The genus Euceratia is reviewed by the re-descriptions of two known species, E. castella and E. securella, and the description of a new species from California, E. intermedia n. sp. Euceratia is compared to other genera of Ypsolophidae. Diagnostic characters are provided for the three species of Euceratia and their distribution in the Nearctic Region is discussed. Imagoes and genitalia of all the species of Euceratia are illustrated.
The Nearctic genus Euceratia was described by Walsingham (1881) for two spiecies, E. castella and E. securella, from California, USA. It was originally assigned to Tineidae, in which most primitive micro-moths were included by authors. Meyrick (1914) and Fletcher (1929) transferred the genus to Plutellidae, based on superficial features. Kyrki (1984) revised the systematic prosition of Euceratia and found that it is associated with Ypsolopha. Kyrki (1990) finally assigned the genus to Ypsolophidae.
The two Euceratia described by Walsingham (1881) are still the only known species of the genus. Both of these share the characteristic labial palpi that Walsingham (1881) highlighted with relatively lengthy description. Heppner (1984) assigned Calantica argentea Busck, 1913 to Euceratia. This Mexican species differs from Euceratia in the characters of labial palpus and genitalia. Friese (1960) assigned the species to Niphonympha (Yponomeutidae). It is known that the larvae of Euceratia are leaf tiers on snowberries (Symphoricarpos) and honeysuckles (Lonicera) of Caprifoliaceae (Powell and Opler 2009).
The aims of this paper are to redescribe Euceratia and its two known species, E. castella and E. securella, and to describe a new congener from California. In addition, the distribution of the three species of Euceratia is discussed and compared with those of other ypsolophid genera.
Materials and Methods
Pinned specimens from five institutional collections were examined. The abbreviations of these and other relevant depositories are as follows:
ANSP: Academy of Natural Sciences of Drexel University, Philadelphia, USA;
BMNH: Natural History Museum (formerly British Museum of Natural History), London, UK;
EMEC: Essig Museum of Entomology, University of California, Berkeley, USA;
MCA: Museum of Comparative Zoology, Harvard University, Cambridge, USA;
USNM: United States National Museum of Natural History, Washington DC, USA;
YPM: Yale Peabody Museum of Natural History, New Haven, USA.
Other abbreviations used in the specimen data include:
BL: black light; Co.: county; ft.: feet; gen. no.: genitalia slide number; Is.: island; mi: mile; Mt.: mount; Mts.: mountains; MVL: mercury vapor lamp; UVL: ultraviolet light; and WL: white light.
Verbatim label data are provided only for primary types. The marks ‘/’ in the label data indicate line breaks. The genitalia slides were prepared according to Clarke (1941), except that chlorazol black was used for staining and euparal resin was used for permanent slide mounting. The wing venation slides were prepared according to Hodges (2005). Terminology follows Klots (1970) for genitalia and Wootton (1979) for wing venation. The names of host plants follow the Plant List ( http://www.theplantlist.org/).
Taxonomic Accounts
Ypsolophidae: Ypsolophinae
EUCERATIA Walsingham, 1881
Euceratia Walsingham, 1881: 310.
Type species: Euceratia castella Walsingham, 1881, by subsequent designation by Fletcher (1929).
Diagnosis. This genus is similar to Phrealcia Chrétien, 1900 in the external appearance and the female genitalia, especially the presence of a band-like, scobinate signum. Euceratia, however, differs from the latter in having the porrect labial palpi longer than the head (as long as or shorter than the head in Phrealcia).
Redescription. Head (Fig. 1) — Vestiture of vertex and frons rough with piliform scales; ocelli present [Walsingham (1881) erroneously stated that Euceratia lacks ocelli]. Antenna filiform in both sexes, annulated, naked ventrally. Labial palpus porreet, 3× as long as head, 1st segment 1/5 as long as 2nd, with piliform-scale tuft ventrally; 2nd segment as long as 3rd, with piliformscale tuft ventrally, scales of tuft longer distally; 3rd segment duster-like with piliform scales. Maxillary palpus short, 1-segmented [Walsingham s (1881) observation that maxillary palpi are absent in Euceratia is not correct].
Thorax — Patagium comprising piliform scales. Forewing subtriangular, with oblique termen; pterostigma spanning near 1/2 of costa and vein R. Forewing venation (Figs. 2 & 3) with Sc reaching margin near 2/3 of costa; R arising from near middle of radius; Rs1–3 reaching margin above apex; Rs1 arising from anterior margin of accessory cell at 4/5 length of cell; Rs1 nearly parallel to Rs2; Rs2 and Rs2 close basally, then divergent; Rs4 reaching margin below apex at the anterior 1/5 of termen; M with 3 branches; M1 and M2 slightly divergent; M, nearly equidistant from M1 and from M3; CuA1 and M3 connate basally (E. securella) or equidistant as CuA1 and CuA, (E. intermedia n. sp.); CuA, nearly straight; CuP vestigial as fold in basal 4/5; basal fork of 1A+2A near 1/4 of length. Hindwing venation (Figs. 2 & 3) with Sc+R1 reaching margin at 4/5 of costa; Rs reaching margin above apex, stalked with M1 in posterior 2/5 (E. securella) or 3/5 (E. intermedia n. sp.); M stem vestigial, as a short branch; M2 nearly parallel to M3; M3 close to CuA1 basally and then divergent; CuA1 nearly parallel to CuA2; CuP present in posterior half (E. securella) or entire (E. intermedia n. sp.); 1A+2A slightly divergent from CuP in distal 1/3, with basal fork 1/5 of length.
