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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Volume 25 Supplement 2

A BlOSYSTEMATIC STUDY OF TWO SPECIES OF CALLOPHRYS ( CALLOPHRYS) IN CALIFORNIA (LYCAENIDAE)

Glenn Alan Go re lick Citrus College, Azusa, California

Table of Contents

Introduction ..................... 3

Acknowledgements ................. k

Historical Review .................. 5

Methods and Materials ................ 6

Geographic and Ecological Distribution ........ 8

Comparative Morphology

Evaluation of Characters: Callophrys dumetorum . . 10

and Callophrys viridis .............. 12

Evaluation of Characters: Callophrys dumetorum

perplexa . . ................... 12

General Biology ................... \k

Biology of Callophrys dumetorum ........... 15

Adult Behavior.................. 15

Li fe H i story................... 17

Biology of Callophrys viridis ............ 21

Adult Behavior.................. 22

Li fe H i story................... 23

Results and Conclusions ............... 26

Literature Cited ................... 30

Appendix (Materials Studied) ............. 32

Figures and photographs ............... 36

INTRODUCTION

The genus Callophrys Bi 11 berg is represented in North America by six subgenera separable on the basis of male genitalia (Clench, 196l). These subgenera are Xamia (one species), Sandia (one species), Inoisalia (eight species), Mitoura (seven species), Cyanophrys (one species) and the nominal subgenus Callophrys (six species)J The six species included in the nominal subgenus are characterized by green on the undersides of the wings, with or without presence of white macu1 a t i ons.

Callophrys (C,) affinis (Edwards) has a light green color usually without any maculations. This species ranges from northern Utah and Wyoming to Washington and British Columbia, and according to Clench the life history is still unknown. Two subspecies C. a. affinis (Edw.) and C. a. washingtonia Clench currently are recognized, separable on the basis of scale color dorsally and ventrally (Tilden, 1963) .

A related species, C. sheridani (Carpenter), is characterized by a solid line of white on the ventral surface of the hindwings. There are three subspecies, C, s. sheridani (Carpenter), C. s. neoperplexa (Barnes & Benjamin), and C. s. newoomeri Clench, separable by the pattern and degree of development of the macular band (Tilden, 1963) - The latter two are confined mostly to Washington and Oregon away from the immediate coast, whereas the nominal subspecies occurs in the Rocky Mountain states.

A third species, C. apama (Edw.) has two subspecies, C. a. apama (Edw.) and C. a. homoperplexa Barnes & Benjamin, the latter characterized by the absence of a macular band. The nominal subspecies is characterized by the maculations on the undersides of the hindwings tinged with black rather than the dominant white seen in other species in the subgenus.

Callophrys oomstocki Henne, a recently described species, occurs in semi-arid to arid habitats from northern Inyo County in California south to the Providence Mountains of San Bernardino County. C. oomstocki appears to be closely related to C. apama, but lacks the tricolored fulvous band found on the undersides of C. apama (Henne, 19^0).

The present study concerns two species found in California, C. dumetorum (Boisduval) and C. viridis (Edwards). The former was previously considered to include two subspecies, C. d. dumetorum (Bdv.) and C. d, perplexa Barnes &

1 A recently described population from the higher elevations of the Sierra Nevada has been named C. lemberti by Tilden (1963), but not enough evidence concerning life history and distribution is known to confirm its specific status and relationships.

I n t rodu ct i on

Benjamin, occurring from British Columbia to Baja California Norte, with perplexa said to be confined to San Diego and Los Angeles counties, and adjacent portions of Baja California. The ventral sides of the hindwings of this species are a light green with a variable number of white maculations.

A third entity was recently described from the Pacific Northwest as C. d. oregonensis , based on restriction of the gray scales to the Cu2 vein on the undersides of the fore-wings as well as on comparative notes on this entity's bi-ology (Gorel i ck, 1970 ) .

The last species, C. viridis (Edw.) which also has a variable number of white maculations, is known only from San Mateo County to Mendocino County, California, along the i m -medtate coast. The deep yellow to blue-green tinge separates C. vividis superficially from C. sheridani and C. dumetorum.

Barnes & Benjamin (1923), Clench (19^) and Tilden (1963) in characterizing differences between species in the subgenus Callophrys , included morphological features such as scale color (wing fringes, forewing costa below, shade of green below, labial palpi, facial hairs, and dorsal ground color) and shape or angle of forewing and hindwing margins. After examining a large series of specimens of the two so-called species, Callophrys dumetorum (Bdv.) and Callophrys viridis (Edw.), the present author found no evidence which seemed conclusive or sufficient to warrant separation of the two as species. These characters vary considerably even within a single popu1 at i on.

Populations of the two were found to be allopatric, indicating that interbreeding probably does not occur. In the absence of evidence of interbreeding, biological characteristics such as differences in female ovipositional substrate and differences in courtship and mating are characteristics which were examined in an attempt to confirm or refute a specific level of differentiation between the two. Comparisons of larval morphology, host specificity, and development time were also used to help determine whether the two are to be considered subspecies or species.

Acknowledgements

Thanks are due to Dr. Jerry A. Powell, University of California, Berkeley, for assistance in the preparation and format of this study, as well as in the reading of the manuscript. Also helpful were Drs. Robert Ornduff and Howell Daly both of the University of California, Berkeley, in reading and criticizing the manuscript. I also wish to thank Paul A. Opler, University of California, Berkeley, for his advice concerning research techniques and choice of study sites. Special thanks also go to Oakley A. Shields, formerly of La Mesa, California, for data and observations concerning

Two species of California Callophrys

the bionomics and behavior of a southern California population of Callophrys dumetorum; John Emmel, San Francisco, California, for observations on a population of Callophrys viridis in San Francisco, and David McCorkle of Monmouth, Oregon, for life history data of C. dumetorum in Oregon and Wash i ngton .

Grateful thanks are also due to the following individuals who allowed the examination of specimens from institutional or private collections In their care: Dr. Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco; David L. Bauer, Bijou, California; Dr. Ernst S. Dornfeld, Corvallis, Oregon; Dr. Stanley G. Jewett, Jr., Portland, Oregon; Lloyd Martin, Los Angeles County Museum, California; David L. Mays, University'of Florida, Gainesville; David V. McCorkle, Monmouth, Oregon; Dr. E.J. Newcomer, Yakima, Washington; Paul A. Opler, and J. A. Powell, University of California, Berkeley; R. 0. Schuster, University of California, Davis; Dr. 0. E. Sette, Los Altos, California; Oakley Shields, formerly of La Mesa, California; Fred T. Thorne, El Cajon, California; Ronald S. Wielgus, Phoenix, Arizona.

My sincere thanks go to Perry E. Turner, Jr., Paul A. Opler and Alfred Blaker, all of the University of California, Berkeley, for photographing the adults and young stages used in the study, and to Dr. Kenneth Hagen, Division of Biological Control, Albany, California, for identification of the hymenopterous parasite reared on Callophrys viridis . John Emmel kindly contributed 28 pupae (20 C. dumetorum, 8 C. viridis) and Dr. John Downey, University of Northern Iowa, Cedar Falls, loaned several Callophrys pupal shells. Dr. R. Bacigalupi, Jepson Herbarium, University of California, Berkeley, determined several plant species mentioned in the text .

HISTORICAL REVIEW

Linnaeus, in 1758, proposed the genus Papilio which included a diversity of butterflies. A single hai rstreak characterized by the complete green undersides of the wings was named Papilio rubi. The name Callophrys was originally propounded by Billberg (1820) to include three groups of ly-caenids based on possession of three tails, two tails and one tail. In the group characterized by one tail only P. rubi was recorded. P. rubi was selected as the type of the genus Callophrys by Scudder in 1875 although the more encompassing generic name Thecla remained in general usage until 191^.'

Boisduval (185 2) described Thecla dumetorum which he believed to be a local race of the European species.

W. H. Edwards (1862) described Thecla viridis, characterized by the uniform deep green wing undersides and by white antennae.

6 Methods and Materials

Barnes & Benjamin (1923), after examining a small series of Callophrys dumetorum from the San Diego area, named the southern California race perplexa on the basis of complete or almost complete absence of white maculations on the undersides. They also believed Callophrys viridis to be conspecific with the typical northern California race of Callophrys dumetorum, rendering C. viridis a synonym of C. dumetorum.

Comstock (1927- plate 50) pictured a male and female of C. viridis from San Francisco which he labeled "c. dumetor-umtl, but c. dumetorum^ in reality, does not occur in San Franc ts co.

Clench (19^) revised the genus Callophrys and cited C. viridis as a species. San Francisco was named as the type locality and a neotype was designated.

