Corresponding author: Tomas Mustelin (
Academic editor: D. Lafontaine
The genus
Mustelin T, Crabo LG (2015) Revision of the genus
The genus
Most species of
The combination of too many names, geographical variation, and the fact that many
Wing pattern and genitalia structure terminology follow
Characters and nomenclature of the genus
The male and female genitalia were prepared using standard methods (
The 658 base pair DNA “barcode” region of the mitochondrial cytochrome
Species | Haplotype | Voucher # | Seq. length | Country | State/Prov. | Exact Site | Lat | Lon | Collectors | Deposition |
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ABI1 | CNCNoctuoidea12188 | 658[0n] | USA | CA | Pine Mountain, Ventura Co. | T. Dimmock |
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ABI2 | CNCNoctuoidea12190 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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ABI3 | CNCNoctuoidea12200 | 658[0n] | USA | UT | Capital Reef, Garfield Co. | P. Opler |
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ABI4 | TMustelin#319 | 609[0n] | USA | CO | John Brown Canyon, Mesa Co | J.S. Nordin |
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ABI5 | CNCNoctuoidea12164 | 658[0n] | USA | WY | Upper Blair P.G. north of Rd 705, Albany Co. | J.S. Nordin |
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ABI6 | CNCNoctuoidea12165 | 658[0n] | USA | NV | 11 mi SW Wells | Lafontaine and Troubridge |
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ABI7 | TMustelin#317 | 658[0n] | USA | WY | Fox Creek, Albany Co | J.S. Nordin |
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ACA1 | CNCNoctuoidea12193 | 658[0n] | USA | CA | Anza Borrego, San Diego Co. | T. Mustelin |
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ACA2 | CNCNoctuoidea12196 | 658[0n] | USA | CA | Anza Borrego, San Diego Co. | T. Mustelin |
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ACA3 | CNCNoctuoidea12197 | 658[0n] | USA | AZ | Hwy 88 12miNE Apache Jct, Maricopa Co. | J. Troubridge |
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ACH1 | CNCNoctuoidea12207 | 658[0n] | USA | WA | Bridgeport |
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J. Troubridge |
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ACH2 | CNCNoctuoidea12209 | 658[0n] | Canada | BC | Mount Kobau |
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J. Troubridge |
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ACH3 | CNCNoctuoidea12212 | 658[0n] | USA | NV | Angel Lake |
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Troubridge and Lafontaine |
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ACH4 | CNCNoctuoidea13376 | 658[0n] | USA | CA | San Bernardino Mountains, San Bernardino Co. |
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T&S Mustelin |
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AET1 | CNCNoctuoidea12175 | 658[0n] | USA | CA | Magalia, Butte Co. | L. Crabtree |
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AET2 | TMustelin#193 | 592[0n] | USA | CA | 2 mi south of Lake Henshaw, San Diego Co | T. Mustelin |
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AET3 | TMustelin#258 | 609[0n] | USA | OR | Illinois River, Josephine Co |
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J. Troubridge |
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AFA1 | CNCNoctuoidea13155 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T.&S. Mustelin |
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AFA2 | CNCNoctuoidea13378 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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AFE | CNCNoctuoidea12170 | 658[0n] | USA | CA | 2miNE of Julian, San Diego Co. | T. Mustelin |
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AFUE1 | CNCNoctuoidea12176 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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AFUE2 | CNCNoctuoidea13380 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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AFUM | CNCNoctuoidea12161 | 658[0n] | USA | CA | Pine Mountain, Ventura Co. | T. Dimock |
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ALI | TMustelin#318 | 609[0n] | USA | CA | Twain Harte, [? County], | Lundgren |
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MU | CNCNoctuoidea12173 | 612[0n] | USA | CA | Inaja Picnic Ground, San Diego Co. | T. Mustelin |
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APE1 | CNCNoctuoidea12127 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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APE2 | CNCNoctuoidea12129 | 658[0n] | USA | CA | Peñasquitos Canyon, San Diego | T. Mustelin |
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APE3 | TMustelin#307 | 609[0n] | USA | CA | Upper Ojai Valley, Ventura Co | T.E. Dimock |
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APS | CNCNoctuoidea12148 | 658[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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ASE | TMustelin#316 | 572[0n] | USA | CA | In-Ko-Pah Gorge, Imperial Co | T. Mustelin and N. Bloomfield |
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ASU1 | TMustelin#22 | 603[0n] | USA | CA | San Diego |