Abdomen — Sternum II (Fig. 4) with apodeme and venula 4× longer than transverse ridge; venula arched in anterior half; transverse ridge present at level of anterior 2/5 of venula. Pleuron VIII (Fig. 5) expanding to posteriorly, semicircular; interspace between pleural lobes slightly emarginated. Male genitalia (Figs. 15–17) with uncus trapezoidal or subtriangular; socius with small spine on apex; subscaphium present; valva subrectangular, round apically, densely setose in distal 3/4; anellus with minute thorns. Phallus with spinulate cornutal zones. Female genitalia (Figs. 18–20) with ovipositor telescopic, with two subdivisions; apophysis posterioris with Y-fork basally; ductus bursae slender; signum band-like, scobinate.
Included species
Euceratia castella Walsingham, 1881
Euceratia intermedia n. sp.
Euceratia securella Walsingham, 1881
Euceratia castella Walsingham
Figs. 6–8, 15, 18
Euceratia castella Walsingham, 1881: 310.
Diagnosis. This species is superficially similar to an yponomeutid species, Eucalantica polita (Walsingham) but differs from the latter in having the longer labial palpi and by the lack of a reddish brown dorsal patch on the forewings. Also, the tergites of Eucalantica possess spiniform setae which are absent in Euceratia.
Redescription. Head — Vertex and frons white. Antenna 1/2 as long as forewing; scape white; first three flagellomeres with white annulation; remaining flagellomeres dark brown. Labial palpus white, sparsely intermixed with dark brown-tipped, white scales.
Thorax — Patagium, tegula and mesonotum white. All legs white or yellowish white. Forewing length 6.8–10.1 mm (n = 10), white, slightly broadened to termen; apex narrowly round; a small black spot present on distal 1/2 of CuP; dark brown scales scattered in various density, depending on individuals; a small black spot at the lower angle of discal cell usually present, but sometimes absent (Fig. 7); dark yellow, intermittent postmedian line and striation in distal area sometimes present (Fig. 8); scales of fringe white with dark brown tip. Hindwing dark brownish gray, suffused with white in basal and posterior areas; marginal scales white, with pale dark brown tip; fringe white, tinged with pale gray in anterior area.
Abdomen — Terga white; sterna yellowish white. Male genitalia (Fig. 15) with uncus subtrapezoidal; socius slender, 4× longer than uncus. Tegumen wide, trapezoidal, 1/2 as long as socius. Valva subrectangular, round apically, densely setose in distal 2/3; sacculus round in basal half, sparsely setose. Vinculum nearly straight anteriorly; saccus elongate, slightly broadened in distal half, as long as socius. Phallus 2.5× longer than valva, straight, slender, slightly broadened anteriorly; two cornutal zones 1/10 as long as phallus. Female genitalia (Fig. 18) with ovipositor 1.5× longer than corpus bursae. Apophyses posteriores 2× longer than apophyses anteriores. Ductus bursae narrow, bowl-shaped near ostium bursae, 3× longer than corpus bursae. Corpus bursae ovate; signum on posterior 3/5 of corpus bursae.
Types. Lectotype [designated by Sohn (2015a)] — male, “Type” [round label with red borders], “Russian R/ Sonoma Co./ CALIFORNIA/ 19.V 1871/ Wlsm”, “Walsingham/ Collection./ 1910-427”, “NG Euceratia/ castella Wlsm/ TYPE” [hand-written], “Euceratia/ castella, Wlsm./ P.Z.S.Lond. p. 311. Tf. 35′13 1881/TYPE ♂ figd. & descr.” [hand-written on label with black borders], detached abdomen in a gelatin capsule attached with the specimen, BMNH.
Paralectotypes — USA: CALIFORNIA: San Francisco: 1♂, 1♀, [no date & collector], MCZ. Walsingham (1881) described Euceratia castella, based on “several specimens near San Francisco and on Mount Shasta.” Three specimens listed here were the only representatives of Walsingham's syntypes of E. castella which could be traced.