Ziegler (1960) , in his revision of the Lycaenidae based on male and female genitalia, combined Mitoura Scudder, Sandia Clench and Ehrlich, and Callophrys Billberg under the latter, making each a subgenus.

Tilden (1963) discussed many diagnostic characters useful in separating the members of the subgenus Callophrys , emphasizing scale patterns and wing shapes.

Gorelick (1970) described Callophrys dumetorum oregon-ensis from the Pacific Northwest based on scale patterns and biological data .

METHODS AND MATERIALS

The study was divided into four stages, the first being the examination of 782 museum specimens (both species). Diagnostic characters were sought other than those having to do with scale patterns in order to separate not only C. dumetorum from C. viridis but also C. d. dumetorum from C. d. perplexa. Dissections of male and female genitalia from both species were done using the technique outlined in Ehrlich and Ehrlich (1961), these specimens being chosen from many different localities for possible variation between popu1 at i ons .

The second stage dealt with obtaining specimens (included in the study) from many 1epidopterists to clarify the geographic range of these species. Weekend trips were taken during the late spring months of 1967 to extend the range of

C. viridis .

The third dealt with the selection of several localities in the San Francisco Bay area where ecological and behavioral data for the two species could be obtained and compared during the spring of 1967. Study sites were the fo11ow i ng:

Two species of California Callophrys

Callophrys viridis:

1. San Bruno Mountains, near Daly City, San Mateo Co.

2. 9 mi. NE of Pt. Reyes lighthouse, Marin Co.

3. Del Puerto Canyon, 22 mi. W. Patterson, Stanislaus Co.

Callophrys dumetorum:

1. San Gabriel Canyon, near Azusa, Los Angeles Co.

2. Sand dunes, 1 1/2 mi. E. of Antioch, Contra Costa Co.

3. Brannan Id. State Park, Sacramento Co.

Observations of C. dumetorum were also made at Dictionary Hill (San Diego Co.) by Oakley Shields (via correspondence). Specific localities within each study site were chosen for capturing, marking and releasing adults in order to study individual behavior.

The San Bruno Mountains population was observed from February 10 to May 20, whereas Antioch and Brannan Island were visited from April k to May 30. Observations were made at the Pt. Reyes study site between April 30 and June 3, and Del Puerto Canyon was only visited twice, in late March and in mid-May. Field observations were made two or three times each week at all but the latter two localities.

Adults of both species occurred at specific hilltop localities in the San Bruno Mountains (C. viridis) and Brannan Island (c. dumetorum) . Marks were made with a black felt pen on the green scales on the wing undersides while the specimen was held within the net. The marks were made so that the specimen could be determined as to date of marking. To avoid loss of individuals immediately after marking, they were held in the net until docile and then released. Marking was done all day for one day at each of the above localities and all recaptured specimens taken throughout the study were recorded as to date and time of day taken. No secondary marking or re-releasing of the recaptured specimens was conducted.

Adults were observed in the lab on several plants including the natural host plants but no courtship, mating or oviposition occurred in the lab.

The fourth stage included the lab rearing of larvae of both species on a series of related and unrelated plants to determine relative specificities and also to note the rate of development on each. The experimental host plants used we re :

1. Lupinus arboreus Sims. (Leguminosae)

2. Vioia sp. (Leguminosae)

3. Lotus sooparius Ottley (Leguminosae)

h. Trifolium obtusiflorum Hook. (Leguminosae)

5. Cytisus monpessulanus L. (Leguminosae)

6. Eriophyllum staeohadifolium Lag. (Compositae)

Geographic and Ecological Distribution

7. Gnaphalium sp. (Compositae)

8. Achillea lanulosa Nutt. (Compositae)

9. Amsinokia intermedia F.& M. (Boraginaceae)

10. Eriogonum fasoioulatum Benth. (Po1ygonaceae)

11. Eriogonum latifolium latifolium Sm. (Polygonaceae)

Eggs of C. dumetorum were obtained at the Antioch and Brannan Island study sites by taking large numbers of Lotus sooparius buds from the field to the lab for microscopic examination. C. viridis eggs were taken from the leaves of Eriogonum latifolium latifolium at Pt. Reyes.

Glass vials were used, one individual per vial, to rear larvae of both species. Each vial was labeled according to date of egg hatching and arranged by locality in a rack for daily observations. Larval development during the 19 6 7 season was not recorded in detail, but successive instars were accurately recorded in the 1968 lab data using head capsules as an instar number indicator. One or two leaves and several flowers, were placed in each vial with the egg or larva and replaced every other day when desiccation or wilting occurred. Records were kept of the larval acceptance of each plant, especially the time taken to feed, the manner and/or duration of feeding, and of larval refusal to feed. The rate of development on each plant was used as an index for feeding success as well as host acceptance. Lab temperatures during the rearings (1967 and 1968) ranged from 72° to 76° F.

Developing larvae were examined under 45x for distinguishing characteristics and an ocular micrometer was utilized at 32x for larval head capsule measurements. Life history photographs were taken (Figures 1-14) and comparative morphology of the immatures were diagrammed (Figures 15"21B). Larvae were fixed in KAAD for 20 minutes and preserved in 95% ethyl alcohol. Many of these larvae, along with all marked and recaptured specimens have been deposited in the California Insect Survey collection at the University of California, Berkeley.

GEOGRAPHIC AND ECOLOGICAL DISTRIBUTION

Callophrys dumetorum (Bdv.) occupies a wide distribution extending from British Columbia to Baja California Norte, eastward to the eastern edge of the Sierra Nevada in California. Sierra Nevada populations show broad ecological tolerance, occurring as high as 5000'. Callophrys viridis (Edw.) is found along the immediate coast in California from the San Bruno Mountains in San Mateo County to Juan Creek, near Westport, Mendocino County (see map 1). Efforts to find this species along the coast in Oregon and the northernmost boundaries of Humboldt County failed. Neither C, dumetorum nor C. viridis occur in Lower Sonoran areas.

Two species of California Callophrys

MAP 1 " Distribution of Callophrys dumetovum and C. vividis in California.

Comparative Morphology

Flight Periods

Populations of C. dumetorum fly early in February in southern California whereas adults in San Francisco Bay area populations are not seen until mid-March. At Kusshi Creek, Yakima County, Washington, flight does not occur until May (Newcomer, 1965)- The C, viridis population studied in the San Bruno Mountains reached a peak flight period on March 7, 1967 whereas the C. viridis population found along the sand dunes at Pt. Reyes on the coast in Marin County did not reach a peak flight period until May 1k, 1967.

Host Associations

Eriogonum latifolium latifolium Sm., the observed food-plant of C. viridis (Brown and Opler, 1967) is only known from the coast of California and Oregon, whereas C. dumetorum females have been observed ovipositing on species of Syrmatium, Hosackia, Lotus and Eriogonum in southern California (Coolidge, 192*0. Newcomer (1965) lists C. dumetorum foodplants in Washington as Eriogonum heraoleoides , E. 00m-positum and E. elatum. The hostplant of a variant population found at China Flat Campground in the Sierra Nevada is probably Eriogonum wrightii Torr. ssp. trachygonum (Torr.) Stokes whereas the foodplant of the Del Puerto Canyon population, on the basis of three Callophrys eggs found on a single stand in March, 1968, is Eriogonum latifolium Sm. ssp. aurioulatum (Benth.) Stokes.

COMPARATIVE MORPHOLOGY Evaluation of characters: Callophrys dumetorum and C. viridis

Genital dissections were carried out using specimens taken in two localities for each species. No differences were discovered between the males of C, viridis and those of C. dumetorum. The male gnathos and valvae appeared to have the same length and the same degree of sc1erotization. The females of both species, too, had no structural differences. Careful examinations of sternites VIM, IX and X yielded no differences in size or shape. The signum was examined for possible differences but the amount of sc1erotization was great in worn specimens while barely present in fresh ones, evidently a function of age. Male and female genitalia of C. lemberti Tilden and c, oomstocki Henne were also examined but showed no apparent structural differences from one another. Therefore these structures do not provide the answer in determining the validity of species.

Androconial scales from the scent pouches of the males of both species were removed with a minute probe and placed on a slide for examination under lOOx. No apparent differences were seen. Labial palpi from both males and females of both species also manifested no structural or scale differences. Of the fourteen characters mentioned by Tilden (1963)

Two species of California Callophrys

for separating the members of the subgenus, ten were insufficient after examination of 150 specimens of both species. Such a character, for example, is the number of white macu-lations present on the undersides of the wings mentioned by Barnes & Benjamin (1923), Clench (19H), and Tilden (1 963) as a good diagnostic character. But according to Ford (19^5), individuals within a single population vary in the number of maculations present and do so even in the European species.

