1969 Journal of the Lepidopterists' Society

PHILOTES OF NORTH AMERICA: SYNONYMIC LIST AND DISTRIBUTION (LYCAENIDAE)

ROBERT L. LANGSTON

31 Windsor Ave., Berkeley, California

INTRODUCTION

49

The genus Philofes Scudder has received more than the ordinary amount of attention in the last two decades, particularly from lepidop­terists in the western United States. During this period several papers dealing with Philotes have been published, and I have amassed consider­able information by examination of museum specimens, correspondence, and exchange with numerous collectors. In the present list, this material is brought together for the genus as a whole, including the known dis­tributions-both by regions and by counties within each state where suf­ficient records are available to warrant their inclusion. As an addendum to the synonymic list, additional notes arc given on intermediate forms, questionable records, allopatry, sympatry, and type localities for each name listed.

Four new subspecies of Philotes have been described since the dos Pass os list (1964). According to my concepts, these newer subspecies, along with those described earlier, fall into "phylogenetic-geographic" sequences, which are parallel in two of the species.

In the arrangement given below species are indicated by numbers, and follow the arrangement as listed by dos Pass os (1964), with the known distributions added.

Synonyms and aberrant forms are also listed as they appear in dos Passos, with only the localities published in the original descriptions added. No new names have been described for the more recent aberra­tions.

Subspecies are indicated by letters. The subspecies of Philotes battoicles and P. enoptes are arranged in a geographical "ring" by allopatric pop­ulations in the following order:

a The nominal subspecies in the Sierra Nevada of California; b Adjacent or contiguous subspecies on the west and east slopes

of the Sierra Nevada; thence c East and north through the Great Basin; thence cl North through the Rocky Mountains; thence e West into the Pacific Northwest; thence f South along the Pacific Coast; thence g East and southeast into the Mojave and Colorado Deserts.

50 LANGSTON: Review of Phi/otes Vol. 23, no. 1

The subspecies of Philotes rita are arranged in the order given by Mattoni (1966), which also indicates a geographical "ring" and a simi­larity of wing-pattern sequence:

a The nominal subspecies in Arizona and southwestern New Mexico; thence

b Northcast into the praIrIe grassland of eastern Colorado; thence c Northwest across the Rocky Mountains into the Great Basin of

Utah; thence d West into the mountains of Nevada; thence e Southwest into the east slope areas and Mojave Desert of Cali­

fornia.

SYNONYMIC LIST AND GEOGRAPHICAL DISTRIBUTION

PHILOTES Scudder, 1876

Type: Lycaena regia Boisduval, 1869 (= Lycaena son oren sis Felder & Felder, 1865)

471 hattoides (Behr), 1867 a. h. hattoides (Behr), 1867

CALIFORNIA: Sierra Nevada; Arctic Alpine. County records: Alpine, eastern Fresno, northwestern Inyo, Mariposa, Mono, eastern Tulare, Tuolumne.

b. h. intermedia Barnes & McDunnough, 1917 CALIFORNIA: Northern ranges; Cascades; Sierra Nevada, west slopes.

Counties: Calaveras, Del Norte, EI Dorado, Nevada, Placer, western Shasta, Sierra, Siskiyou, Tehama, Trinity.

ah. malcolmi Gunder, 1927 CALIFORNIA: American River, Placer Co. (Type).

c. h. glaucon (Edwards), 1871 CALIFORNIA: Cascades, east slopes; Great Basin; Sierra Nevada, east slopes.

Counties: Inyo, Lassen, Modoc, Mono, eastern Shasta. IDAHO: Western; Great Basin.

Counties: Adams, Elmore, Owyhee. NEVADA: Western & northwestern; Sierra Nevada region.

Counties: Douglas, Elko, Humboldt, Lander, Lyon, Ormsby, Storey, Washoe.

OREGON: South-central & southeastern; East of Cascade divide. Counties: Grant, Harney, Klamath, Lake.

d. h. centralis Barnes & McDunnough, 1917 Rocky Mountain areas of ARIZONA, COLORADO, NEW MEXICO, UTAH and WYOMING.

e. h. oregonensis Barnes & McDunnough, 1916 OREGON: North-central & northeastern; Central & southern Cascades. (Limited to higher elevations in Cascades and other mountains in the southern and southeastern regions.)

Counties: Baker, Crook, Deschutes, Gilliam, Jackson, Jefferson, Klam­ath, Lake, Linn, Morrow, Wasco.

1969 Journal of the Lepidopterists' Society 51

\YASHINGTOJ\: Cascades, central & east slopes; Blue Mtns.; Northeastern. Counties: Chelan, Columbia, Kittitas, Okanogan, Stevens, Yakima.