Materials examined. CANADA: BRITISH COLUMBIA: Vancouver Is.: Duncan, 2♀, [no date] (AW Hanham), USNM; Quamichan District [near Duncan]: 1♂, 15–31 May “192” (AW Hanham), USNM; Victoria: 1♂, 13 June 1901 (AW Hanham), USNM; 2♂, 4 June 1910 (AJ Croker), USNM; 1♂, 12 June 1920 (EH Blackmore), USNM; 1♂, 1♀, 5 July 1920, USNM; 2♂, 30 May 1921 (WR Carter), USNM; 4♂, 6 June 1922, USNM; 1♂, 1♀, 6 July 1923 (JF Clarke), USNM; Wellington: 1♂, 1♀, [no date] (GW Taylor), USNM. USA: CALIFORNIA: Alameda Co.: Berkeley: 3♂, 6 May 1921 (EO Essig), USNM; 1♂, 26 May 1921, USNM; 1♂, 18 May 1931 (D Meadows), USNM; Hayward: 1♂, 17 May 1966 (AJ Slater), EMEC; Northeast Oakland: Piedmont Pines: 1♀, 19 June 1962 (PD Hurd), EMEC; Northeast Oakland: Berkeley Hills: 1400 ft.: 1♀, 10 May 1962 (J Powell), EMEC; Tilden Park: 2♀, no date (DL Wagner), rearing no. LII-IV-79, reared from Symphoricarpos, emerged on 1 June 1979 & 8 May 1979, EMEC; 1♀, 17 April 1969 (PA Opler), rearing no. 69D34, reared from Symphoricarpos (?), emerged on 14 May 1969, EMEC; [no specific locality]: 1♂, 1♀, June (CV Riley), reared from Symphoricarpos mollis, USNM; 20♂, 19♀, 11–20 May 1908 (GR Pilate), USNM; 3♂, 2♀, 22 May 1908, ANSP; 1♀, 4 June 1908, USNM. Alpine Co.: Carson River: 8 mi south of Markleeville: 1♂, 22 June 1962 (J Powell), EMEC. Calaveras Co.: Mokelumne Hill: 1500 ft.: 1♀, June [no year], USNM. Contra Costa Co.: Lafayette: 1♂, 2 June 1960 (J Powell), at BL, EMEC; Orinda: 1♂, 4 June 1957 (SF Cook Jr), at light; 1♀, 9 June 1957, EMEC. El Dorado Co.: Blodgett Forest: 14 mi east of Georgetown: 1♀, 21 June 1982 (JA DeBenedictis), at UVL, EMEC; Lake Tahoe: Deer Park Springs: 2♂, 24–30 June [no year], USNM; 2♂, 2♀, 1–7 July [no year], USNM; 3♂, 1♀, 8–15 July [no year], USNM; Pilot Creek: 1♀, 8 July 1967 (WJ Turner), EMEC. Glenn Co.: 5 mi north of Black Butte: 6200 ft.: 1♂, 19 June 1956, EMEC. Inyo Co.: [no specific locality]: 1♀, 1–15 June 1922 (OC Poling), USNM. Kern Co.: Tehachapi Mt. Park: 5600 ft.: 1♀, 18 June 1989 (J Powell), at BL, EMEC. Lake Co.: Summit Elk Mt.: 1♂, 2 July 1949 (CC & UN Lankar), USNM. Los Angeles Co.: “La Puerta Valley”, 1♀, [no date], USNM; Tujunga: 1♂, 14 May 1940 (C Henne), YPM. Marin Co.: Phoenix Lake, 2♂, 7–8 April 1927 (HH Kelfer), reared from Symphoricarpos raceniosus. ANSP. Mendocino Co.: 4 mi south of Hopland: 1♀, 9 May 1961 (J Powell), EMEC. Modoc Co.: Warner Mts.: 3 mi east of Davis Creek: 5500 ft.: 2♂, 8♀, 8–15 July 1922 (AW Lindsey), USNM. Mono Co.: 4 mi east of Monitor Pass: 1♂, 30 June 1962 (J Powell), EMEC. Monterey Co.: Carmel Valley: 1♀, 30 May 1997 (J Kruse & J Powell), at BL, EMEC. Napa Co.: Soda Creek: 2♂, 1♀, 22 March 1933 (Keifer), USNM; 3♂, 3♀, 27 April 1934 (Keifer), USNM. Nevada Co.: Donner Summit: 7200 ft.: 1♂, 11 July 1979 EMEC. Placer Co.: Colfax: 1♀, April [no year] (AH Vachell), USNM; Ward Creek: 2 mi south of Tahoe City: 2♂, 3♀, 26 July 1969 (N Westerland), USNM. Plumas Co.: 1♂, 3♀, 16– 23 June [no year], USNM. San Bernardino Co.: San Bernardino Mts.: 3000 ft.: 1♀, 9 May 1931 (CM Dammers), USNM; 1♂, 16 June 1933, USNM. San Luis Obispo Co.: 3 mi west of Paso Robles: 1♂, 28 April 1968 (D Veils & J Powell), at light, EMEC. San Mateo Co.: San Bruno Mt.: 1♂, 18 March 1983 (JA DeBenedictis), rearing no. JADeB No. 83077-C, reared from Symphoricarpos albus, emerged on 6 April 1983, EMEC; 2♂, 3♀, 1 March 1985, rearing no. JADeB No. 85C2, reared from Symphoricarpos albus, emerged on 1–2 April 1985, gen. no. EMEC-JCS-031 (♀), EMEC; 1♀, 22 March 1985 (JB Whitfield & JA DeBenedictis), rearing no. JEW No. 85C22, reared from