Examination of 596 specimens of C. dumetorum and 186 of C. viridis (total of 782 specimens) yielded the following characters useful in separating the two species:

1. Females of C. viridis are a uniform olive-gray on the dorsal surface, as are the males (fig. 7), varying to a mild olive-brown suffusion in some specimens, c. dumetorum females are dark gray-brown with a variable amount of yellow-brown suffusion within a single population as well as throughout the entire flight range (figs. 8, 9).

2. The color of the ventral surface of the wings appears to be a soft or dull ye 1 1ow-green, varying to olive-green in almost all California populations of C. dumetorum (figs. 10-12). C. viridis , on the other hand, has deep yellow to blue-green undersides varying only with occasional mild yellow-green individuals within

a single population as well as throughout the entire ran ge (figs. 13, 1 A) •

3. The antennae, when viewed dorsally, are black and white in C. dumetorum whereas fresh C. viridis specimens have all white-scaled antennae.

k. In most California populations of C. dumetorum, the green scales on the ventral surfaces of the forewings medial ly never exceed the Cui vein whereas C. viridis populations examined almost always have green scales reaching the Cu] or Cu2 vein of the forewings.

Characters 1 and 2 are evidently the most efficient way to separate C. viridis from C. dumetorum. The third characteristic is useful only when the specimens being compared are fresh. C. viridis antennae tend to lose the white annular scales after a few days of flight, making them appear similar to those of C. dumetorum.

Of 68 randomly selected C. dumetorum specimens (both male and female) examined from all over California, k (5.8%) had the medial green scales on the forewing undersides between the costal and radial veins, 17 (25.0%) had green scales extension restricted to the M j vein, 3 2 (h~f ,\%) had green scales reaching the Mo, 12 (17.6%) had green scales reaching the M3 and 3 (^.h%) showed green scales reaching the Cu 1 .

1 2

Comparative Morphology

On the other hand, 75 examined C. viridis specimens from Pt. Reyes and the San Bruno Mountains showed 5 (6.6%) with green scales restricted to the Mo, 68 (90.1%) with green scales reaching Cui and 2 (3«3%) with green reaching C u 2 .

Evaluation of characters: Callophrys dumetorum perplexa

Other questions raised concern the validity of Callophrys dumetorum perplexa, the atypical race found in southern California. Barnes & Benjamin (1923) recognized this race as a valid one occurring in Los Angeles and San Diego counties southward into Baja California Norte and distinguishable from typical C. dumetorum by "the complete or almost complete absence of white markings on the undersides".

Sixty two specimens of C. d. perplexa were examined from the vicinity of the type locality (San Diego), 53 of which were males. These males included 6 individuals with no white maculations on the wing undersides (11.3%), 8 with

1 (15.1%), 36 with 2 (67.8%) and 3 specimens having 3 (5.8%).

Of the 9 females examined none lacked any maculations,

2 had only 1 (22.2%), 2 had 2(22.2%), and 3 showed 3(33-/*%). The remaining 2 females had more than 3 (22.2%) (see graph 1A).

In comparison, a series of 51 males of C. dumetorum examined from Brannan Island State Park, Sacramento County, varied from none without any maculations, 3 with 1 (5-9%), 35 with 2 (68.6%), 8 with 3 (15-7%), and 5 with more than 3 (9.8%).

Females of the above population also varied considerably. Of the 21 specimens examined, 2 had no maculations (9.5%), 2 had 1 (9.5%), 8 had 2 (38.1%), 6 had 3 (28.6%), and 3 showed more than 3 (14.3%) (see graph IB).

The results of the above comparisons suggest that northern California populations are characterized by more spots but not significantly enough to warrant subspecific recognition. Because the males (and most females) of both populations compared showed that the two-spot condition was most prevalent, clinal variation was not assumed to be the case. These maculations, although variable in number, never vary in scale pattern. All are white outwardly and black inwardly.

The invasion of the brown scales on the undersides of the forewings almost to the costa, thus restricting the green scales to the costal margin area, is a character which is supposed to be typical of C. d. perplexa- No differences could be found, however, concerning the restrictions of green scales in any of the C. dumetorum examined throughout its entire range in California.

Two species of California Callophrys 13

m -+1

i a

GRAPH 1 ~ a comparison of maculation percentages between two California populations of Callophrys dumetorum: A,- Southern California population. B - Northern California population.

General Biology

The color of the scales composing the fringes of the wings was used as a character by Barnes & Benjamin (1923) separating the perplexa form from the typical form of C. dumetorum. These fringes were dark and becoming white outwardly, seen in typical C. dumetorum as well as in C. d. perplexa. This, too, is a variable trait of no significance.

Lastly, Clench (194*0 asserted that C. d. perplexa is characterized by the "presence of fulvous edging (light brown or dull yellow) on the costa of the forewings ventral-lyM. This is a characteristic also seen in the northern California specimens of C. dumetorum-

After review of 554 specimens relative to the above four characters attributed to Callophrys dumetorum perplexa, this writer concludes that they do not validate a distinct southern California subspecies because of their presence in the phenotypes of northern California populations as well. Because of the population variation factor present in California, no nomenc1atoria 1 distinction is required.

The chief enigma encountered during this study was the discovery of two populations of Callophrys (s. str.) that were not clearly either C. dumetorum or C. viridis. A female Callophrys examined from China Flats Campground, El Dorado County, appears to be C. viridis although populations representing the C. viridis phenotype are not otherwise known to occur in the Sierra Nevada. A second interesting population was found in Del Puerto Canyon in western Stanislaus County. Here individuals match the original description of C. viridis with the exception of the black annular scales present on the antennae which fit C. dumetorum. In several specimens from Del Puerto Canyon, the green scales were restricted to the area above the subcostal veins as seen in C. dumetorum. Since these characters were seen even in fresh specimens, the question must be raised as to whether C. dumetorum and C. viridis interbreed. Field work throughout the Bay Area where the two come closest together while remaining allo-patric indicate that they do not.

In summary, the failure of good evidence of interbreeding suggests the two are reproductive1y isolated species, and the failure of comparative morphology to reveal reliable criteria for separation of the two indicates other means of confirming their distinctness must be sought. Biological characteristics are therefore examined in this study.

GENERAL BIOLOGY

Most of the species in the subgenus Callophrys are single brooded, fly only during the spring and feed on a fairly wide range of hostplants. Females of the Palearctic species Callophrys rubi (L.) oviposit on rock-rose (Cistaceae), purging buckthorn (Rhamnaceae), gorse, broom, Dyer's green-weed (Leguminosae), bramble (Rosaceae, dogwood (Cornaceae)

Two species of California Callophrys

and whortleberry (Ericaceae) (Ford, 19^5). Most Nearctic species use members of the buckwheat genus Eriogonum, with C. dumetorum also using the leaves and buds of legumes such as Lotus (some genera recorded in older literature as food-plants for C, dumetorum, i.e., Lotus> Hosaokia and Syrma-tium, recently have been included in the genus Lotus (Munz, 1963)). Larvae feed on the leaves and flowers and appear to derive protection from predators due to their cryptic coloration. Pupation occurs in inconspicuous places at the base of the hostplant (Comstock, 1927).

General adult activity is influenced by weather conditions. Clouds blocking the sun, temperatures below 55° F, wind and rain prevent flight activity. During cold weather, adults lean or flatten themselves against leaves, rocks and other flat surfaces. Presumably this is a thermoregulatory mechanism through increased area for insulation (Clench, 1966). On warmer, sunny days flight activity is seen all day.

BIOLOGY OF CALLOPHRYS DUMETORUM

The biology of C. dumetorum was studied at four localities, the first being San Gabriel Canyon (near Azusa), Los Angeles County, an area characterized by many chaparral species, including Eriogonum fasoioulatum and Lotus scoparius. The second locality was Dictionary Hill, San Diego County, where observations were made by Oakley Shields in early March for several weeks. The third locality, 1 1/2 miles east of Antioch in Contra Costa County, is characterized by a river delta sand dune flora. The last locality, Brannan Island State Park in Sacramento County is also typified by a sand dune flora. Observations were made on seven days between April 8 and May 11 at the latter two localities.