BRITISH COLUMBIA: Southern interior; Kootenays.

f. b. bernardino Barnes & McDunnough, 1916

CALIFORNIA: Coast ranges, central Calif. to Mexican Border; Southern Sierra Nevada; Tehachapi Mtns .; Western Mojave and Colorado Deserts to Pacific Coast.

Counties : Fresno, Kern, lnyo, Los Angeles, Monterey, Orange, River­side, San Benito, San Bernardino, San Diego, San Luis Obispo, Santa Barbara, Tulare, Ventura .

BAJA CALIFORNIA NORTE: U. S. Border south to Cedros Island; Sierra San Pedro Martir.

abo baldyensis Gunder, 1925 CALIFORNIA: Camp Baldy, San Bernardino County (Type).

g. b. martini Mattoni, 1955 ARIZONA: Deserts, west to central; Desert mountains (Ajo, Hualapai Mtns., Prescott Nat. For.).

Counties: Maricopa, Mohave, Pima, Yavapai. CALIFORNIA: Mojave Desert, eastern & northern ; Desert mountains (lvan­pah, Old \Yoman, Panamint, Providence, Sheephole Mtns.).

Counties: Inyo, Kern, eastern San Bernardino.

472 enoptes (Boisduval), 1852

a. e. enoptes (Boisduval), 1852 CALIFORNIA: Northern ranges; Cascades; Sierra Nevada, east & west slopes.

Counties: Alpine, Amador, Calaveras, Eldorado, Fresno, northeast Humboldt, Inyo, Kern, Lassen, Modoc, Mono, Nevada, Placer, Plumas, Shasta, Sierra, Siskiyou, Tuolumne, Trinity, Tulare, Yuba.

NEVADA: Western; Sierra Nevada region. Counties: Douglas, Ormsby, Washoe.

b. e. ancUla Barncs & McDunnough, 1918

COLORADO: Rocky Mountains. IDAHO: Southern, southeastern. MONTANA: Southern; Bitterroot Range, Crazy Mtns. NEVADA: Eastern, northeastern. NEW MEXICO: Northern. UTAH: Northern, northeastern.

Counties: Davis, Duchesne, Juab, Salt Lake, Summit, Tooele. WYOMING: Rocky Mountains; Grand Tetons.

C. e. columbiae Mattoni, 1955 OREGON: Eastcrn; Cascadcs, north-central & east slopes.

Counties: Baker, Clackamas, Harney, Morrow, Umatilla, Wasco. WASHINGTON: Columbia River Basin; Cascades, central & east slopes.

Counties: Chelan, Kittitas, Klickitat, Okanogan, Yakima.

d. e. bayensis Langston, 1964 CALIFORNIA: North Coastal ; San Francisco Bay, North Coast Range.

Counties: Contra Costa, Humboldt, Marin, Mendocino, Solano, Sonoma.

e. e . smithi Mattoni, 1955 CALIFORNIA: Central Coastal; Seashore dunes , Santa Lucia Range.

County: coastal Monterey.

52 LAKGSTON; Review of Philotes Vol. 23, no. 1

f. e. tildeni Langston, 1964

CALIFORNIA; Central Inner Coastal; Mt. Hamilton & Diablo Ranges. Counties: western Kern, eastern Monterey, San Benito, eastern San Luis Obispo, eastern Santa Clara, western Stanislaus.

g. e. dammersi J. A. Comstock & Henne, 1933

ARIZONA: Deserts, west to central; Mountains, central (Prescott Nat. For., Tonto Basin).

Counties: Coconino, Gila, Yavapai. CALIFORNIA; Colorado & Mojave Deserts; Central & desert slopes of Laguna, San Bernardino, San Gabriel & San Jacinto Mtns.

Counties: Los Angeles, Riverside, San Bernardino, San Diego.

473 mojave Watson & W. P. Comstock, 1920 mohave Auctorum

CALIFORNIA: Colorado & Mojave Deserts; Desert slopes of San Bernar­dino, San Gabriel, San Jacinto & southern Sierra Nevada Mtns.

Counties; Inyo, Kern, Los Angeles, Riverside, San Bernardino.

474 rita (Barnes & McDunnough), 1916

a. r. rita (Barnes & McDunnough), 1916 ARIZONA: Mountains, central & southeastern.

Counties: Coconino, Cochise, Pima, Santa Cruz, Yavapai. NEW MEXICO: Southwestern.

County; Grant.

b. r. coloradensis Mattoni, 1966 COLORADO: East of Front Range, prairie grassland

Counties: Chaffee, Cheyenne, Custer, EI Paso, Fremont, Lincoln, Prowers, Saguache.

c. r. pallescens Tilden & Downey, 1955 UTAH: Northwestern, Stansbury Mtns.

