are presented to separate Montiphylax from Philocasca. A summary of the distribution and life history of the Montiphylax species is provided. A key to ease separation of the known North American caddisfly larvae with gill clusters of single filaments is given.


Introduction
The generic placement for the seven Philocasca taxa [P. alba Nimmo, 1977, P. antennata (Banks, 1900) as Stenophylax Kolenati, 1848, P. banksi (Denning, 1941) as Anisogamus McLachlan, 1874, P. demita Ross, 1941, P. oron Ross, 1949, P. rivularis Wiggins & Anderson, 1968, P. thor Nimmo, 1971] has been confused since Schmid (1955) placed Stenophylax antennatus within Philocasca. Wiggins and Anderson (1968) examined the known Philocasca taxa at that time and accepted Schmid's placement of S. antennatus in Philocasca. Two additional taxa (P. thor Nimmo, 1971, P. alba Nimmo, 1977 were subsequently described. Mutch (1981) pointed out the polyphyletic nature of Philocasca. Vshivkova et al. (2007) elevated Philocasca to the subfamily Philocascinae based only on the characters of P. rivularis. In order to clarify the status of Stenophylax antennatus, a new genus, Montiphylax (Limnephilidae, Limnephilinae) is proposed to accept P. antennata, P. thor, and P. alba. Wiggins and Anderson (1968) presented, in detail, the prior problems associated with description and relationships within the Philocasca and its placement within the Limnephilidae. These problems were the result of only five male specimens spread across the four known species. No females, or larvae, had been associated at the time. And, at the time, only Banks and Ross seem to have examined those same specimens, and they disagreed on their generic placement. Wiggins and Anderson (1968), with still only the type of S. antennatus male available, did not acknowledge the significant generic dissimilarities pointed out by Banks and added no further information to support their placement of S. antennatus within Philocasca. In the discussion below we present the detailed history of the confusion, and rationale for the creation of a new genus, Montiphylax, to resolve the confusion. The information is based, in part, on now available female and larval associations.
Also, since Wiggins and Anderson (1968), Philocasca has been removed from the Limnephilinae and placed within the Philocascinae (Vshivkova et al. 2007). Philocasca can be distinguished by many non-Limnephilinae characters: wings very broad, rounded throughout, without obvious color patterns/spotting, etc.; mesonotal warts absent; female without mesal lobe of vulvar scale; larvae with strongly rounded anterior portion of pronotum without transverse groove, and with unique, long, flattened, scale-like setae on head and pronotum. The arrangement and shape of these larval setae are diagnostic for the Philocasca species.
While the spotted wings, female three-part vulvar scale, and pronotal transverse groove of the larva currently place Montiphylax within the Limnephilinae sensu Vshivkova et al. (2007) its further placement is unclear. This has resulted in the need to clarify the larval characters currently used in North American larval keys (Wiggins 1996, Morse andHolzenthal 2008) to separate the genera. We have provided a key to more consistently separate these subfamilies and genera.

Materials and methods
Abdomens were removed, cleared in 10% potassium hydroxide, mounted in glycerin and examined/imaged with the use of Leica stereo and Olympus compound microscopes, Canon DSLR cameras and EOS image capture software. Subsequent images were processed with Zerene image stacking software and Photoshop Elements image editing software. All collected material is preserved in 70% ethyl alcohol. Barcoding was successful for M. antennatus and Barcode of Life Datasystems, Ratnasingham and Hebert (2007), has M. thor barcodes available. Morphological terminology follows that of Wiggins (1996), Wiggins and Currie (2008) and Schmid (1998).

Diagnosis of the new genus Montiphylax
The Montiphylax male genitalia superficially resemble Philocasca; however, examination of head, setal warts, venation, and 9 th and 10 th segments shows no sister relationship. Vshivkova et al. (2007) completely separated Philocasca from the Limnephilinae based on characters of P. rivularis. Philocasca adults are readily separated from Montiphylax adults by the lack of irrorate forewings and metanotal setal warts in Philocasca. Montiphylax females have a mesal lobe on the vulvar scale that Philocasca females lack. Montiphylax larvae lack head and thoracic scale setae that are present on Philocasca larvae. Montiphylax larvae are distinguished from other North American Limnephilidae larvae by the combination of: pronotum not convex in lateral view, with a distinct indentation present between the anterior third to half of pronotum; all gill clusters consist of single filaments; sclerites of metanotal setal area 2 broadly separated; lateral line gills absent, and forked lamellae absent.
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.
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.