Adult Behavior

Callophrys dumetorum adults, flying not more than three or four feet above the ground, alighted repeatedly on both Lotus scoparius and Eriogonum fasoioulatum in San Gabriel Canyon. At Antioch, however, females circled specific stands of Lotus and landed frequently, ignoring the numerous stands of Eriogonum latifolium ssp. present. This lack of interest in E. latifolium by C. dumetorum females was seen throughout the 1967 flight season and indicates that this plant is probably not used as a host. "Aggressive" behavior was displayed by many males, especially when another butterly species entered the area. The possibility, however, that these flights were merely an attempt to elicit female response should not be ruled out. Not infrequently two C. dumetorum males perched within one or two inches of one another and numerous "scuffle flights" or mutual response reactions (whereby males compete for a given territory while in flight) occurred, those individuals involved revolving around one another up to 20 feet in the air, returning to separate leaves

Callophrys dumetorum

on the same plant or to different plants. Such behavior has been seen in many butterly species and is discussed in de-tai1 by Shields (1968) .

At Brannan Island 10 males and 9 females were marked and released at 10:00 A.M. on April 13, but of the 10^ females and kl males taken between April 13 and May 2, only 1 marked male was recaptured.

In March 1967, Oakley Shields marked individuals of c. dumetorum for 18 days to determine whether males attempt to occupy specific territories. The markings occurred on Dictionary Hill in San Diego County, an area characterized by an abundance of Brassica nigra* Of 20 males marked, 7 (35%) were recaptured as follows: 2 after 1 day, 1 after k days, 1 after 5 days, 1 after 8 days, 1 after 9 days, and 1 after 18 days. The worn male taken after 18 days was not in excellent condition when marked, giving some indication of the longevity of individuals of this species.

Courtship behavior, observed several times at Brannan Island during the 1967 season, commenced as early as 10:15 A.M. (PST) when an individual male approached a female in flight for several seconds, "forcing" the female to alight. The female, facing the shrub, perched motionless while the male perched on a leaf behind the one occupied by the female and vigorously vibrated its wings. It then moved in a clockwise direction 180° until facing outward in the opposite direction. The entire courtship process lasted no longer than 15 seconds.

As a summary to both Shields' and the present author's observations, mating occurred on the summit in a given territory assumed by a single male. The territories assumed by males of C, dumetorum during these observations (as shown by the marking study results) seemed to shift around when many males were present.

Mating behavior was witnessed at 12:30 P.M. (PST) at Brannan Island on April 18, 1967 when one pair was observed for 30 minutes. The female faced the interior of the shrub {Lupinus arboreus) with the antennae oriented upwards whereas the male faced the opposite direction, antennae pointing downward. Later, another pair was observed in oopulo on Lupinus from 10:56 A.M. to 11:^5 A.M. In April, 1968 a single mating pair was observed displaying the above posture as late as 3:00 P.M. on Lotus. sooparius . After several observed ma-tings, lab dissections of four females (two of which had been observed mating) were carried out. The two mated females yielded two concentrically-deposited spermataphores in the mouth of the bursa copulatrix indicating that many females mate at least twice.

Oviposition was seen at 10:0 0 A.M. on May 5, 1967 when a female, perched on a Lotus bud, inserted its abdomen well into the bud and deposited a single green egg 90 seconds after

Two species of California Callophrys

ew me

insertion. After turning 180° to face the interior of the same bush, the butterfly flew to another apical bud and ovi posited another egg after only 30 seconds. Rapid wing rubbing ensued, followed by antennal vibration. Only when the antennae touched another bud or leaf surface did the femal^ move onto another part of the plant. The female inspected many buds and leaves before attempting to oviposit and fie off if a satisfactory substrate was not found. At no time did a female C. dumetovum at Brannan Island attempt to oviposit on Eriogonum Zatifolium ssp. No eggs or larvae were collected from Eriogonum fasoiculatum in San Gabriel Canyon thus failing to support Coolidge's claim that females accept the latter as a suitable host. Since E. fasoiculatum is often found in association with Lotus soopavius throughout the California coast ranges, Eriogonum should not be ruled out as a foodplant for some populations.

Life History

The eggs, approximately 0.4 mm in diameter, are ovoid, finely reticulated, green, and are deposited singly on the host plant (figs 1, 15). Eggs were collected from apical buds of Lotus sooparius at Antioch and Brannan Island and kept at room temperature. Emerging larvae were placed on experimental foodplants, and the results of these rearings are presented in Tables 1 and 2.

Morphological changes in each instar in the Brannan Island population corresponded closely to the descriptions given by Coolidge (1924). He showed Callophrys dumetovum to have four larval instars, verified by measurement of the head capsule widths of each larva examined. This technique has been found to be useful; there is apparently no overlap of head capsule width between instars in C. dumetovum and C. viridis larvae. This may be true of most lycaenids since it was also reported by Lawrence and Downey (196 7) in larvae of a plebejine lycaenid, Everes comyntas Godart.

The following measurements were made on a total of 23 C. dumetorum larvae reared and preserved during 1967 and 1968.

Head Capsule Width (mm) Body Length (mm)

I instar: (N=7) 0.15-0.25 0.92-1.54 (avg. 1.17) (avg. 0.20)

II i nstar: (N=l ) 0.40 3 .20

Ml instar: (N=4) 0.70-0.92 4.80-9.10 (avg. 7-59) (avg. 0.80)

IV instar: (N=ll) 1.23 11.8-15.5 (avg. 14.8)

(avg. 1.23)

Table 1 - Host Plant Acceptance and Development of Fourtee

HOSTPLANT Hostplant locality

Egg 1st 2nd 3rd instar instar instar

Lotus sao- Brannan Is, parius buds State Pk. and leaves

Lotus soo-

pav%us leaves

Lotus sco-

parius

leaves

IV-8 IV-13 ? ?

IV-13 IV-17 IV-20 IV-23

Lotus soo-

pavius

leaves

IV-13 IV-15 ? ?

Lotus sco-

pavius

leaves

IV-13 IV-14 IV-17 IV-23

Trifolium San Bruno obtusiflo- Mountains rum flowers

IV-20 IV-28 IV-30 V-9

Refr.

until

IV-2 5

Eriogonum	San Bruno	IV-20
latifolium	Mountains	Refr.
latifolium		until
flowers		IV-25
Eriogonum	Del Puerto	IV-2 0
fascicula-	Canyon	Refr.
tum leaves		until IV-2 5
Lupinus	Brannan	IV-2 0
avboreus	Is. State	Refr.
leaves	Park	until

IV-2£

V-1

IV-2 8 IV-30

IV-27 (Died IV-29)* IV-25

Eriophyllum San Bruno staechadi- Mountains folium buds and leaves

IV-20 IV-27

Refr. (Died

until IV-30)* IV-25

Vioia sp . leaves and flowers

IV-25

IV-29 (Died)*

Cytisus mon-

spessulanus

leaves

IV-25 IV-27 (Died)'

Gnaphalium leaves

IV-27 IV-30 (Died V-6)

Amsinckia inteYmedia> Achillea lan-ulosa leaves

IV-20 IV-28 Refr. (Died V-1.)*

V-15 ?

Callophrys dumetorum

Table 2 - Development of Callophrys dumetorum larvae on Lotus scoparius leaves (1968)1

Egg 1st 2nd 3rd 4th Color Pupa instar instar instar instar changes

IV-7 IV-7 IV-11 IV-16 IV-21 Pink IV-30

IV-26

IV-7 IV-8 IV-11 IV-16 IV-21 Pink IV-30

IV-26

IV-7 IV-8 IV-12 IV-19? IV-25 Pink V-3

IV-29

IV-7 IV-8 IV-11 IV-21 IV-25 Pink V-3

IV-29

1 Eggs collected April 7, 1968, at Brannan Island State Park, Sacramento County, California on Lotus scoparius.

Measurement of the molted head capsule widths at the end of each instar yielded the following ranges:

I instar: 0.15-0.38 mm

I I i nstar: 0.39-0.68 mm

III instar: 0 . 69-1 .10 mm

IV instar: 1.11-1.23 mm

The mature 4th instar larvae of C. dumetorum are uniform light yellow-green with subdorsal and lateral white stripes, turning pink just before pupation (figs. 2, 16). Triordinal crotchets arranged in a mesoseries are present on the prolegs (figs. 18A, B) and the crotchets (fig. 18A) are interrupted at the middle by a spatulate lobe, stated by Fracker (1915) to be typical of lycaenid larvae in general. The body, covered by numerous secondary setae, has uniformly rounded segmentation throughout the entire length. The head capsule isheavily sclerotized from the first, to the third instars but only the ventral portion of the adfrontal sutures (above antennae) and the ocellar area remain heavily sclerotized in the fourth instar(fig. 17).

