Research Article |
Corresponding author: B. Christian Schmidt ( christian.schmidt@canada.ca ) Academic editor: Donald Lafontaine
© 2015 B. Christian Schmidt.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Schmidt BC (2015) Revision of the Lacinipolia vicina (Grote) complex (Noctuidae, Noctuinae, Eriopygini). In: Schmidt BC, Lafontaine JD (Eds) Contributions to the systematics of New World macro-moths VI. ZooKeys 527: 103-126. https://doi.org/10.3897/zookeys.527.9686
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The Lacinipolia vicina (Grote) species complex, previously consisting of L. vicina, L. teligera (Morrison), L. pensilis (Grote), and L. subalba Mustelin is revised to six species: L. vicina (eastern USA), L. teligera (southern Great Plains), L. pensilis (Pacific Northwest and northern Rocky Mountains), L. acutipennis (Grote), stat. rev. (= L. subalba syn. n.) (western North America), L. sareta (Smith), stat. rev. (Canada and western USA) and L. dimocki, sp. n. (California and Pacific Northwest). Lectotypes are designated for L. vicina, L. teligera and L. pensilis.
Cryptic species, Pacific Northwest, California
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 Eriopygini, a diverse, largely New World tribe, the current concept of Lacinipolia is not monophyletic and in need of revision. Lloyd Martin initiated this considerable undertaking in the 1960s, but abandoned the project after the loss of all his notes and type photographs (
Adult genitalia were prepared following the methods of
BMNH
The
Variation of the ‘barcode’ section of the COI gene was compared among 264 specimens representing all six species (Suppl. material
1 | Male | 2 |
– | Female | 7 |
2 | Spines posterior to juxta (‘above’ juxta in slide preparations) pointing antero-ventrally and forming spinose crests or simple patch |
3 |
– | Spines posterior to juxta pointing postero-dorsally and situated on inside surface of spade-like plate (Figs |
6 |
3 | Medial field of ventrally projecting spines located adjacent to juxta (Figs |
4 |
– | No medial spine patch adjacent to juxta(Figs |
5 |
4 | Clasper with thumb positioned at basal third of distance to apex (Fig. |
L. vicina |
– | Clasper with thumb positioned nearly halfway to apex (Fig. |
L. teligera |
5 | Crest of phallus usually with a thin, delicate apically-directed spine (sometimes broken off, in which case base is still evident) (Fig. |
L. acutipennis |
– | Crest of phallus never with a thin, delicate basally-directed spine (Fig. |
L. pensilis |
6 | Clasper with a thumb-like process on ventral margin, clasper flattened and apex rounded; digitus pointed (Fig. |
L. sareta |
– | Clasper without process, shaped like a sinuate spine with a pointed apex; digitus rounded (Fig. |
L. dimocki |
7 | Ostium asymmetrical, like opening of a conch; margin of prevaginal plate straight or slightly convex (Figs |
8 |
– | Ostium symmetrical, opening simple; margin of prevaginal plate strongly convex (Figs |
1 |
8 | Ostium complex 1.4–1.5 × longer than wide; caudal portion of ostial slit gradually curved (Figs |
9 |
– | Ostium complex 1.0–1.1 × longer than wide; caudal portion of ostial slit sinuate (Figs |
10 |
9 | Forewing ground colour highly variable, but medial area concolourous with postmedial and antemedial areas, and antemedial line absent or poorly defined; usually with apical pale area extended through postmedial line into reniform spot; subterminal area usually darker than postmedial area; reniform and orbicular spots often only faintly visible; orbicular spot sometimes flattened and elongated; arid low elevation habitats including shortgrass prairie and sagebrush steppe | L. acutipennis |
