VOLUME 28, NUMBER 2 107 would be assigned, we propose to discuss the relationships between ecology and taxonomy of the insect in a separate paper.

ACKNOWLEDGMENTS We would like to gratefully acknowledge aid, particularly in pinpointing E. editha population locations and flight times, from the following people: Ralph Wells, Fred Thorne, J. Tilden, William Swisher, Michael Smith, Oakley Shields, Harriet Reinhard, Paul Opler, James Mori, Andrew Moldenke, David McCorkle, Sterling Mattoon, Chris Henne, Lawrence Gilbert, Clifford Ferris, Thomas Emmel, Paul R. Ehrlich, Helen Cox, and David L. Bauer. This work was supported by NIH traineeships 000-365-06 through 000-365-11; NSF grants GB 8038, 8174, 19686, 22853, and 35259; and by a grant from the Ford Foundation.

LITERATURE CITED BASSETT, 1. J. & B. R. BAUM. 1969. Conspecificity of Plantago fastigiata of North America with P. ovata of the Old World. Can. J. Bot. 47: 1865-68. DOWNEY, J. C. & W. C. FULLER. 1961. Variation in Plebejus icarioides (Lycaenidae). 1. Food plant specificity. J. Lepid. Soc. 15: 34-42. LABINE, P. A. 1968. The population biology of the butterfly, Euphydryas editha. VIII. Oviposition and its relation to patterns of oviposition in other butterflies. Evolution 22: 799-805. MUNZ, P. A. & D. D. RECK. 1959. A California Flora. Univ. Calif. Press, Berkeley. SINGER, M. C. 1971. Evolution of food-plant preferences in the butterfly, Euphydryas editha. Evolution 25: 383-89. THORNE, F. 1970. Habitat: Euphydryas editha wrighti. J. Res. Lepid. 7: 167. WHITE, R. R. 1973. Community relationships of the butterfly, Euphydryas editha. Ph.D. Thesis, Stanford University.

THE BIOLOGY OF PAPILlO INDRA NEVADENSIS (PAPILIONIDAE) IN NEVADA

THOMAS C. EMMEL Department of Zoology, University of Florida, Gainesville, Florida 32601

AND

JOHN F. EMMEL 1117 9th Street, Santa Monica, California 90403

In the late 1960's, a new race of Papilio indra Reakirt was discovered in at least two isolated mountain ranges of the Great Basin state of Nevada. Described by Emmel & Emmel (1971), Papilio indra nevadensis is differentiated in the adult stage from all other known 108 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

BOX ELDER HUMBOLDT

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TOOELE

LANDER CHURCHILL HEY DA • MILLARD •• •• • BEAVER

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Fig. 1. Map of the known distribution of Papilio indra nevadensis Emmel & Emmel, including closely related segregates in Nevada, California, and Utah referred to in the text. The central area in Lander and Nye counties is the Toiyabe Range referred to in the text. • - closed circles = nevadensis populations in the Toiyabe Range and Toquima Range; 0 - open circle = Humboldt Range, Pershing Co., population; • - closed square = Westgard Pass, Inyo Co., California, population; 0-open square = Pine Valley Mountains, Washington Co., Utah, population; !::::. - open triangles = Grant Range and Quinn Canyon Range, Nye Co., popu­ lations; ... - closed triangles = Spring Mountains, Clark Co., population. VOLUME 28, NUMBER 2 109

indra subspecies by the character combination of a wide postmedian yellow band on the forewing and hind wing, long tails, large size and elongated wings. The purpose of the present paper is to describe the distribution, habitat, behavior, foodplants, and life history of this dis­ tinctive Nevada subspecies.

Distribution, Habitat, and Habits Papilio indra nevadensis has been found in fair numbers in a series of canyons (especially Jett, Kingston, Peavine Creek, Summit, and Twin River Canyons) along the east side of the Toiyabe Range in Nye and Lander counties, Nevada. It also occurs to the east in the Toquima Range (Nye Co.) which runs parallel to the Toiyabe Range. Peter J. Herlan of the Nevada State Museum has taken two P. indra specimens in the Humboldt Range, Pershing Co., which represent either spring brood specimens of nevadensis or a population intermediate in adult characters between typical indra and nevadensis. This mountain range is approximately 100 miles NNW of the Toiyabe Range. To the west of the Toiyabe Range, at Westgard Pass at the south end of the White Mountains in Inyo Co., California, a P. indra popu­ lation is found which appears intermediate between nevadensis and typical indra. It utilizes the same Pteryxia petraea foodplant as nevaden­ sis. Southeast of the Toiyabe Range, several P. indra segregates which show a close affinity to nevadensis have been studied. P. indra larvae were collected on Lomatium parryi (Wats.) Macbr. (Umbelliferae) in the Grant Range and Quinn Canyon Range, Nye Co., in 1969, but the resulting pupae died, so the adult phenotype of these populations is not known. The coloration of these larvae appeared closest to that of P. i. martini. Farther south, in the Spring Mountains of Clark Co., adults and immatures of P. indra have been collected which show characters of both nevadensis and martini. The adults of this popu­ lation are large with elongated wings as in nevadensis. The postmedian band of yellow spots tends to be intermediate in width between that of martini and nevadensis, and on the secondaries it tapers posteriorly as in martini. The color pattern of larvae from the Spdng Mountains appears to be closest to that of martini. Another atypical P. indra segregate is found in the Pine Valley Mountains in extreme southwestern Utah. Adults and larvae of this population seem closest to those of the Spring Mountains' populations. However, pupae from this locality are closest to those of P. i. kaibabensis. The locations of these populations are shown on the accompanying map (Fig. 1). llO JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Figs. 2-5. Habitat and foodplants of Papilio indra nevadensis: 2-3, Jett Canyon, from the east side of the Toiyabe Range, Nye County, Nevada; 4, canyon wall with scattered Pteryxia plants at Jett Canyon; 5, Pteryxia petraea at Kingston Canyon in the Toiyabe Range, Lander County, Nevada.