Upon hatching, first instar larvae began to skeletonize terminal leaves of Lotus scoparius; the second instars did the same. The larvae moved down the stem when the upper leaves died or became completely skeletonized. Mature larvae defoliated the lower branchlets rather than skeletonizing leaves. Larval feeding in general occurred throughout the morning hours in all instars, somewhat actively in the afternoon, tapering off to almost no feeding at all by early evening. Resting larvae positioned themselves on the stem or on the tops of the leaves. Coolidge (1924) stated that the de-

Two species of California Callophrys 21

velopment time from egg to pupa in a southern Cal i fornia population was 32 days with larvae feeding on Hosackia (Lotus)* Rearing results from the Brannan Island population indicated a 30 to 35 day span on Lotus* Although feeding occurred successfully on the flowers of Trifolium obtusiflorum and Eriogonum latifolium latifolium* development from egg to pupa took as long as 39 days. Feeding on Eriogonum fasoioula-tum leaves resulted in one c. dumetorum larva reaching the pupa stage in 36 days.

Unfortunately, a search for field larvae failed and all life history work was carried out in the lab. Thus no parasitism was encountered during the course of the larval rear-fngs and only one mature larva was lost due to bacterial

disease,

The pupae (figs. 3, 20, 21B) are a uniform dark or brown-black with the surface texture of the wing pads excessively sculptured, almost forming rows, in contrast to the dorsal surface. A total of 26 were measured, with the length ranging 8.6-10.0 mm (avg. = 9.27mm).

BIOLOGY OF CALLOPHRYS VIRIDIS

Observations on C. viridis (reported as "Thecla dumetor-umn) were made in the western part of San Francisco by Williams (1910) who first determined the larval hostplant to be Eriogonum latifolium latifolium* In June, 1963 J. A. Powell (in litt*) collected c. viridis larvae in the San Bruno Mountains in flower heads of e. I. latifolium and noted that their pale color with pinkish markings blended quite well with the flowers. Adults emerged in February and March the following season. Further observations on C. viridis, emphasizing oviposition behavior, were made by Brown and Opler (1967) iti the San Bruno Mountains.

C. viridis was studied extensively during the present investigation at two localities in central California. The first is the San Bruno Mountains near Daly City, San Mateo County. This study site, located about 4 miles from the Pacific Ocean, is characterized by steep rocky slopes with dense, diverse coastal chaparral including an abundance of E. I, latifolium* No Lotus sooparius was seen here. Observations were made on 1h days between March 2 and May 12, 1967-

The second study site, nine miles northeast of the Point Reyes lighthouse, Marin County, consists of beach sand dunes and associated flora, including E. I, latifolium. Like the San Bruno Mountains, no Lotus sooparius was found. Cold weather with low clouds and fog was prevalent during the early morning hours (before 11:00 A.M. PST) but most days warmed up considerably by noon when most activity was observed. Observations were made on five days between April 30 and May 28, 1967.

Callophrys viridis

Adult Behavior

As in the case of most Callophrys species, C. viridis activity is greatly influenced by weather. Flight habits and sunning behavior are similar to those seen in C. dume-torum. Observations during the 1967 season were made with difficulty at the San Bruno Mountains study site due to wind and long periods of rain.

The courtship pattern exhibited by C. viridis is like that of C. dumetorum; a female entering a given male territory was courted by one or more males while in flight. But unlike C. dumetorum, ma 1es of C. viridis in the San Bruno Mountains (due to their lesser numbers) were able to maintain a specific territory with a minimum of shifting around. Two females and two males were marked in late March, 1967, a'nd two males were recaptured on the same plants five days later.

Courtship, mating and oviposition behavior of Callophrys viridis was observed at Pt. Reyes. The first two behavioral mechanisms were found to be similar to that observed in C. dumetorum at Brannan Island State Park. Oviposition was witnessed on May }h, 1967, at Pt. Reyes at 10:00 A.M.; a single female landed on many stands of Eriogonum, tapping several leaves on each with the antennae. Finally, after twenty minutes of flying from plant to plant, the female alighted on an Eriogonum leaf; and, after touching the underside of the leaf with the antennae, bent the abdomen underneath and deposited a single green egg near the apical end of the leaf after an estimated 15 seconds. The pilose mat on the leaf underside made it almost impossible to perceive the egg (photo h) .

Host selection by Callophrys viridis females was limited at both localities to Eriogonum latifolium . This may be due in part to the absence of suitable leguminous plants such as Lotus sooparius . Vioia sp. occurs at the San Bruno Mountains site, but was found to be unacceptable to C. viridis larvae in the lab (see Table 3).

John Emmel {in litt.) observed a female C. viridis oviposit on Lotus sooparius at 11:30 A.M. (PST) in the Sunset Heights district of San Francisco, April 27, 1968. The egg was laid within the apical bud as observed during my study of C. dumetorum females. The Sunset Heights locality is characterized by an abundance of E. latifolium latifolium^ and Emmel reared larvae taken from both plants between June 3 and June 10, 1968 at this site (west of the U.C. Medical Center). According to Emmel, the San Francisco Presidio dunes north of the Veteran's Administration Hospital support both Lotus and Eriogonum and both are accepted by C. viridis fema1es.

No Eriogonum in the Pt. Reyes vicinity was yet in bloom while the females were ovipositing. Oviposition on stands

Two species of California Callophrys 23

of E. I. latifolium which have reached the bloom stage, however, has been recorded by Brown and Opler (196 7) in the San Bruno Mountains. In this case the female walked slowly up and down the inflorescence stalk of the hostplant before depositing the egg on a leaf underside. It appears that no matter where the egg is deposited the females of both species of Callophrys display numerous inspection flights before the hostplant is chosen.

Life History

The eggs are identical in size, shape and color to those of Callophrys dumetorum but on Eriogonum are deposited singly on the undersides of leaves near the base of the plant (fig. 4). Both eggs and larvae were collected from the leaves of E. Z. latifolium at Pt. Reyes in May of 1967 and 1968 and young larvae were offered the same series of plants used to test host specificity in C. dumetorum. The results are incomplete, however, due to disease and parasitism encountered during the rearings. The larvae in many cases died before pupation. The results of the 1967 and 196 8 rearings are given in Tables 3 and 4.

Body lengths and head capsule widths were measured in 38 C. viridis larvae as follows:

Head Capsu1e W i d t h (mm) Body Length (mm)

I instar: (N= 10) 0.31-0.38 1.4-2.9 (Avg. 2.25) (Avg. 0.3^)

II instar: (N=ll) 0.40-0.62 3 . 7- 5 . 5 (Avg. 5.1) (Avg. 0.57)

III instar: (N=7) 0.69-0.99 4.9-9.5 (Avg. 7.0) (Avg. 0.83)

IV instar: (N=10) 1.15-1.30 9.8-16.0 (Avg. 12.8) (Avg. 1.25)

Measurements of the molted head capsule widths yielded the same ranges found for C. dumetorum larvae, with possible exception of the upper limit of the IV instar head capsule width.

The first instar larvae examined were light green with setation much like that described by Coolidge (1924) for C. dumetorum* Red spots appeared along the subdorsal ridges during the late third instar and remained in the mature fourth instar, corresponding to the description given by Williams (1910). Mature larvae were variable in color; some were light yellow-green with a small amount of pink varying slightly to red, whereas others, even within a single population were whitish marked predominantly with red (figs. 5, 19). Unlike the uniformly rounded segmentation in C. dumetorum, larvae of C. viridis have pronounced dorsolateral and segmental ridges. The setation, crotchet arrangement and head cap-

Table 3 - Host Plant Acceptance and Development of Fourteen Callophry

HOSTPLANT Hostplant Egg 1st 2nd locality instar instar

Lotus scoparius Brannan Is. V-28

leaves State Park

Lotus scoparius " V-28

leaves (

Lotus orassvfol%us Juan Creek

leaves Mendocino Co. . V-28 Di

Eriogonum latifolium San Bruno V-28 VI-8 latifolium leaves Mountains (Died VI-10)

Eriogonum latifolium " V-28 V-7

latifolium leaves (Died

parasitized)

Trifolium obtusi- " V-28

florum flowers " (Died: VI-3)*

Tvifolium obtusi- " V-28

florum flowers (Died: VI-3)*

Eriogonum fasoicu- Del Puerto VI-8

latum leaves Canyon (Died: VI-10)

Eviogonum fasoiou-latum leaves

Del Puerto Canyon

Eviogonum latifolium San Bruno latifolium

Eviogonum latifolium latifolium flowers

Lupinus avboveus leaves

Eviophyllum staeoh-adifolium leaves

Vioia sp. leaves; Cytisus monspessu-lanus leaves

V-21

V-2c:

V-2t

V-28 (Died: V-30)

V-28 (Died: VI-1)

V-2 8 (Died: VI-1)

Cone 1 us i ons

sule morphology are identical to those in C, dumetorum. The distance between the lateral adfrontal sutures proximal to the clypeus (widest point) were identical for mature larvae of both species (avg. 0.70 mm).

