Micropteryx pardella Walsingham, by original designation.

Epimartyria appears closely allied to the Asian genus Paramartyria as suggested by the similar elongate process arising from the inner base of the male valvae (Fig. 78) and by similar larval chaetotaxy (Hashimoto 2006). More significantly, close affinities of these two northern genera were also indicated from the molecular study initiated by Kobayashi et al. (2000) and Gibbs et al. (2004), based on the 16S rRNA gene. At least one species of the Asian genus Vietomartyria, Vietomartyria nankushana Hirowatari & Hashimoto (Hirowatari et al. 2009), also posseses a similar basal process on the valva, as pointed out by one reviewer. The forelegs of Vietomartyria also lack an epiphysis as do two species of Epimartyria. Epimartyria differs from all other micropterigid genera in possessing a deeply divided phallus, and from Paramartyria in particular by possessing a pair of lateral projections near the apical one third of the distal phallus and in having tergum X divided into dorsal and ventral processes (Hashimoto 2006). Hashimoto (2006) also mentioned the presence of an epiphysis in Paramartyria as one feature that distinguishes the latter from Epimartyria. Two of the three species of Epimartyria lack an epiphysis, but an epiphysis is present in Epimartyria pardella (Fig. 17).

Head (Figs 13–15): Vestiture entirely hairy, scales erect and piliform with acute apices. Antenna (Figs 18–20) 0.75–0.9× length of forewing, slightly longer in male; pedicel enlarged, ~ 1.5× length of first flagellomere; flagellum moniliform, with 46–58 flagellomeres in male, 38–47 in female; flagellomeres mostly sparsely covered with long, piliform scales which exceed the length of their supporting flagellomere; basal 2–3 flagellomeres in male and 5–7 in female covered dorsally with moderately broad scales; a pair of large ascoid sensilla, opposite one another, with ~ 11–16 elongate, curved, sensory branches (Fig. 18) on each flagellomere; a single irregularly shaped and often bilobed multiporus sensillum placodeum (Faucheux 1997) arising between the ascoid sensillae from a shallow pit near the ventral anterior margin of the flagellomere. Compound eyes reduced, interocular index (Davis 1975) ~ 0.35–0.37; interfacetal setae absent. Ocelli present, base moderately elevated. Labrum approximately pentagonal, length ~ 2× that of clypeus. Mandible elongate triangular in form; distal edge truncate. Maxillary palpus elongate, 5-segmented, with main flexions between segments 1 and 2 and between 3 and 4; length ratio from basal segment 1: 1:2.7:2.7:0.9. Labial palpus short, total length ~ equal to that of basal segment of maxillary palpus; 2- segmented; sensory pit (organ vom Rath) present distally on apical segment enclosing numerous sensillae; apices of most sensillae terminating in a cluster of ~ 2- 5 minute acute lobes (Figs 21–22). Proximal prelabial sclerite slender, crescentiform; distal prelabial sclerite broadly triangular. Occipital sulcus incomplete but distinct laterally.

Thorax: Scales of mesonotum broad, appressed. Metanotum mostly naked except for a few long, piliform scales. Tegulae rather sparsely covered with long piliform scales. Forewing length: 4.2–5.5 mm; forewing (Fig. 16) with humeral vein present; Sc deeply bifurcate; R simple; Sc-R crossvein present near fork of Sc; Rs with 4 veins; Rs3–4 fused to ~ basal 1/3; accessory cell present; M with 3 branches; 1A and 2A fused over distal half; 3A extending across base of moderately small jugal lobe. Wing scale morphology of the primitive, generally non-glossatan type (Kristensen and Simonsen 2003) consisting of fused dorsal and ventral surfaces (without internal chambers), and with a herringbone pattern formed by oblique-longitudinal crests overlying a dense layer of transverse microribs (Figs 28–31). Hindwing venation similar to forewing except with Sc and R fused; 1A and 2A completely fused; anal crossvein connecting to CuP near distal 2/3; scales over distal third of hindwing dark fuscous and nearly as broad and iridescent as in forewing; scales gradually becoming more slender, gray, and without iridescence over basal 2/3. Legs (Fig. 17) with tibial spur pattern of 0–0-4; a short epiphysis ~ 1/3 the length of tibia arising slightly beyond its midlength present in Epimartyria pardella; epiphysis absent in Epimartyria auricrinella and bimaculella; pretarsus (Figs 24–27) consisting of a pair of strongly curved claws; a lateral pair of pad-like pulvilli densely covered with long spinose setae; a median arolium with apical surface densely lined with minute grooves (Fig. 27); pseudempodial seta (Fig. 25) with longitudinal grooves.

Abdomen: Cuticle dark brown, sparsely covered with long, piliform scales. A pair of glands present, opening on sternum V in both sexes (Philpott 1925); glands similar to those present in Paleomicroides, Paramartyria, and Neomicropteryx in not protruding and possessing a narrow slit-like opening within a smooth, hyaline area (Kristensen 1984a).

Male genitalia: Tergum X (uncus) ~ half the median ventral length of IX; apex deeply divided nearly half its length into two broad lobes. Sternum X (venter X) variously bilobed, with or without short lateral lobes. Segment IX a completely sclerotized ring, with dorsal median length ~ 1/6 of ventral length. Sternum IX (vinculum) a broad plate with subparallel lateral margins; anterior end as broad or broader than caudal end. Valva with a subacute to rounded apex; base of valva with a long digitate process from mesal surface. Medial plate (juxta) with a slender stalk-like base gradually expanding anteriorly to a small, flat, oval plate. Distal phallus divided into two slender branches ~ half the total length of phallus; shorter dorsal branch of phallus terminating in gonopore (phallotreme) with thickened radial folds; a pair of minute, acute spines present laterally near distal third of dorsal branch; apex of longer ventral branch densely covered with numerous minute flattened scutate processes with rounded apices directed basad; phallobase moderately inflated, as long as or slightly longer than divided branches.

Female genitalia: Abdominal segment IX a complete ring with mid dorsal length ~ 0.5–0.6× the mid ventral length. Segment X consisting of a pair of lateral, setose plates; cloaca ending terminally; X often telescoped into IX and VIII in repose. Apophyses absent. Genital chamber with thickened walls surrounding a variably shaped slerite; caudal end of sclerite furcate. Ductus spermatheca with a moderately enlarged, spindle-shaped reservoir (utriculus) located at varying distances along ductus. Corpus bursae gradually enlarging anteriorly, membranous, with four tridentaform signa equally spaced around middle of corpus bursae; enlarged bases of signa projecting externally beyond wall of corpus bursae, with spinose branches projecting internally.

