a Established in 1962 Edited by WILLIAM HOVANITZ Volume 5 1966 Published at 1160 W. Orange Grove., Arcadia, California, U.S.A. THJl JOURNAL ©F RISEARCHJ ©N THJl LEPIJOOFTIRA C ONT ENTS Volume 5 Number 1 March, 1966 Forelegs of Butterflies I. Introduction: Chemoreception Richard M. Fox 1 A New Species of Epinotia Hubner from British Columbia ( Olethreutidae ) T. N. Freeman 13 Studies in the Life Histories of North American Lepidoptera California Annaphila II. J. A. Comstock and C. Henne 15 New Skipper Records for Mexico H. A. Freeman 27 A Moth Sheet Noel McFarland 29 Cover Photo: Daritis ? hotvardi Hy. Edw. larvae Noel McFarland 36 Rediscovery of Annaphila casta Hy. Edw. in California (Noctuidae) John S. Buckett 37 Studies on the Nearctic Euchloe. Parts I, II Paul A. Opler 39 Butterfly Aggregations Walfried J. Reinthal 51 Volume 5 Number 2 June, 1966 Morphology of the Immature Stages of Everes Comyntas Godart Donald A. Lawrence and John C. Downey 61 A Little-Recognised Species of Heliconius Butterfly John R. G. Turner 97 Comparative Speciation In Two Butterfly Families, Pieridae and Nymphalidae Bjorn Petersen 113 The Butterfly Fauna of a Yellow Pine Forest Community Oakley Shields 127 Volume 5 Number 3 September, 1966 Laboratory Techniques for Maintaining Cultures of the Monarch Butterfly F. A. Urquhart and R. W. Stegner 129 Vital Staining of Colias philociice and C. eurytheme John M. Kolyer 137 The North American Species of the Genus Zeiraphera A. Mutuura and T. N. Freeman 153 Notes on Fades penelope (Saturniidae) Brian O. C. Gardiner 177 Remarks on The Genus Zera Evans in Mexico with a New Record H. A. Freeman 181 Studies on the Nearctic Euchloe Part 3. Complete Synonymical Treatment 185 Part 4. Type Data and Type Locality Restrictions 190 Paul A. Opler Volume 5 Number 4 December, 1966 A New Species of Oncocnemis from the Western United States ( N octuidae : Cuculliinae ) John S. Buckett and William R. Bauer 197 Speciation in the Agathymus (Megathymidae) H. A. Freeman 209 The Eggs and First Instar Larx'ae of Three California Moths John Adams Comstock 215 An Additional Food Plant Record for Papilio thoas autocles R. & J. John Adams Comstock 220 A New Species of Polia Ochsenheimer from California and Notes on Polia discalis (Grote) ( NoctuidaeiHadeninae) John S. Buckett and William R. Bauer 221 Know Your Author — Brian O. C. Gardiner 229 A Gynandromorph of Lycaena gorgon Paul A. Opler 230 The Distribution and Bionomics of Arctic-Alpine Lycaena phlaeas Subspecies in North America Oakley Shields and Johnson C, Montgomery 231 Three Western Species of Polites E. J. Newcomer 243 Know Your Author -■ Bryant Mather 248 Overcoming Difficulties With The Pupae of Euproserpinus phaeton mojave Noel McFarland 249 Speyeria cybele In Mississippi. Argynninae: (Argynnis) Bryant Mather 253 Euphyes dukesi — Additional Record Bryant Mather 254 The Little Known Moth Euxoa sculptilis (Harvey) in Arizona, with Descriptions, Illustrations, and Notes on Euxoa violaris (Grote and Robinson) (Noctuidae- Agrotiinae) J. S. Buckett 255 Discovery of A Larval Hostplant for Annaphila lithosina With Notes on the Species (Noctuidae Amphipyrinae ) John S. Buckett 262 Appendix to Distribution of Lycaena phlaeas Oakley Shields and Johnson C. Montgomery 265 Correction to “A Little-recognized Species of Helicormis Butterfly” John R. G. Turner 267 THE JOURHA.L OF ^'iSEARseHJ @N THE LEFIJPOPTER^A 6" • 76 OS J^yiSec H . Volume 5 Number 1 March, 1966 a quarterly published at 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HOVANITZ THE PURPOSE OF THE JOURNAL is to combine in one source the work in this field for the aid of students of this group of insects in a way not at present available. THE JOURNAL will attempt to publish primarily only critical and complete papers of an analytical nature, though there will be a limited section devoted to shorter papers and notes. QUALITY WORK on any aspects of research on the Lepidoptera is invited. Analytical and well illustrated works are pre- ferred, with a minimum of long description. AUTHORS ARE REQUESTED to refer to the journal as an example of the form to be used in preparing their manuscripts. Illu- strations should be of the best quality black and white, or line draw- ings and should be pre-arranged by the author to fit a reduced size of 4” X 61/2.” Footnotes should be avoided; bibliography should be as indicated. Tables should be set-up for page size as indicated. Manuscripts in good form and requiring little work by the editor will naturally appear first. Authors, who wish drawings made for them, may submit rough sketches and will be billed for the cost, which will be very negligible. 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The personal subscription rate is included in the membership to the Lepidoptera Foundation indicated below. SPECIAL SERVICE TO FOREIGN ADDRESSES: THE JOURNAL will be mailed air mail or registered at cost to the subscriber, if so desired. Second Class Postage Paid at Arcadia, Calif. Journal of Research on the Lepidoptera 5(1) :142, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 FORELEGS OF BUTTERFLIES 1. INTRODUCTION: CHEMORECEPTION. RICHARD M. FOX Carnegie Museum, Pittsburgh, Pa. 15213 This study, planned as the first of a short series on butterfly forelegs, considers some general questions raised by peculiarities of these appendages, briefly reviews evidence for chemoreception by the legs of butterflies and presents a preliminary investigation of an additional chemoreceptive function. Subsequent studies will be concerned with morphologic homologies, taxonomic correla- tions and with function. It is a pleasure to acknowledge the assistance of Albert Lloyd, John Bauer, Richard T. Satterwhite — who also prepared the drawings — and Jean Walker Fox, all of Carnegie Museum, in collecting live material for study, and of Clifford F. Thompson, graduate student at the University of Pittsburgh, in preparing serial sections for me. I am particularly grateful to Dr. Vincent G. Dethier for reading a preliminary version of this paper and for his very useful suggestions. It is also a pleasure to acknowledge the support of my research by National Science Foundation through grants GB-510 and GB-2928. SOME GENERAL CONSIDERATIONS The degree of reduction in size of the prothoracic legs and differences in reduction between the sexes have traditionally been among the principal characters ' used to define the major butterfly taxa •— the four superfamilies as presented by Fox et al. (1965), sometimes ranked as families: Hesperioidea, Papilio- noidea, Nymphaloidea and Lycaenoidea. My reasons for using superfamilies have been discussed elsewhere (Fox et al., 1965: 44-46). The primitive condition probably was one in which all three pairs of legs were of about the same size and each leg had all five tarsal subsegments and a post-tarus bearing claws and cer- tain median structures (see Fox and Fox, 1964: 71-72). Among butterflies, reduction of forelegs occurs in two different ways: 1 2 RICHARD M. FOX /. Res. Lepid. ( 1 ) miniaturization without loss of any part, and ( 2 ) miniaturi- zation along with reduction through fusion in the apparent num- ber of tarsal subsegments, plus the real or apparent loss of the post-tarus and, in a few extreme cases, reduction in proportionate size of the tibia. Reduction is minimal among Hesperioidea and Papilionoidea. The prothoracic legs of both sexes are only slightly shorter than the mesothoracic and metathoracic legs and all “normal” parts of the tarsi and post-tarsi are present. To a slight degree, this is type 1 reduction. In Lycaenoidea the females have the forelegs distinctly smaller than the mid- and hindlegs, but all five tarsal' subsegments and the post-tarsus are present. The males not only have the forelegs quite small, but the post-tarsus is absent and some or all of the tarsal subsegments are fused. Thus females exhibit type 1 reduc- tion while males have type 2 reduction. Foreleg reduction is pronounced among Nymphaloidea, with marked miniaturization in both sexes. In males (Fig. 1) the post- tarsus is never present and, as a result of fusion, the tarsus super- ficially appears to have only one subsegment. In most nympha- loid families the female foreleg (Fig. 2), despite its minute size, retains all five tarsal subsegments and the post-tarsus, though the post-tarsal claws are absent in all but a few of the most primitive species. Again, type 2 reduction is found in males, type 1 in females. Within each superfamily there is, of course, variation in leg ratios; this variation, which has useful taxanomic correlation, will be considered subsequently. Meanwhile, in order to illustrate Figs. 1 and 2. Forelegs of Vanessa atalanta, coxae not shown, drawn to the same scale. Fig. 1, male; fig. 2, female.Fm, femur. Pts, post-tarsus. PtsA, post-tarsal apodeme (tendon). Tb, tibia. Ts, tarsus, with subscript numerds denoting subsegments. 5 (1) 1966 FORELEGS 3 in a general way the comparison among the four major taxa, the legs of both sexes of a typical species of each siiperfamily were measured and the pro- and methathoracic legs expressed as per- centages of the length of the mesothoracic legs (Fig. 3). The reduction of the forelegs of butterflies raises several in- teresting questions, one of which is that of selective value. Many insects sacrifice ambulation in order to convert the fore- legs to some other purpose having a clearly positive selective ef- fect — mantids and mole crickets are among many possible ex- amples. Such cases make it evident that the meso- and methathor- acic legs suffice for ambulation; one may fairly assume that the forelegs can be altered and deprived of ambulatory function with- out thereby creating a negative selective effect. But may one properly conclude from this that the forelegs will actually be altered unless the alteration leads to a positive selective value? Certainly, there is no obvious selective advantage in the little forelegs of so many butterflies: is this a case of modification without either positive or negative selective value? Apparently neutral, nonadaptive characters have been observed in animals and discussions to this point are found in most gener- al books on evolution (e.g., Dodson, 1960: 250-251; Moody, 1953: 319, 328, 353-354). Dodson states that a character with truly neutral selective value can become established ( 1 ) through 3 Fig. 3. Ratio of length of fore- and hindlegs to midlegs of four African species selected at random; meaurements made from Tagiades flesus ( Hes- periidae), Papilio dardanus ( Papilionidae ) , Cymothoe beckeri (Nymphal- idae) and lolaus agues ( Lycaenidae ) . Males shown by solid bars, females by open bars. Lengths of midlegs taken as unity. 4 RICHARD M. FOX /. Res. Lepid. genetic drift, (2) as an incidental effect of a pleotropic gene, ( 3 ) because of linkage with a beneficial gene, or (4) as part of the phenomenon of vestigiality. I have previously commented (Fox, 1956: 24-25) that the strongly aborted forelegs of Ithomiidae and related families might become established merely because foreleg reduction is not itself detrimental to survival for these insects. But Dodson (1960: 250) makes the point that nonadaptivity is difficult to demonstrate and that many characters, which at first seemed to be selectively neutral, have proved upon closer analy- sis to have positive adaptive value. Are the little forelegs of nymphaloids really useless? Have they indeed only neutral selec- tive value or is there some positive value? It is most intriguing that the degree and type of foreleg re- duction differs between the sexes in two of the four superfamilies. Why do the tarsal subsegments and the post-tarsus persist in fe- males in spite of radical miniaturization, whereas in males with equally miniaturized forelegs, the post-tarsus and most of the tarsal subsegments are absent? Are forelegs more useful to fe- male butterflies than to males? If so, what function can be carried out by a miniaturized foreleg but requires all the segmental com- ponents and cannot be performed by a leg having the tarsal sub- segments drastically fused and reduced? SUCROSE CHEMORECEPTION Other than ambulation, the only important function ever dem- onstarted for butterfly legs is the perception of fluids — specifical- ly, water and water solutions of sugars — by means of contact chemoreceptors on the tarsi. Experiments reported by Minnich (1921, 1922a, 1922b), Weis (1930), Anderson (1932), Frings and Frings (1949, 1956), Ku- wabara (1951, 1952, 1953), and Hodgson (1958) used the ‘‘Pro- boscis reaction” to demonstrate that butterflies perceive fluids via tarsi and proboscis. Individual specimens were restrained and the proboscis and legs were put into contact with various fluids; if the proboscis was extended as for drinking, perception was considered confirmed. Most of this work was physiologically oriented and primarily concerned with finding the lowest concentration of sugar in water which would elicit the reaction. Several workers also sought a “negative” reaction to solutions of materials thought to be un- pleasant to the butterfly. In general, little effort was made to discriminate among the pairs of legs as to respective perception. 5 (1) :1~12, 1966 FORELEGS 5 Thus, sensitivity on the midtarsi was reported for papilionids, lycaenids and hesperids studied but there was no indication as to whether or not the fore- or hindtarsi were separately checked. However, Minnich (1921) and Frings and Frings (1949, 1956) obtained selective data on all three pairs of legs for two pierids and six nymphaloids. The pierids studied both showed positive reaction to solutions via the ventral surfaces of the fore- and midtarsi but no reaction via the hindtarsi. The nymphaloids showed positive reaction via the mid- and hindtarsi but negative via the foretarsi. These results suggest a taxonomic correlation of considerable interest. Reported results of experiments with butterflies along these lines are summarized in Table 1. Eltringham (1933), apparently at Minnich’s behest, prepared celloidin sections of the midtarsus of male Vanessa atalanta and reported the presence of trichoid sensilla scattered along the ven- Table 1 Perception of Solutions by Contact Chemoreception Recorded for Butterfly Tarsi Family^ genus & species . Reaction to solutions by Reference Foretarsus Mi-dtarsus Hindtarsus Papilionidae ; Papilio machaon ? + ? 4 PapiUo polyxenes ? + ? 4 Papilio philenor ? + ? 5 Papilio ajax ? + ? 2 Pieridae : Colias philodice + + _ 3 Pieris rapae + 2, 3 Danaidae : Danais plexippus - + + 2, 3 Satyridae : Lethe eurydice + + 4 Cercyonis pegala - + + 2, 4 Nymphalidae : Liminitis arthemis + + 4 Limlnitis archippus ? + ? 2 Yanessa atalanta _ + + 1, 2 Nymphalis antiopa ? + + 1 Phyciodes tharos ? + ? 2 Speyeria cybele ■+ + 4 Lycaenidae ; Lycaena thoe ? + ? 2 Atlides halesus ? + ? 5 Hesperiidae; Polites mystic ? + ■ ? 2 Epargyreus clarus ? + ? 5 positive reaction reported. negative reaction reported. ?, no observa- tion reported. References; 1, Minnich, 1921. 2, Anderson, 1932. 3, Frings & Frings, 1949. 4, Frings & Frings, 1956. 5, Hodgson, 1958. 6 RICHARD M. FOX /. Res. Lepid. tral surface. He described these structures as “extremely slender, thin-walled tubes ... set in sockets of the usual kind” and sup- plied with a trichogen cell, a tormogen cell and with “several sensory cells . . . having a distal process . . . which enters the delicate tube on the edge of its base, and taking a somewhat irregular course, extends right to the distal end, where there in a minute papilla.” The setae were found to be about 150 micra long and 6.2 micra in diameter. Grabowski and Dethier ( 1954 ) ex- amined the mid- and hindlegs of a hesperid {Eiipargijreiis dar- ns), a lycaenid (Everes comyntas), two pierids {Colias eurij- theme, Pieris rapae) and a nymphalid {Phijciodes tharos) and found that “all possess on the tarsi only one type of thin-walled hair. This is blunt-tipped and usually curved, but the wall ap- pears to be of equal thickness throughout.” Foretarsi of females were not examined. The supposition that these trichoid sensilla are the contact chemoreceptors concerned with detection of sugars and certain other solutions was confirmed by electrophysiological techniques by Morita and Takeda (1957, 1959), Morita, Doira, Takeda and Kuwabara (1957) and Takeda (1961). Note that all reports on the morphology of tarsal chemorecep- tors mentioned that these trichoid sensilla are scattered singly, and that no difference between the sexes — if, indeed, females were critically examined — was recorded. Fig. 4. Second {Ft 2 ) , third, fourth and fifth foretarsal segments of Speyeria cybele female in ventral view to show arrangement of trichoid sensilla and spines. 5(1) :1-12, 1966 FORELEGS 7 CLUMPED SENSILLA ON FEMALE FORETARSI Many years ago I noticed that the foretarsi of female ithomids bear clusters of translucent hairs on the ventral surface. More recently, while trying to find some sound morphological basis for recognizing species in Mechanitis, a genus in which male genitalia are valueless for the purpose, I made a comparative examination of the female foretarsi and noticed that these clus- tered hairs are in fact the setae of trichoid sensilla and that the clusters are always placed beneath the spine or spines borne on the preceding subsegment. Since all available information indicates that sucrose sensitivity is not vested in the foretarsi of nymphalids ,and since trichoid sensilla are known to be chemoreceptors, my curiosity was at once aroused. I then made whole mounts of the legs of several hundred butterflies representing all families and, of the Nympha- loidea, the principal subfamilies and tribes. Clustered trichoid sensilla were found on the foretarsi of all females of all groups, but such clusters were not found on foretarsi of males or on mid- or hindtarsi of any butterfly of either sex. (Details of the tech- nique for whole mounts, which is quick, easy and permanent, are given in Cummins, Miller, Smith and Fox, 1965: 151-152.) In order to verify the histological structure of these trichoid sensilla, legs were removed from freshly caught specimens of Speyeria cyhele, fixed in Allen’s modification of Bouin’s fluid, embedded in high melting point (62° ) wax, cut at 7 micra and stained for general examination with Mallory’s triple stain. On whole mounts of foretarsi of female nymphaloids, the tri- choid sensilla (Figs. 4, 5, 6) are seen as groups of blunt-tipped, Fig. 5. Three terminal foretarsal subsegments of S. cybele female photo- graphed in lateral view at lOOx from a whole mount. Fig. 6. Fourth foretarsal subsegment of same, photographed at 200x in lateral view. 8 RICHARD M. FOX J. Res. Lepid. translucent curved setae, each arising from the center of what appears to be a translucent disc or mound. These sensilla are arranged in tight clusters on the \'entral side near the proximal end of a subsegment and beneath the ends of the large, promi- nent spines projecting from the distal end of the preceding sub- segment. The number of sensilla in a cluster varies with the subsegment, the distal subsegments usually having the larger clusters; details of size, number and arrangement differ among various taxa. In Speijeria cijbele a pair of ventral spines arises at the distal margin of each of the first four foretarsal subsegments, one spine on each side of the ventral midline (Fig. 4). Beneath each spine there is a cluster of sensilla on the next distal subsegment; no such cluster is present on the first subsegment. The two clus- ters on the second subsegment each consists of four sensilla; on the third and fourth subsegments each cluster has seven sensilla; on the fifth subsegment each consists of twelve sensilla. The setae (Figs. 5, 6) are 72 to 98.4 micra long and 7.2 to 9.0 micra Fig. 7. Longitudinal section of foretarsus of S. cijbele female, cut at 7 micra, stained with Mallory’s triple stain and photographed at lOOx, show- ing spine of first subsegment and two sensilla of second segment; only basal part of setae included. Fig. 8. Same two setae and associated cells photographed at lOOOx. Fig. 9. Three setae in cross-section photographed at lOOOx, showing posi- tion of dendrite (n) against wall of lumen. Fig. 10. Longitudinal section through tip of seta photographed at lOOOx, showing terminal papilla ( p ) and part of neural denrite ( n ) . 