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ASU2 | TMustelin#15 | 609[0n] | USA | CA | Scissors Crossing, San Diego Co. | T. Mustelin |
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ASU3 | TMustelin#321 | 609[0n] | USA | CA | Laguna Mountains, San Diego Co. | T. Mustelin |
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ATO | TMustelin#284 | 609[0n] | USA | CA | Torrey Pines State Reserve, San Diego | N. Bloomfield |
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PAD1 | CNCNoctuoidea12201 | 658[0n] | Canada | BC | Vancouver Island, Saanichton | J. Troubridge |
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PAD2 | CNCNoctuoidea12202 | 658[0n] | USA | CA | Lee Vining, Mono Co. |
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J. Troubridge, L. Crabo |
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PAD3 | CNCNoctuoidea12206 | 658[0n] | Canada | BC | Kirby Flats | J. Troubridge |
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PAD4 | CNCNoctuoidea12203 | 658[0n] | USA | CA | Laguna Mts, San Diego Co. | T. Mustelin |
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PAD5 | CNCNoctuoidea12204 | 658[0n] | USA | CA | Alamo Mountain, Ventura Co. | T. Dimmock |
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PAD6 | LEP038031 | 658[0n] | USA | CA | 2 mi. E. Bassetts, Hy 49, Sierra Co. | P. A. & E. Opler |
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VMA | CNCNoctuoidea12235 | 658[0n] | USA | CA | McCain Valley, San Diego Co. |
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T. Mustelin |
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This study is based on examination of material, including type specimens and genitalia preparations, in the following collections:
The American Museum of Natural History, New York, New York, USA
The Natural History Museum [British Museum of Natural History], London, UK
Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Ontario, Canada
Field Museum of Natural History, Chicago, Illinois, USA
Los Angeles County Museum, Los Angeles, California, USA
Lars Crabo Collection, Bellingham, Washington, USA
Michigan State University, East Lansing, Michigan, USA
Oregon Department of Agriculture, Salem, Oregon, USA
Oregon State Arthropod Collection, Corvallis, Oregon, USA
Ronald H. Leuschner Collection, now at the McGuire Center, Gainesville, Florida, USA
San Diego Natural History Museum, San Diego, California, USA
Thomas E. Dimock Collection, Ventura, California, USA
Tomas Mustelin Collection, Potomac, Maryland, USA
University of California at Riverside, Riverside, California, USA
National Museum of Natural History [formerly United States National Museum], Washington, District of Columbia, USA
W. F. Barr Entomological Collection, University of Idaho, Moscow, Idaho, USA
James Entomological Collection, Washington State University, Pullman, Washington, USA
Zoological Museum, Helsinki, Finland
1 | Male vesica with apical long spine-like cornutus; posterior half of female ductus bursae membranous |
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– | Male vesica apex lacking cornutus or with minute cornutus; posterior half of female ductus bursae at least partially sclerotized |
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2 | Ampulla of male clasper present; female corpus bursae with four long signa and posterior ductus bursae sclerotized circumferentially |
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– | Ampulla of male clasper absent; female corpus bursae lacking signa and posterior ductus bursae with sclerotized plate in ventral wall |
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1 | Male |
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– | Female |
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2 | Antenna serrate |
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– | Antenna filiform |
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3 | Digitus absent |
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– | Digitus elongate, narrow |
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4 | Ventral cucullus with spike-like process; forewing mottled olive green to olive yellow; California |
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– | Ventral cucullus normal, rounded; forewing not olive; widespread, including California |
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5 | Forewing ground color olive green with yellow tan and black pattern; Kern and Tuolumne counties, California, and north; male valve nearly straight; female genitalia indistinguishable from |
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– | Forewing ground color light olive to olive-yellow; Kern and Tuolumne counties, California and south; male valve bent slightly ventrad at mid-point; female genitalia indistinguishable from |
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6 | Vesica with single cornutus; forewing mottled light yellow tan and gray; deserts of southern California and Arizona |
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– | Vesica with two or more cornuti; forewing not as above; widespread in western North America |
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7 | Valve slightly S-shaped; forewing mottled gray, or gray and tan |
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– | Valve nearly straight; forewing dark, blackish |