The typical semi-arid, lower montane habitat where P. i. nevadensis occurs is exemplified by the Jett Canyon area in the Toiyabe Range. This canyon is located on the eastern slope near the southern end of the range (Fig. 2). The entrance to the canyon is very narrow, with steep walls on both sides of the narrow, 4-wheel-drive road going up the defile. The Canyon bottom is well watered by a permanent stream (Fig. 3) . Typical vegetation within the canyon includes pinyon pine, willows (Salix exigua Nutt.), sagebrush (Artemisia tridentata Nutt. ), Prunus virginiana L. var. demissa (Nutt.) Sarg., Purshia tri­ dentata (Pursh) DC., and H olodiscus boursieri ( Carr. ) Rehd. The umbelliferous foodplants of this butterfly grow on the steep, rocky slopes and canyon walls (Figs. 4, 5). At present, we have only scant data regarding the spring brood. Based on our observations of immatures in June and July, we suspect that the spring brood flies in late May and June. The size of the sum­ mer brood is variable; in 1967, 25 adults were collected in Jett Canyon in one day in August. In 1968, no adults were seen when the area VOLUME 28, NUMBER 2 111

was visited on 10 August, while on 3 August 1969, only one adult was taken. Males were taken feeding on Cirsium species (thistles) and at mud or wet sand, and occasionally they visited blooms of Clematis vines. Females frequented Cirsium flowers and one was observed feeding on a Convolvulus (morning glory); several were flying along the canyon bottom. Other Papilio species actively flying at this time in these Toiyabe Range canyons are P. zelicaon Lucas, P. bairdii bairdii Edwards and P. b. form brucei Edwards, P. multicaudatus Kirby, and P. rutulus Lucas. Only the first species uses the same larval foodplant as P. i. nevadensis. The altitudinal span inhabited by P. i. nevadensis in the Toiyabe Range is 6200 to at least 7200 ft., with most specimens being taken between 6300 and 6800 ft. Undoubtedly hilltopping males ascend to the highest peaks of the Toiyabes, which are over 11,000 ft.

Foodplant and Life History Throughout the Toiyabe Range and in the Toquima Range in central Nevada, Papilio indra nevadensis uses Pteryxia petraea (Jones) C. & R. (Umbelliferae) as a larval host. Females have been observed to oviposit on these plants in the field, and larvae of all five instars have been found on the Pteryxia in these mountain ranges. Pteryxia petraea is also found in the Humboldt Range, Pershing County, and doubtless serves as the foodplant for the P. indra population there. Egg: Globular in shape, smooth, about 1 mm. in diameter, and creamy white; laid singly on underside of foodplant leaf. Early instars black with white and light orange or yellow markings and closely resembling those of P. i. minori and P. i. kaibabensis. Fifth-instar Larva: Length: 40-45 mm. at maturity. Head; Width of head capsule, 4.0 mm. Head capsule pattern in most examples (Fig. 10) distinct from patterns of all other P. indra subspecies. Ground color black. Inverted "V" of light orange occurs on adfrontal margins, and low inverted "U" of similar color occurs laterally. On P. i. indra, minori, kaibabensis, and pergamus, these lateral marks extend dorsally to or near to top of head capsule. In P. i. martini and fordi, they are absent. Body (Figs. 6-9); Ground color black. In more common morph, each segment with narrow transverse cream colored band with pinkish tint, of a width covering two-thirds of anterior half of segment (not all the way to anterior edge) and ending on either side at level of spiracles (thoracic segments) or well below spiracles (abdominal segments). These bands yellowish to light pink in P. i. indra Reakirt (see Emmel & Emmel, 1973), white to pinkish gray in P. i. pergamus Hy. Edwards (see Comstock, 1928, and Emmel & Emmel, 1973), white, bluish white, or pale pink in P. i. fordi Comstock & Martin (see Comstock & Martin, 1955, and Emmel & Emmel, 1973), dull pink or salmon in P. i. martini Emmel & Emmel (see Emmel & Emmel, 1968), and rich bright pink in P. i. kaibabensis Bauer (see Emmel & Emmel, 1967) and P. i. minori Cross (see Emmel & Emmel, 1964). Transverse row of six rather large, yellowish orange spots located just beyond 112 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Figs. 6-10. Larvae of Papilio indra nevadensis: 6, common morph of fifth-instar larva, dorsal aspect; 7, common morph of fifth-ins tar larva, lateral aspect; 8, dark fifth-instar larva, dorsal aspect; 9, dark fifth-instar larva, lateral aspect; 10, head capsule pattern of fifth-instar larva, frontal view. VOLUME 28, NUMBER 2 113