For many years John Heath, formerly employed at the Experimental Research Station at Monks Wood in England, pursued research on the family Micropterigidae, resulting in about 20 papers on this group (Emmet 1987). Heath had partially completed a revision of the genus Epimartyria, but this was never published. We had not viewed a copy of this manuscript until our publication was in review. In his manuscript, Heath recognized an additional new species from New Jersey, based on specimens collected at Essex County Park by W. D. Kearfott. Our studies found no morphological justification for this species.

Because this is the first taxonomic revision of Epimartyria, there remain some gaps in our knowledge about their biology which cannot be answered with available material and evidence.

Adult Epimartyria auricrinella are easily distinguished from those of the other members of Epimartyria in possessing uniformly dark fuscous forewings without the yellowish spots present in those species.

(Figs 1, 4–5). Head: Vestiture light orange brown. Antenna with vestiture of scape and pedicel concolorous with head; scales of flagellum dark brown to fuscous. Labial palpus cream.

Thorax: Dark fuscous with coppery to purplish luster. Tegula concolorous with head. Forewing dark fuscous with coppery or golden to purplish luster dorsally, less iridescent ventrally; fringe paler, more gray. Forewing length: 4.2–5.6 mm. Hindwing with scales over distal third nearly as broad, dark fuscous and iridescent as in forewing; scales gradually becoming more slender, more gray, and less iridescent over basal 2/3; fringe gray. Legs medium to dark brown dorsally, light brown ventrally and at apices of tarsomeres; epiphysis absent.

Abdomen: Piliform scales uniformly brown dorsally and ventrally. Paired glands of sternum 5 with muscle for opening glands originating on anterior edge of sternum 6 and inserted into each gland duct just inside aperature; gland reservoir slightly larger and more ovoid in female, but surrounding layer of secretory cells better developed and 2–3× thicker in female (Djernaes 2011, Djernaes and Sperling 2011).

Male genitalia (Figs 74–78): Caudal lobes of tergum X broadly rounded. Caudal apex of sternum X deeply divided, with apex of lobes acute, recurved; a pair of short, lateral lobes present near base. Valvae moderately long, ventral length nearly half the maximum length of segment IX; apex subacute and bearing a short, slender, recurved spine; a short, triangular, rounded process arising midway from mesal surface; elongate basal process ~ 4/5 the length of valva; distal margin of valva variable within populations from slightly concave to convex (Figs 78a-d). Dorsal branch of phallus cylindrical and smooth.

Female genitalia (Figs 79–80): As described for genus. Caudal end of genital sclerite moderately furcate as in Epimartyria bimaculella; length of furcations ~ 0.2 that of relatively shorter, undivided base.

(33–59). Mature larva up to 5 mm in length. Body approximately hexagonal in cross section; color generally brown, lighter brown ventrally. Integument over dorsal half of body with a honeycomb-like surface of raised ridges (Figs 52, 58); integument of ventral half densely covered with micropapillae (Fig. 56) with an extensive plastron surface laterally (Fig. 52). Primary setae longitudinally ribbed, moderately slender, long, clavate.

Head: Prognathous and capable of being retracted into prothorax. Antenna elongate, slender, 3- segmented, arising posterior of clypeal margin and dorsal to stemmata; second segment the longest, ~ 2× the length of basal segment; all antennal segments without sensory setae except for elongate terminal spinose seta. Five stemmata present, arranged in a tight circle. Adfrontal sutures vestigial, not extending to vertex; adfrontal ridges similarly undeveloped. Ecdysial lines externally indistinct. Tubular spinneret absent; external opening of labial salivary gland circular, relatively large, diameter ~ equal to length of second segment of labial palpus. Cranial setae reduced in length and number and concentrated over anterior third of head; stemmatal setae absent; a single medial (M) seta arising midway between antennae, without homology in other Lepidoptera but possibly homologous to campaniform sensillum in Trichoptera larva (Kristensen 1998). Labrum with 6 pairs of primary setae and numerous spines along anterior margin; seta La 1 arising distad of anterior margin of labrum (Fig. 38). Mandible generally triangular in form with 3 acute cusps, the basal-most cusp the most reduced. Maxillary palpi relatively well developed, 3–segmented, with apical sensillae as in Fig. 46. Labial palpi reduced, 3 - segmented with minute apical segment bearing a long sensillum (Fig. 47). Intersegmental membrane between head and thorax covered with flattened, multidentate, scutate outgrowths (Figs 41–42).

Thorax: Prothorax with 7 primary tactile setae and 4 peg-like microsetae, the latter located along anterior margin of prothorax near the head- prothoracic fold; XD1 and XD2 greatly reduced to peg-like microsetae along dorso-anterior margin of prothorax below D2; L1 posterior to XD1; L2 below L1 and anterior to spiracle. MV1 and MV2 short, peg-like, below SV2 and closer to anterior margin of prothorax; MV2 about 2 × length of MV1. Subdorsal setae absent on all body segments. Meso- and metathorax with 5 primary setae and one microseta (SV2); L1 and 2 well developed and equal in size. Legs with 3 well defined segments and large pretarsal segment; 4- segmented including reduced coxa; pretarsal claw curved, elongate, ~ 1/3 the length of remainder of leg; axial spine at base of claw well developed; femur and trochanter fused, as well as tibia and tarsus; coxa with a bilobed and possibly eversible tactile vesicle located posterior-mesally near base of femur (Figs 48–49);

Abdomen: Segments 1–8 with 4 primary setae and 2 peg-like (L2) to spherical microseta (SV2); segment 9 with only D1 and L1; segment 10 with 2 microsetae, possibly representing D1 and L1. Spiracles peripneustic, located anteriorly in intersegmental fold on segments 1–8; spiracle raised to form a small dome with walls subdivided into ~ 10–12 fimbriated bands. Abdominal segments 1- 8 with short, fleshy, nonmuscular prolegs with rounded apices (Fig. 50); crochets absent in all genera of Micropterigidae.

Hepaticophyta: Lepidoziaceae: Bazzania trilobata (L.) S. Gray.

Unknown.