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8 | Digitus perpendicular to valve; southern California |
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– | Digitus oblique to valve, pointed ~45° toward ventral cucullus; widespread, including southern California |
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9 | Digitus origin near ventral valve with most of it below ventral valve margin; forewing dark brown to black brown; widespread in southern California |
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– | Digitus origin on mid-valve near base of ampulla, barely reaching ventral margin; immediate coast near San Diego, California |
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10 | Aedeagus longer, > 4× as wide as long; smaller narrower-winged species (wingspan ≤ 35 mm); forewing with contrasting light postreniform patch or small black basal dash |
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– | Aedeagus stout, ≤ 4× as wide as long; large broad-winged species (wingspan ≥ 35 mm); forewing without basal dash and usually without postreniform patch |
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11 | Small basal dash present, evident in all but the darkest specimens; forewing a shade of brown; hindwing base gray; widespread in western North America |
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– | Basal dash absent; forewing gray with patches of pale rusty brown; hindwing base white with streaks extending distally; deserts of Southwest and southern California |
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12 | Forewing smooth gray with pale costa; maculation reduced to dark filling of spots and dotted lines; extreme southern California |
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– | Forewing not as above, costa similar to rest of wing; West Coast and parts of Southwest, including southern California |
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13 | Forewing mottled gray brown with conspicuous gray filling of spots, small yellowish postreniform spot, and irregular black to dark gray shade proximal to subterminal line; California, Arizona, and southern Utah and Nevada |
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– | Forewing light or dark but more uniform, filling of spots not strongly contrasting, medial area between reniform spot and postmedial line not significantly lighter than rest of wing or reddish in central and northern California; California, Arizona, Oregon, and Washington |
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14 | Forewing ground color strongly red brown |
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– | Forewing not red brown, sometimes patchy reddish areas near reniform spot in central and northern California |
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15 | Forewing rusty red brown with darker markings; extreme southern California |
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– | Forewing bright red brown, occasionally with darker markings; San Benito County, California |
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16 | Southern California |
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– | Central California to Washington |
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17 | Forewing ground color slightly mottled pale gray brown; male valves oriented ≥ 120° relative to each when mounted flat; female corpus bursae ~5× as long as wide |
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– | Forewing ground color medium to dark brown; male valves oriented at ~90° when displayed similarly; female corpus bursae ~7× as long as wide |
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18 | Forewing ground color slightly mottled brown, usually with evident dark shade preceding subterminal line and often with reddish postreniform spot; male valves oriented ≥ 120° relative to each when mounted flat; female corpus bursae ~5× as long as wide |
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– | Forewing ground color blackish brown, maculation faint; male valves oriented at ~90° when displayed similarly; female corpus bursae ~7× as long as wide |
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19 | Corpus bursae elongate, 5–7× as long as wide, with small cone-shaped appendix bursae; papilla analis without long hair-like basal seta | ||
– | Corpus bursae wider, ovoid, 1.3–1.5× as long as wide, with appendix bursae not as above; papilla analis with sparse or thick hair-like basal setae |
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20 | Forewing mottled olive green to yellow green; California |
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– | Forewing ground color gray or brown; widespread, including California |
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21 | Apex of papilla analis with a thin sclerotized flange |
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– | Apex of papilla analis lacking a flange |
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22 | Hairs at base of papilla analis dense, with expanded ventral patches; Southwest deserts and southern California |
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– | Papilla analis with a sparse single row of basal hair-like setae; widespread, including southern California |