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.
Etymology. Montiphylax from the Latin montis (mountain) and the Greek phylax (guard), referring to the mountainous habitat of this genus.
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, eggshaped with smoothly rounded anterior margin.      Philocasca thor Nimmo, 1971: 147-148, figs 143a, b, 545-547, 653. Notes. 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)  Description. 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 Montiphylax albus (Nimmo, 1977) Figs 2, 3, 4A, B, 5, 9, 10 Philocasca alba Nimmo, 1977: 45-46, 49, figs 102-105, 121;Mutch, 1981: 222-228, fig. 43.
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, approxi-mately 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.
Material examined.

Distribution and life history
Like Philocasca, all species of Montiphylax are known only in the Western Cordillera of North America. Montiphylax is the more northern of the two genera and has been found in locations ranging in latitude from ca. 46 to 54 degrees north. Adults of the three species of Montiphylax have been found at altitudes ranging from 1354m (4,442') in the Cascade Mountains (M. antennatus Whatcom Co. Washington) to 1936m (6,350') in the Rocky Mountains (M. thor Waterton National Park). The larvae of M. antennatus and M. albus inhabit small cold sub-alpine streams. The larva of M. thor is unknown but given the locations the adults have been taken, it seems highly likely the larvae of this species will be found in high mountain streams.
A detailed life history is known only for M. albus. To date this species has only been associated with small, cold sub-alpine mountain streams in the Eastern Slope of the Rocky Mountains of Canada. In these streams the life cycle is three years but the presence of a few intermediate sized larvae suggests that the life cycle is flexible. There are five instars. All instars build cases of detritus, predominately of cone bracts, bark, small bits of conifer needles and wood. Instars I -IV have a case with a non-curved triangular cross-section. The V instar case is cylindrical in cross section and is made principally of the same materials as earlier instars except that some small stones are added. The emergence period is from mid-May until mid-June and the flight period lasts until the last week of July. Egg masses have not been located in the field. Growth of larvae is confined to the ice-free period (June -early November). The larvae are detritivores but moss may play a significant role in the diet of the V instars (Mutch 1981).
The flight periods of M. thor and M. antennatus are not known but adults of M. thor have been collected in July and adults of M. antennatus collected in June/July/early August.
Final instar larvae of M. antennatus build a case that has more mineral content than that of M. albus.

Discussion
Montiphylax forms a closely allied group with Homophylax. Montiphylax, Chyranda, Clostoeca, Homophylax, and Phanocelia all fall within the Limnephilidae Genera Incertae Sedis A of Vshivkova et al. (2007), and they all lack larval forked lateral lamellae. This lack of lateral lamellae is a rare condition within the Limnephilidae. During preparation of the enclosed key we also noted that larvae of Pycnopsyche gentilis group also lack the larval forked lateral lamellae. Adult Montiphylax will track to couplet 21 in Schmid (1998) and then fail, not matching either choice. In Ruiter (2000) Montiphylax tracks to Chyranda. Montiphylax can be readily separated from Chyranda by the presence of irrorate wings in Montiphylax. Montiphylax larvae will track to Homophylax or Philocasca in Wiggins (1996) depending on how strongly sclerotized the lateral hump sclerites are. In Morse and Holzenthal (2008) Montiphylax will key to Homophylax, Pycnopsyche, or Psychoglypha for the same reason. The problem with current larval keys is that the arrangement and size of sclerites around the lateral spacing hump is more variable than presented in the keys. As more larval specimens of these western taxa are examined it has become obvious that the lateral hump sclerite spacing and size are not consistent within the genera, particularly within Psychoglypha. We have provided a new key for the North American Limnephilidae larvae that possess gill clusters comprised of only a single filament. Hopefully this will provide for more consistent results within these taxa and lead to improved larval species associations. The pupae of Montiphylax we have will not key to the limnephiloids in Wiggins and Currie (2008) because of the non-hooked labral setae. It is possible that all our specimens (n = 4) possess broken setae since these setae are often broken on pupal material. Since caddis pupae are still poorly described, further work at the family level will be needed.
Key to the North American Limnephilidae larvae which possess gill clusters consisting of a single filament* 1 In lateral view pronotum distinctly inflated at midlength (Wiggins 1996: fig.20.31)  Metanotal setal area 1 & setal area 2 sclerites large in relation to metanotum, distance between setal area 2 sclerites less than 3 times width of one setal area 2 sclerite, usually much less (Wiggins 1996: fig. 20.14) .............Ecclisomyia * † The larvae of Chilostigmodes remain un-described although it is expected to have single filament gill clusters. Chilostigmodes records occur throughout northern North America from Alaska to Newfoundland.