County; Tooele.

d. "Phi/otes rita subspecies," Clench, 1967 NEVADA; Northeastern, Montello Wells .

County; Elko.

e. r. elvirae Mattoni, 1966 CALIFORNIA; vVestern Mojave Desert; Desert slopes of San Gabriel Mtns. & Sierra Nevada.

Counties; Inyo, Kern, Los Angeles, Mono.

475 spaldingi Barnes & McDunnough, 1917

ARIZONA; Northern; Kaibab Plateau, North Rim. COLORADO; Rocky Mtns.; San Juan Mtns.; Mesa Verde. NEW MEXICO; Northern; Northwestern; Zuni Mtns. UTAH; Central & eastern Rocky Mtns.

476 speciosa (Henry Edwards), 1876

CALIFORNIA; Western Colorado & Mojave Deserts; Southern San Joaquin Valley; West slope of Sierra Nevada.

Counties; Imperial, Inyo, Kern, Los Angeles, Mariposa, San Bernardino, San Diego, Ventura.

1969 Journal of the Lepidopterists' Society 53

477 sonorensis (Felder & Felder), 1865

CALIFORNIA: Coast Ranges, central Calif. to Mexican Border; West slopes of Sierra Nevada; Slopes adjoining wes tern Colorado and Mojave Deserts .

Counties: EI Dorado, Imperial, Los Angeles, Mariposa, Monterey, Orange, Placer, Riverside, San Benito, San Bernardino, San Diego, San Luis Obispo, Santa Barbara. Santa Clara. Stanislaus. Tuolumne. Ventura.

BAJA CALIFORNIA NORTE: U. S. Border south to vicinity of Punta Prieta; Ensenada; Sierra Juarez.

Syn. regia (Boisduval ) , 1869 CALIFORNIA: Los Angeles (Type) .

Genetic form comstocki Gunder, 1925 CALIFORNIA: Los Angeles County-several localties, San Gabriel River, Duarte '( Type).

abo sonoralba Watson & W. P. Comstock, 1920 CALIFORNIA : Los Angeles County; San Diego (Type).

DISTRIBUTION, INTERMEDIATE FORMS , HOSTS, AND TYPE LOCALITIES

Distribution.-The known distribution of the genus Philotes confirms that given in previous lists (Martin & Truxal, 1955; Mattoni, 1955). In addition, more recent data increases the known ranges of most of the species and subspecies. This is partly due to more interest in Lycaenidae in the last two decades, and many more Lepidopterists collecting at un­likely localities and at unusual times of year, even in some well-known areas.

Intermediate Forms.-Within the species that have been broken up into subspecies, by definition it may be assumed that the subspecies are allopatric to each other. However, this is not completely true. In general, each subspecies appears "typical" and different from the others when sampled well within its range or center of distribution. However, in some instances the subspecies become "atypical" on the peripheries of their ranges. Where one meets another this gives rise to intermediate forms and the geographical area can be considered a "blend zone."

Sympatl'y .-In the context of the following notes, the sympatric species are in association either on an ecological or a purely geographical basis.

Ecologically sympatl'ic refers to instanccs where flight periods of two or more species coincide and the opportunity for interspecific rna tings occurs. This is a broad definition, and even though the two or more species may be in the same general vicinity, usually one or more eco­logical barriers exist. For example: a) All of the Philotes are rela­tively weak fliers, and tend to stay close to their specific food-plant; b) The foodplants are almost always different species for the sympatric Philotes (food plants for those proven in recent literature are all different species); c) The adult flight periods may overlap only slightly; and d) The adult flight periods may not coincide at all in some seasons.

54 LA:-JGSTON: Review of Phi/otes Vol. 23, no. 1

Geographically sympatric refers to situations where the adult flight periods of two or more species occur at different times of the year. In many instances the flight periods are separated by several months. Feb.­Mar. & June-July, or May-June & Sept.-Oct. are common patterns in many of the sympatric areas. The greatest spacing in scason in sympatric species appears to be Feb.-Mar. for Philotes sonorensis as opposed to Aug.-Sept.-Oct. for three subspecies of P. enoptes. In many areas, the enoptes adults can be collected in October at the exact spot where sono­rensis was taken the previous February. The adult flight periods for most Philotes are correlated with the blooming season of the foodplants -some species and groups of Eriogonum flowering in the spring, others in mid-summer, and still others in late summer and autumn.

Foodplants.-Most Philotes species and subspecies have a narrow host plant preference. If the life history and/ or the exact host has been re­corded, it is also listed in the following notes, giving the literature source. In some of the older literature there may be general foodplants recorded, such as "buckwheat," "Eriogonum sp.," "stonecrop," or "Sedum." In most instances, these are not repeated, and the literature is not cited.

