Revision of the Lacinipolia vicina (Grote) complex (Noctuidae, Noctuinae, Eriopygini)

Abstract The Lacinipolia vicina (Grote) species complex, previously consisting of Lacinipolia vicina, Lacinipolia teligera (Morrison), Lacinipolia pensilis (Grote), and Lacinipolia subalba Mustelin is revised to six species: Lacinipolia vicina (eastern USA), Lacinipolia teligera (southern Great Plains), Lacinipolia pensilis (Pacific Northwest and northern Rocky Mountains), Lacinipolia acutipennis (Grote), stat. rev. (= Lacinipolia subalba syn. n.) (western North America), Lacinipolia sareta (Smith), stat. rev. (Canada and western USA) and Lacinipolia dimocki, sp. n. (California and Pacific Northwest). Lectotypes are designated for Lacinipolia vicina, Lacinipolia teligera and Lacinipolia pensilis.


Introduction
Lacinipolia McDunnough is currently one of the largest North American noctuid genera with 61 species, and includes another 10 species described from Mexico and Central America. The diversity centers for Lacinipolia are the arid habitats of the American Southwest and Mexico. Like many of the constituent genera of the 6 Clasper with a thumb-like process on ventral margin, clasper flattened and apex rounded; digitus pointed ( Fig. 59); widely distributed, including West Coast states (Fig. 73) (Fig. 69)  Type material. Mamestra vicina: The type material of L. vicina almost certainly consisted of two species, the eastern species known previously as L. imbuna or L. teligera (Franclemont and Todd 1983) Todd (1982). Type locality: Luzerne Co., Pennsylvania. The original type series of Mamestra imbuna probably also included L. sareta from the southern Lake Michigan region, as Smith (1905a) mentions an August specimen from Hessville, Indiana (a suburb of Chicago), but Todd's lectotype designation fortunately restricts the concept of the name.
Diagnosis. Within the eastern North American range of L. vicina, L. sareta is most similar but the two can usually be separated without dissection by the more southern distribution, larger size and bivoltine spring / fall flight (April-May and September -October) of L. vicina (univoltine from late June to early August for L. sareta). In the male genitalia, L. vicina differs most obviously in the arrangement of the spines above the juxta, consisting of two lateral and one medial field of ventrally projecting spines, whereas in L. sareta the spines are directed dorsally and are on the inside of a large, rhomboid plate. Females of L. vicina have an asymmetrical, invaginated ostium, like the opening of a conch, compared to a simpler ostium with a convex prevaginal plate margin in L. sareta.
Although L. vicina is most closely related to L. teligera, L. vicina and L. teligera are not likely to be confused given the range disjunction and more extensive dark fuscous shading of the hindwing in L. vicina. The male genitalia differ in the shape of the clasper, with the apical lobe narrower and more pointed in L. vicina, and the thumb-like lobe situated one third the distance from the base, compared to halfway in L. teligera.
Distribution and biology. Specimens of L. vicina were examined from Massachusetts, New York, Pennsylvania, Virginia and North Carolina (Fig. 69); Forbes (1954) also cites New Jersey (Lakehurst) and Indiana records. The Indiana record (Smith 1905a) may be erroneous given the long-standing confusion with L. sareta, as discussed in the "Type material" section above. Eastern Ohio records of L. teligera from May and September given by Rings et al. (1992) are most likely L. vicina. Moore's (1955) records for Michigan probably all apply to L. sareta based on flight dates and the widespread distribution of L. sareta in the Great Lakes region. There is no clear indication of habitat preference; in North Carolina L. vicina occurs in open oak-hickory forest (B. Sullivan pers. comm.). Despite the relatively broad distribution and apparent lack of specialized habitat requirements, L. vicina records are few. Lacinipolia vicina is apparently bivoltine, flying in spring (April-May) and in late summer to early fall (late August to early October), with later dates farther south. The larvae were described and illustrated by Godfrey (1972) (reared vouchers examined; CUIC), and are probably polyphagous ground dwellers like other Lacinipolia (Wagner et al. 2011).

Lacinipolia teligera (Morrison, 1875)
Diagnosis. Although closely related to L. vicina, the challenge in identifying L. teligera is in separating it from L. sareta in the southwestern Great Plains where the ranges of the two can overlap. Compared to L. sareta, L. teligera is slightly larger with a broader forewing and better-defined, crisper forewing maculation. Reliable identification should be based on genitalic structure, where L. teligera males have a medial and lateral field of short, ventrally directed spines above the juxta, rather than two large flanges laterally on the juxta with dorsally directed spines in L. sareta. The female L. teligera has an asymmetrical, invaginated ostium (like the opening of a conch), whereas that of L. sareta has a simple ostium with the margin of the prevaginal plate convex.