5 (1) 1966 FORELEGS 9 j in diameter at the base. In each clump the setae appear to grade : in length, the longest being distal. The discs or mounds in which the setae are 12.0 to 14.4 micra in diameter and are placed from 4.7 to 9.5 micra apart. Examination of serial sections cut longitudinally (Figs. 7, 8) shows that each disc is a hemispherical mound set in a depression of the cuticular surface and that the whole structure is a tri- choid sensillum much as presented by Eltringham ( 1933 ) , Snod- grass (1935: 515-518) and Dethier (1963: pi. 11), with trichogen cells and, in this case, with apparently three neuron cell bodies. The setae are nearly round in cross-section, hollow, the walls about 1.75 micra thick (Fig. 9). The dendrites of the neurons (n) extend along the dorsal wall of the setal lumen to its tip, where there is a tiny terminal papilla (p) (Fig. 10). As noted previously, Minnich (1921) and Frings and Frings (1949, 1956) showed that pierids studied differ from nympha- lids studied in that sensitivity to sucrose solutions is present on the foretarsi of the former but not of the latter butterflies. In addition, therefore, to the clumped sensilla on the female fore- tarsus, pierids would be expected to have trichoid sensilla in- dividually scattered along the foretarsi of both sexes. Whole mounts of the foretarsi of Pieris rapae confirm this situation. Scattered along the entire ventral surfaces of the foretarsi of both sexes are trichoid sensilla separated from each other by at least 24 micra. Similarly arranged sensilla are found also on the midtarsi (Fig. 11) but not on the hindtarsi of both sexes. Fig. 11. Fifth midtarsal subsegment of P. rapae female photographed at 200x, showing ventral spines and scattered sensilla. Fig. 12. Fifth foretarsal subsegment of P. rapae female photographed at 200x, showing clumped sensilla and paired spines. 10 RICHARD M. FOX /. Res. Lepid. Clumps of sensilla are present only on the foretarsi of females (Fig. 12). The discs are separated by 4.8 to 9.5 micra, otherwise these sensilla are structurally similar to the scattered sensilla. The setae are 55.2 to 60.0 micra long and 4 to 6 micra in diameter at the base.The two clumps on the fifth subsegment each consist of 12 sensilla, those on the fourth of four each and those on the third of two each; there are no clumps on the first or second subsegments. The individually scattered sensilla are, as reported by Eltring- ham ( 1933 ) and others, the contact chemoreceptors apparently concerned with appreciations of solutions: confirmation has been provided by electrophysiological methods and there is correlation of structure and function. By the same token, the clumps of sensilla cannot be involved in detection of solutions, since these structures are present on the foretarsi of females shown not to possess on the foretarsi any sensitivity to fluids. A clue to an additional foretarsal function in female butter- flies was found in the work of Use (1937), who experimented with insect vision and color discrimination. She reported that ovipositing Fieris brassicae females, when placed on a green sur- face, reacted by rapidly drumming on the surface with the fore- legs, but that no such reaction was elicited when the same fe- males were placed on yellow or red surfaces. Use was content to interpret the drumming reaction as a sign that the butterfly recognized the color green, without wondering why recognition should be followed by such astonishing foreleg activity. It occurred to me that the drumming reaction on the part of an ovipositing female might be an attempt by the butterfly to carry the visual recognition of green one step farther — the con- tact chemoreceptive identification of the correct lar\^al foodplant. As a crude preliminary investigation of this line of thought, ovipositing females of Fieris rapae, Colias philodice, Fhyciodes tharos, Speyeria cyhele and Euphydryas phaeton were netted in the field. Wings were clamped together with paper clips and the butterflies were placed on various colored surfaces. When black, white or orange surfaces were used, no reaction was ob- served. When a green surface was used, a form of the drumming reaction was observed in every case. The two species of pierids performed drumming exactly as Use described it for P. brassicae, a repeated, rapid alternating movement of the forelegs against the surface. In the three nymphalids the reaction consisted of a single darting motion, first with one, then with the other foreleg. 5 (1) -.1-12, 1966 FORELEGS 11 In all cases the terminal tarsal subsegments were brought into contact with and scraped along the surface at the end of the stroke. Each female was then placed upon various green leaves, in- cluding the leaf of her foodplant. Each new offering was always drummed. Three females belonging to two species followed what apparently was the butterfly’s recognition of her foodplant by oviposition: two different females of P. tharos laid eggs on leaves of aster but did not do so on other leaves offered; one female of C. philodice oviposited on a clover leaf but not on any other offering. My theory -- certainly not proved at this time — is that the female butterfly, by drumming, abrades with the tarsal spines the surface of the leaf, releasing essential oils and enabling her to detect via the clumped trichoid sensilla the material character- istic of the plant. It seems probable that, during my rough ob- servations, the females who did not follow drumming on the larval foodplant by ovipositing were not ready to do so for other reasons or were inhibted by having the wings restrained. In any event, it does not necessarily follow that just because a fe- male comes into contact with and recognizes her foodplant that she will immediately and invariably oviposit. On the other hand, the instances of ovipositing observed may have been mere eo- incidence and may have had nothing to do with drumming. If, however, the above explanation of drumming turns out to be correct, the clumps of trichoid sensilla on the female foretarsi are also explained and the function which can be performed by miniaturized foreleg only if it is complete and unfused has been found. Experiments are currently under way in our laboratory and are expected to shed additional light on these questions. REFERENCES CITED ANDERSON, A. L. 1932. The sensitivity of the legs of common butter- flies to sugars. Jour. Exp. Zool, 63: 235-239. CUMMINS, K. W., L. D. MILLER, N. A. SMITH & R. M. FOX. 1965. Experimental Entomology. Reinhold Corp., New York, viii -f 176 pp. DETHIER, V. G. 1963. The physiology of Insect Senses. Methuen, Lon- don and Wiley, New York, ix -j- 266 pp. DODSON, E. O. 1960. Evolution: Process and Product. Reinhold Corp., New York, xvi -f 352 pp. ELTRINGHAM, H. 1963. On the tarsal sense organs of lepidoptera. Trans. Ent. Soc. London. 81: 33-36. 12 RICHARD M. FOX /. Res. Lejrid. FOX, R. M. 1956. A monograph of the Itliomiidae, Part I. Bull. Ameri- can Mus. Nat. Hist., Ill: 1-76. FOX, R. M. & J. W. FOX. 1964. Introduction to Comparative Entomol- ogy. Reinhold Corp., New York, xiv + 450 pp. FOX, R. M., A. W. LINDSEY, H. K. CLENCH & L. D. MILLER. 1965. The butterflies of Liberia. American Ent. Soc., Mem. 19: i-ii, 1-438. FRINGS, H. & M. FRINGS. 1949. The loci of contact chemoreceptors in insects. Amer. Mid. Nat., 41: 602-658. FRINGS, H. & M. FRINGS. 1956. The loci of contact chemoreceptors involved in feeding reactions of certain Lepidoptera. Biol. Bull., 110: 291-299. GRABOWSKI, C. T. & V. G. DETHIER. 1954. The structure of tarsal chemoreceptors of the blowflv Phormia regina Meigen. Jour. Morph., 94: 1-19. HODGSON, E. S. 1958. Electrophysiological studies on arthropod chem- oreception, HI. Chemoreceptors of terrestrial and fresh- water arthro- pods. Biol. Bull, 115: 114-125. ILSE, D. 1937. New observations on responses to colours in egglaying butterflies. Nature, 140: 544. KUWABARA, M. 1951. Effects of inorganic ions on tlie tarsal chemore- ceptors of the butterfly Vanessa indica, Zool. Mag., 60: 9. KUWABARA, M. 1952. Effects of inorganic salts on the tarsal chemore- ceptors of the butterfly Vanessa indica. Zool. Mag., 61: 121. , KUWABARA, M. 1953. Effect of successive stimulation on the tarsal chemoreceptors of Vanessa indica. Zool. Mag., 62: 154. MINNICH, D. E. 1921. An experimental study of the tarsal chemore- ceptors of two nymphalid butterflies. Jour. Exp. Zool, 33: 173-203. MINNICH, D. E. 1922a. The chemical sensitivity of the tarsi of the Red Admiral butterfly, Pyrameis atalanta L. Jour. Exp. Zool, 35: 57-81. MINNICH, D. E. 1922b. A quantitative study of tarsal sensitivity to solutions of saccharose in the Red Admiral butterflv. Pyrameis atalanta L. Jour. Exp. Zool, 36: 445-457. MOODY, P. A. 1953. Introduction to Evolution. Harper & Brothers, New York, xii -f- 475. MORITA, H. & K. TAKEDA. 1957. The electrical resistance of tire tar- sal chemosensory hairs of tlie butterfly Vanessa indica. Jour. Fac. Sci. Hokkaido Univ. (series 6), Zool, 13: 465-469. MORITA, H. & K. TAKEDA. 1959. Initiation of spike potentials in con- tact chemosensory hairs of Vanessa. Jour. Cellular Comp. Physiol, 54: 177-187. MORITA, H., S. DOIRA, K. TAKEDA & M. KUWABARA. 1957. Elec- trical response of contact chemoreceptors on the tarsus of tlie butter- fly Vanessa indica. Mem. Fac. Sci. Kyushu Univ., (series E), 2: 119- 139. SNODGRAS, R. E. 1935. Principles of Insect Morphology. McGraw- Hill, New York, ix -|- 667 pp. TAKEDA, K. 1961. The nature of impulses of single tarsal chemorecep- tors in the butterflv Vanessa indica. Jour. Cellular Comp. Physiol, 58: 233-245. WEIS, I. 1930. Versuchc liber die Geschmachsrezeption durch die Tarsen des Admirals Pyrameis atalanta L. Zeifschr. Physiol (Berlin), 12: 206-248. Journal of Research on the Lepidoptcra 5(1) : 13-14, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A NEW SPECIES OF EPINOTIA HUBNER FROM BRITISH COLUMBIA (OLETHREUTIDAE) T. N. FREEMAN Entomology Research Iristitute, Research Branch Canada DepaHment of Agriculture, Ottawa, Ontario During the past few years an undescribed species of Epinotia Hbn. has been causing considerable injury by mining the leaves of the western hemlock [Tsiiga heterophylla (Raf.) Sarg.] in northern Vancouver Island, British Columbia. The description of it is presented here to enable officers of the Canada Department of Forestry to report on its injury and control. Epinotia tsugana Freeman, new species Colour and maculation. — Antenna grey with dark brown annulations. Palpus grey, darker apically. Head pale grey. Thorax grey. Forewing grey, sparsely covered with short, black, trans- verse dashes and light ochreous scales, more noticeable on the apical half; costal margin whitish, with evenly spaced, short trans- verse, black dashes; apical quarter with a transverse, metallic, leaden-grey bar from near the costa almost to the tornus; beyond with a subapical, curved, metallic, leaden-grey bar extending from the costa to the tornus; a narrow, black line extending around the apical margin; fringe grey, paler basally. Hind wing dark grey; fringe light grey, darker basally.Abdomen metallic leaden-grey. Legs grey; tarsi with black bands. Wingspread, — 9.0-9.5 mm. Male genitalia (Fig. 1). -- Uncus well developed; bifid. Socii prominent; truncate at apex, Clasper arcuate; basal half much broader and with a cluster of short, thick spines near the base, Aedeagus short, stout, cylindrical; containing a cluster of six cornuti that are almost as long as the aedeagus. Type material. — Holotype, male, Holberg, Vancouver Island, B.C., 13 May 1966. Type No. 7976 in the Candian National Col- lection, Ottawa, Ontario. Paratypes. — Five males seven females, 2-20 May 1966; two females, 18 June 1965. All paratypes from Holberg, Vancouver Island, B. C. All type material reared from western hemlock by officers of the Candian Forest Insect Survey, Canada Department of Forestry. 13 14 T. N. FREEMAN J. Res. Lepid. Remarks. — The maciilation of E. tsugana somewhat resembles that of Epinotia nanana (Treit. ). On the basis of the male genita- lia however, tsugana is more elosely allied to Epinotia lomonana (Kft). Eig. 1. Male genitalia of Epinotia tsugana. Journal of Research on the Lepidoptera 5( 1 ) : 15-26, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 STUDIES IN LIFE HISTORIES OF NORTH AMERICAN LEPIDOPTERA CALIFORNIA ANNAPHILA IT JOHN ADAMS COMSTOCK and CHRISTOPHER HENNE Del Mar, California and Pearhlossom, California In carrying on studies of life histories of Annaphilas, prior to March 1964, a number of incomplete records and drawings were made which were withheld in the hope that additional informa- tion might be obtained. It now seems advisable to record these, as they may aid workers who are planning to publish more com- plete life histories. Annaphila astrologa Barnes & McDunnough In our published notes prior to March 1964 we were unable to describe and illustrate the ovum of Annaphila astrologa. Later, eggs were obtained, and additional information recorded. Lo- cation of capture was the northeast rim of Bobs’ Gap, Holcomb Ridge, Mojave Desert, Los Angeles County, elevation 4200 feet. The eggs were laid on Emmenanthe penduliflora Benth. Egg; (Fig. 1 A) Spherical, 0.5 mm. wide by 0.4 mm. tall. Color, bright yellow. The surface is covered with 48 to 50 ridges, arising from the base and terminating near the micropylar depression. Several coalesce superiorly. Ridges are topped by a line of pearl-like nodules or expansions. They are closely crowded together, and there are apparently no cross lines between them. Eggs hatched April 9, 1964. FIRST INSTAR LARVA: (One day after hatching) Length, 2.5 mm. Head width approx. 0.4 mm. Jet black. The body is narrower than the head. It is cylindrical and trans- lucent, and is tinged with light green from the first segment to about the eighth. The cauda is light yellow. The cervical shield is distinct, and is spotted with black. Legs, black. Prolegs, distally black. SECOND INSTAR LARVA: Head width 0.7 to 0.75 mm. Mottled yellow-green and black. ^ Illustrations reproduced from water color drawings by J. A. Comstock 15 16 COMSTOCK AND HENNE /. Res. Lepid. 5 Fig. 1. Annaphila astrologa Barnes & McDiinnoiigh. A. Egg, enlarged X 95. rl B. Front view of head, enlarged. I Body, 7 mm. long. Color, deep green, with rows of small brown tubercles, each bearing a colorless seta. Legs tipped with black. Prolegs, yellow with brown crochets. The third instar larva was described and illustrated in our prior paper (1964) but the distinctly marked and spotted head was . reproduces on too small a scale to be clearly defined. It is now pictured on Figure 1 B. 5(1) : 15-26, 1966 ANNAPHILA 17 Head width is 1 mm. It is studded with numerous brown dots and blotches. Those which bear setae are raised. The ground color of the face is yellow with a tinge of green. The antennae and labrum are white. Additional larvae were taken on the Mulholland Highway in the Santa Monica Mountains, May 16, 1965. The pupa illustrated in our prior paper ( 1964) was taken at Red Rover Canyon, south- west of Acton, Los Angeles Coimtv, elevation 3200 feet. May, 1963. Larvae of all the astrologa complex produce their pupal chambers by cutting into pithy wood. A colored figure of the moth was included by Draudt in Seitz, Vol. 7, 47e, 1927. The type locality is Redington, Arizona. REFERENCE COMSTOCK, JOHN ADAMS and CHRISTOPHER HENNE, 1964. Studies in Life Histories of Nortli American Lepidoptera. California Annaphilas. /. Res. Lepid. 3(3): 175-191. Annaphila pseudoastroioga Sala This species was published by Frank P. Sala in 1963. Prior to that he submitted a pupa under the manuscript name which we then illustrated and held for later verification. In addition, we had taken larvae of the same speeies feeding on the flowers and buds of Phacelia miner Thell, at Vasquez Rocks, Sierra Pelona Valley, Los Angeles County, April 1963. There is still need of the ovum to complete the life history, but it is hoped that Sala will include this in his promised life history study. LARVA, INTERMEDIATE STAGE: (Fig. 2 A) Length, 8.5 mm. Head width, 1 mm. Head is glistening light green, speckled with brown, heaviest over the crown. Ocelli, black. Mandibles edged with brown. Body, dark olive-green, with numerous raised white nodules, topped by relatively long setae. Middorsally, a narrow longi- tudinal white stripe, and dorso-laterally a similar stripe parallel- ing it. Below this, the body color gradually changes to a deep rose-purple, which extends downward to the spiracular line where it abruptly gives place to a deep green. This green also covers the entire venter. The spiracles stand on elevated nodules. They are black-centered with white circlets surrounded with rose-purple blotches. The caudal area has a considerable blotch- ing of rose-purple, extending over the dorsum. The white setae 18 COMSTOCK AND HENNE J. Res. Lepid. Fig . 2. Annaphila pseudoastrologa Sala. A. Larvae, intermediate instar, X 10. B. Mature larva approximately X 5. are mounted on elevated nodules with black centers and white circlets. The legs and prolegs are green. Later, the rose-purple invades most of the dorsal surface. MATURE LARVA: (Fig. 2 B) Length, 20 mm. Widtli, through center, 3 mm. Head width, 2.1 mm. Head color is glistening light yellow, heavily sprinkled with light brown blotches and spots over the upper part of 5(1) : 15-26, 1966 ANNAPHILA 19 cheeks and along the sides of the epicranial suture. The front is unspotted. Ocelli, dark brown, resting on yellow bases. Man- dible, black. Setae, soiled white, arising from minute brown nodules. First thoracic segment of body, yellow, with an orange cast, weakly mottled with light brown and white spots over dorsum, strongly blotched with brown laterally. There is a narrow middorsal white stripe. Remaining segments, basically yellow- green with thick spotting and mottling of darker green. As the spiracular area is approached, spotting changes to red-brown, with the black-centered and white-rimmed spiracles standing out in strong contrast. In caudal area, red-brown spotting ex- tends over dorsum. There is a bare suggestion of a longitudinal middorsal stripe, formed by indistinct darker spots. Venter, un- spotted light green. Legs, soiled yellow-green. Prolegs, mottled light green with brown crochets. All setae, relatively short and white, resting on small white papillae with minute black centers. Prior to pupation the larva assumes a deep mottled purplish- brown color above the spiracular line, and the ventral green area becomes heavily mottled with purplish-brown and minute white dots. The stigmatal line becomes more pronounced and is lighter. Thereafter the larva begins to chew its way into soft pithy bark, in which it weaves a pupal chamber. In addition to Fhacelia minor the food plant has been noted as Phacelia parry i Torrey. PUPA: (Fig. 3A, B, C.) Length, 9.25 mm. Greatest width, 3 mm. Fusiform. Head rounded. Eyes relatively small. Maxillae terminate almost at the margin of the wing cases. Antennae, slightly shorter. Cremaster terminates in two short recurved spicules pointing laterally, and two minute spikes pointing caudally. General body color, rich chestnut brown with lighter shade on the wings and abdominal segments. Antennae, nearly black. Wing cases, marked with an assemblage of buried black dots, predominating on forward half of wing, irregularly placed. Body surface texture, predominantly finely rugose, but maxillae and leg cases bear fine raised striations. Other structural features are adequately shown in the illustration. Our first pupa was obtained in 1956 from Frank Sala. It was collected in Latego Canyon above Seminole Hot Springs, Los 20 COMSTOCK AND HENNE /. Res. Lepid. Fig. 3. Annaphila pseudoastrologa Sala. Pupa, A. lateral, B. ventral, and C, dorsal aspect, approxim. X 7. Angeles County, which its author later designated the type locality. The moth is pictured on pages 290 and 292 of Sala’s original 1963 description. REFERENCE SALA, FRANK P. 1963. The Annaphila astrologa Complex, with Description of Three New Species. Jour. Res. on the Lepid. 