Lasionycta uniformis uniformis (Smith)

Crabo, Lars & Lafontaine, Donald, 2009, A Revision of Lasionycta Aurivillius (Lepidoptera: Noctuidae) for North America and notes on Eurasian species, with descriptions of 17 new species, 6 new subspecies, a new genus, and two new species of Tricholita Grote, ZooKeys 30 (30), pp. 1-156 : 69-77

Lasionycta uniformis uniformis (Smith)

Figs 79–81, 171, 227. Map 16

Scotogramma uniformis Smith, 1893b: 101 .

Type material. Holotype ♁ [ USNM, examined]. Type locality: Laggan [Lake Louise], Alberta.

Diagnosis. The nominate subspecies from the Canadian Rocky Mountains is aptly named. Most specimens have a uniform medium-gray forewing with faint markings. Some specimens have olive scales in the forewing spots. Th e dorsal hindwing is uniform gray, with lighter areas along the anterior margin in some specimens, and a white fringe. Th e ventral hindwing is unique in Lasionycta . It is pale whitish gray with a dark-gray chevron-shaped discal spot, very faint postmedial line, and solid dark-gray marginal band.

Lasionycta uniformis is most likely to be confused with L. brunnea with which it occurs. Differences between them are given under L. brunnea . Lasionycta uniformis also occurs with L. promulsa from which it is readily distinguished by the hindwing ground color and pattern. That of L. promulsa is yellow brown with a relatively faint discal spot and marginal band.

Th e four CO1 haplotypes of this subspecies differ by up to 0.8 %.

Distribution and geographical variation. Lasionycta u. uniformis occurs in the Rocky Mountains and Purcell Mountains of southwestern British Columbia north to northeastern British Columbia.

The L. uniformis population from Pink Mountain in northeastern British Columbia is variable and shows intergradation between subspecies uniformis and subspecies multicolor. Some specimens with distinctly marked vividly colored dorsal forewings resembling those of L. u. multicolor show the typical ventral pattern of subspecies uniformis . Conversely, some specimens have the opposite combination of characters with a drab dorsal pattern and pale ventral hindwings with a dark postmedial line and fainter marginal band. This population is assigned to the nominate subspecies based on the most typical form at this location and the proximity of the locale to other L. u. uniformis populations. Farther north and west at Montana Mountain in southwestern Yukon most specimens show characters most consistent with L. u. multicolor and are assigned to that subspecies. Very little material is available from the mountains in northern British Columbia. It is expected that a cline between these subspecies exists in this region.

Specimens have been collected during daytime but it is most common at night, even at the northernmost extent of its range. It has been collected from early July to mid-August.

Lasionycta uniformis multicolor Crabo & Lafontaine , ssp. n. urn:lsid:zoobank.org:act:9FAC8E64-5BF3-4EE8-9C45-6DDE56DA3175

Figs 70–73, 172, 228. Map 16

Type material. Holotype ♁. Canada, British Columbia, Gott Peak , 7100’, 50° 21' N 122° 08' W, 13 July 2001, J. Troubridge. CNC . Paratypes 326 ♁, 77 ♀. Canada.

British Columbia. Same data as holotype (1 ♁) ; same locality as holotype, 29 July 2000, J. Troubridge and H. Hensel (178 ♁, 40 ♀), 30 July 1994, L. Crabo and J. Troubridge (29 ♁), 23 Aug. 1996, J. Troubridge (1 ♁), 29 July 2000, J. Troubridge (3 ♁, 2 ♀) ; 17 Aug. 2000, J. Troubridge (5 ♁, 3 ♀); 6 Aug. 2005, L. G. Crabo (1 ♁), 26 July 2006, L. G. Crabo (74 ♁, 22 ♀); Mission Ridge , 5800’, 50° 45' N 119° 37' W, 10 July 1998, J. Troubridge (1 ♁, 3 ♀), 15–16 July 1994, L. Crabo and J. Troubridge (33 ♁, 7 ♀). AMNH, CDFC, CNC, GBC, JSC, LGC, OSU, TMC, UASM, USNM, WSU .

The type series is restricted to Gott Peak and Mission Ridge in the southern British Columbia Coast Range.