Symphoricarpos, emerged on 5 April 1985, EMEC. Santa Cruz Co.: Big Basin [erroneously given as San Francisco Co.]: 3♀, 18 June 1971 (E Jäckh), USNM. Sierra Co.: 1 mi east of Bassetts: 1♂, 29 June 2002 (J Powell). Siskiyou Co.: 5 mi east of McCloud: 1♂, 7 July 1957 (J Powell), EMEC; 13, 14 July 1962, EMEC; Mt. Shasta City: 23, 24–26 June 1958 (J Powell), EMEC; Shasta Retreat: 1♂, 4♀, 8–15 June [no year], USNM; 3♀, 16–23 June [no year], USNM. Sonoma Co.: 0.5 mi east of Forestville: 1♂, 26 May 1966 (J Powell), EMEC; [no specific locality]: 6♂, 6♀, 10–25 May [no year] (AH Wachell), USNM. Tuolumne Co.: Kennedy Meadows: 1♂, 3 July 1959 (PA Opler), EMEC; Big Oak Flat: 1♀, 12 June 1961 (J Powell), EMEC. Tulare Co.: Mineral King: 1♂, 24–30 June [no year], USNM; 2♀, 8–15 July [no year], USNM; 2♂, 2♀, 16–23 July [no year], USNM; 2♂, 24–31 July [no year], USNM; 1♀, 1–7 August [no year], USNM; Monarch Meadows: 8000 ft.: 1♂, 17 July 1917, ANSP;1♂, 4♀, 8–14 July [no year], USNM; 1♂, 1♀, 16–23 July [no year], USNM; 1♀, 8–15 Aug. [no year], USNM. IDAHO: Boise Co.: Cold Spring Creek: 5 mi west of Idaho City: 1♂, 9 June 1976 (JFG Clarke), USNM. Bonner Co.: Priest River Experimental Forest: 2500 ft.: 2♂, 19–20 June 1979 (DC Ferguson), USNM. Elmore Co.: Sawtooth National Forest Station: 4900 ft.: 1♀, 13 July 1969, USNM. Latah Co.: Moscow Mt.: 1♂, [no date] (CV Piper), USNM. OREGON: Baker Co.: Big Lookout Mt.: 1♂, 4♀, 13 July 1974 (JH Baker), USNM; Spring Creek: 1♀, 7 August 1962 (JH Baker), USNM; 1♀, 7 July 1970, USNM; 1♂, 11 July 1972, USNM. Grant Co.: Ritter: 4200 ft.: 2♂, 3♀, 17–20 July 1962, USNM. Hood River Co.: 7 mi west of Hood, 1♀, 14 June 1963 (RW), at light, gen. no. EMEC-JCS-032, EMEC. Jackson Co.: Kane Creek: 5 air mi south of Gold Hill: 1♂, 22 June 2000 (J Powell & D Rubinoff), at BL, gen. no. EMEC-JCS-035, EMEC; Mt. Ashland: 6650 ft., 2♂, 1♀, 27 July 1990 (R Robbins), USNM. Josephine Co.: 2 mi east of Merlin: 7♂, 2♀, 27 May 1970 (JFG Clarke), USNM. Linn Co.: 8 mi west of Mill City, 2♂, 1♀, July 1963 (SG Jewett), USNM. Marion Co.: Salem: 2♂, 2♀, 2 June 1961 (K Goeden), USNM. Wasco Co.: The Dalles: 2♂, 2♀, 3 June 1970 (JFG Clarke), USNM. Washington Co.: Forest Grove: 1♀, May 1923 (SE Keen), reared from Symphoricarpos sp., USNM; 5♂, 6♀, 1–11 May 1925 (SE Keen), reared from Symphoricarpos albus, USNM; 6♂, 3♀, 24–30 April 1925 (SE Keen), reared from Symphoricarpos albus, USNM; Portland: 3♀, [no date] (CV Piper), USNM; 1♀, 12 June 1931 (JF Clarke), USNM. Yamhill Co.: Williamson State Park: 1♂, 9 June 1962 (CWO Brien), EMEC. WASHINGTON: Clallam Co.: Tumwater Creek: 4 mi southwest of Port Angeles: 1♂, 17 July 2002 (J Powell), at BL, gen. no. EMEC-JCS-037, EMEC. Ferry Co.: Bodie Mt.: 5200 ft.: 3♂, 24 July 1962 (JFG Clarke), USNM; Sherman Pass: 5600 ft.: 1♀, 23 July 1962 (JFG Clarke), USNM. Grays Harbor Co.: Hoquiam: 1♀, 2 June 1904 (Burke), USNM. Klickitat Co.: Lyle: 1♂, 9 June 1931 (JF Clarke), USNM. Mason Co.: Shelton: 1♂, 20 June 1931 (JF Clarke), USNM. San Juan Co.: Deer Harbor: Orcas Island: 1♂, 1♀, 14 July 2002 (J Powell), gen. no. EMEC-JCS-036 (♀), EMEC. Stevens Co.: Kettle Falls: 2600 ft.: 4♂, 3♀, 20 July 1962 (JFG Clarke), USNM. Thurston Co.: Rochester: 1♀, 13 June 1929 (WW Baker), USNM; 1♂, 26 June 1929, USNM. Whatcom Co.: Bellingham: 1♀, 11 June 1922 (JF Clarke), USNM; 1♂, 25 June 1950, USNM; Chuckanut Bay: 1♀, 23 June 1955 (JFG Clarke), USNM. Whitman Co.: Pullman: 1♀, 10 July 1891 (CV Piper), USNM; 1♂, 8 June 1930 (JF Clarke), USNM; 1♀, 6 June 1933, USNM.
Distribution. Canada (British Columbia) and USA (California, Idaho, Oregon, Washington).