2 (4): 289-300. Annaphila vivianae Sala This species was published by Frank Sala in 1963. Prior to that (1956) he sent us pupae with his manuscript name '‘vivianae" attached, which made possible the description and drawings presented here. PUPA: (Fig. 4A,B,C) Length, 9.5 mm. Greatest width, 3 mm. Body color, dark reddish-brown. Wings, lighter. Spiracles, dark brown. Fig. 4. Annaphila viviana Sala. Pupa, A. lateral, B. ventral, and C, dorsal aspect, approxim. X 7. The antennae terminate 0.8 mm. cephalad to the wing margins, and the maxillae extend slightly beyond these margins. The head is rounded and the eyes not prominent. The spiracles are slightly elevated. The cremaster is a rounded knob with a squared plate extension devoid of booklets. Other structural features are shown in the illustration. The range of the species is the southern Sierra Nevada Mts. Type locality, the Lower Kern River Canyon, Kern County, California. It also extends north to Three Rivers and the East Kaweah River, Tulare County. It may in addition be found throughout the range of its specific food plant, Phacelia cicutaria Greene, among the rocky out-croppings in the foothills of the Sierra Nevadas, from Bette County south to the Tehachapi Mountains. An additional foodplant successfully used in the laboratory was Phacelia tanacetifolia Benth. REFERENCE SALA, FRANK P. 1963. The Annaphila astrologa Complex with Descriptions of Three New Species. Jour. Res. Lepid. 2 (4): 289-300. 22 COMSTOCK AND HENNE J. Res. Lepid. Annaphila diva Grote This species was briefly discussed in our 1954 paper. We re- cently secured a mature larva, pupa, and cocoon which makes possible an illustiation and a few additional notes concerning its life history. Our larva was preparing for pupation, which may account for its somewhat darker color and foreshortening in form as compared with William H. Evans’ description in the Rindge and Smith 1952 Revision of the Genus. MATURE LARVA: ( Fig. 5 A and B ) Length, 19 mm. Greatest width through center, 3 mm. Head width, 1.9 mm. Head (Fig. 5 C) ground color, glistening soiled yellow, blotched witli black, particularly over the crown. Ocelli, black. Antennae, white at base, dark on distal two-thirds. Margins of mandibles, black. Body, predominantly striped and blotched with black, in- terrupted with various light lines and spots. There is a middorsal longitudinal line of dull pink, lighter on the anterior segments, gradually darkening toward the cauda. Lateral to this is a wide gray area. On the margin of this is a distinct light pinkish band running from the first to third segment. Beyond this is a wide band of mottled black running caudally, and extending down- ward to the wide spiracular band. This band is dull pink or pinkish-white, and it runs the entire length of the body. The black spiracles are located close to its upper margin. Several round light spots and a few irregular larger blotches occur on the body as shown on the illustration. The cocoon was composed of soil granules. In shape it was oval — length, 10 mm. Width, 5.6 mm. PUPA: (Fig. 5D) Length, 8 mm. Greatest width through center, 2.75 mm. Color throughout, uniform dull orange. Texture, glistening. The maxillae reach to the wing margins, and the antennae are slightly shorter. The wing covers are translucent, allowing the segmental junctures to be dimly seen through them. The caudal segment is evenly rounded, and bears no spines or cremasteric hooks. 1| The species ranges from southern California to British i Columbia. || The larval food .plant is Montia perfoliata (Bonn) Howell, and j the adults have been taken in association with other species of Montia throughout its range. 5(1) : 15-26, 1966 ANNAPHILA 23 Fig. 5. Annaphila diva Grote. Mature larva, A. dorsal, and B. lateral aspects, X approxim. 5. Head of larva, C, X 10. Pupa, D. ventral aspect, X approxim. 8. REFERENCES COMSTOCK, JOHN ADAMS and CHRISTOPHER HENNE, 1964. Studies in Life Histories of Nortli American Lepidoptera. California Annaphils /. Res. Lepid. 3(3): 175-191. EVANS, WILLIAM H., 1952. Rindge and Smith Rev. of the Annaphilas. pp. 238-239. HAMPSON, CEORCE F., 1910. Catalogue of the Noctuidae in the Col- lection of the British Museum, Lepidoptera Phalaenae. 9. p. 481, fig. 299. British Museum (Natural History) London. 24 COMSTOCK AND HENNE /. Res. Lepid. HOLLAND, WILLIAM H., 1908. The Moth Book. p. 246, PI. XXLX, Fig. 20. Doubleday, Page & Cmpany, Ne\\’ York, N. Y. RINDGE, FREDERICK H. and CLAUDE I. SMITH, 1952. A Revision of ( the Genus Annaphila Grote, ( Lepidoptera, Phalaenidoe), Bull, Am. ; Mus. Nat. Hist. 98 ( 1 ) : 187-256. | Annaphila spila Rindge and Smith i Females of this species were observed ovipositing deep in the ! flower bracts of Linanthus montanus Greene, in a meadow off | Mineral King Road, East Fork Kaweah River, Tulare County, ! California on May 3, 1963 at an elevation of 3200 feet. Females ' were also confined from this locality and oviposited readily on growing plants transplanted from the field to the laboratory. ' EGG: (Fig. 6 A) : Nearly spherical, slightly flattened at the base. Diameter, 0.4 mm. j There are from 30 to 32 ridges running from base toward mi- cropyle. These are knobbed along their crests. The ridges are slightly wavy and their knobbed crests are not uniform in size or direction. The grooves between the vertical ridges show minute and barely visible horizontal striae. The micropylar depression is large and deep. Color, bright orange, turning to riddish prior to hatching. FIRST INSTAR LARVA: (A single example, observed May 21, 1963) Length, 2.75 mm. Head width, 0.25 mm. Head, jet black. Body, rich yellow with a tinge of orange. First thoracic segment with a shading of brown, due to minute dots forming a prothoracic shield. There is similar shading on the last caudal segment. There are minute dark dots on the dorsal area, some of them probably giving rise to colorless setae. Legs, shading to black at the tips. Prolegs, concolorous with the body. With only a single larva for observation, unusual care in handling was necessary, so some details were likely not noted. In spite of precaution the larva died. The early instar larvae feed within the bracts and consume the buds of the young plants. They are not readily seen in the field at this stage. The adult larva rests upon the reddish stem of the food plant, feeding externally upon the flowers and buds, its coloration blending with that of the plant. MATURE LARVA: (Fig. 6B) A single example was obtained June 13, 1963. Length, 13 mm. Width through 9th segment, 2 mm. Width of head, 1.4 mm. 5(1) -.15-26, 1966 ANNAPHILA 25 Fig. 6. Annaphila spila Rindge & Smith. Egg, A. approxim. X 100. B. Larva, X 12. Head, clear glistening yellow, the mouth parts slightly darker. Ocelli, black. Dorsal half of body, deep pinkish-rose. The first segment has a middorsal yellow line extending slightly onto the 2nd segment. There is also a short dorso-lateral line on the 1st segment only. A wide stigmatal cream-colored band runs the entire length of the body. The spiracles are placed on its upper edge. Below this spiracular band the body is pinkish-rose as is also the venter. In 26 COMSTOCK AND HENNE J. Res. Lepid. the middle of the venter the rose color becomes gradually lighter, with a suggestion of yellow. Spiracles, small and dark-centered, with a faintly suggested white margin below each one. Legs, translucent yellow, with dark tips. Prolegs, pinkish-rose, except for those on the head and cauda. Our few examples failed to pupate as the flowers of the Linanthus became too dry for the larvae to feed upon and sub- stitute food plants (other species of (Liruinthus) proved un- successful. Journal of Research on the Lepidoptera 5(1) : 27-28, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 NEW SKIPPER RECORDS FOR MEXICO H. A. FREEMAN' Garland, Texas During the process of making a careful systematic study of the Hesperiidae of Mexico my attention was drawn to three species of skippers located in specimens obtained from Dr. T. Escalante, Mexico, D. F., and one species in material recieved from Stallings & Turner, for determination. As there are no pub- lished records of the occurrence of any of these four species from Mexico I am recording them as new records for that country. Astraptes helen Evans This species was described as a subspecies of parisi Williams by Evans, the basic difference being in the genitalia. In this group of Astraptes there are several sibling species which are actually best separated by their genitalia, indicating that we should con- sider helen and parisi as separate species since their genitalia differ considerably. A. helen was described from San Pedro Sula, Honduras. Besides the seven specimens in the British Museum from Honduras, Evans also records Nicaragua and Panama as constituting the range of that species based on material available to him. Among the specimens received from Dr. T. Escalante for study was found a male helen from Paraje Nuevo, Veracruz, Mexico, IV-52. This constitutes a new hesperiid record for Mexico. Cogia mala Evans This species was described by Evans from two males collected in Guatemala during 1904 by A. Hall. From what information available there are no other records for this species. Among Dr. Escalante’s specimens was found a male mala from Acahuizotla, Gro., IX-60. While collecting just north of Tehuacan, Pue., August 15, 1964, I collected five males of this species. This is another new record for Mexico. It was interesting to note that Cogia aventinus Godman & Salvin was found in the Escalante material from Acahuizotla also. Mala and aventinus show con- siderable superficial relationship, however, the larger hyaline ^This research has been made possible by a research grant GB-4122 from the National Science Foundation for which I am deeply thankful. 27 28 H. A. FREEMAN J. Res. Lepid. spots of mala and the difference in the genitalia make these two species easy to separate. Ouleus cyrna (Mabille) This species was described by Mabille from a female, collected in Chiriqui, Panama. Later it was redescribed by Godman & Salvin as simplex and then fasciata from material collected in the same general area as the Mabille specimen. Evans records cyrna from Costa Rica and Panama, based on specimens in the British Museum. In the Escalante Hesperiidae there was found a female cyrna which had been collected at Sta. Rosa, Comitan, Chis., Mexico, IX-63. This is the first time that this species has been recorded from Mexico. Carystoides lila Evans This species was described from L. Sapatoza Region, Chirigua district, Columbia. Evans records it from Honduras, Columbia, Venezuela, and Ecuador. Stallings & Turner collected a female (which is in my collection) lila at Tamazunchale, S. L. P., Mex- ico, VL6-41. Apparently this is the first record for this species from Mexico. LITERATURE CITED BELL, E. L. 1942. New Records and new species of Hesperiidae from Mexico. Sobret. de los Andes de la Escuela Nacional de Ciencias Bio- logicas. VoL 11, No. 4. 455-468. FREEMAN, H. A. 1951. New Skipper Records for Mexico. Field & Lab., Vol. XIX, No. 1, 45-48. DRAUDT, M. 1924. Hesperiidae, in Seitz, Macrolepidoptera of the world. Vol. 5. The American Phopalocera. Stuttgart, vii, 1139 pp., 203 pl. EVANS, W. H. 1951. A catalogue of the American Hesperiidae indicat- ing the classification and nomenclature adopted in the British Museum. Part 1. Introduction and Pyrrhopyginae, London: British Museum. 1-92 pp., 1-9 pis. — 1952. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part II. Pryginae. Sec. I. London: British Museum. 178 pp., pis. 10-25. 1953. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part III. Pyrfinae. Sec. 2. London: British Museum. 246 pp., pis. 26-53. 1955. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part IV. Hesperiinae and Megathyminae. London; British Museum. 449 pp., pis. 54-88. GODMAN, FREDERICK DUCANE, and OSBERT SALVIN 1887-1907. Biologia Centrali-American, Insecta, Lepidoptera-Rhopalocera, II: 244- 637; LII: pis. 112. HOFFMANN, C. C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros Mexicanos, Sergunde parte-Hesperioidae. An. Inst. Biol. Mexico. 12: 237-294. MacNEILL, C. DON 1962. Preliminary report on the Hesperiidae of Baja California. Froc. Calif. Acad. Sci., (4) 30 (5); 91-116. Journal of Research on the Lepidoptera 5(1) ;29-36, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A MOTH SHEET FOR ATTRACTING AND RETAINING LIVE SPECIMENS WITHOUT THE USE OF A TRAP OR TENT-ENCLOSURE. NOEL McFarland South Australian Museum, Adelaide, South Australia For many collectors who have no need or desire to trap and kill the vast numbers of insects coming to their insect-attracting lights, the type of device described below is very much worth the trouble to construct. It is -- in any event — useful, as it ef- fectively increases the power of the sheet-and-light combination to draw in, and “hold”, nocturnal flying insects. It is of particu- lar value to entomologists who are interested in capturing live females, for purposes of obtaining eggs for rearing and life his- tory investigation. This “PARABOLIC MOTH-SHEET” was in constant use, along with several “conventional” set-ups (flat sheets with lights hanging in front of them) for a continues per- iod of 20 months in a single locality (McDonald Forest, 5 miles west of Corvallis, Benton County, Oregon; October 1961 to June 1963). I constructed it at the suggestion of Mr. Gerald Bene- detti, then a graduate student in Mechanical Engineering at Oregon State University. The effect of directing a correctly-located light at a verti- cal sheet, which is held in the shape of a parabolic curve, is such that the whole reflecting-surface (i.e., the whole sheet) appears to be the source of the light.. This results in a very different situation from that observed in the usual (flat) sheet-and-light combination, where there is a circle of fairly intense light at the middle of the sheet only. When the sheet is bent in the form of a parabolic cun^e, the whole sheet appears to “come alive” with the brilliance of the light it is reflecting, provided that the light is placed at the correct distance from the (inside) vertex of the curve. The distance of the light from the vertex is determined by the shape of the para- bolic cun^e in use; see Fig. 1 (A-D) and photographs. The size of the sheet ( or canvas ) used during the trial-period in western Oregon was 16 feet x 7 feet, and it was in operation at ground level, being held in position by 13 vertical poles run- 30 NOEL McFarland J. Res. Lepid. Fig. 1 (A-D) — Diagrams of four possible curves for the Parabolic Moth Sheet. Light source at (L), on a vertical pole. Vertex of curve at (V). Squares are one foot square. Sheet is 16 feet long, shown as if viewed from directly above . 5 (1) :29-36, 1966 MOTH SHEET 31 ning through sleeves which were sewn to the canvas; the poles were in turn inserted into short ground-stakes (see Fig. 2) which were easily hammered into the ground. The placement of these ground-staks determined the accuracy of the resulting parabolic curve made by the canvas. Guy wires from the tops of the poles secured the whole structure in the event of high winds. Two 18-inch, 15"Watt black light tubes (G.E. F15T8/BL) were hung vertically on the light-pole, one facing directly at the inside vertex of the curve, and the other facing in the opposite direc- tion. (The addition of this second, outward-directed light did not seem to affect the results greatly, although it may have in- creased the “attracting-power” of the device). The general ap- pearance of the whole set-up can be seen in the photographs, which are of the original parabolic sheet used in Western Ore- gon. There are various ways and materials for constructing such an apparatus, either with the intention of making it portable (as described above), or heavier and permanent in one location. The important points are to maintain the parabolic curve with reasonable accuracy, so that the entire sheet appears to be throwing out the light, and not to make the sheet too small. The more surface area for incoming moths to land upon, the better. The 16 foot X 7 foot canvas operated in Oregon did a very im- pressive job. Of six black light and sheet set-ups in use in the one locality, the parabolic ( curv^ed ) sheet invariably drew in, and re- tained, far more insects than any of the others; usually it had a greater catch than the totals of the other five combined. Moths coming to the parabolic sheet have a greater tendency to remain and settle there. They often bounce back and forth within the curving enclosure before finally settling, but there is not much loss from individuals simply flying off around either end, or over the top, and not returning. (This is one of the disadvantages of a conventional flat sheet operating with- out any sort of trapping device). Some of the moths will fly around to the unlighted back of the sheet (the outside of the curve) and settle there; it is highly desirable to leave the back unlighted to accommodate such moths, as these are the individuals that would sometimes settle in nearby vegetation after once flying to the light; such species ™ this is sometimes a behavioral difference between species — are usually missed or overlooked at conventional flat sheets. When this occurs it often pays to shake or lightly beat nearby vegetation, which will cause many of the moths to fly back to the sheet, unless it 32 NOEL McFarland J. Res. Lepid. i I Fig._ 2 ~ Diagram of a Ground Stake, which is easily pounded into the ground to j receive the poles that will form the parabolic curve of the sheet. Made of 1 Vi-inch angle | iron. Total length about 21 inches, a = 6-inch metal tube, welded into angle iron 3 ‘ inches below the top; this tubing is a fraction larger than the diameter of the pole it ' is to receive; b = a small triangular plate of iron, welded about one inch below the bottom of the tubing, to stop the pole at a specified height after it is inserted through the tubing; c = tapered point on angle iron makes it easier to drive into hard ground. i is cold, in which case most of them will just drop to the ground. j There are many different parabolic curves that could be tried, | and some may prove to be more efficient than others in “hold- I ing” the moths on the sheet; perhaps also in drawing them to 1 the sheet in the first place. Four possible curves to try are sug- } gested in Fig. 1 (A-D). All comments in this paper are based : on experiences with only one of those parabolic curves (Fig. lA), j the shape of which is determined by the light source being set j at three feet from the inside vertex of the curve. The light i; 5(l):29-36, 1966 MOTH SHEET 33 faces directly toward the inside vertex of the curve, as it hangs on a vertical pole which is set at the specified distance out from the inside vertex of the curve. In my experiments, using the curve shown in Fig. lA, two lights were in operation — one fac- ing the vertex, and the other on the opposite side of the light- pole, facing directly away from the vertex. Other placements of the light ( or of several lights ) on the pole could be tried, and these might give varying results. For spacing all the poles correctly on the ground, the easiest method is to stake out 2 strings at right angles to each other, along the ground, one representing the x-x axis (8 feet long is sufficient), and one representing the y-y axis (16 feet long is sufficient). Measurements are made by going out from the ver- tex, on the x-x axis, and then going either up or down parallel to the y-y axis, to the correct measured distance, for placement of each pair of corresponding opposite poles. (The only single pole, not having an opposite, is the one at the vertex). The sheet should be stretched tightly, so that the end result presents a smooth, wrinkle-free, evenly-curving reflective surface. A soiled, “off-white” canvas or sheet is preferable to a pure white one, for purposes of getting the moths to settle on the sheet with a minimum of flying about. A ground-sheet (“apron”) is very desirable along the bottom of the curve. This can be made in 4 four-foot sections, which are easily snapped on along the bot- tom of the curve after it is set up. ( If it is all in one piece, there are too many wrinkles in a 16-foot curving piece ) . An advantage in having the apron detachable is that it may be removed and washed from time to time, without the inconvenience of taking down the whole structure. After one wet season on muddy ground, the apron becomes the color of the soil, in which case it is rendered practically useless until it is washed. To be of much value, the apron should extend out onto the ground, from the base of the curve, for at least 18 inches all the way around. It is not needed on the back ( outside ) of the curve. If a few pieces of curved bark ( as available ) are placed on the ground around the sheet, it will be found that certain moths (often of the same species) will hide under these, rather than disappearing into nearby weeds or ground litter; in this category are certain noctuids such as some Agrotis, Euxoa, and Ufeus spp., among others. Quite a few moths will settle on the guy wires, especially (for some unexplained reason) on cold nights; thus the guy wires should be rope or cord, not a smooth-surfaced 34 NOEL McFarland J. Res. Lepid. 