(Figs 9–10). The species occurs in shaded locations, in wet swampy woods, boggy ditches, or creek sides where leafy (moss-like) liverworts, the probable larval host, grow. Such habitats can be periodically or seasonally flooded. Larvae possess a plastron which indicates the capacity to live for short periods in a subaquatic environment or, at least in a habitat that is water-saturated. Adults are diurnal and are best obtained by gently sweeping the understory or clumps of liverworts (Landry and Landry 1992). They can be seen perched on low foliage during the day and can be active even in early morning after sunrise (Figs 1–2). Mating was observed in the afternoon between 1200–1700H (JFL pers. obs.). Larvae obtained (by JFL) by placing in a Berlese funnel clumps of the liverwort Bazzania trilobata collected on 3 September 2000 at Lac Brûlé (Quebec) yielded larvae of two different size classes (3.4 mm vs 1.8 mm overall body length). This supports the previous observations by Davis (1987) that larval development probably spans over two years, at least in the northern part of the range, although adults emerge every year. One larva was found on the tip of a liverwort leaflet at the same locality in early October when the air temperature was around 5°C. Adults generally begin to emerge in mid May in the southern part of their range (Georgia, North Carolina) with April 30 being the earliest date recorded (from southern Maryland near Washington, DC). Further north the flight period is gradually delayed, with adults in northern New York and all of Canada active during the summer between mid-June and mid-July.

♂, USA: North Carolina, 1884, H. K. Morrison, Type No. 35325, slide BM 8947 (BMNH).