posterior edge of each cream colored band in dorsal, suprastigmatal, and lateral positions. Thoracic legs and prolegs black; large white patch found laterally on each proleg and at base of each of the other segments. One very dark larva taken at Kingston Canyon (Figs. 8, 9). Here, cream bands and other patches quite whitish and reduced in size and yellow-orange spots on body very reduced (dorsal rows) or absent (suprastigamatal and lateral rows) . Head capsule markings remain similar to those of lighter morpho Pupa: Length: 25-30 mm. Greatest width at wing cases: 7-9 mm. Morphologi­ cally like those of other subspecies of P. indra. General ground color light dull tan, with mottling of various darker and lighter brown marks and lines over entire surface.

DISCUSSION In the adult stage, Papilio indra nevadensis combines several of the key characteristics of P. i. pergamus (large size, elongated wings, long tails) and P. i. fordi (broad yellowish bands on the wings). In the original description (Emmel & Emmel, 1971), the new subspecies was said to be superficially closest to P. i. pergamus. Biologically, however, it differs in being double-brooded, having a generically different food­ plant, and in having major larval and pupal color-pattern differences. There appear to be populations intermediate between P. i. indra and nevadensis to the north and west of the Toiyabes and intermediate between martini and nevadensis to the south. Populations in the Pine Valley Mountains in Utah show a combination of characters of nevaden­ sis, martini, and kaibabensis. Thus the group of central Nevada popu­ lations that are placed under the name nevadensis represent a true geographical subspecies which has departed evolutionarily in both larval, pupal, and adult characters, as well as general biology, from its con­ specific relatives.

SUMMARY The distribution, habitat, behavior, life history and foodplants of Papilio indra nevadensis are described from field work in Nevada, particularly in the Toiyabe Range. The mature larva differs in head capsule pattern, body pattern and coloration from larvae of all other P. indra subspecies. The foodplant is Pteryxia petraea (Jones) C. & R.

ACKNOWLEDGMENTS The present paper is part of a continuing study of evolution in populations of the Papilio machaon complex in North America. We thank the Allyn Museum of Entomology and the Los Angeles County Museum of Natural History for travel funds from 1967 through 1969. Completion of this research was aided by National Science Foundation Grant GB-32151 as part of a study on chromosome evolution in 114 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

Lepidoptera. Peter J. RerIan of the Nevada State Museum and his wife Barbara assisted greatly during field work in Nevada. Oakley Shields and Scott Ellis assisted in field collection of material.

LITERATURE CITED COMSTOCK, J. A. 1928. Early stages of Papilio pergamus. Bull. So. Calif. Acad. Sci. 27: 82-86. --- & L. M. MARTIN. 1955. A new Papilio from California. Bull. So. Calif. Acad. Sci. 54: 142-150. EMMEL, J. F . & T. C. EMMEL. 1964. The life history of Papilio indra minori. J. Lepid. Soc. 18: 65-73. ---. 1966. A new Papilio from the Mojave Desert of California (Lepidoptera: Papilionidae). Entomol. News 77: 57-63. ---. 1968. The population biology and life history of Papilio indra martini. J. Lepid. Soc. 22: 46-52. EMMEL, T. C. & J. F. E:MMEL. 1967. The biology of Papilio indra kaibabensis in the Grand Canyon. J. Lepid. Soc. 21: 41-48. ---. 1971. A new subspecies of Papilio indra from central Nevada (Lepidoptera: Papilionidae). Pan-Pacific Entomol. 47: 220-223. ---. 1973. The Butterflies of Southern California. Los Angeles Co. Mus. Nat. Hist., and Ward Ritchie Press, Los Angeles.

FIELD OBSERVATIONS ON COLlAS ALEXANDRA EDWARDS (PIERIDAE)

SCOTT L. ELLIS 1217 4% St. N.W., Rochester, Minnesota 55901

In recent years there has been vigorous interest in all aspects of the biology of North American species of Colias. Colias alexandra Edwards, a widespread species of the western foothills and mountains has attracted increased attention. Rovanitz (1950a) described its distribution, and ( 1950b) plotted frequencies of the dimorphic females. Ae (1959) in­ duced laboratory crosses between C. alexandra and C. eurytheme Bois­ duval. Masters (1970) and Ferris (1972, 1973) examined the taxonomy of the species. John M. Burns (unpubl.) has studied the electrophoretic variation of esterase in different Colias species, including alexandra. An attempt is made here to augment this work with notes on foodplants, population structure, and behavior in Colias alexandra. Observations were made on 35 C. alexandra populations during 1971 and 1972 in Colorado, Utah, Nevada, Idaho, and Montana. Colias alexandra is widely distributed along the axis of the Rocky