Distribution and biology. Lacinipolia teligera is known from the Great Plains of central Colorado and eastern Kansas southward to central Texas (Fig. 70). Nothing is known of the early stages, although these are undoubtedly similar to those of L. vicina.
Remarks. Lacinipolia teligera is closely related to L. vicina, and the two have previously been considered conspecific (as L. imbuna; Franclemont and Todd 1983). However the two differ structurally as outlined in the L. vicina diagnosis, in addition to a DNA barcode difference of 1.0%. The two species occupy separate ecoregions and different habitats, with teligera in the grasslands of the Great Plains and vicina in deciduous forest of the Appalachian and Atlantic region. Diagnosis. Lacinipolia pensilis is a northwestern montane species that is often confused with L. sareta and also L. acutipennis in parts of the range. Compared to L. sareta, L. pensilis flies later (August to September versus June to early August), and differs considerably in genitalic structure of both males and females as outlined in the key and the L. sareta account.
Separating pensilis from dark forms of L. acutipennis, which are prevalent in montane habitats of the Pacific Northwest, poses the greatest identification challenge in the L. vicina group. Lacinipolia pensilis usually has better-defined forewing markings, richer brown tones in the forewing medial area, and no tendency for streaky pale patches in the forewing apical area; L. pensilis also averages slightly larger with a broader forewing. The spined crest of the male phallus is more robust and usually with more spines, and never has the thin apically-projecting spine that is normally found in L. acutipennis. In montane parts of the Pacific Northwest (interior British Columbia, northern and central Washington) habitat can help to separate the two, with L. pensilis occurring from dry montane woodland to high elevation subalpine forest, whereas L. acutipennis is characteristic of the dry, low-elevation habitats of the major intermontane valleys. See also remarks in the L. acutipennis account.
Distribution and biology. This species occurs in the western cordilleran region from central British Columbia and western Alberta southward to at least Washington and central Utah. The distribution pattern suggests it may occur farther south along the Cascade-Coast Ranges through Oregon, and further work is needed to establish the southwestern range limits. The larval description and host plants require clarification since the information given by Crumb (1956) and Godfrey (1972) was probably based on both L. acutipennis and L. pensilis. The larvae likely are ground-dwelling, general feeders on shrubs and herbs. (Grote, 1880) Diagnosis. Lacinipolia acutipennis is a western steppe / grassland species that shows considerably greater regional phenotypic variation than others in the L. vicina group. In more mesic habitats (including higher elevations) of the Pacific Northwest and central Rocky Mountains L. acutipennis is replaced by the very similar L. pensilis. The two occur sympatrically in many transitional habitats, mostly dry montane woodlands at moderate elevations. Although phenotypes of L. acutipennis from the most arid habitats (e.g., Figs 34-39) can be distinguished from L. pensilis with relative ease, many northern L. acutipennis populations in the Pacific Northwest are dark, well-marked and very similar to L. pensilis, which makes identifying the two very difficult and led previous workers to conclude that they represent the same species. Compounding this difficulty is the lack of conspicuous genitalic differences that are otherwise typical of the genus. Despite the identification difficulties in the Pacific Northwest, other sympatric populations of L. acutipennis and L. pensilis have clearly different phenotypes. Differences are most pronounced in Great Basin populations (L. pensilis,Figs 25,27 and L. acutipennis, and in the northern Rockies/Great Plains (e.g., Montana L. pensilis, like those in Figs 23,24,and L. acutipennis,. The two differ in male genitalia structure as discussed below. These differences, in addition to a minimum 2.5% divergence in DNA barcodes (Fig. 75), show that (at least) two species are involved.

Lacinipolia acutipennis
Similar phenotypes of L. acutipennis and L. pensilis differ in the shape and size of the forewing, which averages more acute and smaller in L. acutipennis; the brown tones of the medial forewing are more muted in L. acutipennis compared to L. pensilis, giving an overall lower contrast in tone of the medial area with the grey-black antemedial and postmedial areas; the white spot in the anal angle is often more prominent in L. acutipennis, particularly in females; the forewing apex has a more contrastingly pale diffuse area that usually extends farther towards the reniform. In the male genitalia of L. acutipennis, the spinose crest of the phallus usually has a thin, delicate apically-directed spine (which is sometimes broken off, in which case the spine base is still evident), which is absent in L. pensilis; this thin spine is sometimes absent also in L. acutipennis, but in such individuals the entire crest is small and with fewer, smaller cornuti ( Fig.  61c) compared to L. pensilis (Fig 62).