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23 | Papilla analis covered with short needle-like setae, medial dorsal margin smooth; forewing patchy light yellow tan and gray |
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– | Papilla analis rugose, scale-like, dorsal medial margin irregular; forewing gray with few light marks |
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24 | Ground color of forewing dark smoky brown to nearly black; appendix bursae bluntly rounded and mediolaterally compressed |
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– | Ground color variable, brown to gray; if dark brown then more light scaling including filling of lines and ochre postreniform patch; appendix bursae not as above, asymmetric |
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25 | Ground color of forewing a shade of brown (including gray brown), but lacking extensive gray areas |
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– | Forewing gray or gray with light tan or rusty accents |
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26 | Forewing mottled light gray, with at most minor patches of olive or tan scales |
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– | Forewing gray with extensive tan or rusty-tan scales |
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27 | Forewing mottled light gray, markings include a thin black basal dash; Pacific Coast near San Diego, California; female genitalia unknown |
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– | Forewing mottled light and medium dark gray to darker medium gray, black dash absent; widespread, including in southern California; appendix bursae broad based but short; corpus bursae lacking signa |
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28 | Forewing pattern longitudinally streaked, with rusty patches that are most prominent in fold and distal to reniform spot; bursa copulatrix with corpus bursae and appendix bursae nearly equal in size; desert Southwest and southern California |
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– | Forewing pattern mottled gray and tan or rusty tan, not streaky; appendix bursae much smaller than corpus bursae; widespread in western North America |
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Fifteen
Two species previously associated with
The name
The
All species in the
The larvae of
The
A large and broad-winged
The Latin name
The male genitalia of
The
The name
This is a large species with perhaps the broadest forewing in the group, wingspan 40.0±1.0 mm (n=12; range 39–42 mm). The forewing is smooth gray with a slightly bluish sheen when fresh, a diffuse and faint dark reniform spot, postmedial line of black dots, and a pale-cream costa in fresh specimens. The hindwing is barely a shade paler than the forewing.
This species can be identified by the combination of large size, smooth gray forewing with pale costa, and southern California distribution. It is most likely to be confused with
The name
The
This species is endemic to southern California. All records are from San Diego County from an area between Boulevard-Manzanita near the Mexican border north to Lake Henshaw at altitudes of 800–1600 m. It flies in open oak forest, foothill chaparral, and in the mountain-desert transition zone.
The name
This species is similar in size and shape to
Separating
Prior to its description in 2006, this species was thought to represent the southern California form of
The mixed red and black population from San Benito County is a unique phenomenon. The red color might be due to a gene mutation, but could also be adaptive. Many noctuids that feed on
The name
The
A medium-sized noctuid with a wingspan of 35.5±1.8 mm (n=8; range 33–39 mm) with a powdery dark olive-green forewing produced by a mixture of black, green, and yellow scales.
The relatively few specimens of
This species was described in the genus
Most
There is striking variation in this species, both within populations and over larger distances. Specimens from the coastal region of the Pacific Northwest are fairly uniform with a warm dark brown forewing with distinct maculation (Fig.
The male genitalia of
Most specimens of
This species is widespread in western North America west of south-central Alberta, Wyoming, and Nebraska. Along the Pacific Coast it occurs from northern Mexico to south-central British Columbia. It can be found from sea level to altitudes over 2000 m in a variety of habitats from dense forest to shrub desert. In the south the flight begins in March at low elevations and in April to July in the mountains. In the north the flight begins later and lasts into August. The larva is green with a white and red lateral stripe. It feeds on numerous woody shrubs, including
Despite the superficial variability of
The species name
An average or slightly smaller than average
The male genitalia are similar to those of
This species occurs in deserts of Arizona, California and Baja California, Mexico. Most specimens are from the western edge of the Colorado Desert in San Diego, Imperial, and Riverside counties, California, but there are colonies throughout the Colorado, Mojave, and Sonora deserts. Like many desert insects, the flight period depends on winter rainfall and is early, generally early March to April. The food plants and immature stages are unknown.