Type Localities.-The type localities of all of the species and subspecies are given. For the earlier described entities the reader is referred to Comstock & Huntington (1958-1964) where each name is listed alpha­betically giving the type locality, location of type in museums, original description, additional references and synonyms. For these I have listed the locality essentially verbatum as given by Comstock & Huntington, adding only the county where appropriate.

PHILOTES BATTOIDES (Behr)

Philotes battoides battoides is the high elevation subspecies in the Sierra Nevada of California. Some specimens from mid-elevations on the east slopes show intcgration into glaucon, whereas no tendencies toward intermedia on the west slopes have been observed. In much of its range, battoides is ecologically sympatric with P. enoptes enoptes. Type: Mineral King, Tulare Co., Calif., elev. 11,000 ft.

P. b. intermedia is primarily in the northcrn California mountains and the west slopes of the Sierra Nevada. However, there are a few speci­mens labeled with localities in Kern and Los Angeles countics that are on deposit in some museums. These are old specimens (30 to 60 years old) and although they are intermedia more recent specimens from the same areas have been bernardino which is very common in southern California (Tehachapis, Mt. Baldy, etc.). Throughout most of its range, intermedia is ecologically sympatric with the much commoner P. enop·tes enoptes. Type: "Shasta County, Calif."

1969 Journal of the Lepidopterists' Society 55

P . h. glaucon is the subspecies represented in the Great Basin, with a wide range in several states. Some specimens show integration into oregonensis at higher elevations, particularly in southeast Oregon and along the east slopes of the Cascades. In various parts of its range, glaucon is ecologically sympatric with P. enoptes enoptes and columhiae, and is usually geographically sympatric with P. rita elvime.

Foodplant: Eriogonum umbellatl1m (sens. lat. )--Calif., O. Shields; Ore., C. Crowe; Idaho, R. Langston. Type: "Nevada."

P. h. centmlis is the subspecies represented in the Rocky Mountains, and appears to show a clinal variation from north to south. Although recorded from the "Rocky Mountain States," the actual localities of speci­mens examined were so scattered that county listings would be super­fluous. If additional specimens along with more distributional data had been available, the variation in centmlis might seem more logical. In parts of its range, centralis is ecologically sympatric with P. enoptes ancilla and P. spaidingi.

Foodplant: Eriogonurn umbellatl1m (Scott et al., 1968). Type : Salida, Chaffee Co., Colo.

P. b. oregonensis is the high elevation subspecies in southern Oregon, but the elevation at which it occurs decreases with increased latitude as it ranges north through Washington into British Columbia. Some speci­mens show integration into glaucon at lower elevations, particularly in southeast Oregon and along the east slopes of the Cascades . In a con­siderable portion of its range, oTegonensis is ecologically sympatric with P. enoptes columbiae. At some of the localities for oregonensis cited by Leighton (1946), columbiae has been taken commonly. Since columbiae was described later (Mattoni, 1955 ), there is a probable mixture of the two in older collections. Type: Crater Lake, Klamath Co., Ore.

P. b. bernardino is the common spring and early summer subspecies in cismontane southern California, but ranges both north and south of this locus as noted in the synonymic list. With little variation, it occurs from the immediate coast to the tops of the higher mountains in certain counties (e.g., Los Angeles, San Diego). Although their flight periods are usually earlier, P. moiave, P. speciosa, and P. sonorcnsis are sympatric with bernardino in portions of its range. It is also geographically sym­patric with the late-season flying P. enoptes tildeni & dammeTsi, and P. rita eivirae.

Foodplants: Eriogonum fascicl1latum fasciculatum in coastal southern California; E. f. foliolosum throughout most of its range; and E. f. poli­folium & flavoviride in the deserts and east slope areas (Langston, 1965).

LANGSTON: Review of Philotes Vol. 23, no. 1

Type: Camp Baldy, San Bernardino Mts., Calif.

P. b. marlini is a desert subspecies. Although the typical insect is quite distinct, intermediates to bernardino have been taken in the western parts of its range. In the western part of its range, marlini is ecologically sym­patric with P. mojave, and it is geographically sympatric with the later­flying P. enoptes dammersi in California and Arizona, and with P. rita rita in Arizona.

Foodplant: Eriogonum fasciculatum polifolium; and the earlier known distribution are given by Mattoni (1955). Description and Type: Oat­man, Mohave Co., Arizona (Mattoni, 1955) .