Etymology. The name multicolor is derived from the Latin and has the same meaning in English. It refers to the vivid color and myriad forms of this subspecies.

Diagnosis. Lasionycta u. multicolor is a protean subspecies from the Pacific Northwest. Its forewing is gray to dark gray with variable patches of yellow or blue gray in the basal, postmedial, and terminal areas producing a number of color forms that appear mottled gray, blue gray, yellow, yellow green, or nearly black. Th e variability is most marked in the British Columbia Coast Range. Th e hindwing postmedial line is darker than the marginal band.

This subspecies is partially sympatric with L. caesia and L. gelida , but can be separated from both species by features of the hindwing and genitalia. Lasionycta u. multicolor has a pale-gray ventral hindwing with a narrow postmedial line and lighter patchy marginal band lacking dark scaling on the veins. That of L. caesia has a wider less distinct postmedial line, a marginal band of similar darkness to the line, and dark scaling on the veins. Lasionycta u. multicolor is distinguished from L. gelida by the ground color of the dorsal hindwing, gray with broad dark marginal band in L. u. multicolor and pale whitish-gray with thinner dark marginal band in L. gelida . Males of L. u. multicolor have a corona that is mostly single. Those of L. caesia and L. gelida are compound with several rows of setae. Th e female genitalia are similar, but the oviposi- tors of L. u. multicolor are normal for the species-group whereas those of L. gelida are large and rounded apically.

Th is subspecies has four CO1 haplotypes differing by up to 1.2 %.

Description. Head – Antenna of male biserrate and fasciculate, individual segments triangular, 1.5–2.1× as wide as central shaft. Antenna of female filiform and ciliate. Dorsal segments slate gray proximally, mostly white with scattered slate-gray scales distally. Scape luteous off-white. Eye normal size. Palpus covered with slate gray and luteous off-white scales. Frons luteous off-white centrally, gray laterally. Top of head a mixture of cream and white-tipped gray hair-like scales. Thorax – Vestiture hair-like luteous off-white and white-tipped gray scales, appearing a uniform slightly luteous gray. Legs covered with slate-gray and luteous scales. Tarsal segments slate gray with distal ring of luteous scales. Wings – Forewing length: males 14–16 mm (expanse 31–36 mm); females 14–16 mm (expanse 33–36 mm). Forewing covered with medium- to dark-gray, light to dark ochre-yellow, and pale bluish-white scales; appearing patchy medium to dark gray with very variable blue-gray, light- to dark-yellow, or greenish-gray patches, most prominently proximal to antemedial line, adjacent to postmedial line, and in terminal area. Lines dark gray, single. Basal and antemedial lines undulating to jagged. Medial line dark gray, weak to moderately dark. Postmedial line scalloped between veins, convex from costal margin to fold, then oblique to posterior margin. Subterminal line uneven, pale, same as terminal area in vividly colored specimens. Spots dark gray, less conspicuous than lines. Orbicular spot variable in size, round to ovoid, filling usually same color as basal area, with or without a dark-gray ocellus. Reniform spot kidney shaped, variable in prominence, evident mostly due to darker black medial part and filling; filling similar to basal area in color peripherally but usually less vivid than filling of orbicular spot, with variable dark-gray central filling in entire spot or limited to lower part. Claviform spot absent. Fringe same color as terminal area, weakly to strongly checkered with gray between veins. Ventral forewing gray, with variable suffusion of pale-luteous scales in fold, distal to discal spot, and distal to postmedial and subterminal lines. Costa darker gray, usually mixed with yellow scales. Discal spot dark gray, thin and small. Postmedial line dark gray, variable, usually thick and undulating but occasionally scalloped between veins or only evident near costa. Postmedial line pale, preceded by a wide dark-gray shade. Fringe yellow gray, checkered with darker gray between veins. Dorsal hindwing gray with darker markings. Discal spot chevron- to half-moon shaped. Postmedial line thick, similar to discal spot in darkness, undulating. Marginal area similar to discal spot and postmedial line, broad with diffuse inner margin. Fringe light gray and pale yellow proximally, pale yellow distally. Ventral hindwing luteous off-white with dusting of gray scales. Discal spot prominent, dark gray, arrowhead shaped. Postmedial line located near discal spot, slightly lighter gray than discal spot, thick and slightly ill defined, nearly even with slight undulation near anal angle. Marginal band gray, similar in darkness to postmedial line or slightly lighter, interrupted by patchy suffusion of ground color, moderately wide with diffuse inner margin. Fringe two toned, proximal row luteous off-white with scattered light-gray scales, distal portion uniform luteous off-white. Abdomen – Light gray, luteous distally in males. Male genitalia – (Fig. 172) Genital capsule and aedeagus as in L. leucocycla species-group and L. phoca sub-group descriptions. Valve approximately 7–8× as long as wide with moderately constricted neck. Digitus moderately long. Cucullus moderate size, usually with pointed apex, with corona comprised mainly of a single row of setae, partially double at apex. Vesica with 0–3 basal cornuti (N = 7). Female genitalia – (Fig. 228) Ovipositor lobe, segment VIII, and bursa copulatrix as in the L. leucocycla species-group and L. phoca sub-group descriptions.