5(1) :29~36, 1966 MOTH SHEET 35 material. These and other refinements can be included to suit the intentions of the collector. To experiment with other parabolic curves for the sheet (i.e., where the light is at some distance from the vertex other than the examples illustrated by Fig, 1.) the following general equa- tion, for determining any points on a proposed parabolic curve, can be used: y^' = 4 AP. We are solving for P, which is a point on the proposed parabolic curve, when y is a known distance up or down the y-y axis from its intersection (V) with the x-x axis, and A is a KNOWN distance of the light source (L) from the vertex (V), along the x-x axis. This is plotted on graph paper. Lay out the y-y axis and the x-x axis on the graph. When the curve is drawn in, it is to be symmetrical about the V-x axis, with its vertex touching V. An example follows, to show the steps in finding a point on the proposed curve, when y is set ( for example ) at 6 feet up or down from the vertex, along the y-y axis, and the distance (A) of light source to vertex is set at 3 feet: y^ = 4 AP (solving for P) 12 P = 36 P = 3 feet In other words, a point on the proposed curve, either 6 feet above or below the x-x axis, is located 3 feet to the right of the y-y axis. Repeat similar calculations again for several other points on the same proposed curve, setting y at, say, 1, 2, 3, 4, 5, and 8 feet, and keeping the distance (A) of light source to vertex the same in each calculation. Place appropriate dots on the graph after each calculation; when finished, the series of dots may then be connected, resulting in the desired parabolic curve. Using graph paper divided into ^A-inch or ^A-inch squares, it is only necessary to solve for about 6 or 7 points on a curve that is to be in the vicinity of 16 feet long. This will give a curve sufficient- ly accurate to produce the results described above. Anything more precise is not practical to attempt setting up (or to main- tain) under field conditions. ACKNOWLEDGMENTS I am indebted to Mr. Gerald Benedetti of Livermore, Califor- nia, for the original suggestion of trying a parabolic curve, and for plotting the curve that was used; to Mr. Lloyd M. Martin of 36 NOEL MgFARLAND /. Res. Lepid. the Los Angeles County Museum, California, for many helpful suggestions on the construction of various types of portable collecting sheets, frames, and ground stakes. REFERENCES McFarland, a. N. (1963) — The Macroheterocera ( Lepidoptera ) of a mixed forest in Western Oregon. Thesis submitted to Oregon State University, Entomology Department, Corvallis, Oregon. U.S. DEPT OF AGRICULTURE (1961) — Agricultural Research Ser- vice. Response of insects to induced light. Presentation papers (ARS 20-10). 66 pp. COVER PHOTO Daritis ? howardi Hy. Edw. (PERICOPIDAE) Penultimate instar, filled-out, resting on Eupatorium stem. (Length: 80 mm. when filled-out in final instar). Collected at approx. 7200 ft. elev., on the W. slope of Cabezon Peak, Sandoval Co., NEW MEXICO; 11 OCT. 1958 (half grown); S. L. VanLandingham, collector; on Brickellia sp.(?) Reared to the pupal stage by Noel McFarland, on a substitute plant, Eupatorium rugosum, in Lawrence, Kansas. Color transparencies by Carl W. Rettenmeyer. Larva should be hanging from the branch rather than standing upright. References: 1) Dyar (1900): Proc. ent. soc. Washington 4:407. 2) A. Seitz (1940): Macrolep. of the World 6 (Text): 425,427,447. 3) N. McFarland (1961): Jour, Lepid. Soc. 15(3):172-174. Journal of Research on the Lepidoptera 5(1) : 37^38, 1966 1160 W. Orange Grove Ace., Arcadia, California, U.S.A. 91006 © Copyright 1966 REDISCOVERY OF ANNAPHILA CASTA HY. EDW. IN CALIFORNIA (NOCTUIDAE) JOHN S. BUCKETT University of California Davis, California After the preparatiox of the manuscript dealing with a re- evaluation of Annaphila casta (Biickett, 1965), more specimens have become available for study. In addition to the two Cali- fornia specimens cited by Rindge and Smith ( 1952 ) , there are now a number more California specimens available. In the authors and Mr. Baiters joint private collection, there are 3 males and 12 females that were collected in central California; all the specimens are in good to excellent condition. During the autumn of 1964, Mr. Noel LaDue of Sacramento brought some diurnal moths to Mr. Bauer, and eventually to me also, for determination. Much to our astonishment, some of the moths were the very scarce, and long sought A. casta. The speci- mens were collected earlier the same year on 10 May, near Plantation, Sonoma County, California. During the following spring of 1965, Mr. Bauer, Mr. M. R. Gardner and I made several attempts to collect A. casta near Plantation, but failed on our earlier attempts. It was not until the exact date of 10 May, 1965 that Mr. Bauer, Mr. LaDue, and I succeeded in collecting speci- mens of A. casta. Subsequent collecting expeditions were made, one on the 12th of May and another a week later; howei^er, no specimens of A. casta were taken, or even seen. It seems rather odd that the species is apparently so restricted in its period of flight! Yet the same trend was observed in the spring of 1966. It was very difficult to determine just where the specimens were coming from, and therefore it was equally diffi- cult to locate a probable larval host plant. The specimens collected on 10 May 1965 were observed floating from the skies between the upper canopies of the redwoods (Sequoia sempervirem Endl. ) , to alight in a meadow during the morning rays of the sun. Their actions were typical of species within the genus, especial- ly the slow waving up and down of the wings while the specimen was alighted. There was apparently no particular preferred 37 38 JOHN S. BUCKETT /. Res. Lepid. alighting spot, as specimens were taken from the ground, grass, and low growing young redwoods. During the spring of 1965, little attempt was made to determine the larval host plant(s) of casta. During 1966, however, Mr. and Mrs. Chris Henne were in the vicinity of Plantation, and ardent attempts were made to determine a larval host plant relationship, but with no positive results. To date there is apparently no knowledge as to the larval host plant ( s ) of A. casta. While collecting A. casta during 1965, it was noticed that the moth occurred only in one very limited area. Other adjacent areas with virtually the same environment as this one particular meadow yielded absolutely nothing, nor were any specimens of casta even seen. As yet, there is no explanation for this acute endemicity, although it is believed with additional biological information gained in relation to casta, a logical hypothesis might result. Buckett (op. cit. ) discussed various possibilities for the cor- ! rect type locality of A. casta. It was (and still is) believed that the type locality of the species is in Oregon, not in California ( for | evidence to support this idea, refer to the aforementioned paper ) , One fact that is supported by the recent collections of casta is the probable validity of the labels on the two specimens contained | in the American Museum of Natural History in New York City. I These two specimens are labeled: “Sonoma Co. May Cal.” and “Sonoma Co. May Calif.,” and because they are each labeled differently there is the suggestion that the specimens were prob- ably collected during different years. Judging from the evidence available in the literature, the Oregon type locality, in all prob- ability is the correct one. It is hoped that in the future more work will be done in attempting to solve the life history of A. casta. Also, it would be of academic interest to determine whether or not casta has a continuous vs. discontinuous distribution between its known i localities (namely Sonoma and Mendocino Counties, California, and Benton and Douglas Counties, Oregon). I would like to extend my appreciation to those individuals that took part in this project: Mr. Bill Bauer; Mr. Mike Gardner; Mr. and Mrs. Chris Henne; Mr. Noel LaDue. LITERATURE CITED BUCKETT, J. S., 1965 A reevaluation of Annaphila casta (Noctuidae). Jour. Res. Lepid. 4(3): 199-204. I RINDGE, F. H., and C. I. SMITH, 1952. A revision of the genus Annaphila Grote. Bidl. Amer. Mus. Nat. Hist. 98(3) :191-256. Journal of Research on the Lepidoptera 5(1) : 39-50, 1966 1160 W , Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 STUDIES ON THE NE ARCTIC EUCHLOE PART 1. INTRODUCTION PAUL A. OPLER Department of Entomology University of California, Berkeley, California This paper will serve as the introduction for a series of papers which will present the results of an inv^tigation primarily con- cerned with systematic placement of Nearctic members of the genus Euchloe Hiibner, The genus Euchloe is composed of but- terflies known as “marbles” which are members of the family Pieridae. The larvae of these insects feed upon species of Cmci- ferae. Populations of one or more species are found throughout most of the Neaxctic Zoogeographic Region but occur most fre- quently in somewhat xeric habitats in the western United States and Canada. The taxonomic relationships of the Palaeartic and Ethiopian members of the genus are not considered in this study. Brief references will be made to the Old World entities in order to clarify important points relating to Nearctic members of the genus. The author hopes to correct several faulty systematic concepts which have been applied to the Nearctic members of Euchloe for many years. These misunderstandings were due to the fact that the adults w«e identified by a few superficial external characteristics. To correct the above stated situation, an attempt has been made to arrive at a more meaningful systematic treatment of the genus which will reflect probable phylogenetic relation- ships. This study presents the results of an analysis based on mo’rphological, distributional, and ecological characteristics of all Nearctic members of the genus. Based on this analysis a workable systematic framework is presented. Four spedes of Nearctic Euchloe are recognized: E. ausonides (Lucas), E. creusa (Doubleday), E. olympm (Edwards), and E. hyantis (Edwards). These are believed to be phylogenetically and morphologically divisible into two species groups. One is comprised of E. ausonides, E. creusa and E. olympm, and will be referred to as the E. ausonides group. The other is composed of E. hyantis and its several segregates; this group will be re- ferred to as the E, hyantis complex. iThe bulk of the materia! presented in this series of papers was part of a thesis pre- sented in partial fulfillment of the requirements for the degree of Master of Arts at San Jose State College, San Jose, California. Dr. J. W. Tilden of the above institution was research advisor. 39 40 PAUL A. OPLER /. Res. Lepid. Two new combinations will be presented as subspecific desig- nations of E. hyantis in a later paper. Below are the titles of the papers which are anticipated for this series. Part 1: INTRODUCTION Part 2: CHRONOLOGICAL LITERATURE REVIEW AND BIBLIOGRAPHY Part 3: COMPLETE SYNONYMICAL TREATMENT Part 4: TYPE INFORMATION Part 5 DISTRIBUTION Part 6: ADULT MORPHOLOGY AND KEY TO IDENTI- FICATION Part 7: LARVAL MORPHOLOGY AND KEY TO IDEN- TIFICATION Part 8: BIONOMICS Part 9: DISCUSSION AND SUMMARY ACKNOWLEDGMENTS The author expresses his gratitude to J. W. Tilden of San Jose State College who gave direction to this study. F. M. Brown, H. K. Clench, C. F. dos Passes, Paddy McHenry, J. A. Powell, and N. D. Riley contributed advice and information which made the construction of the synonymies and the writing of the portions which dealt with the type material an easy task. P. R. Ehrlich, T. H. Emmel, C. L. Remington, O. E. Sette, and Fred Thorne contributed invaluable information on the biology and the mor- phology of the genus. A. B. Klots, C. D. MacNeill, L. M. Martin, as well as Tilden gave freely of tlieir knowledge on biological concepts with special regard to members of the genus Euchloe. I am also grateful for the advice on special techniques, equipment, and methods which was given by R. C. Ballard, J. G. Edwards, W. E. Ferguson, C. D. MacNeill, J. A. Powell, C. L. Remington, and J. W. Tilden. I greatly appreciated the suggestions of Leroy Parks, E. S. Ross, and Ronald Stecker about details concerning the format of these papers. My sincere tlianks to Peter Herlan, Nevada State Museum, MacNeill, California Academy of Sciences, Martin, Los Angeles County Museum, Powell, California Insect Survey, University of California, Berkeley, and F. H. Rindge, American Museum of Natural History who allowed me to examine or borrow the specimens ofEuchloe in collections under their care. The following individuals kindly loaned specimens from their personal collections: Richard Brown, Concord, Cali- fornia, John Lane, Los Angeles, California, R. L. Langston, Kensington, California, R. E. Stanford, Los Angeles, Cdifomia, and Fred Thorne, El Cajon, California. E. J. Newcomer of Yakima, Washington collected many specimens of Euchloe from that state for the author’s examination. C. W. Sharsmith of San Jose State College kindly confirmed or corrected the author’s determination of plant specimens. Margaret S. Bergseng of the University of California Herbarium determined two species of Cruci- ferae which were collected during the summer of 1965. My special thanks are extended to J. W. Tilden, H. T. Harvey, and C. W. Sharsmitli, who read the thesis upon which these papers are based and helped to correct errors. Any errors which occur in these papers are the responsibility of the writer. 5(1) :39~50, 1966 NEARCTIC EUCHLOE 41 PART 2. CHRONOLOGICAL REVIEW OF THE LITERATURE AND BIBLIOGRAPHY The FmsT portion of this paper will deal with the develop- ment of the concepts concerning the specific status of the Euchloe of the Nearctic Region. The second portion is a com- plete bibliographic listing of all of the literature which is cited in the papers of this series. CHRONOLOGICAL REVIEW OF THE LITERATURE In this review opinions of all authors which contain contribu- tions to development of concepts concerning the status of Nearc- tic Euchloe are given. Personal interpretations are made by this writer for the sake of bringing clarity to past opinions which, in the view of the author, are often illogical or nebulous due to the presentation of insufficient information. As an aid in evaluation of this study a brief synonomy of the names that have been proposed for Nearctic Euchloe is pre- sented: Euchloe (Euchloe) 1. ausonides (Lucas), 1852 Var. flavidalis Comstock, 1924 Var. semiflava Comstock, 1924 ab. boharti Doudoroff, 1930 i color adensis) ab. hemiflava Field, 1936 A. ausonides coloradensis (Hy. Edw. ), 1881 (belia belioides) race montana Verity, 1908 B. ausonides mayi Chermock & Chermock, 1940 2. olympia (Edwards), 1871 A. olympia rosa (Edwards), 1871 3. creusa Doubleday), 1847 Var. elsa Beutenmuller, 1898 (belia) var. orientalides Verity (Partim.), 1908 4. hy antis (Edwards), 1871 (belia) var. pseudo ausonides Verity, 1908 A. hy antis lotta Beutenmuller, 1908 New Combination (creusa) ab. pumilio Strand, 1914 (belia) var. orientalides Verity (partim.), 1908 B. hyantis andrewsi Martin, 1936 New Combination Doubleday & Hewitson (1847) named Euchloe creusa as the first entity of the genus for North America based on specimens collected in the Rocky Mountains of Canada by Lord Derby. [Westwood died before date of publication] No description was given, and the identity of the form was based only on a type specimen [not designated] and an inadequate plate figure. Lucas (1852) described Euchloe ausonides on the basis of specimens collected in California by Lorquin. No figure was included, but lengthy descriptions of both sexes were given. Boisduval (1852) mentioned the name ausonides in a paper on the butterflies of California. Boisduval was credited as the 42 PAUL A. OPLER /. Res. Lepid. author of the name in many citations due to a misunderstanding of publication dates. Dos Passos (1962) gave detailed informa- tion concerning the dates of publication of the papers in ques- tion on the basis of which Lucas is credited with the authorship of ausonides. Edwards (1863) redescribed ausonides from California, Can- ada, and Colorado. It is interesting that this description could be applied to any of the Nearctic entities with the possible ex- ception of Euchloe olympia. Edwards (1871) described Euchloe hyantis from specimens collected at Mendocino City, Mendocino County, California by R. H. Stretch. Strecker (1878) treated ausonides as a variety of Euchloe aus- onia, a Palaearctic form. Henry Edwards (1881) described Euchloe ausonides colora- densis. This name was proposed as a new species according to the taxonomic usage which was in practice at that time. The description included was just sufficiently detailed to establish the identity of the insect. The type was collected by T. L. Mead in June, 1871 while he was in Colorado with the Wheeler Geo- graphical Expedition. Beutenmuller (1898) wrote the only revision of the Nearctic Euchloe prior to the present study. In this revision the author revised his opinions from those expressed in his paper of 1897. He introducted two new names, Euchloe lotta described as a specific entity, and Euchloe creusa elsa, which has been sub- sequently treated as a synonym of creusa. The name lotta was proposed for the entity found in the Artemisian Biome and deserts of the southwestern United States. Previously, the name creusa had been applied to these populations, which Beutenmuller correctly realized were not conspecific with creusa. Although Beutenmiiller’s concept had an insight into relationships in- volved, he not only did not explain the reasons for his arrange- ment of the entities within the genus, but incorrectly considered the name hyantis to be a synonym of creusa. Butler (1899b) published a paper in response to the work of Beutenmuller (1898) in which Butler stated that all Nearctic entities were forms of ausonides. Beutenmuller (1899) stated, in a reply to the comments of Butler ( 1899b ) , “I concluded that it would be best to allow the species to remain distinct until more light could be obtained on the subject.” He then proceeded to elaborate on his concept per- taining to the identities of creusa and hyantis lotta. 