Host plants. Caprifoliaceae — Lonicera spp., Symphoricarpos albus (L.) S. F. Blake, and S. mollis Nutt. (Powell and Opler 2009: this study). Miller and Hammond (2003) listed Quercus garryana Douglas ex Hook. as a host plant of E. castella. This record seems questionable. Boulton (2004) extensively surveyed the caterpillars feeding on Garry oak (Q. garryana) and common snowberry (Symphoricarpos albus) from British Columbia and found that the larvae of E. castella were collected exclusively from the latter plant.
Remarks. This species shows some variations in the forewing patterns. These variations appear not to be correlated with gender, habitat, or flight time. The larvae are found in March and April and the adult moths are flying from late April to August (Powell and Opler 2009). It is the most widely distributed species of Euceratia (Fig. 21). The collecting locality of the lectotype specimen was given as “Russuian R[iver], Sonoma Co.” which, however, was not one of the type series Walsingham reported, i.e. “near San Francisco and Mt. Shasta.” There is no evidence that the Russian River specimen was recognized by Walsingham as one of the “Types.” It can be argued that the “near San Francisco” record (Walsingham 1881) is equivalent to the Russian River of Sonoma County given on the type label.
Figs. 1–5.
Generic characters of Euceratia. 1. Head of E. securella, lateral view. 2. Wing venation of E. securella. 3. Wing venation of E. intermedia. 4. Abdominal segment I and II of E. securella, tergite I in left, sternite II in right. 5. Abdominal segment VIII of E. securella, tergite and pleural lobes in left, sternite in right.
Figs. 6–14.
Adults of Euceratia. Figs. 6–8. E. castella. 6. male, holotype, California, Sonoma Co. 7. male, California, Mono Co. 8. male, California, San Mateo Co. Figs. 9–10. E. intermedia. 9. male, paratype, California, Del Puerto Canyon. 10. female, paratype, California, San Luis Obispo Co. Figs. 11–14. E. securella. 11. female, California, Lake Co. 12. male, California, San Diego Co. 13. female, California, Orange Co. 14. female, California, Humboldt Co.
Euceratia intermedia
Sohn et Powell,
new species
Figs. 9, 10, 16, 19
Diagnosis. This species is similar to its two congeners, E. castella and E. securella, but differs from them as follows: (a) the posterior half of forewing is densely irrorated with brown scales in E. intermedia and E. securella but sparsely intermixed with dark brown dots or yellowish brown mottling in E. castella; (b) dark brown markings are present on the forewing of E. securella but absent on those of E. intermedia and E. castella; (c) in the male genitalia, the saccular angulation is less profound in E. intermedia and E. securella than in E. castella; (d) the apex of the valva is more narrowly round in E. intermedia than in E. castella and E. securella; (e) the phallus is straight in E. castella but slightly curved in E. intermedia and E. securella; and (f) in the female genitalia, the signum of E. intermedia is longer than that of E. securella, but shorter than that of E. castella.
Description. Head — Vertex and frons white. Antenna 1/2 as long as forewing: scape white: first three flagellomeres with white annulation: remaining flagellomeres dark brown. Labial palpus with 1st segment white; 2nd segment brownish white laterally, white mesally, a few scales with dark brown tips; scales of 3rd segment white, with dark brown tips.
Thorax — Patagium, tegula and mesonotum white. Fore- and midleg with coxa brownish white; femur and tibia dark brownish gray on exterior surface, white on interior surface; tarsomere dark brownish gray with a white ring distally. Hindleg brownish white; tarsomeres with white ring distally. Forewing length 9.2–11.1 mm (n = 10), white, tinged with pale brownish gray in distal area; apex slightly acuminate; dark brown scales scattered sparsely in basal area, more densely to termen; a small black spot present on distal 1/2 of CuP; scales of fringe brownish gray with brownish black tip. Hindwing brownish gray, paler to base; fringe pale brownish gray.
Abdomen — Terga gray; sterna pale gray. Male genitalia (Fig. 16) with uncus elliptical, extending posteriorly, bifid apically; socius slightly broadened to base, slightly curved in distal 1/5. Tegumen trapezoidal, as long as uncus. Valva rectangular, slightly broadened distally, angulate at distal end of costa, narrowly round apically, densely setose in entire area; sacculus convex. Vinculum slightly convex anteromarginally, angulate laterally; saccus broadened distally. Phallus 2.3× longer than valva, slightly curved at basal 1/3, broadened in basal 1/5; cornutal zone 1/10 as long as phallus. Female genitalia (Fig. 19) with ovipositor 2× longer than corpus bursae. Apophyses posteriores 2× longer than apophyses anteriores. Ductus bursae 3.3× longer than corpus bursae, slender, slightly broadened to corpus bursae, granulate in posterior 1/8; antrum near ostium bursae, thimble-shaped. Corpus bursae globular; signum on posterior half of corpus bursae.