An undescribed species of Apanteles (Hymenoptera: Bra-conidae) was recovered from one second, two third and one fourth instar larvae during the 1967 reari ngs . Thompson (19^*0 recorded no parasite record for any species of Nearc-tic Callophrys , but several ichneumonid and tachinid species are known to parasitize c. rubi in Europe.

Pupae were examined from Pt. Reyes and the San Bruno Mountains and their lengths are: (N=l^) 8.2-9.7 mm (avg.

9.2 mm).

These pupae, unlike those of C. dumetorum described above, are brown with a mottled appearance owing to interspersed pale areas, particularly on the wing covers. Both species exhibited finely sculptured ridges, but these appear

less pronounced on the lighter colored pupa of C. viridis

(figs. 6, 2 1a).

RESULTS AND CONCLUSIONS

Laboratory and field study during the spring of 1967 and 1968 on Callophrys viridis and C. dumetorum produced interesting similarities between the two. Host selection (based on host availability) in the field proved to be of little use in separation of the two species, while larval characteristics provided the most valuable diagnostic information. The results of the study are summarized as follows:

1. Laboratory inspection of 63 San Diego County Callophrys dumetorum specimens showed that the four characters used most frequently to define the so-called subspecies C.

d. perplexa were found to be present in northern California populations in varying proportions. The degree of difference does not appear to warrant treatment of C. d. perplexa as a valid subspecies.

2. The percent difference in green scaling on the wing undersides between C. viridis and C. dumetorum is consistent enough to distinguish between them; ground color on the female uppersides and the quality of green color on the undersides are also useful in separating the two entities.

3. At no place in the respective distributions of the two species has interbreeding been found to occur, although allopatric San Francisco Bay area populations of both species fly during the same months.

k. The number of individuals marked and released for each species was too small to be of statistical significance, but the number of recaptures along with observations of male

Table 4 - Observations on the Development of Callophr

HOSTPLANT	Hostplant locality	Egg	1st instar		2nd instar
Eviogonum latifolium latifolium leaves	Pt. Reyes	V-10	V-ll		V-15
Eviogonum latifolium latifolium leaves	tt		V-10	(P.	V-l4 arasitized)
Eviogonum latifolium latifolium leaves	M		V-10	(P.	v-:U arasitized)
Eviogonum latifolium latifolium leaves	tl	V-10	V-lk		V-29? (Died)
Eviogonum latifolium latifolium leaves	11		V-10		v-m
Lotus scopavius leaves	Brannan Is. State Park				V-23
Lotus scopavius leaves	H				V-23

Conclus i ons

concentration points indicate that courtship behavior for both species is identical. Territories, while sustained by individual males for short times, shift around from day to day when numerical density is high, but appear to be maintained over several day periods in sparse colonies.

5. Aerial courtship and reverse male-female orientation on the host and non-hostplant substrates were seen to occur at least twice by individuals of both species.

6. Mating was observed twice during the morning hours in both s_pecies; courtship behavior and oviposition also were witnessed in morning and not commonly during afternoon. Undisturbed males and females of both species remain in cop-ulo for at least 30 minutes.

7. Females of C. dumetorum select Lotus scoparius for oviposition even when Eriogonum latifolium is present; while C. viridis females select Lotus and/or Eriogonum when both are present in the same locality. Females of both species spend much time examining and probing the substrate before egg deposition, evidently in relation to the structural quality of the host plant.

8. In the laboratory, larvae of C. dumetorum fed successfully on Lotus scoparius , Eriogonum latifolium, E. fas-oiculatum and Trifolium obtusiflorum; C. viridis larvae fed successfully on the first three but failed to accept Tri-folium.

9. Larvae of both C. dumetorum and C. viridis matured more rapidly on Lotus scoparius than on any other host.

10. Immatures provided the best morphological■characters for the separation of these two species. Most noteworthy are the characters given in Table 5.

As stated by Clench (1963), Callophrys is an interesting subgenus because of the difficulty in discriminating between the species comprising it. The results of the present study indicate that the morphology of the larvae lend strongest evidence for confirmation of species integrity in C. dumetorum and C. viridis. It appears that C. viridis is not conspecific with C. dumetorum as believed by many earlier workers, but rather a species which is ecologically a 1 lopat -ric. Females' host acceptance, especially the preference for Lotus scoparius by both species, indicates that the two are closely related. It is possible that C. viridis is a more specialized offshoot of C. dumetorum as suggested by the more restricted distribution and acceptable host substrates (as seen in the laboratory). But as Clench (1963) asserted, it is also possible to think of C. viridis as conspecific with Callophrys sheridani (Carpenter) and because relatively little has been recorded on the biology of the latter species, this hypothesis remains to be tested. Further work of a biological nature is needed to clarify confusing issues plaguing this subgenus.

Two species of California

Table 5 - Criteria used to separate the immature stages of Callophrys dumetorum and Callophrys viridis

STAGE

MATURE LARVA:

a. Subdorsal white stripes

b. Subdorsal ridges with red spots

c . Ground color

d. Subdorsal and segmental ridges

PUPA

Color

Wing pad sculpturing

CONDITION

C. dumetorum

present absent

green to yel-ow green

absent

dark brown-black

pronounced

C. viridis

absent present

yellow-green to whitish

present

light brown, mottled paler

weak

LITERATURE CITED

Barnes, Wm. and F. H. Benjamin, 1923. "Callophrys dumetorum" . Contrib. Nat. Hist. Lepid. North America, 5:64-69.

Billberg, C. J., 1820. Enumeratio insectorum in Museo Bill-berg, p . 80 .

Boisduval, Jean, 1852. Lepidopteres de California. Ann. Soc. Ent. France, 10 (2) :29 1 .

Brown, R. M. and P. A. Opler, 1967. Biological observations on Callophrys viridis (Lycaenidae). J. Lepid. Soc., 21 (2) : 1 13-1 14.

Clench, H., 1944. Notes on lycaenid butterflies: a. The genus Callophrys in North America. Bull. Mus. Comp. Zool., 94(6):217-299.

1961. In Ehrlich & Ehrlich, How to know the butterflies. Wm. C. Brown Co., Dubuque, pp. 200-211.

1963. Callophrys (Lycaenidae) from the Pacific Northwest. J. Res. Lepid., 2 (2) : 151 - 160.

1966. Behavioral thermoregulation in butterflies. Ecology, 4 7 (6) : 1022-1034.

Comstock, J. A., 1927. Butterflies of California. McBride Printing Co., Los Angeles. Plate 50, pp I68-I69.

Coolidge, K. R., 1927. Life history studies of some California Rhopalocera (Lepidoptera). "The life history of Callophrys dumetorum Boisduval". Trans. Amer. Ent. Soc. , 50(4) -.329-335.

Edwards, W. H.,1862. Descriptions of certain species of

diurnal lepidoptera found within the limits of the United States and of British America. Proc. Acad. Nat. Sci . Phi la., 14:223 .

Ehrlich, P. R. and A. H. Ehr1ich, 1961. How to know the butterflies. Wm. C. Brown Co., Dubuque.

Ford, E. B., 1945. Butterflies. N. N. Collins Co., London, pp 129, 221 .

Fracker, S. B., 1915. The classification of 1epidopterous larvae. 111. Biol. Monographs, 2 (1 ) :120-122.

Gorelick, G. A., 1970. A new subspecies of Callophrys

{Callophrys) dumetorum from Washington and Oregon (Lycaenidae). J. Res. Lepid., 7(2):99"104 ["I968"].

Henne, C., 1940. Two new species of Lepidoptera from California. Bull. So. Calif. Acad. Sci., 3 9 ( 1) :71-72 -

Inter-Society Color Council, National Bureau of Standards, Color-name chart illustrated with centroid colors, sample #2106. Circular 553 (supplement).

Lawrence, D. A. and J. C. Downey, 1967. Morphology of the

immature stages of Everes oomyntas Godart (Lycaenidae). J. Res. Lepid. , 5(2) :Gh ["1966"]

McCorkle, D. V. (contributor), 1965- In: News of the Lepi-dopterists1 Society, "Annual summary". No. 3, p 5.

Munz, P., 1963. A California Flora. U. California Press p. 8^9.