-Metanotal sclerites small, distance between setal area 2 sclerites greater than 3 times width of setal area 2 sclerite (Wiggins 1996: fig. 20.7)  Setae present on metatergal membrane between setal area 2 sclerites (Wiggins 1996: fig. 20.11)  Head and pronotum covered with small spines (Wiggins 1996: fig. 20.19 Banks (1900) described Stenophylax antennatus from a Mt. Rainier, Washington, specimen. At the time he pointed out it was not a true Stenophylax but was waiting for more specimens with which to describe a new genus. Stenophylax antennatus was not mentioned again until the major review of the Limnephilidae (Milne 1935) where Milne placed S. antennatus within Anisogamus based on the forewing shape with long apical cells. Denning (1941) described Anisogamus banksi from a single Wallace, Idaho, male and mentioned that Banks (in lit.) thought it was near Pseudostenophylax edwardsi. Denning's A. banksi type information is: male, Wallace Idaho, 29 April 1938, Otto Huellemann; deposited at University of Minnesota collection. This locality and collector are the same as mentioned by Banks (1943), for a specimen of S. antennatus, although the date is different. Denning's (1941) aedeagal figure (Fig. 11a) is upside down. The Denning holotype was examined during this study, is pinned and in good condition with the cleared abdomen and separated aedeagus within a separate microvial on the pin. Ross (1941) created the genus Philocasca for a single male (P. demita) and placed it within the Limnephilidae, Limnephilinae, Stenophylacini. This placement was based on his conclusion of a close association between P. demita and Anisogamus, and followed Milne's (1935) conclusion. [Until the recent description of a second species (Graf et al. 2015), Anisogamus was a monospecific genus based on A. difformis (McLachlan 1867) known from only a small area in Europe and thought to be most closely related to Stenophylax.] The P. demita type information is: Boyer, Oregon, Hood Craven Cabin, September 30, 1933, J.A.A. The Boyer type locality is questionable as Hood Craven Cabin is not at Boyer. By 1933, Boyer, Oregon, had been relocated from the original locality in Lincoln County to its current location along Highway 18 (Benton County Museum 1981) in far southeast Tillamook County (45.06990 -123.72656) at ca. 600 feet altitude. Ross (1941) Alexander (1954) indicates Hood Craven Cabin is the same as Saddle Mountain, Lincoln County, at 3,000 feet. Saddle Mountain is actually 2,220 feet elevation, and located ca. 60 miles SSW of Boyer. Wiggins and Anderson (1968) indicated that Boyer is in Lincoln County, possibly following the Alexander information, or the original post office location. It is likely an error was introduced by connecting Hood Craven Cabin with Boyer. Banks (1943) indicated that the type of Stenophylax antennatus Banks was a male and provided a re-description and clear, representative figures. He also indicated he had another male from Wallace, Idaho, 12 June (Huellemann) -the same locality/ collector (although different date) as Denning's Anisogamus banksi. Banks (1943) did not mention Denning's (1941) Anisogamus banksi description, or where he thought S. antennatus should be placed. Banks (1943), in his following description of Drusinus frontalis [moved to Eocosmoecus Wiggins and Richardson, 1989], has a note indicating: "We have no species closely congeneric with Anisogamus, and [Anisogamus] edwardsi and [Anisogamus] atripennis are better in Drusinus". [Drusinus Betten (1934) is a synonym of Pseudostenophylax; see Schmid (1955).] Then Banks (1943), in a discussion focused on Philocasca, disagreed with Ross' conclusion that Philocasca was close to Anisogamus and proposed P. demita may be a member of Phacopteryx Kolenati (1848) [currently a synonym of Anabolia Stephens (1837); see Betten andMosely 1940, Schmid 1950]. Banks also stated that Milne's placement of Anisogamus disjuncta, and Asynarchus costalis as Anisogamus was incorrect, resulting in Banks' conclusion that no North American taxa belonged within Anisogamus. In the following paragraph, Banks (1943) created the genus Clostoeca for Clostoeca sperryae and moved Anisogamus disjuncta to Clostoeca. It is clear that Banks had concluded S. antennatus did not belong within Philocasca Ross. Ross (1944), without comment, moved Anisogamus banksi to Philocasca along with P. demita and left S. antennatus as Stenophylax, apparently agreeing with Banks (1943). Ross (1949) described Philocasca oron and pointed out (although did not illustrate) the aedeagal structure was typical for the genus. The primary characters Ross listed to separate P. oron from P. demita were width and thickness of mesal cerci processes (thin and curved ventrad) and reduced spur count (1-2-2  Wiggins and Anderson (1968) noted the collection of a 1-3-4 spur count P. oron specimen. During