Distribution and geographical variation. Lasionycta u. multicolor occurs from Montana Mountain in southwestern Yukon, south in the British Columbia Coast Range to the Cascades in southern Washington. The southern populations extend east to the Okanagan Trench. Populations from the Coast Range and northern Washington are variable whereas other Washington specimens are more uniformly gray. Those from mountains east of the Coast Range in British Columbia are very dark, resembling the eastern species L. phoca .

Lasionycta uniformis fusca Crabo & Lafontaine , ssp. n. urn:lsid:zoobank.org:act:5963F711-8C0D-4C26-B6DC-CA2F3B5DAB80

Figs 76–78, 173, 229. Map 16

Type material. Holotype: ♁. USA, Colorado, Estes Park , 11 mi WNW, 11,000’, 25 July 1967, D. F. Hardwick. CNC . Paratypes 21 ♁, 4 ♀. USA. Colorado. Same data as holotype, (2 ♁) ; Estes Park , 12 mi WNW, 11,600’, 22–30 July 1967, D. F. Hardwick (11 ♁, 3 ♀) ; Mt. Evans , 3450 m., timberline, 17 July 1993, K. Mikkola (2 ♁) ; Chaffee Co., Cottonwood Pass , 19 July 1982, G. Balogh (1 ♀) ; Park Co., Pennsylvania Mt., near Fairplay , 11,500’, 11, 17 July 1979, P. G. Kevan (4 ♁) ; Guanella Pass Co., 11,000’, 17 July 1998, D. E. Bowman (2 ♁). CNC, GBC, LGC, MZHF .

Th e type series is restricted to Colorado.

Etymology. The name is derived from the Latin fuscus meaning dark. It refers to the dark, drab coloration of this subspecies relative to L. u. multicolor.

Diagnosis. Lasionycta u. fusca occurs in the central Rocky Mountains. Like subspecies multicolor it differs from the nominate subspecies in having more distinct forewing markings. The ventral hindwings are similar to those of subspecies multicolor, although the postmedial line of L. u. fusca is often wider. Th is subspecies is less colorful than L. u. multicolor, being gray, greenish gray, or brown gray depending on location. The individuals of each population are less variable than those of L. u. multicolor, although there is variation between populations in different mountain ranges.

Lasionycta u. fusca occurs with L. discolor in Colorado and Wyoming and pale specimens resemble this species. Th e dorsal hindwing of L. u. fusca has a sinuous postmedial line and lacks white distal to it, whereas L. discolor has a weakly dentate line with white between the line and marginal band. Th e ventral hindwing of L. u. fusca is pale gray and lacks dark veins, whereas that of L. discolor is nearly pure white with dark veins. Lasionycta u. fusca tends to have fewer cornuti on the male vesica (up to three), whereas L. discolor has up to six. Females are structurally indistinguishable. Brownish populations of L. u. fusca can be mistaken for L. promulsa but have a larger arrowheadshaped ventral hindwing discal spot than this species. Th e brown populations are also similar to L. brunnea but are easily distinguished by locality since L. brunnea occurs from northeastern Washington and further north in the range of L. u. uniformis .