CANADA: NOVA SCOTIA: Baddeck: 1 ♂, 23 Jun 1936; 1 ♂, 30 Jun 1936, T.N. Freeman, specimens # CNCLEP00077282–00077283, slide MIC1825 (CNC); Parrsboro: 1 (abdomen missing), 12 Jul 1944, J. McDunnough, (CNC). ONTARIO: Ottawa: 4 ♂, 19 Jun 1906, C.H. Young, slide USNM 16615 (USNM, CNC); 5 ♂, 20 Jun 1906, C.H. Young, slide USNM 34372, specimens # CNCLEP00077266–00077268, CNC slide MIC1822 (CNC, USNM); 1 ♂, 27 Jun 1906, slide USNM 98008 (USNM); 1 ♂, 12 Jun 1946, G.S. Walley, specimen # CNCLEP00077269 (CNC). Black Lake, N of Burgess Township: 1 ♂, 22 Jun 1974; 2 ♂, 1 ♀, 14 Jun 1975, J.A. Downes, specimens # CNCLEP00077277–00077280 (CNC). Moosonee: 1 ♀, 18 Jul 1934, G.S. Walley, specimen # CNCLEP00077288 (CNC). Orillia: 3 ♂, 26 Jun 1926; 1 ♂, 3 ♀, 2 Jul 1926, C.H. Curran, specimens # CNCLEP00077276–00077270–00077276, CNC slide MIC1823 (CNC). Thunder Bay: 1 ♀, Jul 1945, H. S. Parish. QUEBEC: Havre-Saint-Pierre: 4 ex., 3–17 Jul 2010, malaise trap, C. Bélanger. Gaspé Peninsula: Mont Albert: 2 ♂, 1 ♀, 18 July 1940, A. E. Brower, slides USNM 16621, 17501, SEM slide 18394 (USNM); 11 ♂, 1 ♀, 19 Jul 1940, 2♂, 22 Jul 1940; side Mt. Albert: 2 ♂, 2 ♀, wing slide USNM 16157, slides USNM 18396, 18408, 98007, SEM slide 18430, (USNM, CNC). Mansonville: 1 ♂, 18 Jun 1928, W.J. Brown, specimen # CNCLEP00077286, CNC slide MIC1824 (CNC). Ste-Agathe-des-Monts, Lac Brûlé, 46.0903°N, 74.26°W, 370 m: 11 ♂, 6 ♀, 26 Jun 1991, afternoon sweeping in swampy ditch with liverworts and mosses at edge of spruce forest, J.-F. Landry, specimens # CNCLEP00076615–00076631 (CNC); Lac Brûlé, 46.0909°N, 74.2756°W, 370 m: 9 ♂, 3 ♀, 8 Jul 1992, 2 ♂, 16 Jul 1992, day sweeping on shaded liverworts near boggy marsh, J.-F. Landry; specimens # CNCLEP00068799–00068800 (CNC, USNM); Lac Brûlé, 46.0903°N, 74.26°W, 370 m: 3 ♂, 2 ♀, 1 Jul 1993, afternoon sweeping liverworts and mosses, J.-F. Landry, specimens # CNCLEP00067565–00067569 (CNC); Lac Brûlé, 46.0885°N, 74.2789°W, 370 m: 1 ♂, 4 Jul 1993, day sweep in mixed forest, J.-F. Landry, specimen # CNCLEP00076570 (CNC); Lac Brûlé, 46.0909°N, 74.2756°W, 370 m: 4 ♂, 2 ♀, 7 Jul 1993, day sweeping in shaded spruce-birch forest swamp, J.-F. Landry, specimens # CNCLEP00076571–00076576 (CNC); Lac Brûlé, 46.0885°N, 74.2789°W, 370 m: 1 ♂, 9 Jul 1993, at mercury light in mixed forest, J.-F. Landry, specimens # CNCLEP00076577 (CNC); Lac Brûlé, 46.0885°N, 74.2789°W, 370 m: 1 ♂, 1 Jul 1996, sweeping ferns at forest edge in afternoon, J.-F. Landry, specimens # CNCLEP00076577 (CNC); Lac Brûlé, 46.0885°N, 74.2789°W, 370 m: 1 ♂, 30 Jun 1997; 1 ♂, 1 Jul 1997, day sweep in mixed forest, J.-F. Landry, specimen # CNCLEP00076578 (CNC); Lac Brûlé, 46.0812°N, 74.2833°W, 370 m: 2 ♀, 1 Jul 1997, sweep in forest trail ca 18:00H, sunny, J.-F. Landry, specimens # CNCLEP00076580–00076581 (CNC); Lac Brûlé, 46.0919°N, 74.2756°W, 370 m: 13 ♂, 5 ♀, 2 Jul 1997, in swampy wood perched on vegetation 11:30H–12:30H overcast just before thunderstorm, J.-F. Landry, specimens # CNCLEP00076582–00076599 (CNC); Lac Brûlé, 46.0921°N, 74.2756°W, 370 m: 1 ♂, 1 ♀, 2 Jul 2000, afternoon sweeping liverworts and low vegetation in forest swamp, specimens # CNCLEP00067787–00067788, CNC slide MIC5756, DNA barcoded (CNC); 14 ♂, 5 ♀, 8 Jul 2002, day sweeping shaded liverworts near boggy marsh, specimens # CNCLEP00007712–00007720, 00068787–00068788, CNC slides MIC5753, MIC5755, MIC5757, MIC5758, MIC5760, MIC5762, MIC5763, 9 DNA barcoded (CNC); 6 ♂, 29 Jun 2003, in mixed forest swamp day-sweeping herbaceous and shrub vegetation, J.-F. Landry, specimens # CNCLEP00002816–00002821, CNC slide MIC5761, DNA barcoded (CNC); Lac Brûlé, 46.0881°N, 74.2788°W, 370 m: 1 ♂, 1 ♀, 4 Jul 2004, day sweep in forest swamp with liverwort, J.-F. Landry, specimens # CNCLEP00006682–00006683, CNC slide MIC5754, 1 DNA barcoded (CNC). Gatineau Park, Ramsey Lake, Hopkin’s Hole, 45.6025°N, 76.1079°W, 245 m: 12 ♂, 3 ♀, 11 Jun 1991, afternoon sweeping in forest swamp, J.-F. Landry, specimens # CNCLEP00076600–00076614 (CNC). Gatineau, Masham Township, 45.68°N, 76.05°W: 1 ♂, 26 Jun 1974; 1 ♀, 30 Jun 1974, D.M. Wood, specimens # CNCLEP00077284–00077285 (CNC). UNITED STATES: GEORGIA: Rabun Co: Chattahochee National Forest, Tate Br. Campground: 1 ♀, 16–17 May 1970, O. S. Flint, Jr. (USNM). KENTUCKY: Powell Co: 1 ♂, 23 Nov 1909, 1 ♂, 25 May 1924 (USNM). MAINE: Aroostook Co: Round Mountain: 1 ♂, 20 Jul 1956. Piscataquis Co: Greenville: 1 ♂, 9 Jul (USNM). Franklin Co: West Farmington: 1 ♂, 29 Jun 1966, A. E. Brower (USNM). Hancock Co: Acadia National Park, Mt. Desert Island: 1 ♂, 30 Jun 1933, (USNM). Penobscot Co: Passadumkeag: 1 ♀, 25 Jun 1938 (USNM). Piscataquis Co: Baxter State Park, Mt. Katahdin, Hunt Trail, 2400 feet: 1 ♂, 17 Jul 1948, bushes by brook, A. E. Brower, slides 16388, wing USNM 29861 (USNM). Sagadahoc Co: Woolwich: 1 ♀, 29 Jun 1965, A. E. Brower, slide USNM 33917 (USNM). MARYLAND: Montgomery Co: Cabin John: 1 ♂, 30 Apr 1921, A. Busck (USNM). MICHIGAN: Keweenaw Co: Isle Royale National Park: 2 ♂, 10 Jul 1957, R. W. Hodges (USNM). Emmet Co: Wilderness State Park, 45.7119°N, 84.9402°W, 180 m: 6 ♂, 30 Jun 1992, 17:00–18:00 hrs sweeping liverworts on banks of shaded stream in oak-pine forest with thuja, J.-F. and B. Landry, specimens # CNCLEP00068781–00068786, CNC slides MIC5752, MIC5764, DNA barcoded (CNC). NEW HAMPSHIRE: Rockingham Co: Hampton: 1 ♂, 6–11–1904, S.A. Shaw (USNM). NEW JERSEY: Essex Co: Essex Co. Park: 1 ♂, 3 Jun 1900, W. D. Kearfott (USNM). Essex Co: 1 ♂, 3 Jun; 7 ♂, 3 ♀, 8 Jun 1907, W. D. Kearfott, slides USNM 18409, 91794, 91795 (USNM). NEW YORK: Essex Co: [Keene]: Table Top Mountain, 3500 feet: 2 ♂, 1 ♀, 21 Jul 1940 (USNM). NORTH CAROLINA: Swain Co: Great Smoky Mountains National Park: Oconaluftee River at Towstring Road: 1 ♂, 11 May 1970, SEM slide USNM 17565 (USNM). Smokemont Campground and nearby: 2 ♀, 11–14 May 1970, slide USNM 33920, head slide 16614 (USNM). Whitewater River at rt. 171: 1 ♂, 18 May 1970, O. S. Flint, Jr. (USNM). PENNSYLVANIA: Dauphin Co: Inglenook: 1 ♀, 30 May 1911 (USNM). SOUTH CAROLINA: Pickens Co: Clemson, Wildcat Creek: 1 ♂, 25 Apr 1968, P. Carlson, J. Morse (USNM). TENNESSEE: Sevier Co: University of Tennessee Field Station, 35.739°N, 83.4235°W, 503 m: 1 ♀, 22 May 2005, afternoon sweeping vegetation along forest creek, J.-F. Landry, specimen # CNCLEP00016403, CNC slide MIC5759, DNA barcoded (CNC). VIRGINIA: Falls Church: 1 ?, 1908, A. Busck, wing slide USNM 91787 (USNM). WEST VIRGINIA: Pendleton Co., Smoke Hole State Park, Briggs Run: 3 ♂, 2 ♀, 28 May 1977, [sweeping low vegetation 8–11 AM], D. & M. Davis, slide USNM 20690 (USNM).

(Fig. 32).Epimartyria auricrinella occurs widely over eastern North America, in Canada from Nova Scotia to Ontario, and in the U.S. from Maine to Michigan and south to Tennessee and Georgia.