5 (1) :39-50, 1966 NEARCTIC EUCHLOE 43 . Browning (1901), in a faunal paper on the Rhopalocera of the Salt Lake City, Utah, region, figured a specimen of hy antis lotta. He correctly applied the name lotta to the specimen, but incorrectly assumed it to be a variety of ausonides, which can be seen immediately by referring to the above mentioned plate. Franck (1905) stated that he believed the name hyantis to represent a subspecific entity of ausonides and that the name coloradensis was a synonym of hyantis. No basis was given for this idea. The probable identity of Euchloe illustrated by Wright ( 1906 ) indicates that in only one or two cases is it possible that the collection data are correct. Coolidge (1908) attempted to present an arrangement of Nearctic entities which would eliminate the wide range of dif- fering treatments of the genus. He placed lotta and hyantis as subspecific names under creusa and placed coloradensis as a synonym of ausonides. This arrangement, in an only slightly modified form, was restated by Barnes and McDunnough ( 1916 ) , and gradually assumed the position as the '‘correct” treatment of the Nearctic members of the genus. This concept was given no bilogical or morphological basis. Verity (1908) attempted to enforce upon the new world mem- bers of the genus Euchloe a biological phenomenon which was well documented in the Palaearctic Region, i.e., the occurrence of two morphologically distinct seasonal forms for each entity. Several unnecessary new names and some very unwieldy nomen- clatorial combinations resulted. The types of ausonides and creusa were figured in this work. Strand (1914) described an aberration hyantis lotta from Oregon by the name pumilio. He incorrectly used the name in combination with creusa. Barnes and McDunnough (1916) gave a lengthy discussion of their opinions of the relationships of North American mem- bers of Euchloe. Although the authors reinforced the erroneous arrangement initiated by Coolidge (1908), they corrected the mistake by Verity concerning the presence of seasonal forms in the Nearctic Region by placing all of Verity’s names in syn- onymy. They also correctly considered the name elsa to be a synonym of creusa, and raised coloradensis from synonymy as a subspecies of ausonides. Barnes and McDunnough (1917) presented the arrangement of the Nearctic entities of the genus Euchloe which was to be followed for many years. The following citations repeated the 44 PAUL A. OPLER /. Res. Lepid. treatment of Barnes and McDunnough: Barnes and Benjamin (1926), Comstock (1927), and dos Passos (1964). Comstock (1924) described two varietal color forms of fe- male aiisonides, which were collected from the poplations which occur in the vicinity of San Francisco Bay, California. Comstock evidently did not know that one of these color forms, semiflava, is nomal for the females of this species, since the hindwing of the female in other species does not show a contrast in pigmentation with the forewing. The variation in pigmenta- tion of this group will be presented in greater detail in a later paper of this series. Coolidge ( 1925 ) described the life history of Euchloe hyantis lotta on the basis of material collected in the western Mojave Desert of California. Coolidge repeated his mistake of an earlier paper (Coolidge, 1908) by associating the name hyantis with Euchloe creusa. Comstock (1927) in “Butterflies of California” introduced two somewhat erroneous concepts which were perpetuated because of the popularity which this book attained. Although Comstock figured specimens of aiisonides coloradensis from Colorado, he stated in the text that this subspecies “is occasionally taken in the high mountains of California”. This statement led to the practice of referring to all specimens of Euchloe collected in the Sierra Nevada as '‘Euchloe aiisonides coloradensis”. Some small isolated populations of ausonides are to be found in the Sierra Nevada; however, most population of Euchloe in that range are hyantis. The latter situation and the fact that Comstock’s treat- ment of hyantis was somewhat nebulous led to misunderstand- ings of hyantis. Doudoroff (1930) gave to a melanic specimen of Euchloe ausonides the name hoharti. Klots (1930b) reported E. creusa from Teton County, Wyo- ming. Specimens from this collection have been examined by the author and proved to represent hyantis lotta. Holland ( 1931 ) figured the type of hyantis and a specimen which may represent the type of Euchloe hyantis lotta. Holland treated the names ausonides and coloradensis conspecifically, and considered the names creusa, lotta, and hyantis to represent separate species. His statement of the distribution of the latter entities showed that he had no clear concept of their relation- ships. 5(1) ;39-50, 1966 NEARCTIC EUCHLOE 45 Field (1936) described a male color form of Euchloe ausonides from Utah in which the ‘upper surface of the secondaries [was] suffused over with yellow”. Not only did Field incorrectly at- tribute this specimen to the name coloradensis, but he may have been incorrect in assuming that it was a male. Martin (1936) described a population of Euchloe from the San Bernardino Mountains of southern California. He gave to this race the name andrewsi in honor of the collector, R. H. Andrews. This population should be referred to as a subspecific category of Euchloe hyantis rather than of Euchloe ausonides as it has been treated. Chermock and Chermock ( 1940 ) described Euchloe ausonides mayh a distinctive population from the Riding Mountains of Manitoba. Brooks (1942) erroneously considered ausonides coloradensis to occur in Manitoba. Leighton ( 1946 ) reported creusa and Euchloe creusa hyantis' as occurring in eastern Washington. The insects referred to by these two names were probably specimens of Euchloe hyantis lotta. Newcomer (1964) followed this procedure in reporting creusa from Yakima County, Washington. Bauer (1953) applied the name creusa to hyantis lotta from Arizona. Brown ( 1955 ) used the name creusa to represent specimens of hyantis lotta from western Colorado. Brown showed that he was aware of difference between hyantis lotta and ausonides colora- densis by his list of differentiating characteristics in the appear- ances of the two entities. Powell (1958) and others have used the combination ausonides andrewsi to refer to specimens of Euchloe from the peninsular ranges of Baja California Norte. Populations of Euchloe from the peninsular ranges of southern California, and Baja California Norte, referrable to hyantis, do not represent the names andrewsi or lotta. Ehrlich and Ehrlich ( 1961 ) used the names Euchloe ausonides, Euchloe olympia, and Euchloe creusa to represent all of the entities of Euchloe found in the Nearctic Region. The authors state that Euchloe creusa “is only doubtfully distinct from Euchloe ausonides." Garth and Tilden (1963) followed the usage of Comstock (1927) in reporting “Euchloe creusa hyantis" and Euchloe aus- onides coloradensis from the central Sierra Nevada of California. 46 PAUL A. OPLER J, Res. Lepid. Storer and Usinger (1963) used the name “Euchloe coloraden- sis'’ in referring to a Euchloe found in the Sierra Nevada of California. Dos Passos (1964) adds the dates of the original citations for all of the names involved, his arrangement is little changed from that of Barnes and McDunnough ( 1917 ) LITERATURE CITED ANDERSON, E. E. 1904. Catalogue of British Columbia Lepidoptera. Provincial Museum, Victoria, B.C., 56 pp. ARNHOLD, F. R. 1952. Notes on collecting Anthocharis midea and Euchloe olympia. Lep. News 6(6-8): 99-100. AXELROD, D. I. 1959. in: Munz and Keck, A California Flora. Uni- versity of California Press, Berkeley & Los Angeles, pp. 5-9. BARNES, W. & F. H. BENJAMIN. 1926. Checklist of the diurnal Lepi- doptera of boreal America. Bull. So. Calif. Acad. Set. 25(1): 3-27. BARNES, W. & J. H. McDUNNOUGH. 1916. Contr. Nat. Hist. Lepid. North America 3(2): 59-62. - 1917. Checklist of the Lepidoptera of boreal America. Decatur, Illinois. 392 pp. BAUER, D. L. 1953. Butterflies at water holes in central Arizona. Lep. News 7(5-6): 146-147. BEUTENMULLER, W. 1897. Minutes of October, 1897 meeting of the New York Entomological Society. J. New York Ent. Soc. 5: 208. 1898. Revision of the species of Euchloe inhabiting Amer- ica north of Mexico. Bull. Amer. Mus. Nat. Hist. 10(13): 235-248. 1899. Notes on the American forms of Euchloe, Hubner. Can. Ent. 31(3): 56. BIRD, C. D. 1956. Collection of Rhopalocera from the Whiteshell Forest Reserve, Manitoba. Lep. News 10: 107-108. BISERTE, F. 1960. Chromatographie en chimie organique et biologique (E. Lederer, ed. ), vol. 2. Masson & Sons, Paris. 876 pp. BOISDUVAL, J. A. 1852. Lepidopteres de la Californie. Ann. Soc. Ent. France 2: X, 275-324. 1869. Lepidopteres de la Californie. Ann. Soc. Ent. Bel- gique 12: 11-94. BROOKS, G. S. 1942. A revised checklist of the butterflies of Manitoba. Can. Ent. 74(2): 31-33. BROWN, F. M. 1955. The Butterflies of Colorado, part IV, Pieridae, Papilionidae. Proc. Denver Mus. Nat. Hist. 6: 177-236. BROWNING, G. W. 1901. Collecting in the vicinity of Salt Lake City. Ent. News 12: 297-304. BUTLER, A. G. 1882. On the butterflies collected by Lord Walsingham in California. J. Linnaean Soc. 16: 462-474. 1899a. Notes on the genus Euchloe, Hubner, a genus of Pierinae. Entomologist 32: 1-3. - 1899b. Notes on the American forms of Euchloe Hubn. Can. Ent. 31: 19. CHANG, V. C. S. 1963. Quantitative analysis of certain wing and gem- talia characters of Pieris in western North America. Jour. Res. Lepid. 2(2): 97-125. CHERMOCK, F. H. & R. L. 1940. Some new diurnal Lepidoptera from the Riding Mountains and the Sand Ridge, Manitoba. Can. Ent. 72(4): 81-83. COCKRELL, T. D. A. 1899. On the origin of the genus Anthocharis, Bdv. {^Euchloe, Hb. ). Ent. 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F. 1962. The authorship of three scientific names of Nearctic Rhopalocera variously credited to Boisduval or Lucas. /. Lep. Soc. 16(1); 45-46. 1964. A synonymic list of Nearctic Rhopalocera. Lep. Soc. Mem. 1: 1-145. DOUBLEDAY, E. & J. W, WESTWOOD. 1847. The genera of diurnal Lepidoptera. Illustrated W. G. HEWITSON [Westwood died before the publication date]. Longman, Brown, Green and Longmans, Lon- don. 1: 250 pp. DEFOLIART, G. R. 1956. Annotated list of southeastern Wyoming Rhopalocera. Lep. News 10(3-4): 91-101. DOUDOROFF, M. 1930, A new aberration of Euchloe ausonides. Pan- Pac. Ent. 6(3): 143. DYAR, PL G. 1894. Notes on Pieris and Anthocharis. Can. Ent. 26: 110. 1902. A list of North American Lepidoptera and key to the literature of this group of insects. Bull. U. S. Nat. Mus. 52; 1-723. EDWARDS HENRY. 1880. Pacific Coast Lepidoptera. No. 22: 11 pp., No. 28: 2 pp. Henry Edwards, New York. 1881. On some apparently new forms of diurnal Lepidop- tera. Papilio 1(4): 50-55, EDWARDS, W. 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Microent. 25(1): 1-89. ELROD, J. E. & F. I. MALEY. 1906. The butterflies of Montana. Bull Univ Montana 30(biol. ser. ): 174 pp. FIELD, W. D. 1936. New Nortli American Rhopalocera. J. Ent. & Zool. 28(2): 18-26. FRANCK, G. 1905. Notes of Anthocharis at meeting of Brooklyn Ento- mological Society. Ent. News 16: 28-29. GARTH, J. S. 1934. Butterflies of tlie Boundary Hill Research Reserve, Yosemite National Park, Calif. Bull. So. California Acad. Sci. 33(3); 131-135. 1935. Butterflies of Yosemite National Park. Bull. So. California Acad Sci. 34(1): 37-75. GARTH, J. S. & J. W. TILDEN. 1963. Yosemite butterflies. Jour. Res. Lepid. 2(1): 1-96. GEDDES, G. 1883. List of diurnal Lepidoptera collected in the North- west Territory and the Rocky Mountains. Can. Ent. 15(12): 221-223. GOOD, P. M. & A. W. JOHNSON. 1949. Paper Chromatography of Pterins. Nature 163(4131): 31. GROTE. 1900. The descent of the Pierids. Froc. Amer. Phil. Soc. 39: 4-67. GUNDER, J. D. 1930. Butterflies of Los Angeles County, California. Bull. So. California Acad. Sci. 29(2): 39-35. HADORN, E. & H. K. MITCHELL. 1951. Properties of mutants of Drosophila melanogaster and changes during development as revealed by paper chromatography. Proc. Nat. Acad. Sci. 37 : 650-665. HERRICH-SCHAFFER. 1867. Prodromus systamatis Lepidopterorum. Corr.-Blatt. Zool. Min. Ver. Regnsh. 21: 138-144. HOLLAND, W. J. 1898. The butterfly book. Doubleday, Page, & Co., Garden City, New York. 382 pp. 1931. The butterfly book, rev. ed. Doubleday & Co., Gar- den City, New York. 424 pp. HOVANITZ, W. & V. C. S. CHANG. 1962. Three factors affecting lar- val choice of food plant. Jour. Res. Lepid. 1(1): 51-61. KIRBY, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera. John Van Voorst, London. 690 pp. KLOTS, A. B. 1930a. A generic revision of the Euchloini (Lepidoptera, Pierinae). Bull. Brooklyn Ent. Soc. 25: 80-95. 1930b. Diurnal Lepidoptera from Wyoming and Colorado. Bull. Brooklyn Ent. Soc. 25: 147-170. 1956. in: Tuxen, Taxonomist’s glossary of genitalia in in- sects, Lepidoptera. Ejnar, Copenhagen, pp. 97-111. LEIGHTON, B. V. 1946. The butterflies of Washington. Univ. Wash- ington Pub. Biol. 9(2): 47-63. LUCAS, P. H. 1852. Description de nouvelles especes de Lepidopteres. Revue et magazine de zoologie 4 (2nd ser.): 324-245. MAEKI, K. & C. L. REMINGTON. 1960. Studies of tlie chromosomes of Nordi American Rhopalocera, 2. Hesperiidae, Megatliymidae, and Pieridae. J. Lep. Soc. 14(1); 37-57. MARTIN, L. M. 1936. A new southern race of Euchloe ausonides (Di- urnal Lepidoptera), Bull. So. California Acad. Sci. 35(2): 94-95. 5 (1) :39-50, 1966 NEARCTIC EUCHLOE 49 MARTIN, L. M. & C. H. INGHAM. 1931. An annotated list of diurnal Lepidoptera of Huntington Lake region, Fresno County, California. Bull. So. California Acad Sci. 29(3): 115-134. MARTIN, L. M. & F. S. TRUXAL. 1955. A list of North American Lepi- doptera in the Los Angeles County Museum. Los Angeles Co. Mus. Sci. Ser. No. 18, ZooL, No. 8: 35 pp. McDUNNOUGH, J. H. 1938. Check list of the Lepidoptera of Canada and the United States of America, part 1, Macrolepidoptera. Memoirs So. California Acad. Sci. 1(1): 272 pp. MEAD, T. L. 1875. in: Wheeler, United State geographical surveys west of 100th meridian. 5( zoology): 1021 pp. Washington, D.C., U. S. govt, printing office. 1877. Notes on certain California diurnals. Psyche 2: 179- 184. MEINERS, E. P. 1956. Lepidoptera collecting at Ranken, Missouri. Lep. News 10(5): 163-168. MERRITT, J. R. 1952. Butterflies and hilltops. Lep. News 6(6-8): 101-102. MILLER, A. H. 1951. An analysis of the distribution of the birds of California. Univ. California Puh. Zoology 50: 531-644. MORRISON, H. K. 1883. List of species of butterflies collected in Arizona by Mr. H. K. Morrison, in 1882. Papilio 3(1): 9-10. MOSCHLER, B. 1878. Catalogue of the Lepidoptera north of Mexico. Stett. Ent. Zeit. 39: 299, 309. MUNZ, P. A. & D. D. KECK. 1959. A California flora. 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E. 1958. in: Hogue, Minutes of the fifth annual Pacific slope section meetings of the Lepidopterist’s Society. Lep. News 12(5-6): 209-212. SHULL, G. A. 1907. Life history and habits of Anthocharis (Synchloe) olympia Edw. Ent. News 18(3): 73. SKINNER, H. 1898. A synonymic catalogue of the North American Rhopalocera. Amer. Ent. Soc., Philadelphia. 99 pp. — — ... 1902. Lepidoptera of Beulah, New Mexico. Trans. Amer. Ent. Soc. 29: 36-38. 1903. in: Smith, Check list of the Lepidoptera of boreal America. Amer. Ent. Soc., Philadelphia. 136 pp. 50 PAUL A. OPLER /. Res. Lepid. SMITH, J. B. 1891. List of the Lepidoptera of boreal America. Amer. Ent. Soc., Philadelphia. 124 pp. SPULER, A. 1908. Die schmetterlinge Europas. Stuttgart, Germany. Band 1, 385 pp. STORER, T. I. & R. L. USINGER. 1963. Sierra Nevada natural history, an illustrated handbook. Univ. California Press, Berkeley & Los An- geles. 374. pp. STRAND, E. 1914. Nordamerikanische insbesondere Californische Lepi- doptera. Archiv. F. 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Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 BUTTERFLY AGGREGATIONS WALFRIED J. REINTHAL 4026 Sequoyah Ave., Knoxville 19, Tenn. Probably a good many collectors have, at one time or an- other, during their field trips seen butterflies accumulating, by the dozens and hundreds, in very small places. This is particu- larly true of the tropics. Species belonging to certain families, particularly the Papili- onids,, Pierids, Nymphalids, and Lycaenids are known to have this habitual tendency. The habit to aggregate in large numbers has obviously devel- oped from the feeding peculiarities of these insects. Feeding peculiarities, on the other hand, are greatly influenced by an ir- resistible attraction that certain odors impose on these butter- flies. Very often it is just a natural dampness caused by rain or running water on roads, river banks, lakes, and sea shores, or other wet places that may attract these thirsty insects. Odors, sometimes very obnoxious to the human nose, caused by the de- caying plants or animal bodies, human or animal excrement and urine, remnants of food, drinks, or of other materials, may at- . tract butterflies in smaller or larger numbers. In the following paragraphs I would like to report on a few cases drawn from my past experiences in the field. PAPILIO GLAUCUS CANADENSIS IN ADIRONDACKS. It was during a collecting trip in the Adirondacks, upper New York State, in June 1956 when this observation was made. The Papilio glaucus canadensis were out in large numbers. Only fresh males were flying and the very first females had just started to appear. The shoulders of the dirt roads were actually swarming with these attractive yellow fliers, and particularly some of the dips on the road, where more moisture had accumulated, and also some damp roadside ditches seemed to attract the hungry insects. 51 52 WALFRIED J. REINTHAL /. Res. Lepid. On Schroon River Road, I drove on a gravel road running through a sparsely populated area and through a partially shady woods. At some distance from a small puddle in the middle of the road I stopped my car. My attention was attracted by a kind of activity on the damp gravel where some fifty males of cana- densis were sitting side by side, all with wings closed, and en- joying the presence of moisture. I approached the spot as cau- tiously as possible, moving the last few feet on “all fours,” fixed my camera at a distance of about two feet and took a few pic- tures. Not one specimen became disturbed. It was pleasantly quiet there without any movement in the air. Next day I concentrated my attention on another dirt road leading from Indian Lake to Cedar River Flow. It was a nice warm day, June 17th, and the canadensis was again swarming along the sides of the road. I was looking for a larger aggregation of canadensis to get a few more pictures. Smaller groups, consisting of about twenty to thirty specimens, were observed. Then, came what I was really looking for. On a dry gravel roadside, just about a foot or two from the very mar- gin of the road, I noticed a large yellow spot. Low grass, sedges mostly, but no flowers, sparsely covered this roadside. It was on this yellow spot, measuring about one and a half feet in diameter, that I noticed about one hundred and twenty canaden- sis sitting, side by side, with wings closed. As I approached cau- tiously, a few flew away but I managed to take two pictures. Then I covered the spot with my eighteen inch collecting net. The entire aggregation flew up, and my large net was filled with frightened Papilionids. I started taking them out one by one, examining and counting the specimens, and afterwards giv- ing them their freedom. Many escaped my counting but, never- theless, I counted eighty-five in all. \'\hthout exception they .were all fresh males, not a single female was among them. I examined the spot closely and concluded that either the presence of mois- ture or some curious odor was the reason for the aggregation. PAPILIO GLAUCUS IN CATSKILLS. One interesting observation comes from the Catskill Mountains in New York State. It occurred in July 1956. After collecting for half a day in the picturesque Big Indian Valley, I was approach- ing the end of the valley on a gravel road where it joins the high- way. It was about three o’clock in the afternoon, and I was getting hungry. 