The CO1 DNA of L. u. fusca appears to be less variable than that of L. u. multicolor. Th e three haplotypes differ by up to 0.8 %.

Description. Head – Antenna and eye similar to subspecies multicolor in males and females. Scape cream with a few gray scales. Palpus covered with mostly cream and a few gray scales. Frons white centrally, gray laterally. Top of head covered with white, black, and white-tipped black hair-like scales. Thorax – Vestiture of white, gray, and luteous (in specimens with luteous scales on forewing) hair-like scales, appearing uniform medium to dark gray with a luteous tint in specimens with yellow on forewings. Legs with gray to dark-gray and white to luteous off-white scales. Tarsal segments dark gray with few white scales, ringed distally with white. Wings – Forewing size same as subspecies multicolor. Ground color a mixture of medium- to dark-gray, white, and pale- to dark-luteous scales, appearing patchy greenish gray, dark gray, or slightly brownish gray; yellow scales more diffuse and forming less solid patches than in subspecies multicolor. Basal, antemedial and postmedial lines weakly double with light-gray to luteous filling. Basal and antemedial lines irregular. Postmedial line dark gray, moderately prominent and darkest near costa. Postmedial line moderately scalloped between veins. Subterminal line pale gray to luteous, irregular, preceded by an indistinct dark-gray shade. Spots dark gray. Orbicular spot round to oval, slightly variable in size, variably filled with ground color, whitish gray, or luteous, without or with a dark-gray ocellus. Reniform spot relatively inconspicuous, kidney shaped, filling usually pale whitish gray peripherally and ground color to dark gray centrally, medial part and adjacent pale filling most prominent. Claviform spot absent. Terminal line dark gray, thin, most evident between veins. Fringe light to medium gray or luteous, weakly checkered with darker gray between veins. Ventral forewing uniform light to medium gray centrally with lighter luteous off-white to light brownish gray from fold to posterior margin and distal to subterminal line; costa a mixture of dark-gray and luteous scales. Discal spot prominent, dark gray with anterior half thickest and darkest, slightly smaller and lighter than hindwing discal spot. Subterminal line dark gray with anterior part darkest, prominent in all but the lightest specimens but slightly lighter than hindwing subterminal line, broad and somewhat indistinct, slightly scalloped in some specimens. Distal wing slightly darker due to dusting of dark-gray scales, condensed to a weak subterminal line in some specimens. Fringe light gray to pale luteous, checkered with dark gray between veins. Dorsal hindwing ground color a mixture of light- and medium-gray scales with very faint brown tinge, lightest near anal angle, with slightly darker markings. Discal spot arrowhead shaped. Postmedial line broad, slightly ill defined and undulating, not scalloped. Marginal band broad and poorly defined medially. Fringe pale gray to luteous gray proximally, pure to luteous white distally. Ventral hindwing faintly luteous to brownish white with dusting of gray scales. Discal spot dark gray, darker than other ventral markings, thick and arrowhead shaped. Postmedial line prominent in all but lightest specimens, but lighter than discal spot, usually wider than in subspecies multicolor, slightly indistinct and undulating near posterior margin. Marginal band paler gray than postmedial line, with diffuse inner margin interrupted by pale scales. Fringe luteous off-white, proximal row patchy light gray in most specimens. Abdomen – A mixture of light-gray to luteous and medium-gray scales. Long scales covering male genitalia uniformly pale gray to luteous. Male genitalia – The male genitalia are indistinguishable from those of L. u. multicolor. Female genitalia – Th e female genitalia are indistinguishable from those of L. u. multicolor.

Distribution and geographical variation. Lasionycta u. fusca occurs from central Colorado and northern Utah to the Beartooth Plateau on the Wyoming-Montana border. Th e population of each mountain range tends to have a distinct appearance. Colorado specimens are dark greenish gray, those from the Snowy Range in southern Wyoming are lighter greenish gray, Beartooth Plateau populations are gray, whereas populations from the Uinta Mountains, Utah and the Big Horn Mountains, Wyoming are brown gray. Most examined specimens are from mid- to late July.