Chaetotaxy:Because the larvae of Micropterigidaelacksome thoracic and abdominal setae present in higher Lepidoptera, determining the homology of those setae present is subject to uncertainty. Various assumptions have been made as to which setae are present, based in part on their position to longitudinal muscle groups or to various body ridges (Hashimoto 2001, 2006, Gibbs 2010), as well as to the generally accepted chaetotaxy of glossatan Lepidoptera (Hinton 1946, Stehr 1987) which was followed by Davis (1987). Greatest uncertainty persists with the prothoracic chaetotaxy, where the number and relative development of setae can vary between different genera of Micropterigidae. Hashimoto (2001) concluded that the XD (of the prothorax) and the SD groups are absent in Micropterigidae, with the possibility that the two most dorsal of the four pairs of peg-like microsetae along the anterior margin of the prothorax in Epimartyria could be vestiges of one or more of these groups. The more ventral of the two microsetae along the anterior margin of the prothorax are believed to represent MV1 and MV2 present in most Lepidoptera, but homology of the dorsal pair is questionable. Because microsetae in this region are not known to occur on the prothorax of other Lepidoptera (Hinton 1946), we have considered the dorsal pair to be homologous to XD1 and XD2 as suggested by Hashimoto (2001). Hinton (1946) briefly discussed the possibility that the XD group in higher Lepidoptera might be homologous to the microsetae of other body segments, but he argued that long tactile setae along the front margin of the prothorax represented instead a special setal group essential for protecting larvae, especially in those species with prognathous, retractable heads. In Micropterigidae it appears as if this protection has been compensated by several of the prothoracic tactile setae being directed strongly forward (Fig. 33). It may also be possible that several proprioreceptor (microscopic) setae homologous to those of higher Lepidoptera do not occur in Micropterigidae, and that all or most of the relatively stout microsetae present may represent greatly reduced tactile setae. Such reductions have occurred with the abdominal L2 and SV2 setae of Epimartyria (Fig. 33), D1 and D2 of Austromartyria (Gibbs 2010), and D2 of Agrionympha (Gibbs and Kristensen 2011). All microsetae of Epimartyria auricrinella are similar to the prothoracic microsetae in being relatively stout and greatly reduced it length (Fig. 33). Consequently, in this study we have largely followed the protothoracic chaetotaxy proposed by Hashimoto (2006) for the closely related genera Paramartyria and Neomicropteryx. The number and distribution of the prothoracic microsetae have not been well studied or illustrated in most genera of Micropterigidae. Better resolution of the prothoracic setal homology might become possible as larvae of more genera are discovered and studied.

Prolegs: The larval prolegs of Micropterigidae, which occur on abdominal segments 1–8 and 10, differ in their morphology from those of all other Lepidoptera where crochet- bearing prolegs are typically present only on segments 3–6 and 10. Hinton (1958) also reported muscles to be lacking in micropterigid prolegs, although this probably should be examined further in some genera such as Micropterix where the prolegs appear more developed and with more melanized, acute, clawlike apices (Figs 66–68). The anal prolegs of Micropterix are also distinct in forming a relatively broad, trilobed sucker (Fig. 66; Zeller-Lukashort et al. 2007).

Integumental specializations: Larvae of Micropterigidae often occur close to the ground in habitats more likely to be subjected to periodic flooding and drying. As an adaptation to such conditions, the larvae have developed an unusual cuticular morphology in the form of a physical gill, or plastron (Thorpe 1950, Davis 1987), which provides extensive air–water interface for gaseous exchange. The aquatic larvae of several species of Crambidae have also developed special gills and plastron cuticles for breathing underwater (Wichard et al. 2002).

An extensive plastron area has been observed in Epimartyria (Davis 1987), and similar cuticular structures with various modifications appear in other genera of Micropterigidae examined. The plastron in Epimartyria auricrinella extends as a broad band laterally around the body between the level of the lateral (L1) and subventral (SV1) setae and then dorsally over the posterior margin of the prothorax (Fig. 33). The abdominal spiracles are located near the dorsal margin of the band at the extreme anterior edge of the segment (Fig. 52). The surface of the integument within this zone is densely covered with minute, irregularly shaped micropapillae (Figs 52, 55). Radiating out between adjacent micropapillae are dense series of even smaller ridges, aligned ~ 0.4–1.0 µm apart. Each ridge bears a single row of elongate, erect, knobby microtubercules ~ 0.2–0.4 µm in diameter and ~ 0.8–1.2 µm in length (Fig. 56). These minute structures are believed to help form an air film around that portion of the body (when submerged) that excludes water under normal hydrostatic pressure. Minute openings in the epicuticle are visible between the ridges (Fig. 57). These lead internally into an unusually complex, multichambered exocuticle reported by Kristensen (1990, 1998) in the larvae of Sabatinca and Micropterix. The basal layer of the exocuticle was found in these genera to possess small pores in each chamber which opened into a fluid- filled space between the exo- and endocuticle. Kristensen hypothesized the function of these unique cuticular specializations may be to assist in maintaining a water balance for larvae in a habitat subjected to periodic drying. An extremely thin, extracuticular pellicle covers much of the dorsal, lateral, and part of the ventral larval trunk to which small particles may adhere. The function and origin of the pellicle remain unknown. Immediately beneath the abdominal pellicle of Epimartyria auricrinella the exocuticle is divided into a series of honeycombed chambers (Fig. 58) resembling the condition Kristensen discovered in the larvae of Sabatinca and Micropterix.

The multidentate, scale-like cuticular outgrowths (Figs 41–42) covering the intersegmental membrane between the head and prothorax of Epimartyria auricrinella may further assist in a respiratory function. These structures superficially resemble the plastron scales present in certain Coleoptera (Hinton 1969, 1976). The spiracles in Epimartyria (Figs 53–54) are also modified to prevent water entry. Each spiracle is raised into a small dome with finely divided, fimbriated walls. The spiracles in later instars of Neomicropteryx larvae are also conical with fimbriated walls, but the spiracular walls of the first instar are completely fused (i.e., solid) (Hashimoto 2006) as they are in later instars of Sabatinca and Micropterix (early instars not examined). Spiracles of the first instar of Epimartyria have not been examined but may be similar to those of Neomicropteryx.

The larval plastron of Neomicropteryx nipponensis Issiki is similar to that of Epimartyria auricrinella in possessing a dense zone of minute, irregularly shaped micropapillae interconnected by dense radiations of smaller ridges bearing rows of knobby microtubercules (Figs 60–62). Scutate outgrowths also arise from the intersegmental head-prothoracic membrane (Fig. 63) as in Epimartyria. It is likely that larvae of all members of the northern hemisphere group of micropterigid genera proposed by Kobayashi et al. (2000) and Gibbs et al. (2010) have developed similar plastron specializations. In contrast, the external surface of the exocuticle of Micropterix (Figs 70–72) possesses a more extensive, regular arrangement of micropapillae, each ~ 10–20 µm in diameter, with ~ 6- 8 relatively stouter, often bifurcate, arm-like ridges radiating from a central disk. The ridges in Micropterix do not continue with those of adjacent ridges, but the extremities of each ridge are densely covered with microtubercules. Minute openings of variable size are present in the exocuticle of Micropterix (Figs 71–72), similar to those observed in Epimartyria and Neomicropteryx.