Newcomer, E. J., I96A. Butterflies of Yakima County, Washington. J. Lepid Soc, 1 8 ( ^ ) : 2 2 5

1965. In: News of the Lepidopterists' Society (contributor), "Annual summary". No 3, p ^.

Shields, 0., 1968. Hilltopping. J. Res. Lepid., 6(2):69~178.

Thompson, W. R., 19^. A catalogue of the parasites and

predators of insect pests. Imperial Parasite Service, Belleville, Ontario, Canada. Sec. 1, Pt. 5 (Lepidop-te ra) .

Tilden, J. W., 19 6 3 . An analysis of the North American species of the genus Callophrys. J. Res. Lepid., 1(A): 281-300.

Williams, F. X., 1910. Butterflies of San Francisco, Calif. Ent. News, 21(1) :36-3 7.

Ziegler, J. B., i960. Preliminary redefinition of North

American hairstreak genera. J. Lepid Soc., 14(1) 19- 2 3 -

APPENDIX Materials Studied

Callophrys viridis

Arranged alphabetically, 106 males and 80 females as follows:

CALIFORNIA. Marin Co.: Point Reyes, 2M, 3F, IV-8-51 (CD. MacNeill), 2M, 111-30-56 (W.R. Bauer), 2F, V-16-58 ("J. Powell); Point Reyes, 9 mi. NE lighthouse, 4M, IF, VI-18-66" (R.S. Wielgus), 3F, IV-30-67, 2M, 8F, V-7-67, 7M, 12F, V-14-67 Call G.A. Gorelick). Mendocino Co.: Fort Bragg, 18.6 mi. N, 2M, VI-12-67 (G.A. Gorelick); Juan Creek, betw. Rockport and Westport on Hwy.IM, 4F, VI-9-67 (G.A. Gorelick); Point Arena, 2M, IF, VI-12-67 (_G.A. Gorelick). San Francisco Co.: Lone Mountain, San Francisco, 2M, 5F, III-6-17-1910 (F.X. Williams); Presidio, San Francisco IM, IF, IV-5-26 (no collector), IM, IF, IV-18-29 (no collector), IM, IV-10-38 (L.I. Hewes); San Francisco, IM, V-28-05 (F.X. Williams), IM, IV-22-16 (K. Coolidge), 2M, IV-26-16 (K. Coolidge), 3M, IV-22-31 (A. Kusche), 2M, IF, IV-19-34, 3M, IF, IV-27-34, 6M, 1P3 v-8-34, IM, 2F, V-13-34 (all W. Hovanitz); Twin Peaks, San Francisco, IF, IV-4-60 CN. LaDue), 5M, 3F, IV-2-60 (R. Stanford}, 2M, IF, LV-13-61 (P.A. Opler). San Mateo Co.: San Bruno Mountains, 4M, 11-28-61, 4M, III-10-61, 2M, 3F, III-I7-6I (J. Powell), 2M, III-28-6I (N. LaDue), IF, V-5-61 (J. Powell), IF, III-9-62 (J. Powell), IM, 3P, 111-31-62 (J.W. Tilden), 9M, 8F, Iv-6-7-62 (J. Powell) 3M, IV-10-62 (J. Chemsak, J. Powell), 2M, 2F, 11-26-28-63 (J. Powell), IM, IV-28-63 (R.L. Langston), 2F, VI-4-63, emerged 11-24-64, 111-16-64 (reared from E. latifolium latifolium) (J. Powell, #63F8), IF, III-17-66 (A.J. Slater), 4M, 111-19-66 (P.A. Opler), 9M, IF, T.II-25-66 (P.A. Opler, IF, IV-13-66 (J. Powell), 5M, 2F, IV-16-66 (P.A. Opler), 4M, IF, IV-24-66 (R.S. Wielgus), 2M, III-2-67 (.G.A. Gorelick), 2M, III-7-67, IM, IF, III-22-67 CG.A. Gorelick).

Variant populations: El Dorado Co. : China Flat Campgrounds, 5 mi. S of Kyburz, lF,VI-5-6l (N. LaDue). Stanislaus Co. : Del Puerto Canyon, 22 mi. W of Patterson, 2M, III-5^3 TR.L. Langston, J. Powell), IM, IF, 11-22-64 (R.L. Langston, J. Powell, IM, IF, IV-6-49, in oopulo (V.M. Stern) (Adobe Creek); IF, 111-24-67 (G.A. Gorelick), IM, IV-9-67 (P.A. Opler).

Callophrys dumetorum

Arranged alphabetically, 270 males and 242 females as follows:

BAJA CALIFORNIA. Baja California Norte: Tajo Branch of Cantillas Canyon, Sierra Juarez, IM, 111-23-67 (J. Powell).

CALIFORNIA. Alameda Co.: Berkeley Hills, 1400', NE Oak-

land, IM, IV-17-62, IM, IV-15-64 (J. Powell); Oakland, NE, IM, III-9-30 (G. Held), IM, V-l-37, 2F, IV-9-38 (C.W. Anderson). Contra Costa Co.: Antioch, 4F, 111-18-57 (P.A. Opler); Antioch, 1 mi. E. IM, 111-19-60 (R. Stanford); Antioch, 1.5 mi. E, (Little Corral) IM, IV-19-60 (P.A. Opler), 6M, 2F, IV-20-67 (G.A. Gorelick; Mt. Diablo, IM, IV-9-61 (P.A. Opler). Fresno Co.: Coalinga, 23 mi. W, IF, V-6-63 (R.L. Langston); Huntington Lake, IM, V-22-36 (L. Martin). Kern Co.: Havilah, 3000r, 2 mi. N, IM, IV-28-64 (J. Powell); Walker Pass, 1 mi. W, IM, IV-26-64 CJ. Doyen); Weldon, 16 mi. S, 7M, IV-26-64 (R.L. Langs-ton j J. Powell). Lake Co. : Bear Creek, 3 mi. E, 2F, VI-24-67 (P.A. Opler); Middletown, 2500', 4 mi. WNW, IF V-15-58 (O.E. Sette); Whispering Pines, 3 mi. SE, IM, IV-4-62 (R.L. Langston). Los Angeles Co.: Altadena, IM, III-7-26 (J.A. Comstock); Azusa, 2M, IV^-1^ (C.I. Smith); Bob's Gab, nr. Littlerock, 2M, IV-2-54, IF, IV-7-55, 2M, 2F, IV-9-60 (P.A. Opler), 2M, IV-2-66 (R. Stanford); Cobal Creek Canyon, 2M, III-I9-6O (K. Hughes); Desert Springs, IM, V-5-56 (J. Powell), IF, V-7-63 (R.L. Langston); Fish Canyon, 2M, III-7-37 Cno collector); Glendale, 3M, IF, 111-26-44 (D.L. Bauer); Griffith Park, IM, IV-10-21 (J.A. Comstock), 2M, IV-14-60 (G.A. Gorelick); Mint Canyon, IM, IV-16-23 (J.A. Comstock); Pasadena, 1200', 2F, IV-15-11 (P. Grinnell, Jr.): San Gabriel Canyon, IM, III-1-25 (J.A. Comstock), 2M, 111-17-61 (K. Hughes), 2M, 3F, 111-20-67 (G.A. Gorelick); San Rafael Hills, IM, 11-13-41, 2M, III-9-41 (D.L. Bauer). Marin Co.: Alpine Lake, IM, IV-25-58, IM, IV-17-61 (J. Powell); Carson Ridge, IM, IV-16-61 (J. Powell); Lake Lagunitas, IF, V-2-59 (C.A. Toschi). Mariposa Co.: Bear Valley, 2 mi. N on Hwy 49, IM, 3F, IV-15-61 (P.A. Opler). Mendocino Co.: Fort Bragg, 27 mi. E, nr. Willits, 5M, IF, VI-9-67 {.G.A. Gorelick). Monterey Co. : Hastings Reservation, nr. Jamesburg, 3M, IF, V-3-58 [~J . Powell); Partington Canyon, IM, IF,IV-21-5I (P.A. Opler); Stone Canyon, IM, IF, IV-21-19 (E.P. VanDuzee). Napa Co. Angwin, IM, 2F, IV-19-45 (D.L. Bauer). Plumas Co.: Dixie Game Refuge, 5800!, 2M, V-17-64 (D.L. Bauer). Riverside Co.: Hemet Lake, IM, IF, IV-28-49 (CD. MacNeill); La Sierra College (campus), IF, 111-12-39 (D.L. Bauer); Lake Mathews, south shore, 2M, III-I9-67 (.J. Lane); Nightingale, IM, IV-28-49 (CD. MacNeill); Perris, 4M, III-4-53 (T.M. Blackman); Sage, 5 mi. S, 2M, IV-I6-65 (J. Powell, D. Veirs). Sacramento Co.: Brannan Island State Park, 50M, 150F, IV-7-to V-7-67 ICA. Gorelick). San Benito Co . : Bitterwater, 5 mi. S, IM III-3O-59 CC.W. O'Brien); Hollister, 24 mi. SE, IM, 3F, IV-7-62 ("O.E. Sette); Paicines, 5 mi. SW (Lime Kiln Rd.), 7M, 2F, 111-24-66 (A.J. Slater, J. Powell, R.L. Langston); Pinnacles Nat Tl . Monument , 7 mi. W of Jet., IM, IV-9-66 (P.A. Opler). San Bernardino Co.: Desert Spring, 1 mi. E, IM, 3F, IV-7-65 (R.L. Langston); Fontana, 8 mi. n (Lytle Creek), IF, IV-14-65 (C.A. Toschi); Highland, NE (City Creek), IF, IV-17-65 (G. Buckingham); Lucerne Valley, 5 mi. SW, 4M, IF, IV-14-64, 3M, IV-15-65 (R.L. Langston); Redlands, IM, 111-15-31 CCM. Dammers); Upland, IM, IV-9-57, IF, IV-28-57 (T.M. Black-man). San Diego Co.: Alpine, 4M, ,111-31-61 (R.L. Langston); Alpine, 5 mi. E, IM, IF, 111-31-61 (J. Powell); Anza-Borrego Desert State Park, 1000 Palms Canyon, IM, III-4-34 (J. Creel-