Type. Holotype — male, “HOLOTYPE/ Euceratia intermedia/ SOHN & POWELL ” [red label circumscribed with black lines], “[Stanislaus Co.] Del Puerto C[an]y[o]n./ 20 mi. W. Patterson/ Calif[ornia] IV.30.63”, “J. Powell/ Collector”, detached abdomen in a gelatin capsule attached with the specimen, EMEC. Paratypes (15♂, 3♀) — USA: CALIFORNIA: Kern Co.: Havilah: 1♂, 8–15 June [no year], USNM. Marin Co.: Alpine Lake: 1♂, 28 May 1960 (J Powell), EMEC. Placer Co.: Cisco: 1♂, 1 June 1905 (AH Vachell), USNM; Colfax: 2♂, 1♀, May [no year] (VH Vachell), USNM. San Benito Co.: 2 mi northeast of New Idria, 1♂, 24 April 1964 (J Powell), wing slide no. SJC-W003, EMEC. San Luis Obispo Co.: 2 mi northeast from Pozo, La Panza Camp, 1♂, 29 April 1962 (J Powell), EMEC; Pozo: 1♂, 1♀, 27 April 1962 (J Powell), gen. no. EMEC-JCS-034 (♀), EMEC. Santa Barbara Co.: 2 mi south of Buellton: 1♂, 12 May 1965 (J Powell), EMEC; Los Prietos: 1♂, 23 April 1966 (AJ Slater), gen. no. EMEC-JCS-033, EMEC. Stanislaus Co.: Del Puerto Canyon: 20 mi west of Patterson: 1♂, same as holotype, EMEC; Del Puerto Creek: 900–1200 ft.: 1♂, 14 April 1973 (M Chinn), “Insect Survey Specimen #” 154079, EMEC; 1♂, 13 April 1980 (K Standow), “Insect Survey Specimen #” 274968, EMEC. Tulare Co.: 5 mi northeast of Springville, 1♂, 14 May 1963 (CA Toschi), EMEC; Camp KEEP near Paso Park: 1440 m: 1♂, 23–24 June 1995 (J Powell), at BL, EMEC. Tuolumne Co. (?): Yosemite: 1♂, 5 June 1924 (HG Dyar), USNM.
Distribution. USA (California).
Host plants. Unknown.
Etymology. The species name is derived from two Latin terms ‘inter’ and ‘medius’ meaning “between” and “middle” respectively and refers to the overall appearance of this new species, which is intermediate between Euceratia castella and E. securella.
Figs. 15–17.
Male genitalia of Euceratia. 15. E. castella. 16. E. intermedia, paratype. 17. E. securelia. Scale bars = 0.5 mm.
Figs. 18–20.
Female genitalia of Euceratia. 18. E. castella. 19. E. intermedia, paratype. 20. E. securella. Scale bars = 1 mm.
Euceratia securella Walsingham, 1881
Figs. 11–14, 17, 20
Euceratia securella Walsingham, 1881: 311.
Diagnosis. This species is similar to a European species, Phrealcia eximiella (Rebel), but differs from the latter in having longer labial palpi, a more slender phallus, and a narrower ductus bursae.
Redescription. Head — Vertex brownish white, sparsely intermixed with pale brown scales; frons pale brown. Antenna 2/3 as long as forewing; scape and first three flagellomeres brownish white, occasionally peppered with dark brown; remaining flagellomeres dark brown, with brownish white annulation. Labial palpus with 1st segment white, tinged with pale brown dorsally; scales of 2nd and 3rd segments pale brown, darkened to tips laterally, brownish white, with dark brown tips mesally.
Thorax — Patagium brownish white; tegula brownish white in inner half, pale or dark brown in outer half; mesonotum brownish white, occasionally peppered with dark brown, tinged with dark brown medially. Foreleg with coxa dark brownish gray on exterior surface, pale brownish gray on interior surface; femur to tarsus dark brownish gray; each tarsomere with a white ring terminally. Midleg with coxa brown on exterior surface, brownish white on interior surface; femur to tarsus brownish gray dorsally, pale brown ventrally; each tarsomere with a white ring terminally. Hindleg with coxa to femur brownish gray; femur pale brownish gray, sparsely intermixed with gray scales; each tarsomere brownish gray, with a white ring terminally. Forewing length 9.1–11.6 mm (n = 10), white to pale brownish gray; an oblique black bar connecting basal 1/3 of dorsum with terminal 1/3 of lower margin of discal cell present in variable width, depending on individual; a small black spot present at the lower end of discal cell; dark brown costal strigulae sometimes present (Figs. 13, 14); dark brown tornal patch variable in size, depending on individual; fringes dark brownish gray, irregularly intermittent with white. Hindwings brownish white; marginal shade brownish gray; fringe pale brownish gray, tinged with yellowish gray in inner margin.
Abdomen — Terga brownish gray; sterna brownish white. Male genitalia (Fig. 17) with uncus rectangular, convex posteriorly, bifid apically; socius broadened basally, falcate apically. Tegumen rectangular, as long as uncus. Valva slightly curved to costa, slightly broadened to apex, densely setose in distal 4/5; sacculus broadly round in basal 1/2. Vinculum nearly straight anteromarginally, nearly angulate laterally; saccus slightly broadened in distal 1/3. Phallus 2.3× longer than valva, slightly curved medially, broadened in anterior 1/7; cornutal zone 1/10 as long as phallus. Female genitalia (Fig. 20) with ovipositor 1.2× as long as corpus bursae. Apophyses posteriores 2.2× longer than apophyses anteriores. Ductus bursae slender, funnel-like near ostium bursae, granulate between posterior 1/9 and 2/9, 2.3× longer than corpus bursae; antrum at posterior 1/17 of ductus bursae, short, cylindrical. Corpus bursae elliptical; signum 1/4 as long as corpus bursae.