5(1) :51-59, 1966 AGGREGATIONS 53 Stopping my car under a shady birch, I was ready to open my lunch box when I saw a movement. It was in a dry pasture, and I had to climb a stone fence to get closer. What attracted me was fresh cow dung on which a large variety of butterflies were feeding. I covered the fresh brown mass trying not to smear my net. After I sorted the insects from my net there were about fifty Papilio g. glaucus, four Limenitis arthemis, and two Limenitis a. astyanax. This is a good example that the \animal excrements sometimes attract a large number of certain butter- flies. This time they were Papilionids and Nymphalids. PAPILIOXIDS IN THE SMOKIES. Speaking of the Papilionids, another experience comes to my mind. Collecting in the Smoky Mountains of Tennessee on May 15th, 1960, there were many philenor, troihis, and glaucus flying in the afternoon along the gravel road in one of the canyons. The philenor outnumbered the other two, and there were very few glaucus. There seemed to be more specimens around the picnic places and on the sections of the road where water was dripping from the canyon walls. I had to descend to the river in order to wash my hands, and while walking down my at- tention was attracted to a small hole between the rocks. In the hole, which was no larger than eight inches in diameter, there were a dozen or more butterflies sitting. Most of them were quietly sucking, however, a few were moving around restlessly. I covered the hole with my net and caught all of them and ex- amined them. There were: eight glaucus, three troilus, six philen- or, three Erynnis horatius, and three Epargyreus c. clarus in this social company. There was very little moisture in the hole. I found four glaucus wings and what looked like the remnants of their decaying bodies. It was my impression that someone had stepped on a few glaucus, which were sucking on moisture, and that this was responsible for the hole. The possibility also exists that a reptile, like a lizard, partly consumed the glaucus. I stepped down to the stream and continued my observation of the place for any further development. After awhile, a male glaucus was flying above the middle of the stream. As it ap- proached me it veered sharply in the direction of the hole, where it landed with the velocity of a bullet. Another male glaucus repeated the same maneuver a few minutes later. No doubt, all 54 WALFRIED J. REINTHAL /. Res. Lepid. the above insects were attracted to the place by the odor of the decaying bodies of the dead butterflies, an odor not perceptible to the human nose but very much perciptible to the fine olfac- tory organs of these butterflies. This last incident reminds me of another day of collecting in the Adirondack Mountains of New York. After catching my first glauctis canadensis in the morning, I squeezed its body and discarded the specimen on the road. I remained at the same place to catch other glauctis that were flying by. I collected about twenty males in one hour, either sitting on or flying around the dead specimen. It was amazing to see even other specimens, which were flying by on the opposite side of the road, suddenly crossing the road to land on or near the dead specimen. In this case again, there was no doubt that it was not the yellow colora- tion of the dead specimen that attracted the males so persistently but the odor of the dead body. I repeated this procedure in the afternoon but it seemed to work only in the forenoon and only with freshly squeezed specimens. When a dozen males, which were squeezed in the morning and dried during the day, were placed in the same area early next day these bodies did not at- tract any butterflies. HAIRSTREAKS. These are another group of butterflies which occasionally tend to aggregate in one place particularly preferable to them from the standpoint of feeding. I have had several cases where I en- countered large numbers of one or more species of Strymon sitting on one large flowering plant. These are my observations. On a trip through the Oklahoma Panhandle to New Mexico several stops were made along the highway in Dewey County, south of Soiling, Oklahoma. In those places the large yellow spots on the roadsides indicated the presence of one of the most desirable plants. Butterfly Milkweed {Ascelepias tuherosa), to hairstreaks and to many other diurnals. It was in June 1952, and the plant was in full bloom. I approached one yellow cushion, about two feet in diameter, and I noticed that it was so densely covered with Strymon that there was hardly a place for additional ones. An estimation was made that there were more than two hundred specimens on that plant. From one comer of the plant I was able to gather the following: Ten males of Strymon titus mopsus, twenty-nine males and two females of Strymon Ontario autolycus, and twelve 5 (1) :51-59, 1966 AGGREGATIONS 55 specimens of Strymon f. falacer, all together, fifty-three hair- streaks. A more careful study of other similar plants in that area disclosed that titus mopsus made up about twenty percent of the entire number of specimens sitting on one plant. The rest being about equally divided between the other two Strymon, namely the falacer and Ontario autolycus. Besides tlie hairstreaks there were a few Melitaea i. ismeria and a few Hesperids. It also should be pointed out that the titus mopsus specimens were all fresh, Ontario autolycus very fresh to fresh, and those of falacer fresh to worn. Four days later, while returning the same way, a stop was made at the described place of the Butterfly Milkweed and the entire picture repeated itself. Same species of hairstreaks were collected on the plant, except that they were not as fresh this time. Besides the Strymon, there were a few Nathalis iole and Vanessa cardui. In similar surroundings I had a commensurate experience in April 1953 on a highway four miles west of Mineral Wells, Palo Pinto County, Texas. The flowering Butterfly Milkweed plants on the roadside were covered with hairstreaks. I chose one of the more spectacular plants and after covering it with my net I had it full of Strymon. Some four or five dozen were collected. These made a small compromised part of all that were feeding on the plant. Unfortunately, I was not immediately cognizant of what species these Strymon represented. After it was checked at home, it turned out to be Ontario autolycus, a species having its main distribution in Oklahoma and Texas, and in parts of some adjacent states. These are quite desirable to collectors. There were only single specimens obseorved on plants other than Butterfly Milkweed in that area. Only a few were collected while flying in a nearby oak thicket. The next year I returned to the same place to get more of the species. Unfortunately, the entire area, including that plant life, was changed because of the construction of a new road. The third case of a mass occurrence of Strymon was observed in a different biotype involving another kind of plant. This time it was in June 1958 in Boiling Springs State Park, Woodward County, Oklahoma. The species involved was Strymon a. alcestis, and the plant that attracted this species was the Hemp Dogbane (Apocynum camnahinum). The plant was in full bloom at that time. The weather was very hot and humid: 105 degrees; with about 80% relative humidity. At this particular time of the year the above mentioned temperature and humidity are not unusual 56 WALFRIED J. REIXTHAL J. Res. Lepid. for this part of the country. Following a female Asterocampa an- tonia in a small forest glade not far from the Park spring I dis- covered a dense growth of Hemp Dogbane. On almost every plant several Strymon were feeding or flying from blossom to blossom. I collected several alcestis males, partly worn, and females, nearly fresh. Besides alcestis there were only a few other butterflies present. Namely, Strijmon cecrops and Atalopedes campestris. The most recent observation involving one of our most com- mon Lycaenids, Celastrina a. pseudargioliis, was made in March 1961 in the Great Smoky Mountains National Park. In two dif- ferent canyons, one on the Tennessee and the other on the North Carolina side of the Park, this species, which had been very plentiful that year in the Smokies, was seen aggregating in groups of fifty to three hundred specimens. These groups were found accumulating in spots on the gravel roads. What exactly attracted the “blues” in such numbers can only be guessed at. Possibly remnants of food or drinks left by pic- nickers, or it could have been gasoline from a leaky car. Any- way, the “blues” were so absorbed in their activities that one could almost step on them without disturbing them. In one place a few other species were associating with the “blues,” but only as single specimens. Namely, Papilio g. glaucus, Graphium mar- cellus, and Erynnis juvenalis. SKIPPERS. Hesperids seldom tend to accumulate in one place in great numbers, but there are exceptions. I have seen Erynnis, sucking by the dozens, on damp spots on the roads, particularly in the morning and forenoon. But what I observed in April 1960 at Fall Creek Falls State Park, Tennessee, was something unusual. The fresh males of juvenalis were plentiful on the Park roads, particularly on road bifurcations and in parking areas. When I came to a camping area resented for organized groups, where a number of cabins and a larger building with a kitchen were occupied by some biology students, I observed the follow- ing: around the mess hall, where some food was being cooked, swarms of Erynnis juvenalis, estimated at about two hundred, were flying around the damp foundation of the building. They were on both the shaded and the sunny sides of the building. Whether or not it was the foundation dampness or some kitchen odor which attracted them I do not know. I lean toward the in- terpretation that the dampness and the peculiar odor of the 5(1) .-51-59, 1966 AGGREGATIONS 57 foundation was the main attraction. A dozen specimens could be netted at one sweep. All of them were fresh males and in a large majority belonged to jiivenalis. Only two fresh Erynnis persius males and a few Erynnis brizo males were among them. Another time, the following observation was made on a gravel road in the Smokies of Tennessee, May 1960. After leaving my car on a stream bank, I saw accumulations of Erynnis icelus in spots no larger than a foot wide. I tried to count the specimens aggregated in one of these places. There were twenty to thirty icelus sitting close together in one spot, no wider than three or four inches, and seemingly feeding on something. A closer ex- amination did not reveal anything in particular which could have attracted the skippers. There was no dampness present. The gravel was entirely dry. But, people had been eating in this area, and possibly some spilled juice, Coca-Cola, beer, or rem- nant of food dropped in this place were attracting the insects. LIBYTHEANA IN RIO GRANDE. Libytheana bachmanii is a species that one usually catches just a few specimens at a time in places where the Hackberry, its food plant, grows. Seldom does one see large numbers of this species in one place. In October 1951 while on a highway three miles east of Laredo, Texas, I tried to catch a few fresh Kricogonia lyside that were flying around in large numbers and I stepped into swarms of bachmanii. All the grass along the roadside and in the shallow ditch, mostly on the south side of the road, was covered with fresh specimens. There was a very light southerly breeze present. My first impression was that the butterflies were migrating. But the specimens seemed to be very fresh, as if they had just emerged, and not flying around or going in any one direction. They seemed to be disturbed only by my footsteps in the grass. It seemed that in some spots there were more of them congre- gated, and stepping into those spots, I caused thirty to forty specimens to fly around for a short while and then settle down again in the grass. At times, when the breeze got stronger, swarms of bachmanii flew across the road in a northerly di- rection. Perhaps it was a migration, and the butterflies were resting there from the previous day’s flight. This mass occur- rence of Libytheana continued for miles and miles with chang- ing density and there were millions of specimens along the highway. 58 W'ALFRIED J. REINTHAL }. Res. Lepid. I moved farther east as the daytime advanced. The southerly breeze increased, and swanns crossing the road containel a large number of Kricogonia and single specimens of fresh Phoebis agarithe maxima. Mass occurrence of hachmanii continued throughout that day, while I traveled a distance of about ninety miles toward Rio Grande City. Of course, they were a nuisance to my collecting! The next day, as I continued my way toward McAllen, there were still a large number of specimens flying around but not the swarms of the previous day. Was this a migration from Mexico or just an unusual mass occurrence of this species that year along the Rio Grande? I do not know. XYMPHALIDS L\ COLORADO. I have seen large numbers of Nymphalids, Limenitis archippus ivatsoni, L. arthemis astyanax, Anaea andria, Asterocampa celtis, A. clyton, A. texana, and A. antonia in Oklahoma and Texas gathering on fallen rotten fruit, particularly decaying peaches and pears. The fig orchards in Louisiana, for instance, are a good place to collect Nymphalids like Polygonia and Asterocampa. The following observation was made on different butterfly species and on different bait during a collecting trip in July 1949 in Walnut Gulch, in Gunnison County, Colorado at an eleva- tion of about 9000 feet. Pursuing the interesting Euphydryas maria alena in that nar- row gulch, I noticed a small aggregation of some twenty but- terflies, Melitaeini and Lycaenini, sucking on a small amount of black excrement of an undetermined origin. When I returned five days later, after it had rained several times, my attention was again drawn to the above mentioned spot. I was fortunate enough to get most of the butterflies I found sitting on the same bait. The ones I collected consisted of: Melitaea palla calydon, Phyciodes camillus, P. tharos pascoensis, and Lycaena amyntula, with single specimens of Thorybes nevada and Erynnis persius fredericki among them. The bait was kept under observation for thirty minutes, and the same species kept coming and alighting on the excrement. Other species were flying at the same time along the dirt road and around the same spot but none of them seemed to be in- terested in the bait. These species were Papilio rutulus, P. bairdi brucei, Colias alexandra, C. scudderi, Pieris napi macdunnoughi, and Oeneis idileri reinthali. 5 (1) :51-59, 1966 AGGREGATIONS 59 REFERENCES BROWN, F. MARTIN. 1955a. Golorado Butterflies, part 2, Danaidae, Heliconidae, and Nymphalidae. Denver Mus. Nat. Hist. Proc., 4:33-112. 1955b. Golorado Butterflies, part 3, Libytheidae, Riodini- dae, and Lycaenidae. Ibid., 5:113-176. 1956a. Golorado Butterflies, part 4, Pieridae and Papilioni- dae. Ibid., 6:177-236. 1956b. Golorado Butterflies, part 5, Hesperiidae Megathy- midae. Ibid., 7:237-331. CLARK, AUSTIN H. 1932. The Butterflies of the District of Colum- bia and Vicinity, Smithsonian Institution Bull. U.S. Nat. Mus., 157, 337 pp. CLARK, AUSTIN H. and LEILA F. 1951. The Butterflies of Virginia, Smithsonian Misc. Call., 116 (7): 1-239. FIELD, WILLIAM D. 1940. A Manual of the Butterflies and Skippers of Kansas, Lepidoptera and Rhopalocera. Bull. Univ. of Kans., 39: 1-328. HOLLAND, WILLIAM D. 1931. The Butterfly Book, (revised edition). Doubleday & Co., Garden City, N.Y., 424 pp. KLOTS, ALEXANDER B. 1951. A Field Guide to the Butterflies, Houghton Mifflin Co., Boston, Mass., 349 pp. LAMBREMONT, EDWARD N. 1954. The Butterflies and Skippers of Louisiana, Tulane Stud. ZooL, 1:125-164. MATHER, BRYANT and KATHERINE. 1958. The Butterflies of Mis- sissippi, Tulane Studies ZooL, 6:, 2:63-109. V ][ *1 •'*i j ’ ' * f , 1 3t • SJ.-.,** fV ' »*Vi> Xvi ■life. ,w •■ ’M'l. ' ’■ 1 •- . ,'f' < ■ ' % ■’It fa V\ .<■ J !’r t 'ii'i* I- j ♦■' yi ,1%. . ■ ni''^', ' ', , i. -.iVyw-fl '.I ■- "U. «:■ \ :-V.r';>> ' A\ i ? ^ Ci ^,' '■ t ‘ ^ V'' ■ ’ ■ ■• '•''■’ 1^-" f>:. ■■%•• ^ . I ir-‘ - - . ,■ . X '.. -v / 'AW ., ; ■■ ' U' . .^-y ■ r i .S . - 'ft-' fi ' . . • iterii f ■ i. ■%•';,* : I ■■'t'>'i ■ Hi • 'i'*^ ",■ ' ' HI ^ ; T.;'< I i. ■■■' y .K, •■ tS'J i'.; ••^Oisl': NOTICES CHANGE OF ADDRESS: When sending a change of address, please include both old and new addresses. This will greatly facilitate your changeover. ZIP CODE: If your zip code is not present, or is incorrect, on your address plate, please inform us as the Post Office requires it on all domestic mailings. BUTTERFLIES OF THE DELAWARE VALLEY, by Arthur M. Shapiro. Special Publication of the American Entomological Society, Philadel- phia, Pa. i-vi, 1-79, 1 3 b. and w. plates, keys and checklist. ^1.50 from publisher at 1900 Race St. , Philadelphia, Pa. AUTOMERIS AURANTIACA, living ova or cocoons urgently required to complete research project, contact Brian O. C. Gardiner, 18, Chesterton Hall Crescent, Cambridge, England. LEPIDOPTERA OF FLORIDA , by C. P. Kimball. 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If in good shape and typewitten in the form used in the Journal much time and labor will be saved, COLOR PRINTS: Submit color preferably in the form of 35mm pos- itive transparencies in order to save on cost and labor. THJl JOUI^NJAL OF RESEARCH on THE LEFIJDORTERA Volume 5 Number 1 March, 1966 IN THIS ISSUE Forelegs of Butterflies I. Introduction: Chemoreception Richard M. Fox 1 A New Species of Epinotia Hubner from British Columbia ( Olethreutidae ) T. N. Freeman 13 Studies in the Life Histories of North American Lepidoptera California Annaphila II. J. A. Comstock and C. Henne 15 New Skipper Records for Mexico H. A. Freeman 27 A Moth Sheet Noel McFarland 29 Cover Photo: Daritis P howardi Hy. Edw. larvae Noel McFarland 36 Rediscovery of Annaphila casta Hy. Edw. in California (Noctuidae) John S. Bucket! 37 Studies on the Nearctic Euchloe. Parts I, II Paul A. Opler 39 Butterfly Aggregations Walfried J. Reinthal 51 THE JOU^HA.L @F RsESlARsCHJ @NJ THE LEFiJ©©FTEFsA Volume 5 Number 2 June, 1966 ywmm. !' .“•id'' Journal of Research on the Lepidoptera 5 (2) :61-96, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 MORPHOLOGY OF THE IMMATURE STAGES OF EVERES COMYNTAS GODART (LYCAENIDAE) DONALD A. LAWRENCE and JOHN C. DOWNEY Carhondale Community High School, Carhondale, Illinois and Southern Illinois University, Carhondale, Illinois Knowledge of the external morphology of the immature stages of butterflies is quite incomplete. Most of the past work has dealt with moths, and on groups which many researchers consider the more “generalized” immature Lepidoptera. The more “specialized” groups, including the Lycaenidae, have been neglected. It is hoped that this detailed study of one species will provide a morphological platform on which to launch more intensive comparative studies within the family. To that end we have arranged the description of the larval morphology by individual structures rather than the more conventional treat- ment of describing each instar. This makes more emphatic the onto genic changes in addition to any individual variations. The latter must be recognized before the former become meaningful. Everes corny ntas Godart is common throughout the Nearctic region and its range extends southward as far as Costa Rica, Eggs and larvae are found on white clover {Trifolium repans L.), red clover (T. pratensis L. ) and lespedeza {Lespedeza sti- pulaceae Maxim.). Pupae have also been found on lespedeza and curly dock (Rumex crispus L. ). Details of the population dynamics of this species will be published elsewhere. PREVIOUS DESCRIPTIONS OF THE IMMATURE STAGES Edwards (1876) gives an account of the early stages of Everes comyntas and gives measurements of the egg, larval stages and pupa. No illustrations were included. Scudder (1889) describes and gives three illustrations of the egg, including two views of the micropyle (loc. cit., Vol. 3, PI. 68, figs. 5 and 12) and an oblique view of the entire egg {loc. cit., Vol. 3, PL 65, fig. 20). The latter appears to us to have too few reticulations in the surface. The same author has illustrated a dorsal view of the first instar larva, several head capsules, and pupae, all of which are accurate but, partly due to small size, lack specific details. Dethier (1941) pictures the larval antenna of comyntas in his 61 62 LAWRENCE AND DOWNEY /. Res. Lepid. Fig. 1. Immature Stages of Everes cornyntas Godart. A— B. Egg; A. Lateral view. B. Dorsal view of micropylar area. C- — D. First instar larva; C. Dorsal view. D, Lateral view. comprehensive paper on the antennae of Lepidoptera. The im- mature stages are superficially described or illustrated in sev- eral other more general works, but these are overly simplified and are usually not diagnostic either of the species, or in some cases, the family. 