PHILOTES ENOPTES (Boisduval)

Philotes enoptes enoptes occurs in California and Nevada in the Cas­cades and Sierra Nevada, from Siskiyou to Kern County. However, there are a few specimens labeled with localities from Los Angeles and River­side counties that are on deposit in the museums. These are old speci­mens (undated, or more than 40 years old-Dodge, Friday collrs.). The late-season specimens are dammersi (before its description), and early season (May-June) specimens of enoptes have not been collected in southern California in recent years. P. enoptes eno1Jtes is quite distinct from all of its subspecies throughout most of its range, although integra­tion into bayensis has b een noted in northeastern Humboldt County.

Foodplant: Adults have been associated with E. latifolium nudum & saxicola-Calif., Nev. (R. Langston). Type: "California. May in dry sections."

P . e. ancilla is the subspecies represented in the Rocky Mountains. Except for Utah, the actual localities of specimens examined from the other states were so scattercd that county listings would be superfluous. Further collecting may show a blend into columbiae along the western periphery of its range. In portions of its range, ancilla is ecologically sympatric with P. battoides centralis and P. spaldingi. In northwestern Utah it is geographically sympatric with the later flying P. rita pallescens. Type: Eureka, Juab Co. , Utah.

P. e. columbiae is the subspecies represented in the Pacific Northwest. Although there may be a blend into ancilla along the eastern parts of its range, at present it appears to be distinctly allopatrie to enoptes enoptes to the south. In many parts of its range, columbiae is ecologically sym­patric with P. battoides glaucon & oregonensis. The description and earlier known distribution are given by Mattoni (195t5). Type : Columbia River near Brewster, Okanogan Co. , Wash. (Mattoni, 1955).

1969 jou.rrUll of the Lepidopterists' Society 57

P. e. bayensis is the subspecies represented around San Francisco Bay and the North Coast Range of California. It is allopatric to all other Philotes species and subspecies, except for a possible blend into enoptes enoptes in the northeastern corner of Humboldt County.

Foodplants: Eriogonum latifolium auriculatum & nudum; detailed dis­tribution given by Langston (1964, 1965), and life history (Langston & Comstock, 1966). Type: China Camp, Marin Co., Calif. (Langston, 1964).

P. e. smithi is known only from the immediate coast of Monterey County, California, and is allopatric to all other subspecies of enoptes. It is geographically sympatric with the spring-flying P. sonorensis.

Foodplant: Eriogonum parvifolium; detailed distribution given by Langston (1964, 1965). Type: Burns Creek, State Highway 1, Monterey Co., Calif. (Mattoni, 1955).

P. e. tildeni is found along the Inner Coast Range of central California, and is allopatric to all other subspecies of enoptes. Normally a late­season flier, it is geographically sympatric with the earlier P. battoides bernardino and P. sonorensis.

Foodplants: Eriogonum latifolium in dictum & saxicola; detailed dis­tribution given by Langston (1961, 1965). Type: Del Puerto Canyon, 22 miles W of Patterson, Stanislaus Co., Calif. (Langston, 1964).

P. e. dammersi is the subspecies represented in the desert and adjacent mountain areas of southern California and Arizona. At present it appears to be completely allopatric to all other subspecies of enoptes. As a fall flier it may be ecologically sympatric with P. rita rita in Arizona. In various parts of its range it is geographically sympatric with the earlier flying P. battoides bernardino & martini, P. mojave, P. speciosa, and P. sono·rensis.

Foodplants: Eriogonum elongatum, E. wrightii trachygonum; life his­tory given by Comstock & Henne (1965). Type: Snow Creek, Riverside Co., Calif. (Comstock & Henne, 1933).

PHILOTES MOJAVE Wats. & Comst.

Philotes mo;ave is found in the Colorado and Mojave Deserts of Cali­fornia, and the bordering mountain slopes to the west. As a spring flier, it is ecologically sympatric in various parts of its range with P. battoides bernardino & martini, P. speciosa, and P. sonorensis. It is also found in the same geographical areas as its closest ally (on the basis of similarity of genitalia) P. enoptes dammersi, and with P. rita. elvirae, but these are both late-season fliers.

58 LANGSTON: Review of Philotes Vol. 23, no . 1

Foodplant: Eriogonum pusillum; detailed distribution and life history are given by Comstock (1966). Type: "Mojave Desert, California."

PHILOTES RITA (B. & McD.)

Philotes rita rita is found in Arizona and southwestern New Mexico. It is allopatric to all of its other subspecies. Mattoni (1966), speculates that its range may extend into Mexico, the lack of records being an artifact of poor collecting. As a fall flier it may be ecologically sympatric with P. enoptes dammersi, and it is geographically sympatric to the spring-flying P. battoides martini in the eastern portions of the latter's range.

Foodplant: "Close to Eriogonum wrightii"; and some of the classic localities are given by Mattoni (1966 ). Type: "So. Arizona."