Two phenotypes have been recognized as separate species, L. doira of the Great Basin (Figs 49-51) and L. subalba of southern California (Figs 43-45). Clinal phenotypic variation, lack of diagnostic structural characters, and similarity in DNA barcodes, lead me to treat -doira and -subalba as regional forms.

Distribution and biology.
Lacinipolia acutipennis is a western species common throughout xeric, low elevation habitats of western North America. The core range includes the dry, western portions of the Great Plains, the Great Basin, and the western intermontane valleys north of the Sonoran zone, from southern Saskatchewan and Alberta southward to northern Arizona and New Mexico. Reports from Wisconsin (cited in Forbes 1954), Texas and southern Arizona (Hampson 1905) are probably misidentifications of L. sareta. Crumb's (1954) records from Nebraska and Kansas are plausible; the easternmost specimens I examined were from Watford City in western North Dakota. In the intermontane valleys west of the Rocky Mountains L. acutipennis occurs from southern British Columbia to southern California and northernmost Arizona and New Mexico (Fig. 72). All Pacific Northwest specimens examined from subalpine habitats and from sites west of the Coast Ranges proved to be L. pensilis.
The larval description and host plants require clarification since the information given by Crumb (1956) and Godfrey (1972) was probably based on both L. acutipennis and L. pensilis. The larvae likely are general feeders and may ascend shrubs to feed. Lacinipolia acutipennis flies in late summer with most specimens recorded from mid-August to late September.
Remarks. The name acutipennis has historically been associated with the taxon L. sareta (i.e. L. vicina of authors) rather than L. pensilis. This apparently stemmed from the fact that historical L. acutipennis specimens from western Nevada (the type locality of L. acutipennis) and adjacent northeastern California had been wrongly associated; a series from Truckee, California, examined by Lloyd Martin (and probably others before him, including McDunnough) consists of male L. sareta and female L. acutipennis, but only the male L. sareta were previously dissected. Female L. sareta from the northern Sierra Nevada and especially Nevada are considerably paler. Comparison of the type female of L. acutipennis to all other L. vicina-group taxa occurring in the region of the type locality shows that L. acutipennis is a dark female of the low-elevation taxon previously treated as a form of L. pensilis.

Lacinipolia sareta
Diagnosis. Lacinipolia sareta is the most common and widespread species in the L. pensilis group, and most of the identification difficulties are in separating it from L. pensilis and L. acutipennis in the West. This is most reliably done based on genitalia, where males lack the ventrally projecting, paired spinose crests above the juxta that are found in L. acutipennis and L. pensilis; females of L. sareta have a simple ostium with a strongly convex prevaginal margin, compared to those of L. pensilis and L. acutipennis which have an asymmetrical, conch-shaped ostium with a straight prevaginal margin. Lacinipolia sareta flies earlier in the year (mostly June-July) than L. pensilis and L. acutipennis (August-September), although the southernmost L. sareta populations in Arizona, New Mexico, and Texas fly again in late September-October after an initial May flight.
The remaining species (L. vicina, L. teligera, and L. dimocki) can, for the most part, be distinguished from L. sareta by geographic distribution; in Washington, Oregon and California, where the range of L. sareta overlaps that of L. dimocki, L. sareta is smaller and has a duller white hindwing, in addition to the genitalic characters given under dimocki. From eastern Colorado and New Mexico through western Oklahoma and northern Texas L. sareta overlaps with L. teligera; characters given in the keys and the L. teligera diagnosis will separate the two. The range of L. sareta might overlap with that of L. vicina in the East (from the Great Lakes region eastward through New York and New England), where the smaller size, different flight period and genitalic differences given under L. vicina will reliably separate the two.
Distribution and biology. Lacinipolia sareta occurs throughout western North America from the southern Yukon and Northwest Territories to Texas, Arizona and California; it undoubtedly also occurs in northern Mexico. It ranges eastward across the southern boreal region to at least Quebec, with an unverified record from Maine (Forbes 1954). Most or all records of L. vicina for Michigan (Moore 1955) probably apply to this species, but L. sareta is not known from Ohio (Rings et al. 1992) where it would be expected in sandy habitats along Lake Erie. Although found in a huge variety of woodland, steppe and prairie habitats, L. sareta particularly favours sandy soils and can be abundant in dune and beach habitats. Crumb (1956) describes the grounddwelling, polyphagous larva (as L. vicina). Godfrey (1972) illustrates the larva, and states that Arizona and Montana larvae are identical.
Etymology. This species is named in honour of Thomas E. Dimock for his contributions to the knowledge of southern California moths. His efforts to collect research specimens provided most of the type series of L. dimocki.