Adultsof Epimartyria bimaculella most resemble those of Epimartyria pardella in possessing dark fuscous forewings marked by pale golden spots. A total of two yellowish spots occur in bimaculella, with only a single large costal spot present beyond the middle of the forewing. Four spots are present on the forewing of pardella, with two of these located across the distal third of the wing on the costal and dorsal margins respectively.

(Figs 2, 6–7). Head: Vestiture similar to Epimartyria auricrinella, light orange brown. Antenna with vestiture of scape and pedicel concolorous with head; scales of flagellum mostly pale golden brown, becoming darker, more fuscous over distal third. Labial palpus cream.

Thorax: Dark fuscous with coppery to purplish luster. Tegula concolorous with head. Forewing mostly dark fuscous with coppery to purplish luster dorsally, marked with two pale yellowish spots; the largest, irregularly oval to rectangular spot extends from the costa approximately halfway across the distal third of wing; a second smaller, more slender spot extends diagonally from about midway along dorsal margin to midway on discal cell; a slight suffusion of pale yellowish scales may be sometimes evident at the base of the forewing, but only seldom does this occur; forewing less iridescent ventrally; fringe pale yellow along termen, fuscous along dorsal margin. Forewing length: 4.6–5.3 mm. Hindwing mostly gray, becoming darker and slightly iridescent toward apex; fringe gray. Legs medium to dark brown dorsally with a slight purplish luster, light brown to cream ventrally; epiphysis absent.

Abdomen: Piliform scales dark brown dorsally, paler brown ventrally.

Male genitalia (Figs 81–85): Tergum X similar to Epimartyria auricrinella, broadly bilobed. Caudal apex of sternum X deeply divided, with apex of lobes acute, only slightly curved; a pair of short, lateral lobes present near base. Valva moderately long, ventral length ~ half the maximum length of segment IX; apex subacute and bearing a short, slender, recurved spine similar to Epimartyria auricrinella; a short but broader and more triangular, rounded process arising midway from mesal surface; elongate basal process ~ 4/5 the length of valva; distal margin of valva variable within populations from slightly convex to ~ straight. Dorsal branch of phallus cylindrical and smooth.

Female genitalia (Figs 86–87): As described for genus. Caudal end of genital sclerite moderately furcate as in Epimartyria auricrinella; length of furcations ~ 0.3 that of moderately long, undivided base.

Unknown.

(Figs 6–7, 11). At the type locality, specimens were captured by sweeping low lying vegetation or during diurnal flight along a shaded seepage in a Douglas Fir–Western Red Cedar forest where leafy liverworts grew. Adults were also observed perching on lower parts of plants such as Salmonberry (Rubus spectabilis Pursh) no more than approximately 25 metres from the liverwort habitat (D.G. Holden, pers. comm.). In different parts of the range, specimens were collected from late April to mid August, with most records in June. Late records (July and August) are from higher elevations.

♂, CANADA: BRITISH COLUMBIA: Belcarra, 49°17'59.11"N, 122°55'30.88"W, Alt. 25 m., 8 Jun 2008, visual sweep, Dave G. Holden, specimen # CNCLEP00067716, CNC slide MIC5768, Barcode of Life Project, leg removed, DNA extracted, digital image captured, (CNC).

CANADA: BRITISH COLUMBIA: Belcarra Park, 49.3107°N, 122.9263°W, alt. 13 m: 2 ♂, 24 May 2009, day sweep, Dave G. Holden, specimen # CNCLEP00076632–00076633 [both DNA barcoded] (CNC); 13 ♂, 1 ♀, 1 Jun 2009, day sweep, Dave G. Holden, specimens # CNCLEP00076634–00076639, 00077846–00077853, CNC slides MIC5765, MIC5767, MIC5766, MIC5570, MIC5571 [all DNA barcoded] (CNC); 2 ♂, 2 Jun 2009, day sweep, Dave G. Holden, specimens # CNCLEP00076640–00076641 [both DNA barcoded] (CNC). Maple Ridge, Univ. of British Columbia Research Forest, 49.277679°N, 122.553870°W, 259 m: 1 ♂, 1 Jun 2011, visual sweep, Dave G.Holden (CNC). Fraser Mills: 6 ♂, 7 ♀, 11 June 1921, L. E. Marmont, SEM slide USNM 18431, slides USNM 33919, 98001, 98002, 98004; 27 ♂, 7 ♀, 15 Jun 1922, E. H. Blackmore collector, slides USNM 17503, 18410–18411, 34282, 91785, 97991, 98000, 98003, 98005, 98009–98017 (BMNH, CNC, USNM). Squamish, Diamond Head Trail: 1 ♂, 12 Aug 1963; 2 ♂, 14 Aug 1963, W.R.M. Mason, specimens # CNCLEP00077292–00077294, CNC slide MIC1826 (CNC). Mt Seymour, 49.337368°N, 122.957695°W, 292 m: 1 ♂, 21 Jun 2011, visual sweep, Dave G.Holden (CNC). Glacier National Park, Loop Trail, 1140 m, 51.254°N, 117.538°W, 1 ♀, 16 Jul 2010, malaise trap, specimen #10BBCLP-2914 [DNA barcoded], CNC slide MIC5769 (BIO). UNITED STATES: Washington: Clallam Co: Olympic National Park, sweeping on Soleduck Trail to Deer Lake, 1000 m: 3 ♂, 15 Jul 1998, D. R. Davis, slide USNM 34302 (USNM). Olympic Peninsula, Port Angeles, 245m, 48, 07924°N, 123, 42990°W ± 50m: 3 ♂ , 1 ♀, 20 Jun 2010; 1 ♂, 20.6.2010, 15:45h, Hausenblas and Zeller-Lukashort (ONPS). Olympic Peninsula, Sol Duc Hot Springs Rd, 390m, 48.06385°N, 123.99565°W ± 50m: 1 ♂, 21 Jun 2010, 13:00h, Hausenblas and Zeller-Lukashort, slide AP-Nr 42/2010 Christof Zeller (ONPS). Olympic Peninsula, Kalaloch, 10m 47.61131°N, 124.37588°W ± 50m: 1 ♂, 23 Jun 2010, 17:00h, Hausenblas and Zeller-Lukashort (ONPS). Olympic Peninsula, Hoh Rainforest Rd, 130m 47.81641°N, 124.05161°W ± 50m: 21 ♂ , 3 ♀, 22 Jun 2010, 16:15h, Hausenblas and Zeller-Lukashort (CZC). Grays Harbor Co: Elma: 46.9738°N, 123.2945°W, yel st trp, 1 ♀, 27 Jun 2011, G. Kohler, WSDA 978–1008A (WSCAD). King Co: Asahel Curtis picnic area: 47.3951°N, 121.4677°W, 1 ♂, 27 Jul 2011, hand col, C. Looney, WSDA W666–1129A, B (WSCAD, USNM). Stevens Pass, Hwy 2, 14.5 km E Skykomish, 645 m., 47.7143°N, 121.1722°W: 1 ♂, 8 Jul 2010, afternoon sweep, J.-F. Landry and D.G. Holden, specimen #CNCLEP00082605, CNC slide MIC5739 [DNA barcoded] (CNC). Mason Co: Skokomish River Rd: 47.3019°N, 123.1858°W, 4 ♂ , 2 ♀, 17 Jun 2011, hand col, C. Looney, WSDA W666–1131A-E (WSCAD, USNM). Pierce Co: Fort Lewis: 1 ♂, 29 May 1951, R. Schuster, Essig Museum slide 0152, (UCB). Snohomish Co: East Arlington Co. Park: 1 ♂, 29 Apr 1979, L. Massell, e. Lepidozia liverwort, slide USNM 98006 (USNM). 6 mi. E of Verlot: 1 ♂, collected 26 Mar 1979, emerged 2 May 1979, reared from liverwort, “Jungermannia obovata” L. Russell (USNM). Thurston Co: Evergreen State College, 47.0791°N, 122.9750°W, 2 ♂, 25 Jun 2011, hand col, C. Looney & E. Lagasa, WSDA 666–1130A, B (WSCAD, USNM).