P. r. coloradensis is found in eastern Colorado, and is allopatric to all other rita subspecies. The types and othcr specimens were taken in gently rolling prairie grassland, further east than any other known Philotes in North America. Mattoni (1966) speculates that this new subspecies probably ranges "east into Kansas, north into Nebraska, and south into New Mexico."

Foodplant: Eriogonum effusum; detailed distribution given by Mat­toni (1966). Type: 7 miles south of Kendrick, Lincoln Co., Colo. (Mat­toni, 1966).

P. r. pallescens is found in northwestern Utah, and is allopatric to all other rita subspecies. Although atypical populations have been found in Duchesne Co., Utah, and other areas (i .e., Nevada) the typical insect is known only from the general vicinity of the type series. It is geograph­ically sympatric with the earlier-flying P. enoptes ancilla.

Foodplant: Eriogonum sp.; detailed distribution given by Tilden & Downey (1955). Type: Little Granite Mtn., Dugway Proving Grounds, Tooele Co. , Utah (Tilden & Downey, 1955 ) .

"Philotes rita subspecies" is based on a single male specimen from Elko County, Nevada. Clench (1967) states that "In the sum of its characters it is closest to pallescens, ... " It is further stated that one "character suggests a relationship to elvirae." Although I have not seen the speci­men, it is no doubt an intermediate between the two. Further late sum­mer collecting in Nevada, will probably reveal rita populations across the state, connecting with the California colonies of elvirae in Inyo and Mono counties.

P. r. elvirae is found in the east slope and Mojave Desert areas of Cali­fornia, and is allopatric to all other named rita subspecies. In the northern

1969 Jou.rnal of the Lepidopterists' Society 59

part of its range it has occasionally been taken with p, battoides glaucon if flying late due to higher elevation or a retarded season. In the southern areas it comes almost into the range of P. enoptes dammersi, which is also a late-season flier. In various portions of its range, elvirae is geo­graphically sympatric with the earlier-flying P. enoptes enoptes, P. bat­toides bernardino, P. speciosa, and P. sonorensis. The life history is re­corded by Comstock & Henne (1967).

Foodplants: Eriogonum plumatella and detailed distribution are given by Mattoni (1966). Type: 3.5 mi. SW. of Pearblossom, Los Angeles Co., Calif. (Mattoni, 1966).

PHILOTES SPALDINGI B. & McD.

Philotes spaldingi is generally distributed in the central and southern Rocky Mountain States. Localities of specimens examined were so scat­tered that county listings would be superfluous. Morphologically, spald­ingi is distinct from all other species of Philotes, but superficially resem­bles Plebejus melissa, especially on the underside. In portions of its range, spaldingi is ecologically sympatric with P. battoides centralis and P. enoptes ancilla.

Foodplant: Eriogonum racemosum (Scott et al., 1968). Type: Provo, Utah Co., Utah.

PHILOTES SPECIOSA (Hy. Edw.)

Philotes speciosa is found most commonly in the western Colorado and Mojave Deserts of California. Morphologically, speciosa is distinct from all other species of Lycaenidae. As a spring flier, it is ecologically sym­patric in portions of its range with P. mojave, P. sonorensis and P. bat­toides bernardino, although thc latter is usually on the wing later in the season. It is geographically sympatric with the late-season fliers P. enoptes dammersi and P. rita elvirae. The ljfe history is recorded by Comstock & Dammers (1932).

Foodplants: Oxytheca perfoliata, O. trilobata & Eriogonum reniforme; and detailed distribution are given by Thorne (1961). Mature larvae have also been found on flowers of Etiogonum ?pusillum (Thorne, 1967). Type: Havilah, Kern Co., Calif.

PHILOTES SONORENSIS (F. & F.)

Philotes sonorensis is found most commonly in cismontane southern California, but ranges both north and south. Except for different num­bers of spots and their locations, plus other variability (comstocki, so­noralba, etc.), that seems to occur in varying percentages within the pop­ulations, there appears to be no other elinal or consistent variation in

60 LANGSTON: Review of Philotes Vol. 23, no. 1

this species throughout its extensive range. Morphologically, sonorensis is very distinct from all other species of Lycaenidae.

In most seasons its major adult flight is in February and March (both in the north and south), but may extend into Mayor June at higher elevations, or if the season is retarded by long periods of rain or snow. Its flight period is essentially over before the appearance of the other Philotes. Although it may overlap in time of adult Hight with P. mojave and P. speciosa, sonorensis can be considered only geographically sym­patric with P. battoides intermedia & bernardino, P. enoptes, smithi, til­deni & dammersi, and P. rita elvirae. The life history is recorded by Comstock and Coolidge (1930).