5 (2) :61-9e, 1966 IMMATURE EVERES 63 THE EGG Figure IB is a lateral view of the egg which shows its typical lycaenid form. It is of turban shape, flat on the bottom and less noticeably truncate on top. The sides and top are covered with reticulations which are slightly elevated from the surface. Rounded prominences are found at the intersections of this net- work of ridges, giving the egg the appearance of an echinoid test. The diameter is 0.6 rrim; the height, 0.3 mm. The micropylar area (Fig. IB) is only slightly depressed into the central vertical axis of the upper surface. Its central cavity consists of either three or four petal-shaped cells, open at their bases. These central cells are surrounded by other cells whose walls are of similar structure except that each cell has a com- plete wall. They vary from semi-round to hexagonal in shape. From one to three of these cells may occur between the petal- cells and the transition zone (see below). The walls of all cells in the micropylar area are uniform in thickness and appear to have a slight greenish tint under high magnification. Various unknown inclusions showing this same tint occur in most of the peripheral cells of the area. This region of the egg is of the same general type as other Nearctic Plebejini but is diagnostic of this species. There is a small transition zone between the area of the micro- pyle and the elevated reticulations which characterize the dor- so-lateral margins and sides of the egg. Although the lateral limits of this zone are often difficult to judge, the region is dis- similar to the other areas in that there is no regularity or uni- formity to the walls of the opaque cells involved. Radiating out from the transition zone is a network of raised, solid ridges arranged in tolerably regular oblique rows running across the top and down the sides of the egg. These are connected by another series of slender ridges which also run in an oblique direction, but at right angles to the first ridges. As indicated above, there is a slight prominence or knobby elevation at the intersection of each ridge. The knobs are largest and most obvious on the sides and obsolescent near the transition zone and on the ventral surface. The ridges appear to be con- tinuous with the walls of the cells in the transition zone, but they are marked by their elevation, prominence, knobbed inter- sections and regularity. The surface of the egg opposite the micropyle contains hexagonal cells, defined by their walls, which are uniform in thickness and, except for their regular shape, resemble the cells in the transition zone on the dorsal 64 LA\\ RENCE AND DOWNEY /. Res. Lepid. surface. The areas between the walls are transparent in hatched eggs, though the entire area may be obscured by the substrate of maternal origin used to attach the egg to a surface. In the living egg the thickened ridges appear white in color while the depressed areas between them are light green, reflect- ing the embryonic contents. A slight darkening of the punctua- tions occurs before hatching, or if the egg is parasitized. The general appearance of the egg, size, reticulations and micropylar area are distinguishing features which help separate this species from allied forms. THE LARVA CRANIUM: The terminology of the sclerites and sutures of the cranium, especially as concerns the lepidopterous larva, has undergone much revision in the past two decades. Short (1951) reviews the various studies concerned with determining the na- ture of the sclerites and sutures of the cranium. One can only conclude from this review that no set of terms has as yet been worked out which would be satisfoctory to all entomologists. The system of Hinton (1947) has been used herein with slight modifications when referring to specific sclerites or sutures. The cranium of E. comyntas lacks conspicuous lateral cleav- age lines ( frontal sutures of Hinton or adfrontal sutures of many authors), see Fig. 2 A. Other lycaenids including Plebejus icar- ioides (Bdv. ) and Plehejiis melissa (Edw. ) also lack these frontal sutures. Most moth larvae have the sutures, although as Ripley (1923) points out, they may not appear except in the mature larvae. These lines are not mentioned by Clark and Dickson (1956) in any of their detailed studies on numerous species of South African lycaenids. Measuring the head capsule width is a reliable method for determining the four larval stages of this species. There is no overlap of head capsule width between instars. The ranges of head capsule width are as follows (measured between widest points ) : First Instar .186— .207 mm. Second Instar .285— .313 mm. Third Instar .492— .519 mm. Fourth Instar .855— .923 mm. In addition, a nearly constant I to I ratio was obtained in all instars by dividing the width of the fronto-clypeal apotome (taken as the distance between seta Cl on the right side and Cl on the left) by its length (taken as the distance between the anteclypeus (ac) and apex of the triangle formed by the lat- eral adfrontal sutures (adl) (See Fig. 2A). 5 (2) :61-96, 1966 IMMATURE EVERES 65 A B Fig, 2. A — B. Chaetotaxy of head capsule, first instar larva; A. Frontal view, B, Lat- eral view. C. Head capsule, fourth instar larva, ventral view. D. Hypopharynx, spin- neret and associated structures, third instar larva, lateral view. Key to lined ab- breviations: ac=anteclypeus, adl=:lateral adfrontal suture, adm=median adfrontal suture, am=arm of mentum, an=;antacoria, bs=basistipes, car=:cardo, cl=clypeus, cs=cervical sclerite, ds=dististipes, fg=fusulus, ga=galea, hp=hypopharynx, la= labium, lp=labial palp, ma=mandible, me=niaxillulae, mn=mentum, mp=maxil- lary palp, oc=ocellus, pfr=palpifer, pgr=palpiger, ptg=postgena, sm=submentum, sp=spinneret. Other lettering refers to specific setae, lenticles or ocelli. 66 LAWRENCE AND DOWNEY /. Res. Lepid. Crumb ( 1929 ) found many positive differences in the arrange- ment of the head setae and punctures in different species of Noctuidae, with a strong tendency toward generic homogeneity. The same author cautions, however, that the degree of individual variation necessitates the examination of a series of specimens before positive conclusions are drawn. In the discussion below of the setae and punctures, the “normal” position or arrangement is given only after numerous larvae of each instar were studied. Chaetotaxy of the cranium-. Even more than the cranium, lar- val chaetotaxy has afforded much discussion in the literature, often involved with attempts either to homologize or to compare the systems used in specific cases by various authors. Thus Peter- son (1951) gives a table comparing names used by Forbes and Heinrich; Mutuura ( 1956 ) tabulates and compares the systems of Fracker, Gerasimov and Hinton with his own; and Hasenfuss (1960) compares the chaetotaxy systems of Gerasimov, Hinton and Bollmann. McKay (1963) discusses some problems and briefly compares certain setae of Mutuura and Hinton, but uses the system of the latter. The reader is referred to these authors for specific details on nomenclature and homology. We have used the terminology of Hinton (1946). For ease in following the discussion of the setae below, the reader is asked to refer to Figs. 2A, 2B and 3H. Setae and punctures of the left side only are discussed unless otherwise indicated. Frontal (A) and Clypeal (C) Setae and Punctures: In the first and second instar the longest seta, Gl, is found in the lateral angles of the isosceles triangle formed by the lateral ad- frontal sutures. Just medial to Gl is a puncture, here called Ga, not mentioned by Hinton. About two thirds of the way to the midline from Gl is another seta, G2, slightly shorter than Gl. Above G2 on the frons but nearer the midline is a puncture Fa. No other setae or punctures are found within the triangle on the first or second instars. With each succeeding instar Ga is pro- portionately nearer Gl than G2. Another, smaller, clypeal seta is found on the third instar just medial to Ga. Anywhere from one to four additional setae may be found above the frontal puncture and within the triangle. Three clypeal setae not found on the first instar are found on the fourth instar between G2 and Ga. Their position is variable within this area and some of them may be absent. As few as six and as many as eleven setae may be present above the frontal punctures and within the triangle on the fourth instar. Adf rental Group ( AF ) : In the first instar lateral to and paral- Fig. 3. — E. Right mandible; A. First instar, oral aspect. B. Fourth instar, oral aspect. C. same, line drawing of optical section. D. First instar, edge, posterior aspect. E. Fourth instar, edge, posterior aspect. F^ — G. Maxilla; F. Left maxilla frontal- dorsal view. G. Right maxilla frontal-dorsal view. H. Chaetotaxy of the left subocel- lar area, head capsule, first instar larva. I — J. Antenna, frontal-lateral view. I. Fourth instar. J. Second instar. K. Schematic section through meso-thorax of first instar larva showing lateral ridge and position of major setae on several adjacent segments. ling the lateral adfrontal suture are three setae and a puncture. These are called respectively from the lower-most to the upper- most; AF3, AFl, AF2, and AFa. The setae are roughly equal in length with AF2 sometimes shorter. AF3 is usually found on a horizontal line with the frontal punctures Fa. AF2 and AFa are variable in relation to one another and have been found with their positions reversed on later instars. No change in 68 LAWRENCE AND DOWNEY /. Res. Lepid. arrangement and no additional setae or punctures are found within the area of this group on any of the succeeding instars. Anterior Group (A): Two setae and a puncture make up this group on the first instar with the longer seta (Al) just above the antenna. A2 is caudal to Al and is about one-fifth to one- fourth its size and is associated with ocellus IL The puncture Aa is caudal and somewhat lateral to A2. It is more closely associated with ocellus II in the first instar than in later instars. Puncture Aa is about equidistant from ocellus I and II in the second and third instars, but on the fourth instar it is usually closer to ocellus L An additional seta is found within this group in some third instars. In individuals of the fourth instar as many as three additional setae may be found. Ocellar Group (O): The ocellar setae are about equal in size. Seta 01 is usually equidistant from ocellus II and III. Seta 02 is caudal to 01 and ocellus I, but usually nearer to ocellus VL Puncture Oa is caudal to ocellus V and below ocellus VI, but with each succeeding instar takes a more caudal position with respect to the latter. Rarely there is a third seta just caudal to ocelli IV and V on the third instar. This additional seta is constantly present on fourth instars. Siihocellar Group (SO): On the first instar S02 is the longest seta of this group and is located just behind ocellus V. It is slightly longer than seta Al. Seta SOI is found just lateral to the antenna between two punctures, SOd and SOe, and is a third to half the size of S02. Seta S03 is caudal to S02 and is about equal to it in size. There are five punctures in this group, but only SOa is named by Hinton. This is located below S02. The others have, been mentioned in relation to SOI; the other two, SOb and SOc, which are closely associated are lo- cated below SOa. Puncture SOc is peculiar in shape and is much longer than wide in comparison with other punctures. It is very constant in position and may have taxonomic significance. Ap- parently most larvae in the Lepidoptera do not possess this punc- ture, at least it is lacking in most figures and illustrations. The second instar has the same arrangement as the first. Ad- ditional subocellar setae variable in number and position can be seen on the third and fourth instars. Lateral (L) and Genal (G) Setae and Punctures: On the first instar puncture La is considerably behind ocellus L Seta G1 is a very minute seta located on the gena. Behind G1 and above it 5 (2) :61-96, 1966 IMMATURE EVERES 69 Fig. 4. A=— D. Labra of larvae, showing setal pattern, facial aspect on left and oral aspect ( epipharynx ) on right of page. A. First instar. B. Second instar. C. Third in- star. D. Fourth instar right half of labrum only. is the puncture Ga, The distance between GI and Ga is quite variable, but positional relationships are constant. No additional lateral or genal setae and/or punctures are found on the later stages. The distance between Gl and Ga on the third and fourth instars is much more constant, however. 70 LAWRENCE AND DOWNEY /. Res. Lepid. Posterior Group (P): Hinton’s posterior group is made up of two setae and two punctures, but only one puncture of this group is found on E. corny ntas. This particular puncture is called Pb and is so closely associated with the setae and punctures of the vertex that it would seem to belong to this group. Puncture Pb is found on all instars in the same position. Setae and Punctures of the Vertex (V): Behind Pb and ex- tending caudally is a group of three setae and a puncture; VI, V2, Va, and V3 in order. The setae are all minute and about the same size as Gl. They are constant in position on all instars. Medial to VI and V2 is a puncture for which Hinton shows no homology unless it represents a puncture formerly belonging to the adfrontal or posterior group, which has migrated in a caudal direction. It is here considered as Vb. Appendages of the Head. The Labrum, Epipharynx: the lab- rum, Fig. 4A to 4D, does not differ greatly from that of any other lepidopterous larva. It is notched, but this is not unique and the depth of the notch is subject to considerable individual variation. The typical first instar labrum bears six pairs of setae on its facial surface (see Fig. 4A) and three pairs of broader more bladelike setae on the oral surface, more properly termed the epipharynx. (Unless otherwise indicated the setae and punctures in the following discussions are those of the left side only. ) The setae are labeled following the system used by Forbes (1911). Seta i is located on the labrum near the midline, about halfway between the distal margin and the anteclypeus. A puncture is found lateral to, and somewhat behind, seta i. Setae ii is the longest seta of the group and is located lateral to seta i. Of the four marginal setae, seta in is the most posterior of the series and is on the lateral margin; seta iv is the longest of this group and setae v and vi are both subequal to in and are the smallest in the series. The arrangement of the setae and punctures on the labrum of the second, third, and fourth instars is the same except the possible inclusion of one or two additional setae in the fourth instar. The first instar larvae has one, and the other instars two, punc- tures on the epipharynx. The blade-like setae do not vary in structure and change but slightly in position between instars. The apices of these peculiar structures are directed toward the mid-line and when seen from below, appear to project slight- ly into the pre-oral cavity. Each appears to have a circular clear 5 (2) :61~96, 1966 IMMATURE EVERES 71 Fig. 5. A- — D. Distal proiegs of larvae showing arrangement of crochets; A. Medial view of proleg, first instar. B. Medial view, anal proleg, first instar. C. Medial view of proleg, fourth instar larva with fleshly spatulate lobe belov/ crochets. D. Ven- tral view of same. E' — J. First instar Thoracic legs; E, G, I, lateral views of left pro-, meso- and methathoracic legs. F, H, J, medial views of right pro-, meso- and methathoracic legs. K— N. Type G setae from first (K), second (L), third (M), and fourth (N) instar. O — R. Longest seta on dorsum of fourth abdominal segment from first ( O ) , second ( P ) , third ( Q ) , and fourth ( R ) instar larvae. S— microsetae from first instar larvae. T- — W. Spiculate setae from second (T), third ( U ) , and fourth ( V ) instar larvae and from the pupa ( W ) . Y — example of major seta from head of pupa from location shown, in X. 72 LAWRENCE AND DOWNEY J. Res. Lepid. area near, but not at the base, which perhaps represents the j internal out-pouching of the integumentary wall The function of these setae is unknown, but they would certainly appear to '' be useful in holding or helping direct the food between the | mandibles. [ A pair of muscle scars (marked "m” in Fig. 4A) are prominent i features of the proximo-mesial surface of the epipharynx. j The Antennae: The antennae (Figs. 2A and 2B, 31 and 3J) do f not differ radically from those of any other lepidopterous larva. ' They are located on the head, ventral and medial to the ocelli. ! Ferris (1943) pointed out that the antennae of lepidopterous i larvae have lost their point of articulation with the antennal ' segment and arise from a basimandibular membrane, no longer | within the boundaries of the sclerotized cranium. In E. comyntas | larvae, the bases of the mandibles and antennae are in close proximity and the latter often remained attached to the former when they were teased from the cranium. The membranous area at the base of the antennae is called the antacoria. From the antacoria arise the three main segments which comprise each antenna. On the second segment are a long and a short seta, two conical projections, and one small setadike projection. A puncture is located on this segment midway between the long seta and the proximal margin of the segment. On the third seg- ment there is one conical projection and two smaller seta-like projections. A very minute projection, not easily seen, is also located on the third segment. Dethier (1941) pictures a lan/al antenna of E. comyntas. The only differences noted between his drawing and our specimens are that the small seta on the second segment is much larger, and the seta-like projections are much smaller than the cor- responding structures on the specimens examined during this study. Measurements were taken to determine the antennal lengths of the various instars. Exclusive of the long seta and including only the three segments, the lengths in microns were: first in- star, 16.6; second instar 32.2; third instar, 51.6; and fourth in- star, 100. The lengths in microns for the long seta were 58.2, 74.6, 100, and 161 microns respectively. There is some variation in antennal length between individuals t of the same instar, and also, a degree of variation in ratios i; of component segments of the antennae between instars. Seg“-,J; ment 2 became proportionately longer than segment 1 and 3 ! with each succeeding instar. 