Additional specimen examined, excluded from type material: CANADA: BRITISH COLUMBIA: Glacier National Park, Loop Trail, 1140 m, 51.254°N, 117.538°W, 1 ♀, 16 Jul 2010, malaise trap, specimen #10BBCLP-2914 [DNA barcoded] (BIO).

(Fig. 32). Epimartyria bimaculella is known from northwestern Washington and southern British Columbia. Most British Columbia records are from the southwesternmost corner in the periphery of the Vancouver area, reflecting a more intense collecting effort in that region. One record from the Rocky Mountains of Glacier National Park, BC suggests a significantly broader distribution.

The species nameis derived from the Latin bi; (two, double) and maculella (little spot) in reference to the two, small, pale yellowish spots present on the forewings.

Adults of Epimartyria pardella most resemble those of Epimartyria bimaculella in possessing dark fuscous forewings marked by pale golden spots. Four spots are present on the forewing of pardella with two of these located across the distal third of the wing. In contrast, a total of two yellowish spots occur in bimaculella, with only a single large costal spot present beyond the middle of the forewing. In the male genitalia, the caudal lobes of sternum X (uncus) are more simple than those of the other members of Epimartyria in consisting of more shallow, rounded lobes compared to being curved and more slender in the males of auricrinella and bimaculella.

(Figs 3, 8). Head: Vestiture light orange brown. Antenna with vestiture of scape and pedicel concolorous with head; scales of flagellum pale golden yellow. Labial palpus pale brown to cream.

Thorax: Dark fuscous with coppery to purplish luster. Tegula concolorous with head. Forewing dark fuscous with coppery to purplish luster dorsally, marked with four, pale yellowish spots; the largest, irregularly rectangular and slightly oblique spot extends from costa approximately halfway across the distal third of wing; a smaller, more oval spot opposite costal spot on dorsal margin; an oblique basal spot arising midway along dorsal margin and extending halfway across wing to base of radial vein; a fourth, smallest spot at base of wing; forewing less iridescent ventrally; fringe pale yellow along termen, more gray along dorsal margin. Forewing length: 5.0–5.5 mm. Hindwing mostly gray, becoming darker and slightly iridescent toward apex; fringe gray; fringe light to dark gray. Legs medium to dark brown dorsally, paler brown ventrally and over tarsomeres; epiphysis present, ~ 1/3 the length of foretibia and arising slightly beyond its midlength.

Abdomen: Piliform scales light to dark brown.

Male genitalia (Figs 88–92): Tergum X with more slender caudal lobes. Caudal apex of sternum X (gnathos) not deeply divided, with short, triangular caudal lobes and without accessory lateral lobes. Valva short, ventral length less than 1/3 the maximum midventral length of segment IX; apex rounded and bearing a short, stout subapical spine; mesal surface smooth, without median process; elongate basal process nearly equal to length of valva. Dorsal branch of phallus more depressed, with subapical margins bearing short, paired spines, gonopore with less thickened radial folds than in previous two species.

Female genitalia: (Figs 93–96): As described for genus. Caudal end of genital sclerite deeply furcated; length of furcations exceeding length of short, undivided base.

Egg. White; dimensions 0.44 × 0.44 mm. Tuskes and Smith (1984) report the ova are flattened, circular and smooth when first deposited but soon become spherical and covered with numerous minute projections similar to those reported for Micropterix calthella (L.) by Lorenz (1961). The eggs were observed to hatch in 21 days at 22°C.

Larva. Not examined. The following description has been summarized from Tuskes and Smith (1984): Body length 4.3 to 4.6 mm; width 1.4 mm; height 1.2 mm. The body tapering at both ends with highest and broadest point at abdominal segment 4. Dorsal and lateral surface brown to dark brown, ventral surface light brown.

Head. Length 0.5 mm, diameter 0.27 mm. Brown. Antennae prominent, 3-segmented and situated on small tubercles located on dorsal lateral portion of head. Stemmata with 5 facets and located near base of antenna. Labrum simple with a pair of 3-segmented palpi. Mandibles simple and dark brown. Head diameter of first and second instar larvae 0.11 and 0.22 mm, respectively.