man); Banner, 3 mi. E (Two Mile Hill), 2M, IF, 111-23-41, IM, 111-22-42, IM, IF, IV-8-44, IM, 111-13-57, IM, 111-13-59, IM, 111-11-61 (F. Thorne); Banner, 7 mi. SE (Box Canyon), 2M, 111-17-53 (F. Thorne); IF, 111-23-53 (J. Powell). Descanso Ranger Station, IM, 111-31-61 (R.L. Langston); El Cajon, IM, 111-16-35 (CM. Brown); El Cajon, 2.5 mi. SE (800'), 3M, 111-29-52, IM, III-7-53, 2M, 111-15-53 (F. Thorne); Jacumba, 3 mi. W, IM, 111-25-60 (R. Stanford), 2M, 111-21-64 (F. Thorne), 3M, 111-22-66 (0. Shields); Lakeside, 2 mi. NE, 2M, 111-29-61, 2M, IF, 111-13-63 (J. Powell); La Posta Creek (Hwy. 80), 2M, IV-13-63 (P. Thorne); Mission Gorge, 9M, IF, II_27-52, 2M, III-6-52, 3M, 111-21-52, 3M, II-9-53, 3M, II-12-53, 2M, 11-14-53 (J. Powell), IF, 111-23-60 (R. Stanford); Mount Laguna Jet., 1 mi. N, 4F, 111-26-61 (R.L. Langston, E.E. Lindquist); Mount Laguna Jet., 3 mi. N (Scove Canyon), 2M, 111-27-61 (.R.L. Langston); Point Loma, IM, 11-12-34, IM, 11-12-35, IM, 11-17-35 (F. Thorne); Ramona, 9-5 mi. NNE (Black Mountain, 4000'), IM, V-9-65 (0. Shields); San Diego, IF, IH-il-33, IM, 111-18-33, IF, 11-12-35 (F. Thorne); Warner's Hot Springs, IM, IV-5-50, 2M, 111-23-51 (J. Powell). San Vicente Reservoir, IM, 111-19-49 (J. Powell); Scissor's Crossing, 1 mi. W, IM, 111-28-53 (J. Powell), 2M, IV-3-60 (K. Hughes). San Luis Obispo Co. : Atascadero, 4 mi. W, IF, V-2-62 (J. Powell); Pozo, 6 mi. NE (Black Mountain, 3300'-3600'), IM, IF, V-l-62 (C.A. Toschi, R.L. Langston); Pozo, 12 mi. NE (La Panza Camp), IM, 2F, IV-29-62 (R.L. Langston, J. Powell); Simmler, IM, 111-20-40 (J.W. Tilden, G.S. Mansfield). Santa Barbara Co.: Jalama Beach, 3M, 7F, IV-22-66 (R.L. Langston, J. Powell, A.J. Slater); Sta. Cruz Is., Prisoner's Harbor, 2F, IV-29-66 (R.L. Langston; Sta. Cruz Is., Upper Central Valley, 6F, IV-26-66 (R.L. Langston, J. Powell). Santa Clara Co.: Alum Rock Park, IM, 11-12-64 (R.L. Langston), 2M, IV-21-66 (P.A. Opler), IM, 11-30-67 (G.A. Gorelick); Capitancillos Lake, IM, 111-14-64 (P.A. Opler); New Almaden, IM, IV-17-64 (.P. A. Opler); Palo Alto, 2M> 30 (w. Hovanitz); San Jose, 3M, IF, V-6-17 (K. Coolidge). Santa Cruz Co_. : Boulder Creek, 6 mi. E, 5F, V-17-64 (P.A. Opler); Mt. Ben Lomond, 2M, V-16-65 (P.A. Opler); Santa Cruz Mtns., 2M, V-15-33, 2M, III-14-36, IV-5-36 (J.W. Tilden), IM, V-1901 (J.G. Grundel), IM, IV-6-31 (no collector). Sierra Co.: Sattley, 1 mi. W, IF, VL-17-67 (G.A. Gorelick). Solano Co.: Green Valley, IM, IV-8-53 (A. Telford). Tulare Co_. : Fairview, 9 mi. S, 3M, IF, IV-27-64 (W. Turner, J. Powell, P. Rude); Johnsondale, 2 mi. E, IM, IV-27-64 (J. Powell); Sequoia National Park, 2M, IF, IV-16-30, IM, V-16-30 (L.I. Hewes). Ventura Co.: Gorman, 5 mi. S, Hungry Valley, IM, IV-10-60 (G.I. Stage); New Cuyama, IF, 111-28-57 (R.P. Allen); Sespe Canyon, IF, IV-13-37 (G. Held). Yolo Co. : Rumsey, 2 mi. NW, 9M, 5F, IV-12-62 (J. Chemsak, J. Powell).

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Explanation of figures

Fig. 1 - Eggs of Callophrys dumetorum on Lotus scoparius bud, Brannan Is. St. Pk., Sacramento Co. (6.A. Gore 1i ck)

Fig. 2 - C. dumetorum larvae and pupa (same data as above)

Fig. 3 - C. dumetorum pupae (ventral , dorsal views) , Pope Creek, Napa Co. (J. Emmel)

Fig. k - Egg of Callophrys viridis on Eriogonum latifolium latifolium leaf underside, Pt. Reyes, Marin Co. (G.A. Gore 1 i ck)

Fig. 5 - C. viridis larvae feeding on Lotus scoparius

Fig. 6 - C. viridis pupae (ventral , dorsal views) , San Bruno Mountains, San Mateo Co. (J. Emmel)

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Explanation of figures

Fig. 7 " Callophrys viridis, dorsal view, Marin Co. (G.A. Gore 1 i ck)

Fig. 8 - C. dumetorum, dorsal view, Sacramento Co. (G. A. Gorel i ck)

Fig. 9 - C. dumetorum, same data as Fig. 8

Fig. 10 - C. dumetorum, ventral view, Sacramento Co. (G.A. Gore 1 i ck)

Fig. 11 - C. dumetorum, same data as Fig. 10

Fig. 12 - C. dumetorum, ventral view, San Diego Co. (J. Powell)

Fig. 13 - C, viridis , ventral view, Marin Co. (G.A. Gore lick)

Fig. 1*1 - C. viridis , same data as Fig. 13

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Fig. 15 - Callophrys dumetorum egg

Fig. 16 - Mature, 4th instar C. dumetorum larva with schematic dorsal view of abdominal segments VII, VI I I

Fig. 17 - Head capsule of mature C. dumetorum larva

Fig. 18A - Lateral view of 4th abdominal proleg showing crotchet arrangement

Fig. 18B - Medial view of 4th abdominal proleg

Fig. 19 - Mature, 4th instar C. viridis larva with

schematic dorsal view of abdominal segments VII, VI I I

Fig. 20 - C. dumetorum pupa, dorsal view

Fig. 21A - C. viridis wing pad sculpturing (ventral view)

Fig. 21B - C. dumetorum wing pad sculpturing (ventral v i ew)

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