– | Forewing ground colour varying in saturation but consistent in tone, with medial area containing brown tones that are lacking in grey-and-black postmedial and antemedial areas; antemedial line usually well defined; pale apical area not extended through postmedial line; subterminal area not darker than postmedial area; reniform and orbicular spot conspicuous and paler than ground; orbicular spot never highly flattened and elongated; low to high elevation woodlands, particularly dry, montane pine and Douglas-fir woodland | L. pensilis |
10 | Basal half of hindwing conspicuously lighter than marginal portion and forewing (Fig. |
L. teligera |
– | Basal half of hindwing nearly as dark as marginal portion and forewing (Fig. |
L. vicina |
11 | Ductus bursae highly flattened dorsoventrally, with pronounced ribbon-like oblique fold (Fig. |
L. sareta |
– | Ductus bursae moderately flattened dorsoventrally, with slight oblique fold; corpus bursae 3–4 × diameter of ductus bursae (Fig. |
L. dimocki |
Lacinipolia adults. 1 L. vicina ♂ NC, Watauga Co., Beech Ck. bog 2 L. vicina ♂ PA, Beaver Co., 6 mi SW Darlington 3 L. vicina ♀ PA, Beaver Co., 6 mi SW Darlington 4 L. teligera ♂ TX, Lampasas Co., Lampasas R. 8 mi S Rte. 190 5 L. teligera ♂ TX, Travis Co., Austin 6 L. teligera ♀ TX, Lampasas Co., Lampasas R. 8 mi S Rte. 190 7 L. sareta ♂ ON, Manitoulin Is., Dominion Bay dunes 8 L. sareta ♂ BC, [Lillooet], Kirby Flats Rd. 9 L. sareta ♀ ON, Manitoulin Is., Sheguindah 10 L. sareta ♂ AB, Waterton Lakes NP, Blakiston Ck. fan 11 L. sareta ♂ AB, Manyberries, Pakowki Dunes 12 L. sareta ♀ AB, Manyberries, Pakowki Dunes 13 L. sareta ♂ CA, Mono Co., Lee Vining 14 L. sareta ♂AZ, Cochise Co., Huachuca Mtns, Ash Cyn. Rd. 15 L. sareta ♀AZ, [Maricopa Co.], Congress 16 L. dimocki ♂CA, Plumas Co., Jackson Ck., DNA voucher # CNCNoctuoidea7972 17 L. dimocki ♂ Holotype, CA, Ventura Co., Cuyama Valley, Apache Cyn. 18 L. dimocki ♀Paratype, CA, Ventura Co., Cuyama Valley, Apache Cyn. 19 L. dimocki ♂ WA, Klickitat Co., Simcoe Butte 20 L. dimocki ♂ WA, Klickitat Co., Munson Prairie 21 L. dimocki ♀ WA, Yakima Co., South Fork Ahtanum Cr. 22 L. pensilis ♂ BC, Squamish, Diamond Head Trail 23 L. pensilis ♂ BC, [11 km WSW Invermere], Watch Peak 24 L. pensilis ♀ BC, [11 km WSW Invermere], Watch Peak 25 L. pensilis ♂ UT, [Utah Co.,] 12 mi N Provo 26 L. pensilis ♂ WA, Yakima Co., Bethel Ridge 27 L. pensilis ♀ UT, Salt Lake City.
Lacinipolia acutipennis adults. 28 ♂ BC, [Lillooet], Kirby Flats Rd. ♂ BC, Savona, 2 mi SW 30 ♀ BC, [Lillooet], Kirby Flats Rd., DNA voucher # CNCNoctuoidea7978 31 ♀ BC, [Lillooet], Kirby Flats Rd. 32 ♂ WA, [Okanogan Co.], Tonasket, 8 mi S 33 ♀ CA, Plumas Co., Jackson Ck. 34 ♂ WA, Douglas Co., Jameson L. 35 ♂ WA, Grant Co., Dodson Rd. 36 ♀ WA, Yakima Co., South Fork Ahtanum Cr. 37 ♂OR, Crook Co., Suplee 38 ♀ OR, Lake Co., Alkali L. 39 ♀ OR, Baker Co., Burnt River, 20 mi S 40 ♂ CA, Ventura Co., Cuyama Valley, Apache Cyn. 41 ♂ CA, [Monterey Co.], Pinnacles Nat. Mon. 42 ♀ CA, [Monterey Co.], Pinnacles Nat. Mon. 43 ♂ [subalba paratype] CA, San Diego Co., S rim Peñasquitos Cyn. 44 ♂ [subalba paratype] CA, San Diego Co., S rim Peñasquitos Cyn. 45 ♀ [subalba paratype] CA, San Diego Co., NAS Miramar 6 46 ♂ AB, Dinosaur PP 47 ♂ MT, [Philips Co.], Malta, 19 mi NE 48 ♀ WY, [Albany Co.], Laramie 49 ♂ UT, [Juab Co.], Eureka 50 ♂ ID, Owyhee Co., [Castle Ck. Rd.] 51 ♀ UT, Grand Co., Thompson Cyn. 52 ♂ CO, Adams Co., Bennett 53 ♂ CO, Larimer Co., Flatiron Reservoir 54 ♀ CO, Larimer Co., Flatiron Reservoir.