Foodplants: The larvae are known to feed on various types of stone­crop (Family Crassulaceae). In older literature it was recorded gen­erally from Sedum (e.g., Comstock, 1927; Comstock & Coolidge, 1930). In the northerly parts of its range it has been associated with Dudleya cymosa cymosa, setchellii, and minor (Langston, 1965), and in southern California with Dudleya lanceolata, and probably other sed urns and stone­crops.

Type: "Sonora, Mexico." There are no records of P. sonorensis from the state of Sonora, Mexico as known by its prescnt boundaries. Accord­ing to Brown (1967), butterflies named by the Felders all are found within the present limits of California and their true type localities must lie somewhere from Los Angeles southward into extreme northern Baja California.

ACKNOWLEDGMENTS

I am greatly appreciative for the free access to museum and institu­tional material made available for this study. The museums have been cited in my previous papers, particularly for specific records and deposi­tion of types. A high percentage of the records and specimens were sent on loan or exchange by individual collectors, which greatly enhanced the data available for this study. Rather than list the institutions and/ or home addresses of the contributors (most of which are included in the "Mem­bership List of the Lepidopterists' Society"), I wish to take this oppor­tunity to list the individuals, and add the states or areas from which most of their material originated.

R. Albright (Ore.) R. P. Allen (Calif.) P. H. Arnaud, Jr. (Nev., N. Mex.) D. L. Bauer (Ariz., Calif., Nev., Ore.,

Wash.) K. S. Brown, Jr. (Calif.) R. M. Brown (Calif.)

C. R. Crowe (Ore. ) T. W. Davies (Calif., Nev. ) E. J. Dornfield (Ore.) J. C. Downey (Calif., Nev., Utah, Wyo.) J. F. Emmel (Ariz., Calif., Colo.) T. C. Emmel (Calif.) C. Henne (Calif.)

1969 Journal of the Lepidopterists' Society

P. J. Herlan (Nev.) K. C. Hughes (Ariz., Calif.) R. J. Jae (Colo.) S. C. Jewett (Ida., Ore., Wash. , ·Wyo.) N. L. La Due (Calif.) J. Lane (Calif., Ida., Nev., Wyo.) R. L. Langston (Calif., Ida., Nev., N.

Mex., Wash.) M. Lundgren (Calif.) C. D. MacNeill (Calif., Nev.) J. W. MacSwain (Calif.) L. M. Martin (Ariz., Calif.) R. H . T. Mattoni (Ariz., Calif., Colo.) E. J. Newcomer (Brit. Col., Ore., Wash.) P. A. Opler (Calif., Nev., Ore.)

J . A. Powcll (Baja Calif., Calif., Ore. ) J. J. Renk (Calif., Colo., Wyo.) D. C . Rentz (Calif.) O. E. Sette (Ariz., Calif., Nev. ) O. A. Shields (Ariz., Calif.) A. J. Slater (Calif.) R. E. Stanford (Calif.) W. J. Steele (Calif.) O. R. Taylor, Jr. (Colo.) F. 1'. Thorne (Ariz., Calif.) J. W. Tilden (Ariz., Calif. ) S. Van Campen (Calif.) D. Veirs (Calif.) T. P. Webster, III (Calif.) R. S. Wielgus (Calif.)

61

Numerous specimens were examined which havc been deposited in museums, many of those listed above being major contributors . In addition, the museums contain specimens that were already on deposit before the inception of this study. Some of the contributors, many now deceased, were as follows: C. W. Anderson, W. J. Barnes, C. & R. Bohart, V. L. Clemence, J. A. Comstock, H. E. Cott, T. Craig, C. M. Dam­mers, E. A. Dodge, M. Doudoroff, W. A. Evans, D. Frechin, F. W. Friday, J. D. Gunder, 1'. R. Haig, G. H. IIeid, L. 1. Hewes, ] . C . Hopfinger, W. Hovanitz, A. Koebele, J. A. Kusche, C. 1. Smith, J. Strohbeen, E. P. Van Duzee, E. C. Van Dyke, F. X. Williams and W. G. Wright.

BROWN, F. M., 1967. 21 (4): 271-274.

LITERATURE CITED

Lorquin's localities "Sonora" and "Utah." J. Lepid. Soc.,

CLENCH, H. K., 1967. Further distribution records and taxonomic notes on Philotes rita (Lycaenidae). J. Lcpid. Soc., 21 (2): 141-142.

COMSTOCK, J. A., 1927. Butterflies of California. Pub!. by author, Los Angeles; 334 pp., 63 pl.