Types. Lectotype [designated by Sohn (2015a)] — male, “Type” [round label with red borders], “head of Dry Creek/ Mendocino Co./ CALIFORNIA/ 24.V. 1871/Wlsm”, “Walsingham/Collection./ 1910-427”, “Euceratia/ securella Wlsm/TYPE Cala” [hand-written], “- If — fwing/ TYPE COLL./ Cab. 12 Dr. 5”, “Euceratia/ securella, Wlsm./ PZS. Lond. p. 311. pt 35′14 1881./ TYPE ♂ figd & descri.” [handwritten on label with black borders], left forewing missing, BMNH. Paralectotypes — USA: CALIFORNIA: Sonoma: 1♂, [no date & collector], MCZ. Walsingham (1881) described E. securella, based on 17 specimens collected from Sonoma Co., California. We were able to locate only two of those.
Materials examined. CANADA: BRITISH COLUMBIA: Victoria: 1♂, 13 June 1903, USNM; 1♂, 1♀, 28 June 1921 (WR Carter), USNM; 1♀, 21 July 1921, USNM; 1♂, 24 June 1922, USNM; 1♀, 7 July 1922, USNM; 5♂, 3♀, 16–19 June 1923, USNM; 4♀, 12 July 1923, USNM. USA: CALIFORNIA: Alameda Co.: Del Valle Lake: 1♀, 2 February 1984 (J Powell), rearing no. 84820, reared from Symphoricarpos albus, emerged on 6 March 1984, EMEC; Northeast Oakland: Berkeley Hills: 1400 ft.: 1♂, 17 April 1962 (J Powell), EMEC; 1♂, 10 May 1962, EMEC; Tilden Park: 1♂, 28 May 1979 (DL Wagner), EMEC. [no specific locality]: 1♂, 28 March 1914 (S Jose), USNM. Contra Costa Co.: Lafayette, 1♀, 20 April 1969 (PA Opler), reared from Symphoricarpos albus, EMEC; 2 mi southwest of Moraga, 2♂, 8 June 1979 (DeBenedictis & Wagner), EMEC; Orinda: 1♀, 20 May 1957 (SF Cook, Jr), EMEC; 1♀, 4 June 1957, EMEC. Humboldt Co.: Areata: 1♂, 30 June 1969 (J Powell), at light, EMEC; 1♂, 15 July 1969, EMEC; Blair's Ranch: Redwood Creek: 1♂, 10 June [no year] (HS Barber), USNM; Kneeland: 69 Prairie Lane: 1800 ft.: 1♀, 14 July 2001 (RS Wielgus), at UVL and MVL, EMEC; 1♀, 24 July 2001, EMEC; 1♀, 12 July 2002, gen. no. EMEC-JCS-056, EMEC; 1♀, 29 July 2002, EMEC; 1♂, 10 July 2005 (RS Wielgus), at UVL & WL, EMEC; Scotia: 1♀, 20 May [no year] (HS Barber), USNM. Lake Co.: Adams Springs: 1♀, 31 May 1982 (JA DeBenedictis), EMEC; 1 mi northeast of Cobb: 1♀, 5 June 1980 (J Powell), EMEC. Marin Co.: 2 mi southeast of Inverness, Inverness Ridge, 1♀, 15 May 1970 (JA Powell), at light, EMEC. Mendocino Co.: 1 mi north of Piercy: 2♂, 4♀, 20–23 May 1976 (Dietz, Chemsak & Powell), EMEC. Monterey Co.: Carmel Valley: 1♀, 30 May 1997 (J Kruse & J Powell), at BL, EMEC; near Jamesburg: 1♂, 3 May 1958 (J Powell), at light, EMEC. Napa Co.: Diamond Mt.: 3 air km south of Calistoga: 520 m: 1♂, 1♂, 21–23 May 1993 (J Powell), at light, gen. no. EMEC-JCS-058 (♂), EMEC. Orange Co.: Silverado: 1♂, 30 March 1968 (J Powell), EMEC; Silverado Canyon: Santa Ana Mts.: 1650 ft.: 1♂, 12 May 1979 (GA Marsh), at WL, EMEC; 2♀, 26–28 May 1979, EMEC. San Diego Co.: 3 mi east of Julian: 4♂, 14–15 June 1998 (N Bloomfield), at BL, USNM; MCAS Miramar: San Clemente Canyon: 2♂, 5 April 1998 (N Bloomfield), wing slide no. SJC-W004, EMEC & USNM; 1♂, 21 April 1998, EMEC; NAS Miramar 2: 1♀, 15 March 1997 (N Bloomfield), USNM; NAS Miramar 9: 1♀, 22 March 1997 (N Bloomfield), at BL, gen. no. USNM-115067, USNM; NAS Miramar 10: 1♂, 1♀, 15 April 1997 (N Bloomfield), USNM; San Clemente Canyon: USMC Air Station: 4♂, 2♀, 13–15 May 2011 (N Bloomfield), at BL, USNM. San Francisco Co.: [no specific locality]: 1♀, 20 July 1971 (E Jäckh), USNM. San Luis Obispo Co .: Alamo Creek, 16 air km east of Nipomo, 1♀, 8 April 2004 (J Powell), at BL, EMEC; 12 mi northeast of Pozo, La Panza Camp, 1♂, 2 May 1962 (J Powell), EMEC; 2♀, 2 June 1962, at light, gen. no. EMEC-JCS-055 & 062, EMEC; 1♂, 25 April 1968 (D Veirs), EMEC; Pozo: 1♀, 4 May 1962 (J Powell), EMEC; York Mt.: 8 mi west of Templeton: 2♂, 27 April 1968 (P Opler), EMEC. Santa Barbara Co .: Santa Cruz Island: Central Valley: 2♀, 25–28 April 1966 (J Powell), at light, EMEC; Field Station Headquarter: 1♂, 21–24 May 1984, EMEC; Islay Canyon Road: 165 m: 1♀, 1 May 2000 (J Powell), at BL, EMEC; La Cascada: 1♀, 23 April 2001 (J Powell), at BL, EMEC; 2♀, 1 May 2000, at BL, EMEC; Prisoners Harbor Creek: 50 m: 1♂, 