This below-average-sized
The male uncus is unique in that the subbasal segment is expanded and dorsoventrally flattened to an elongate rhomboid shape with a slight constriction at the end of the swollen segment. The valve is most similar to those of
Males of
This is a species of the lower mountain-desert transition zone and high desert and has been collected in the Mojave, Colorado, and Sonora deserts of southeastern California, Nevada, Arizona, and Baja California. It flies during the desert spring, between March and early May depending on winter rainfall. Its host plants and immature stages are unknown.
This is the smallest and rarest
Males of
This species is unlikely to be confused with any other
This species is known only from the south side of the sea level salt marsh estuary of the Torrey Pines State Reserve. The habitat is most likely salt marsh, although it could be coastal chaparral. The capture date of April 21 is in line with the spring flight period of most
This is a slender-winged small to medium-sized
The male valve of
No other
In their original description of
A medium-sized
The male genitalia of
The female has a papilla analis covered with short needle-like setae and sparse basal hairs. The corpus bursae is fairly short, 1.5× as long as wide, with strong signa and the appendix bursae is box shaped and laterally compressed.
The simple valve of
The type material of
This is a small dark
Males of
Superficially,
This is a narrow-winged smaller
The male valve is strap-like with a slight S-shape, with a very weak sacculus, slightly expanded cucullus with a rounded apex, rod-like straight ampulla oriented parallel to the costa, and no digitus. The vesica is similar to that of
Similarities between the male genitalia of this species and
Given the geographic variability of
In contrast to the variation in habitus of this species, the genitalia are uniform. Similarly, the variation of
The name
The structural differences of
Several similar features of
A medium-sized noctuid, wingspan 34.9±1.3 mm (n=25; range 32–37.5 mm), that resembles strongly a narrow-winged
Differences in genitalia between
The
Neighbor-joining
This species is widely distributed along the Pacific Coast from northern Mexico to southwestern British Columbia. It is found mostly west of the divides of the Sierra Nevada, Cascades, and British Columbia Coast Mountains but there are at least three colonies east of these mountains: Inyo and Mono County, California; Klamath and Lake counties, Oregon; and interior British Columbia near Lillooet.
As described above,
The name is derived from
The genitalia differ greatly from those of
The higher classification of this genus is enigmatic. Although
The ridge on the ventral male aedeagus and plate in the ventral wall of the female ductus bursae might be engaged during copulation.
A medium-sized or slightly smaller than average stout species, wingspan 31.9±1.5 mm (n = 25; range 29.5–35 mm). The forewing is granular mossy green, occasionally yellowish green, with mottled dark-gray to black and light-green pattern that obscures all but the darkest parts of the lines and spots. The darkest areas are a small patch at the base of the trailing edge of the wing, the cell and fold in the medial area, a bar on the costa preceding the subterminal line, and terminal area opposite the reniform spot. The relatively small reniform spot and round orbicular spot are filled with peripheral whitish and central green scales. The basal, antemedial and postmedial lines are double, black filled with green, and the subterminal line is green; all are sinuous and appear incomplete. The fringe is checkered green and gray. The hindwing is light brown gray with a darker border.
This species can usually be identified by superficial appearance alone. If in doubt, the male genitalia described under the genus description are diagnostic.
This revision should help clear up confusion in the genus
Although our work is based primarily on structural evidence, we also accessed a large pre-existing data base of
Neighbor-joining
We are grateful to J. Donald Lafontaine for photographs of type specimens, barcode sequence similarity trees from BOLD, invaluable advice and comments on the manuscript, to Jeremy deWaard, Ontario Biodiversity Institute for the loan of specimens, to Thomas E. Dimock for numerous specimens, and the late Ronald H. Leuschner for advice, specimens, original descriptions, and the recognition of putative new species of