Thorax: Prothorax distinctly narrower than mesothorax. Prothoracic shield with 10 peg-like setae, 8 on the anterior and lateral border and 2 dorsally. Mesothorax with 8 setae, 6 on dorsal and lateral anterior portion of gray brown pigmented area, and 2 just ventral to this pigmented area. Setae of metathoracic segment similar to those of mesothorax except subdorsal (D2) seta is greatly reduced in size. All thoracic segments have additional small micro-seta just dorsal to each true leg. Thoracic legs brown, with 3 segments (excluding coxa) and simple claw.

Abdomen: Abdominal segments Al to A8 (and T2 and T3) with serrated knobs which form a dorsal and lateral ridge; areas between ridges concave. The mid-dorsal area concave with a small dark depression present on posterior of segments T2 to A8. Segments Al to A8 each with one dorsal seta (length 0.18 mm) atop dorsal ridge. Segments Al to A8 with reduced, almost microscopic subdorsal (D2) seta (length 0.04 mm) and prominent lateral seta (length 0.12 mm) on lateral ridge. Dorsal. subdorsal and lateral setae occur in brownish pigmented area which has rough and wrinkled appearance. Dorsal and lateral intersegmental area constricted and may contain series of 8 to 20 microscopic dots. Cuticle ventral to lateral ridge smooth and light brown. Series of brown dots form pattern around protuberance that usually support a small seta. Conical ventral prolegs occur on segments Al to A8, with a small, sclerotized protuberance present on ventral surface of each. Segments A9 and Al0 fused and with enlarged simple sucker. Spiracles posterior and ventral to lateral setae.

Hepaticophyta: Conocephalaceae: Conocephalum conicum (L.) Dumort.; Pelliaceae: Pellia species, with the latter host preferred from rearings (Tuskes and Smith 1984).

Not examined. Exarate, decticous; white to light brown.

Not examined. Brown, oval in general form, measuring 5.5 × 4.5 mm; primarily of silk with small fragments of vegetation attached.

(Fig. 8). Tuskes and Smith (1984) observed the eggs of Epimartyria pardella to be deposited in June on the underside of liverwort thalli singly or in small clusters of up to five eggs. They are white, measuring ~ 0.40 × 0.44 mm, and are flattened, circular, and smooth when first deposited but become spherical within a short time and covered with a series of small projections. First instar larvae ~ 0.75 mm long were reported to emerge in about 21 days (at 22°C). The larvae are rather inactive (in captivity) and are usually found on the underside of the thalli during the day. When disturbed or inactive the head may be withdrawn into the thorax. Pupation occurs within a thin walled, tightly woven brown cocoon close to the ground and attached to vegetation with strands of coarse silk.

Adults begin to emerge in late May with the flight season ranging from late May to mid- July and peak flight activity in June at the Prairie Creek State Redwood Park locality. Tuskes and Smith reported the adults to be relatively inactive, remaining motionless for hours and seldom travelling more than 30 cm. They are known to be diurnal and most active between 0900 and 1930 h. Adult feeding by Epimartyria pardella has not been reported, but the adults were observed drinking water by lowering their heads to the moisture. Moths can die in less than two days if deprived of moisture but may survive in captivity from 9 to 18 days when provided with water.

Tuskes and Smith concluded that Epimartyria pardella possessed a two year life cycle similar to that proposed for Epimartyria auricrinella (Davis 1987). In captivity eggs deposited in June 1981 became adults in June 1983. In the field they frequently collected second instar larvae during the adult flight period.

♂ (present designation), “OREGON: Klamath Co: nr. Redwood Creek, Coast: 26 June 1872, Wlsm. 90591; B.M. Genitalia Slide No. 25352; Epimartyria (= Micropteryx Wlsm.) pardella Wlsm. PARATYPE; Lectotype ♂, Epimartyria pardella Wlsm. (BMNH).” The lectotype has been selected from a series of five syntypes collected “on the borders of the forest of “redwood” (Taxodium sempervirens) near the coast, in southern Oregon, at the beginning of June 1872” (Walsingham 1880).

UNITED STATES: CALIFORNIA: Humbolt Co: Arcata: 1 ♂, 11 Jun 1969, slide USNM 16613, wing slide USNM 18441 (USNM). Fern Canyon: 1 ♀, 18 Jun 1977, N. J. Smith, slide DRD 4529 (UCB). Fern Canyon, Prairie Creek State Park: 3 ♂, 19 Jun 1981, Ann & Paul Tuskes; 1 ♂, 20 Jun 1981, P. Tuskes, 3 ♂, Ann & Paul Tuskes (USNM). Kneeland: 69 Prairie Lane: 1 ♂, 17 Jun 2001, 2 ♂, 18 Jun 2001, 1 ♂, 29 Jun 2001, 1 ♂, 30 Jun 2001, R. S. Wielgus, diurnal flight in wet meadow below house, slide USNM 34278 (USNM); 1 ♀, 22 Jun 2003, R. S. Wielgus, slides DRD 4528, 4529 (UCB). Trinity Co: Forest Glen: 2 ♀, 25 May 1973, J. Doyen (UCB). OREGON: Klamath Co: nr. Redwood Creek, Coast: 1 ♂ (lectotype), 26 June 1872, Wlsm. 90591; B.M. Genitalia Slide No. 25352, (BMNH). Multnomah Co: Benson State Park, Multnomah Falls: 2 ♂, C. V. Piper, 1 ♂, wing slide USNM 91788 (USNM).

(Fig. 32). Epimartyria pardella is known from northwestern California and northern Oregon. California localities and the type locality in Oregon are near the coast in redwood forests. The most northern Oregon locality occurs in the Columbia River valley.

Information included in this report on the immature stages and life history of Epimartyria pardella has been quoted or summarized from the thorough study of this species by Tuskes and Smith (1984) at the Prairie Creek State Redwood Park, California. In addition to possible color differences, two major morphological differences noted in their description of the larva of Epimartyria pardella from that observed for Epimartyria auricrinella include: (1) 10 versus 11 (in auricrinella) peg-like setae on each side of the prothorax, and (2) D2 of metathorax and abdominal segments 1–8 greatly reduced in pardella (as reported also in Austromartyria, Gibbs 2010, and for the abdomen in Agrionympha, Gibbs and Kristensen 2011). Although examples of the larva, pupa, and cocoon of this species were reportedly deposited in the collections of the California Academy of Sciences, San Francisco, CA by Tuskes and Smith (1984), attempts to locate and borrow this material for study were unsuccessful. The skeletomuscular anatomy of the male genitalia of Epimartyria pardella was reviewed by Kristensen (1984b).