research for this paper, DER has also noted inconsistent spur counts in P. demita and P. rivularis. Often one hind leg preapical spur is totally absent while the second preapical spur is reduced to a nub. We have been unable to locate fresh Clatsop County, Oregon, Philocasca specimens for DNA analysis. The male and female differences presented to separate P. oron and P. demita are minor variations in shape of a couple structures. A larva distinguishable from P. demita has not been found, although the larvae of the other Philocasca species are readily separated. We suspect P. oron is a synonym of P. demita. The type locality is in an area that has been extensively clear-cut and developed for residential/recreational housing over the intervening years (C Kerst, pers. comm. 2018). Schmid (1955) was the first to formally move Stenophylax antennatus to Philocasca. It is not clear why Schmid did this as he indicated he had not examined any of the Ross Philocasca taxa and that the S. antennatus male was admittedly different from the rest of the Philocasca. Schmid also indicated that females, that would have improved his decision, were not available to him. Schmid (1955), in the discussion of Stenophylax, referenced a paper (Schmid 1955 in press) where he reviewed the genus Stenophylax. The Schmid (1955 in press) Stenophylax paper was essentially completed in 1952 but not published until 1957 (see Schmid 1957). Therefore, Schmid had the Stenophylax information in 1955. In the 1957 (1952) paper Schmid did not mention S. antennatus or his 1955 placement of S. antennatus within Philocasca. This appears to indicate Schmid no longer agreed with his own 1955 placement of S. antennatus within Philocasca.
It is fascinating that this long, convoluted taxonomic history resulting in the placement of Stenophylax antennatus within Philocasca appears to be based on the one male type from Mt. Rainier National Park, Washington, and the description of another S. antennatus male by Banks (1943) from Wallace, Idaho (see discussion under M. thor above). The Banks (1943) description of the irrorate wings, "dark crescentic lobe" dorsally on the aedeagus, and upturned spines (parameres) are clearly not characteristic of Philocasca.
Subsequent authors (Flint 1966, Wiggins and Anderson 1968, Wiggins 1977, Anderson et al. 1982, Wiggins 1996, Schmid 1998) followed Schmid's 1955 placement of Stenophylax antennatus within Philocasca, rather than Banks (1943). Wiggins and Anderson (1968) summarized the known Philocasca species. They provided re-descriptions and illustrations for the holotypes of P. demita, P. oron, P. antennata, and P. banksi males. They also provided new descriptions and figures for the P. demita female and larva; P. oron female; and P. rivularis male, female and larvae. There is also a description for an unknown Philocasca larva. Additional specimens representing this unknown Philocasca species have been examined from Idaho and Montana and, with the association of the Stenophylax antennatus larva, we conclude the unknown larva of Wiggins and Anderson (1986) is P. banksi. Nimmo (1971) described P. thor from a single Alberta collection. The first author examined this holotype and compared it to three additional Alberta P. thor males in the Barcode of Life DNA voucher collection, Guelph. They are the same. The female and larvae of P. thor have yet to be associated. Nimmo (1977) described P. alba from three collections in southern Alberta; including, as paratypes, material the second author collected.
During his studies on litter processing and insect life history within a small Alberta stream, Mutch (1981) discovered that the primary insect litter processor was P. alba. Mutch also noted that the larva associated with P. alba was not similar to the other known Philocasca larvae. He conducted a review of Philocasca (Mutch 1981) and concluded that the seven Philocasca species belong within two separate groups, suggesting that P. antennata, P. alba, and P. thor belonged within a different genus. Further studies planned at that time to resolve the issue ceased until now. The first author compared the P. alba holotype to material from this southern Alberta population and they are the same, resulting in associations of larvae, pupae, males and females for P. alba. Schmid (1998) provided an additional description and illustrations of the P. thor male and the P. demita female.
It was not until the Blinn and Ruiter (2013) collections of P. antennata from Mt. Baker, Washington, that P. antennata material other than the holotype was located. This is likely due to the difficulty of collecting high altitude, isolated habitats early in the season. The Mt. Baker material contained males, females and associated P. antennata larvae and also led to the conclusion that P. antennata, P. alba, and P. thor did not belong within Philocasca.