Diagnosis. This western species was previously included with L. sareta (L. vicina of authors), but it is a cryptic, mostly parapatric species that replaces L. sareta from the Washington coast ranges southward through California. The two occur sympatrically in south-central Washington, and possibly elsewhere along the interface of the Great Basin-Coast Ranges and Sierra Nevada. Externally L. dimocki is larger with an overall paler, less contrasting forewing pattern and usually a lighter, more pearly-white hindwing. The male genitalia differ in having a sinuate, tine-like clasper rather than the flattened, two-lobed clasper of L. sareta; also the ventral swelling of the phallus is much more pronounced in L. dimocki. Females can be difficult to separate from those of L. sareta; in addition to the forewing characters mentioned above, L. dimocki is generally larger overall and with a less sinuous, less dorsoventrally flattened ductus bursae and a relatively larger corpus bursae.
Description. Head. Antenna of male appearing filiform, but slightly serrate under magnification; antenna of female filiform; dorsal scaling grey; scape, and vertex with a mix of dull-white and dark-grey scales, these spatulate and bifid apically; frons with thin white, strap-like scales, bordered by transverse band of dark-grey scales at dorsal margin; labial palpi with mix of dull-white and dark-grey scales; 3 rd segment 0.4× length of 2 nd segment. Thorax. Vestiture of light-grey scales tipped with dark-grey apex; tegula and patagium with subterminal border of black scales, border of the tegula diffuse, but that of patagium forming distinct black prothoracic line; caudal margin with slight tuft; legs with mix of light-and dark-grey scales, tarsi with slight banding pattern formed by border of lighter scales along distal margin of each tarsal segment. Wings. Average forewing length of males 15.0 mm (n = 9, range 14.2-15.8 mm), females 15.1 mm (n = 9, range 13.8-16.9 mm); forewing ground colour pale grey, medial area pale grey brown; antemedial and postmedial line incomplete or absent, when present then best developed toward anal margin and fading out towards costa, antemedial line double, sometimes with slightly paler grey infill; postmedial line double, often forming pale, indistinct crescent opposite claviform spot; subterminal area with diffuse dark shading in subapical and anal areas, latter sometimes with a small white crescent; basal dash black and crisp; orbicular spot slightly oblong to slightly kidney shaped, with incomplete, thin black border and interior slightly paler than ground colour; reniform   spot with incomplete thin black border, interior slightly paler than ground, with indistinct, darker inner ring; claviform usually distinct, forming a thin black, open V; fringe dark grey, pale grey at vein terminus resulting in indistinct striping; male hindwing bright, slightly pearlescent white with terminal third of veins, and thin diffuse margin fuscous; female hindwing duller white overall with more extensive fuscous shading on veins and marginal area. Abdomen. Vestiture light grey, first four segments with slight dorsal tufts of darker grey scales; tuft of 4 th segment most prominent. Male genitalia. Uncus slender, 10-11× longer than wide, evenly tapered from base to apex, with sparse long setae directed basad; valve with extreme subapical constriction forming a narrow neck, such that apex consists of strongly spatulate cucullus; valve abruptly angled caudoventrally beyond apical third; cucullus anvil shaped, interior surface densely covered with fine long hairs; corona consisting of a single row of flattened marginal spines, and a cluster of spines in tip of caudoventral lobe; sacullus with membranous, rectangular flap (possibly a modified editum), which is densely covered in long setae; clasper forming a long, simple sinuate tine, extending to, or slightly beyond, costa; digitus a simple flattened lobe, 2× longer than wide; juxta with two lateral, rounded triangular plates flanking phallus, these with short, straight dorsally directed spines on inner surface; phallus with ventral swelling 2/3 from base; apical third curving ventrad slightly; phallus with small, broad-based, thorn-like dorsal cornutus at apical ¾; vesica directed leftventrad, then coiling dorsad and forming extended spiral through one rotation; vesica with small medial patch of spinules, and larger preapical patch extending slightly along axis of vesica. Female genitalia. Bursa copulatrix unisaccate; ductus bursae moderately sclerotized and dorsoventrally flattened, 5× longer than wide; corpus bursae globose, membranous and slightly corrugated, lacking signa; appendix bursae slightly coiled, with ductus seminalis situated preapically; ostium bursae extending caudad as an invaginated slit; prevaginal margin convex and slightly rounded-conical; terminal segments telescopic, with posterior apophysis twice as long as anterior apophysis; papillae small, narrow and lobe-like, membranous and moderately setose.
Distribution and biology. The early stages and larval food plants are unknown, but like other species in the group, larvae of L. dimocki probably are ground-dwelling and polyphagous on herbaceous plants. It occurs from the east slope of the Washington Coast Ranges to southern California. data and information from the Barcode of Life Data (BOLD) system. Molecular analyses were carried out through grants from the National Science and Engineering Research Council of Canada and Genome Canada through the Ontario Genomics Institute.