Lacinipolia female genitalia. 63 Lacinipolia vicina MA, Barnstable,
Mamestra vicina Grote, 1874a: 156.
Mamestra imbuna Smith, 1905a: 201, syn. rev.
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 (
Mamestra imbuna: Male lectotype (
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.
Specimens of L. vicina were examined from Massachusetts, New York, Pennsylvania, Virginia and North Carolina (Fig.
As defined here, L. vicina is the same species later described by
Mamestra teligera Morrison, 1875: 215.
Morrison’s original description was based on two specimens, and Wilterding (1997) discussed two Morrison specimens in the
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.
Lacinipolia teligera is known from the Great Plains of central Colorado and eastern Kansas southward to central Texas (Fig.
Lacinipolia teligera is closely related to L. vicina, and the two have previously been considered conspecific (as L. imbuna;
Dianthoecia pensilis Grote, 1874b: 199.
described from at least 1 male and 1 female syntype; the following male [BMNH] is here designated as lectotype: “Dianthoecia / pensilis ♂ / Type Grote” [red-bordered handwritten label]; “Type” [round, red-bordered, typed label]; Vancouver I / Grote Coll. / 81 – 116”; “5597”; Noctuidae / Brit. Mus. slide / No. 4912 ♂” [blue type-written label]; the following label will be added: “Lectotype / Dianthoecia / pensilis Grote, 1874 / desig. by Schmidt 2015.” Type locality: Victoria, [Vancouver Island, British Columbia, Canada].
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.
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
Mamestra acutipennis Grote, 1880: 214.
Mamestra doira Strecker, 1898: 7, syn. rev.
Mamestra ascula Smith, 1905b: 257, syn. rev.
†Polia pensilis ab. indistincta Strand, 1917: 28, unavailable; infrasubspecific name.
Lacinipolia subalba Mustelin, 2000: 13, syn. n.
Mamestra acutipennis: type female (BMNH; examined); type locality: Nevada. Mamestra doira: type female (
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
Neighbour-joining tree of representative mtDNA barcode haplotypes in species of the Lacinipolia vicina group. Sample size and locality are given in brackets, with number of specimens indicated after two-letter state/province abbreviation. Lacinipolia sareta variation is divided into five haplogroups, A–E. Voucher specimen data is given in Suppl. material
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.
Two phenotypes have been recognized as separate species, L. doira of the Great Basin (Figs
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
The larval description and host plants require clarification since the information given by
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.
Variation in the DNA barcodes (Fig.
Mamestra sareta Smith, 1906: 229.
lectotype male (
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.
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 (
The vast geographic range and considerable DNA barcode variation suggest that L. sareta could be a cryptic species complex. Alternatively, DNA barcode variation simply may not be fully congruent with species limits in the group, a phenomenon that occurs in about 10% of Noctuoidea (
Holotype ♂. California: Ventura Co., Cuyama Valley, Apache Canyon, 0.6 mi E of Hwy. 33, 34.751193°N, 119.399772°W, 3497’, 19.Jun.2009, T. E. Dimock [
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.
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.
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; 3rd segment 0.4× length of 2nd 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 4th 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 left-ventrad, 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.
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.
I thank Jason Dombroskie (Cornell University, Ithaca, NY) and Michael Pogue (Systematic Entomology Laboratory, National Museum of Natural History, Washington, DC), for the loan of specimens, and Jocelyn Gill (
The spine field above the juxta and even the aedeagal cornuti can often be examined without dissection by carefully removing the terminal abdominal hairs with a small brush, especially if the valves have previously been spread by squeezing the base of the genital capsule when specimen is still fresh.
Table S1. Specimen data for mtDNA barcode vouchers
Data type: data spreadsheet
Explanation note: Haplogroup numbers refer to those given in Fig.