1966. Life history of Philotes mohave (Lepidoptera: Lycaeninae) . Trans. San Diego Soc. Nat. Hist., 14 (10): 133-135.

COMSTOCK, J. A., & C. COOLIDGE, 1930. The life history of Philotes sonorensis Felder. Bull. So. Calif. Acad. Sci., 29 (l): 17-21.

COMSTOCK, J. A., & C. HENNE, 1933. A new Iycaenid from Southern California. Bull. So. Calif. Acad. Sci., 32 (1): 23-26.

1965. Notes on the life history of Philotes enoptes dammersi. BulL So. Calif. Acad. Sci., 64 (3) : 153-156.

1967. Notes on the life history of Philates rita elvirae (Lepidoptera; Lycaenidae). Bull. So. Calif. Acad. Sci., 66 (2): 99-102.

COMSTOCK, W. P., & E. 1. HUNTINGTON, 1958-1964. An annotated list of the Ly­caenidae ( Lepidoptera: Rhopalocera) of the Western Hemisphere. Part I: Genera. Jour. N. Y. Ent. Soc., 66 (1958): 103-118. Part II: Species. Jour. N. Y. Ent. Soc., 67 (1959): 59-95, 163-212; 68 (1960): 49-62, 105-122, 176-186,232-240; 69 (1961): 54-58,105-118,157-176,191--200; 70 (1962): 39-46,100-118,177-179; 71 (1963): 45-57, 115-119, 189-197,262-264; 72 (1964): 62-64, 120-130, 173-192.

nos PASSOS, C. F., 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc. Mem. No . 1: v + 145 pp.

LANGSTON, R. L., 1964. Philotes of central coastal California (Lycaenidae). J. Lepid. Soc., 17 (4): 201-223 ("1963").

62 ELLIS: After apacheana Vol. 23, no. 1

1965. Distribution and hosts of five Philotes in California (Lycacnidae). J. Lepid. Soc., 19 (2): 95-102.

L~NGSTON, R. L., & J. A. COMSTOCK, 1966. Life history of Philotes enoptes bayensis (Lepidoptera: Lycaenidae). Pan-Pac. Ent., 42 (2): 102-108.

LEIGHTON, B. V., 1946. The butterflies of Washington. Univ. Wash. Pubs. in Biology, 9 (2): 47-63.

MARTIN, L. M., & F. S. TRUXAL, 1955. A list of North American Lepidoptera in the Los Angeles County Museum. Part I, Butterflies (Suborder Rhopaloeera). L. A. Co. Museum, Science Series, No. 18, Zoology, No.8: 35 pp.

MATTONI, R. H. T., 1955. Notes on the genus Philotes (Lycaenidae: Lepidoptera) 1. Descriptions of three new subspecies and a synoptic list. Bull. So. Calif. Acad. Sci. , 53 (3): 157-165 ("1954" ).

1966. Distribution and pattern variation in Philotes rita. J. Res. Lepid., 4 (2): 81-101 ("1965").

SCOTT, J. A., S. L. ELLIS, & D. EFF, 1968. New records, range extensions, and field data for Colorado butterflies and skippers. J. Lepid . Soc., 22(3): 1.59-171.

THORNE, F. T., 1961. Extensions of range and a new host plant of Philotes speciosa. Bull. So. Calif. Acad. Sci., 60 (1): 37-39.

1967. North American annual summary for 1966. Zone 1: California, Arizona, Nevada. News Lepid. Soc., 9 (3): 4-5 (15 April 1967).

TILDEN, J. W., & J. C. DOWNEY, 1955. A new species of Philotes from Utah. Bull. So. Calif. Acad. Sci., 54 (1): 25-29.

A SEARCH FOR THE TYPE LOCALITY OF SPEYERIA NOKOMIS APACHEANN

SCOTT ELLIS

Hotchkiss, Colorado

The first specimens of Speyeria nokomis apache ana (Skinner) were brought back by the Wheeler Expedition of 1871, and were collected by Ferdinand Bischoff, a naturalist attached to the Expedition as a hospital orderly. The section of the Survey containing Bischoff spent much of the field season of 1871 in Nevada and east-central California. A careful analysis of Bischoff's itinerary (Brown, 1957) during this period shows that the collector's residence at Fort Independence in the Owens Valley of California from the 18th of July to the 10th of August corresponds to the flight period of apacheana in present-day Inyo County. Since Bisch­off probably was limited in his explorations by a combination of the summer heat and the limited transportation at his disposal-walking or horseback, and the fact that Bischoff did not encounter the Round Valley colonies of apacheana further north, it has been assumed by Brown (1957)

1 This is a by-product of a field trip supported through special permission by funds from N. s. F. grant GB-2741 to F. Martin Brown for the study of Edwards's type specimens.