Remarks. This subspecies was previously known as L. discolor , now recognized as a synonym of Lasiestra klotsi Richards.

Lasionycta uniformis shasta Crabo & Lafontaine , ssp. n. urn:lsid:zoobank.org:act:E6C1581B-1051-44FA-86D2-2EFD92E0BD20

Fig. 75. Map 16

Type material. Holotype ♁. USA, California, Siskiyou County, Mount Shasta, Panther Meadows , 7500’, 23 July 1965, E. and I. Munroe. CNC . Paratype 1 ♁. USA.

California. Same locality as holotype except elevation 7600’, 20 Aug. 2000, J. Troubridge, Databased for CNC Noctuoidea #10484, Barcodes of Life Project, University of Guelph, DNA #Noctuoidea 10484 (1 ♁). CNC.

Etymology. Th e name refers to Mount Shasta. It is a noun in apposition.

Diagnosis. Lasionycta u. shasta is paler than other subspecies. Th e forewing is light blue gray with pale-gray mottling. It is the only subspecies of L. uniformis that consistently has dark veins on the ventral hindwing (rarely present in L. u. uniformis ). Lasionycta u. shasta is unlikely to be confused with any other Lasionycta in its range.

The CO1 sequence of this subspecies is most similar to two of the L. u. multicolor haplotypes.

Description. Only known from males. Head – Antenna of male similar to subspecies multicolor. Scape luteous off-white. Palpus luteous off-white with a few gray scales. Frons white centrally, light gray laterally. Top of head mostly white with a few gray and white-tipped gray scales. Thorax – Vestiture cream and white-tipped gray hair-like scales, appearing uniform luteous light gray. Legs with gray and luteous off-white scales. Tarsal segments gray with few white scales, ringed distally with white. Wings – Forewing length 14 mm (expanse 32 mm). Forewing ground color a mixture of gray, white, and luteous scales, appearing mottled gray with light-gray patches, most prominently basal to antemedial line, distal to reniform spot, and in subterminal and terminal areas. Lines dark gray. Basal and antemedial lines uneven. Medial line dark gray, moderately prominent. Postmedial line evenly scalloped between veins, excurved from costa to lower end of cell, then oblique to posterior margin. Subterminal line uneven, pale gray, preceded by slightly patchy dark-gray shade. Terminal line dark gray, evident between veins. Spots dark gray. Orbicular spot round, filled with light bluish gray and a dark-gray central ocellus. Reniform spot kidney shaped, inconspicuous, filled with light gray peripherally and dark gray centrally. Claviform spot absent. Fringe slightly lighter gray than ground color, weakly checkered with darker gray between veins. Ventral forewing uniformly gray, suffused with light gray from fold to posterior margin and with pale-luteous scales distal to reniform spot and postmedial line. Discal spot dark gray, prominent but lighter than hindwing discal spot. Postmedial line dark gray, wide and slightly ill defined, scalloped between veins. Marginal area heavily suffused with dark gray. Terminal line dark gray, interrupted on veins. Fringe luteous white, checkered with gray between veins. Dorsal hindwing ground slightly peach-tinted off-white, suffused with gray scales, especially basal to postmedial line. Markings dark gray. Discal spot arrowhead shaped. Postmedial line prominent, scalloped. Marginal band solid with indistinct inner margin. Hindwing fringe pale gray proximally, nearly white distally. Ventral hindwing luteous off-white with dusting of gray scales, heaviest basal to postmedial line. Markings and veins dark gray. Discal spot darkest gray, arrowhead shaped. Postmedial line similar in prominence to that of subspecies multicolor, undulating. Marginal band similar in darkness to postmedial line, with indistinct slightly scalloped inner margin. Fringe pale gray and white proximally, white distally. Abdomen – A mixture of light- and medium-gray scales. Long scales covering male genitalia luteous light gray. Male genitalia – Indistinguishable from those of subspecies multicolor.

Distribution. Lasionycta u. shasta occurs on Mount Shasta in the Cascade Range of northern California. It might be more widely distributed in northern California and Oregon in the southern Cascades or Klamath Mountains.