1 May 1966 (J Powell), EMEC; 29, 5 May 2000 (J Powell & M Myers), EMEC; University of California Field Station: 76 m: 2♀, 1–7 May 2000 (J Powell & P Hart), at BL, EMEC. Santa Clara Co.: Herbert Creek: 3 mi west of New Almaden: 1♂, 1♀, 28 April 1969 (PA Opler), gen. no. EMEC-JCS-059 (♂), EMEC; 1♀, 19 April 1989 (J Powell & M Prentice), EMEC; New Almaden: 1♀, 28 September 1965 (P Opler), EMEC; [no specific locality]: 1♂, 1♀, [no date], USNM. Sonoma Co.: 1 mi southeast of Plantation, 1♂, 6 June 1979 (J Powell), EMEC; Plantation: 1♀, 30 June 1967 (P Opler), EMEC; [no specific locality]: 9♂, 4♀, 10–25 May [no year] (AH Vachell), ANSP & USNM. OREGON: Clackamas Co.: near Wilsonville: 1♂, 16 June 1982 (DL Wagner), EMEC. Jackson Co.: Kane Creek: 5 air mi south of Gold Hill: 1♀, 22 June 2000 (J Powell & D Rubinoff), at BL, EMEC. Lane Co .: near Dexter: Elijah Bristow State Park: 1♂, 1♀, 15 June 1985 (J Powell), EMEC. Yamhill Co.: Williamson State Park: 1♂, 1♀, 25 June 1975 (J Powell & P Opler), EMEC. WASHINGTON: Whatcom Co.: Bellingham: 2♂, 25 June 1950 (JFG Clarke), USNM.
Distribution. Canada (British Columbia) and USA (California, Oregon, Washington).
Host plants. Caprifoliaceae — Lonicera spp., including L. involucrata (Richardson) Banks ex Spreng., and Symphoricarpos albus (L.) S. F. Blake (Miller and Hammond 2003; Powell and Opler 2009).
Remarks. This species shows some variations in the forewing patterns. These variations appear to be unrelated to gender, habitat, or flight time. Distribution of this species (Fig. 23) largely overlaps with that of Euceratia castella but is rather congregated toward the west and extends slightly farther south. Both species often coexist in foothill canyons (Powell and Opler 2009).
Discussion
The wing venation and genital features of Euceratia strongly suggest their association with Ypsolophinae. In addition to Euceratia, Ypsolophinae currently includes four three genera: Bhadorcosma Moriuti, Phrealcia Chrétien, and Ypsolopha Latreille. No phylogenetic relationships of these five genera have been proposed. Interestingly, all genera except for Ypsolopha possess the same type of signum which is a simple, scobinate band without any transverse ridge. The signum of Ypsolopha possesses a single or two transverse ridges. This may indicate that Ypsolopha is more advanced than other genera, including Euceratia.
Of the ypsolophine genera, Ypsolopha occurs worldwide, while other four genera are locally restricted. The distribution of Euceratia is limited to the New World, in contrast to those of Bhadorcosma and Phrealcia which span the Palearctic and Oriental Regions and Madagascar. It is notable that Euceratia shows several similarities with Phrealcia. The latter genus currently includes eight species occurring in Eurasia (Sohn 2015b, in press). Superficial resemblance between Euceratia securella and Phrealcia eximiella seems especially interesting but their relatedness needs to be verified with phylogenetic approaches.
No clear geographical separation was found for the three species of Euceratia. Of those, Euceratia intermedia occurs in the smallest range, exclusively California. Euceratia castella shows the broadest distribution, extending from British Columbia to southern California. Powell and Opler (2009) observed that E. securella ranges farther south than E. castella. Our data, however, show that both species have just a minor difference in their southernmost limits in distribution. Euceratia securella is, however, absent in Idaho where E. castella occurs. Near sympatry of three congeners of Euceratia presents an intriguing question as to what factors have led to their speciation.
Acknowledgements
We express our cordial thanks to Donald R. Davis (U.S. National Museum of Natural History), Lawrence Gall (Yale Peabody Museum of Natural History), Kevin Tuck (retired, Natural Histoty Museum, London), and Jason Weintraub (Academy of Natural Sciences, Philadelphia) for allowing the first author to examine the collections under their responsibility.