Lasionycta uniformis handfieldi Crabo & Lafontaine , ssp. n. urn:lsid:zoobank.org:act:9A6FA52E-051A-43F6-8086-E478F2A22C84

Fig. 74. Map 16

Type Material. Holotype ♀. Canada, Quebec, Gaspé-Ouest Co, Parc de la Gaspésies, Mont Albert, 3 July 1987, 1070 m, Yves-Pascal Dion, Slide CNC No. 9901, Database # CNC LEP 00053369 . CNC . Paratype 1 ♀. Same locality as holotype, except elev 3500–3750’, 21 July 1933, W. J. Brown, Slide Female CNC No. 9956. CNC.

Etymology. This subspecies is named in honor of Louis Handfield in recognition of his contributions to the study of Lepidoptera in Quebec.

Diagnosis. Lasionycta u. handfieldi occurs on Mount Albert in the Gaspé Peninsula of Quebec. It is hoary dark olive gray due to a suffusion of luteous scales on a black background. Th e dorsal hindwing ground color has a faint peach-colored tint. This subspecies is most likely to be confused with L. phoca , which occurs farther north in Labrador and Quebec. Lasionycta u. handfieldi is greenish and has a pale-brown ventral hindwing with separate line and spot. Lasionycta phoca is blackish gray and has a darkgray ventral hindwing with the postmedial line touching the discal spot.

Lasionycta u. handfieldi represents a curious disjunct occurrence of an otherwise western species. The Gulf of St. Lawrence harbors other disjunct western plants and animals, likely remnants of a Pleistocene glacial refugium.

The worn paratype was correctly associated with L. uniformis by Rockburne and Lafontaine (1976).

Description. Known only from females. Head – Antenna of female filiform and ciliate. Dorsal antennal segments dark gray proximally, luteous distally. Scape luteous with a few gray scales. Eye normal size. Palpus covered with luteous and gray scales. Frons luteous off-white centrally, dark gray laterally. Top of head covered with luteous and dark-gray scales. Thorax – Vestiture hair-like with black, olive-gray, and luteous scales, appearing dark olive gray. Legs covered with mixture of luteous and gray scales. Tarsal segments dark gray with luteous ring of scales distally. Wings – Forewing length 15 mm (expanse 35 mm). Forewing ground color a mixture of black, gray-olive, luteous, and white scales, appearing hoary greenish gray; medial area, apex, and fold distal to subterminal line lighter greenish yellow due to a heavy suffusion of yellow scales. Lines and spots black. Basal line evident only at costa. Antemedial line thick and jagged. Medial line incomplete, most evident near costa and posterior margin. Postmedial line sawtooth shaped, drawn inward between veins, gently excurved from costa to lower cell, then oblique to inner margin. Subterminal line pale luteous, preceded by faint dark-gray shade. Orbicular spot obsolete, evident as a yellow streak. Reniform spot inconspicuous, evident mostly due to light yellow peripheral and black central filling. Claviform spot absent. Fringe luteous, weakly checkered with gray olive between veins. Ventral forewing uniform dark gray centrally with patchy suffusion of light brownish-gray scales in fold, distal to reniform spot, and in subterminal and terminal areas. Discal spot a dark-gray bar, much thinner and lighter than hindwing discal spot. Postmedial line dark gray, moderately thick and ill defined. Postmedial line pale, preceded by a gray shade of similar darkness to postmedial line. Fringe luteous, weakly scalloped with gray between veins. Dorsal hindwing ground color peach-tinted pale gray, palest at costa and at anal angle, moderately suffused with dark-gray scales. Markings dark gray. Discal spot large, arrowhead shaped. Postmedial line weakly scalloped, undulating near anal margin. Marginal band broad, diffuse proximally. Fringe two-toned with luteous proximal and white distal components. Ventral hindwing pale brownish gray with dusting of gray scales, appearing weakly peppered with dark-gray markings. Discal spot nearly black, weakly arrowhead shaped. Postmedial line ill defined due to suffusion with lighter scales, undulating. Marginal band indistinct, heavily suffused with lighter scales. Fringe nearly uniform faintly brownish off-white. Ab-

domen – Abdomens of both type specimens removed for genitalia dissection. Female genitalia – Female genitalia indistinguishable from those of L. u. uniformis .