5 (2) :61-96, 1966 IMMATURE EVERES 73 The Mandibles: The mandibles, Fig. 3A to 3E, have anterior and posterior articulations with the head capsule. The former occupies most of the flattened region on the side of the mandible opposite the incisor edge. The posterior articulation is by means of a rounded, knob-like condyle which is a convenient landmark on the posterior mesial edge of the mandible. The first instar mandible (Fig. 3A) has sixth teeth, the most posterior of which is somewhat elevated and associated with the condyle. From the latter prominence a ridge, or retinaculum, runs across the mandible to the anterior surface. Sclerotized crests run from the teeth to the retinaculum and, particularly with the second and fourth tooth from the posterior end, give the impression of distal linear folds. Two setae, one long and one short, are present on the posterior lateral margin. Later instars (Fig. 3B) have seven teeth, the distal incisor points varying somewhat in shape. Three of the middle points are almost finger-like, while the more anterior and posterior teeth in the series may be reduced and blunted. It would appear that the retinaculum becomes gradually ele- vated in progressive instars, particularly along the posterior parts of the mandible. Teeth are either incorporated in this retinacu- lum (the most posterior tooth in the first instar mandible ap- pears closely associated with the prominence) or develop there, so that the retinaculum of the fourth instar larva is characterized by two prominent teeth. The appearance and ontogeny of these structures is best seen in Figure 3D (first instar) and 3E (fourth instar). The two molar teeth are suggestive of the basal molar parts observed in mandibles of some Coleoptera. They are at a different level than the dentation of the distal margins. Of some interest is the fact that Crumb (1929) noted that de- cided peculiarities in the mandible were “invariably associated” with a wide departure in position and arrangement of setae and punctures on the head. We were not able to notice any such correlated anomalies. A little degree of orientation may help the reader in locating the next four head appendages. If the head capsule of a fourth instar larva is viewed from the ventral surface (Fig. 2C) a pair of maxillae are conspicuous on either side of a centrally located labium. Closer inspection of the maxillae shows four distal sclero- tized rings, the three most distal of which are the maxillary palps. Actually the terminal portions make up an outer palpus. There is a less conspicuous inner lobe of the palpus (the galea) on the dorso-mesial surface. Details of these structures are given be- 74 LAWRENCE AND DOWNEY /. Res. Lepid. low. The fourth sclerotized band from the tip of the maxillae is | the palpifer. The latter attaches to the stipes, the distal parts ( dististipes ) of which are membranous. The proximal area of ! the stipes is outlined on its distal and mesial extremities by a 1 sclerotized band. The proximal arm of this band points to a small [ triangular sclerite, on the posterior-mesial surface of the cardo. The latter segment is rather membranous and appears to lie j obliquely to the stipes, between the latter and the postgena. Between the maxillae, on a line between the palpifers, is the spinneret. Since it points ventrally, it is not as apparent in this view as the sclerotized rings at its base. The lighter sclerotized ring completely surrounding the terminal part of the spinneret is the fusuliger. Lateral to it, sclerites of the palpiger can be seen running to the small pair of labial palps. The palps are anterior I and lateral to the spinneret. The membranous area in front of the labial palps is the hypopharynx. Posterior to the spinneret, between the dististipes, is a dark ; sclerite, the mentum. This envelopes the posterior portions of the spinneret, and lateral arms run anteriorly on either side of the hypopharynx. The submentum is membranous and is just posterior to the mentum between the basistipes. The Maxillae: The maxillae. Figs. 3F and 3G (stipes not shown), vary but little between instars in structure. The stipes, j Fig. 2C, particularly the sclerotized portions, shows the most ; variation in respect to setal pattern. On the first and second in- ' stars there are two long setae on the stipes with a puncture between. A variable number of setae are located on the third and i fourth instar stipes, but always more than two. The punctures on the maxillae are constant in position and number. One puncture is found laterally on each of the two ^ proximal segments of the maxillary palpus (mp). They are barely visible on Fig. 3F. Two puncture-like structures are lo- cated distally on the frontal surface of the third segment of the maxillary palpus, and proximally there is a sensory plate (sensil- I lum placodeum pi. on Fig. 3F), which is just opposite the galea. The galea ( ga ) has a puncture on the ventral surface. The Labium: The labium, Fig. 2D, although variable in size, does not differ in structure or setal pattern on any of the instars. The labial palps (Ip) are distinctly three-segmented in most other lepidopterous larvae, but appear to be only two-segmented in E. comyntas. The basal segment of the labial palpus is roughly equal to the fusuliger (fg) of the spinneret in height. In most specimens it is one third and never more than one half 5 (2) :61~96, 1966 IMMATURE EVERES 75 inm I 23456 789 10 c Fig. 6. A — B Diagrammatic drawings of the fourth abdominal segment of a first instar larva, left side: A. Position of setae and len tides; LG=:lateral groove (furrow), SP=spiracle. For explanation of other lettering see text. B. Color pattern; R=:Red, G=:Green; l-12=areas of color. C. Setal map, left side of first instar larva; thor- acic legs, prolegs and setae on the ventral side are shown at bottom of drawing. 76 LAWRENCE AND DOWNEY /. Res. Lepid. the height of the fusuhis ( fs ) as measured on its posterior aspect. The palpiger (pgr) bears no setae, but on the distal border of the mentum (mn) there is a pair of setae located one on either side of the spinneret. The two arms of the mentum extend along- side the paraglossae (me) anteriorly and then abruptly curv^e in a dorso-caudal direction. The paraglossa bears many spine- like processes which are somewhat larger than those on the hypopharynx (hp). The Spinneret: The spinneret, Fig. 2D, is made up of an elong- ated fusulus (fs) and a wedge-shaped base called the fusuliger (fg). Except for a pair of punctures on its anterior surface the spinneret has no vestiture. The structure of the spinneret is the same in all instars. The measurements of the spinneret vary considerably within an instar, but the ratio between the length of the spinneret and labial palp fall in approximately the same numerical range in all instars, 2.3 to 3.4. Crumb (1929) found the spinneret varied considerably between genera of noctuid moths, particularly its relative length compared to the basal joint of the labial palp, as well as characters of the tip. The Hijpopharijnxi The hypopharynx, Fig. 2D, forms the floor of the mouth and is flanked on either side by the fleshy lobes of the paraglossae (me). It begins as a raised, broad, flattened area anterior to the labial palpi. Its anterior spiny area in the mouth could be called a lingua, but posteriorly in the mouth cavity it narrows and forms a sort of trough between the paraglossal lobes. Small slender projections are sparse at the anterior and become more numerous posteriorly just before the trough. No noticeable differences occur in the hypopharynx between any of the instars. BODY OF THE LARVA. The Neck: The neck is long and retractile into the prothorax. Except for a pair of cervical scler- ites it is membranous. The sclerites (Fig. 8G) are located in the lateral areas of the anterior portion of the neck. There are only slight differences between the sclerites of individual instars ex- cept for gradual increase in size. Setal Types: Four main types of setae were found on the first instar larva. The first type, here referred to as major setae, are all rather long, finely serrate, and have pointed tips. Examples of major setae are shown in Fig. 50 to 5R, and 5Y, and the loca- tion of specific setae on a particular segment is shown on Fig. 6A. All setae on the latter figure, and in the following discussion, are designated by letter. On Figure 6A, the following are major setae, A, B, H, I, J, K, L, M, N and P. Seta A on abdominal seg- ment 4 was 258 microns in length. 5 (2) :61-96, 1966 IMMATURE EVERES 77 Lesser setae, see Fig. 5S, are similar to the major setae struc- turally but differ in size; for example, D on abdominal segment 4 is 9.7 microns. A third type, here designated microscopic setae, are very minute and not easily seen. They are usually located an- teriorly on each segment and are often hidden in the folds be- tween segments. Seta C on the dorsum of the second abdominal segment measures 4 microns. Seta O is ventral in location and is the only other seta of this type given a letter symbol. No serious attempt was made to locate the microscopic setae on succeeding instars. A fourth type of short stout setae is represented by setae G and F. These will be called pegged setae, see Fig. 5K to 5N. Both G and F are clavate and short. Seta G on abdominal seg- ment 4 was found to be 29 microns. Drawings of the longest major seta on the dorsum of the fourth abdominal segment on each instar and on the pupa are shown on Fig. 50 to 5R and 5Y. Here it can be noted that the only difference is a slight increase in diameter and length as follows: first instar, 258 microns; second instar, 343 microns; third instar, 386 microns; and fourth instar, 386 microns. In sharp contrast is the length of this seta on the pupa, 644 microns. The setigerous tubercle on the pupa (Fig. 5Y) is always smooth and wider than tall. Contrary to what might be expected these setae are not proportionally larger with each succeeding instar as is the case with the spiracles on the larvae, which roughly follow a geometric progression in size with each succeeding instar. A pointed projection is sometimes found on the side of the seti- gerous tubercle of the larvae, giving the prominence a starlike appearance from above. Little correlation could be noted with presence or absence of these projections on the tubercles and particular types of setae (except in the spiculate setae discussed below); for instance, the tubercles of 5L and 5M and 5Q and 5R. Many pegged setae are present on the second instar. They tend to be concentrated in the lateral area of all segments though in no recognizable or consistent pattern. Their setigerous tuber- cles have either one pointed projection or none. Fig. 5L shows one of these setae on the fourth abdominal segment. Its length is 48.4 microns. Similar setae are found on the third and fourth instars but the number of projections on most of the setigerous tubercles is four or more. These pegged setae on the third in- star are merely blunt at the end rather than clavate. Fig. 5M shows one of this type on abdominal segment 4, which measures 71 microns. Similar types on the fourth instar are more pointed. 78 LAWRENCE AND DOWNEY J. Res. Lepid. GLAZED EVE- PIECE SCULPTURED EVE-PIECE PROTHORAOC LEG ANTENNA HM K LABRUM PILIFER MAXILLA MESOTHORAOC LEG ABDOMINAL SEGMENT 5 SPIRACLE MESOTHORACIC WING ANAL OPENING Fig. 7. A— C. Pupa, showing setal pattern and sclerites; A. Ventral view. B. Lateral view. C. Dorsal view. 5 (2) :61-96, 1966 IMMATURE EVERES 79 Fig. 5N shows one on the fourth abdominal segment which measures 100 microns. No pegged setae were found on the pupa. A fifth setal type here called a spiculate seta (Fig. 5T to 5W) is not found on the first instar. It differs from the other types in that tiny spicule-like projections occur on the distal third of the seta. At low magnification these distal projections give the im- pression of pine trees, see Fig. 81 and 8J. Spiculate setae on the second, third, and fourth instar larvae have the same length and apparently the same number of spicule-like projections. Spiculate setae on the second instar are located only near the honey gland. Those on the fourth instar differ in that their seti- gerous tubercles have pointed projections, whereas the setigerous tubercles of the second instar spiculate setae are smooth. The fourth instar larvae has spiculate setae on other areas of the body in addition to their normal position. On one specimen they occurred near the second abdominal spiracle and on an- other they were near the first abdominal spiracle. Length meas- urements in instars 2 to 4 are: 101.3, 103.3, and 93.6 microns respectively. Measurement of a spiculate seta on the pupa is 116.3 microns. Again it can be seen that size of a particular type of seta could not be correlated with the increase of size of other structures in succeeding instars. From the above discussion of setae, it can be judged that there is considerable variation in number, kinds, size, and loca- tion of these structures from one instar to the next. In addition, there is variation in setal pattern (except in first instar larvae) between different individuals in the same instar. The short more or less clavate setae are the most variable in position, but the spiculate setae, though often found near the honey gland, may also occur on other areas of the body. Setal Map of the First Instar Larva: The body of the larva is covered by many tubercles, each of which may or may not bear a single seta. Tubercles not bearing setae are here called lenticles, and when the word seta is used the fact that it is in all cases borne on a tubercle is understood. Certain of the setae and lenticles show a definite pattern of arrangement on all instars. The pattern is best seen on the body of the first instar larva (Figs. 1C, ID) and is indicated on the setal maps (Figs. 6A, 6C). The extreme “hairiness” of the later instars makes setal maps impractical. The map (Fig. 6C) shows the left side of a first instar larva. Roman numerals designate thoracic segments, and arabic num- erals the abdominal segments. Large circles represent lenticles 80 LAWRENCE AND DOWNEY J. Res. Lepid. and short lines represent setae. The left thoraeic legs are shown below the map from a medio-ventral view, and the prolegs from a ventral view. A prothoracic shield on the dorsum of the prothorax and an anal plate on the dorsum of the ninth and tenth abdominal segments are indicated on the map. Figure 6A is an enlarged partially schematic view of the fourth abdom- inal segment. Setae are indicated by letters of the alphabet. Setae on other segments are given the same letter as those on the fourth abdominal segment having a similar position. Not all setae could be designated in this manner, especially those of the prothorax and the ninth and tenth segments. A furrow (LG, Fig. 6A), is present on each segment starting on the" dorsum and passing posterior to the edge of the dorsal lenticle and just posterior to the spiracle and ending in the vicinity of the lateral ridge. This furrow varies only slightly from segment to segment. Although it is a convenient landmark, the furrow is not shown on the setal map, Fig. 6C. In plotting a setal map, the rectangles are drawn first, and setae and lenticles are then added. For convenience, the rec- tangles are made equal in size, with the result that horizontal distances are sometimes not in proper proportion. This would occur when some segments, for example, the prothorax, is much wider than another segment. Its setae would nonetheless be placed in the same horizontal distance as a subequal segment. Vertical distance, on the other hand, is easier to indicate on the map, and except for slight distortion of spatial relationship at the dorsal surface (top of the rectangle), the setae are in their proper relative positions. Except for the prothorax all segments of a first instar larva are about equal in width (see Fig. 1C, ID). All segments from the mesothoracic to the third abdominal segment are roughly the same height. Although somewhat schematic, Figure 3K gives the cross-sectional shape of the first instar larvae. Terminology used to designate setee and their location is partially that of Clark and Dickson (1956). The reader is re- ferred to the typical segment drawn on Fig. 6 A, where the following main regions are labeled. Some of the regions, i.e., the lateral ridge, might be more distinctive in Fig. 3K. The area between the mid-dorsal line and the lower edge of lenticle T is called the dorsal area. That between the lower edge of the spiracle and lenticle T is referred to as the lateral area. The ridge below the spiracle is called the lateral ridge and the ridge be- low this is called the sublateral ridge. The leg or proleg is below 5 {2) :61~96, 1966 IMMATURE EVERES 81 >PMp Fig. 8. A — B Dorsal view of honey gland and eversible tubercles; A. Abdominal seg- ments 7, 8, and 9 in fourth instar larva, showing position of organs in relation to each other and spiracles. B. Same, second instar larvae. C — F. Dorsal view of pro- thoracic shield of first (C), second (D), third (E) and fourth instar larvae. Setae a and b can be traced ontogenetically Ijy structures at their base (see text). G. Cervical sclerite, 3rd instar larva. H. Lateral view of extended eversible tubercle from fourth instar larvae. I. Partially schematic anterior view of honey gland of fourtii instar larva. Clusters of spiculate setae are prominent around the gland; the inverted tubercles on segment 8 appear on each side. J. Dorsal view of same. 82 LAWRENCE AND DOWNEY J. Res. Lepid. the sublateral ridge. Where neither proleg nor leg occur on a segment, the area below the sublateral ridge is called the venter. The following section is a description of the specific setae and lenticles on each segment of the first instar larvae and on the prolegs. Only the setae for the left half of the larvae are men- tioned. Later instars are not treated in as much detail as the first. Prothorax: A prothoracic shield, Figs. 8C to 8F, is located on the dorsum of the prothorax. It is roughly diamond-shaped with the anterior corner being so rounded as to make the anterior half approach a semicircle in shape. A major seta is located near the anterior corner just to the left of the mid-dorsal line on the shield. Slightly posterior and somewhat ventral to this seta is a lenticle lying just inside the margin of the shield. Pos- terior to and above the lenticle is another major seta. On the posterior half of the shield are located two more setae; the first on the posterior margin of the shield near the lateral corner and the second dorsal and slightly posterior to the first. The onton- geny of certain structures on the shield is discussed below. The bases of four setae on the lateral surface of the prothorax describe a diagonal line parallel to the anterior margin of the shield. The most posterior of the group is located just below the lateral corner of the prothoracic shield. Two other setae to- gether with a lenticle and the prothoracic spiracle respectively describe another diagonal on the lateral ridge with the spiracle near the intersegmental line. Two more setae, one anterior and dorsal, the other posterior and ventral, are located on the sub- lateral ridge. These have been designated K and L respectively and both occur also on the other thoracic segments and on abdominal segments 3, 4, 5 and 6. Near the midventral line is a seta here called P that is repeated on all segments. This occurs on every thoracic and abdominal segment and is indicated by the lowest setal symbol on the map Fig. 6A. Mesothorax: Six dorsal setae and a lenticle are present. Just to the side of the mid-dorsal line on the anterior third of the segment is a major seta and ventral to this a lenticle called T. A lenticle in roughly this same relative position is found on the methathorax and the first eight abdominal segments. A micro- scopic seta C is found anterior and slightly ventral to this len- ticle and varies in position vertically up and down in front of the lenticle on the metathoracic and first eight abdominal seg- ments. In a ventral and slightly posterior position is found an- other major dorsal seta. Posterior to the intrasegmental furrow 5 (2) :61-96, 1966 IMMATURE EVERES 83 and midway between the lenticle and dorsal midline are two more major setae. The anterior one is called A and the posterior B. This pair of setae is found to be present on the metathorax and first six abdominal segments. A group of four setae is found on the lateral ridge. Setae K, L, and P, are also present. Metathomxi Setae A, B, C, K, L, and and lenticle T are present. A pegged seta G is found midway between the lenticle and the lateral ridge. Four other setae are present on the lateral ridge. Abdomen: First Segment. Setae A, B, C, G, L, and P, and len- ticle T are present. A lesser seta called D is found just dorsal to G. Anterior to the furrow and situated midway between the dorsal lenticle and the spiracle is a pair of pegged setae. The anterior is G, and the posterior, F. Both are found together on the first six abdominal segments, and on other segments F ap- pears to be in the furrow. A lenticle is present below and just anterior to the spiracle and not quite on the lateral ridge. The three setae present on the ridge are designated from the most posterior as H, I, and J. This group is found on the first nine abdominal segments. Second Segment. Setae A, B, C, F, G, H, I, J, and P are pre- sent in their normal positions. A lenticle is present dorsal to, and somewhat posterior of, seta L, which is the only seta on the sublateral ridge. Seta D could not be found. Third Segment. Setae A, B, C, D, F, G, H, I, J, K, and P are present in their normal positions though G is dorsal as well as anterior to lenticle T. Three major setae are present on the pro- leg; an anterior lateral seta called M, a posterior lateral seta called N, and on the medial side of the proleg a seta P. A micro- scopic seta ( O ) is located in an anterior position on the medial surface of the proleg. The setal pattern on the other prolegs is the same except that O could not be located on the anal proleg. Fourth Segment. All setae found on abdominal segment 3 are found on abdominal segment 4 in identical positions. Fifth Segment. Other than D, which could not be found on this segment, and the fact that C is directly anterior to lenticle T instead of dorsal, this segment is identical in vestiture to ab- dominal segment 4. Sixth Segment. A lenticle called S located dorsal and posterior to lenticle T makes its appearance on this segment. In other re- spects the segment is identical in vestiture to abdominal segment 4. 84 LAWRENCE AND DOWNEY J. Res. Lepid. Seventh Segment. Lenticle S is located dorsal and anterior to lenticle T on a vertical line with the spiracle. Both lenticles are posterior to the furrow. K is absent. A lenticle occurs on the sub- lateral ridge. Setae O and F are present but the proleg setae M and N are not. Setae B and G are not present, but the re- maining setae are as on segment 4. Eighth Segment, Lenticle S is the only lenticle present. Pre- sent are seta A, C, H, I, J, L and P only. Ninth Segment. Setae H, I, J, L, O and P are the only setae to be found on this segment. No spiracle or lenticle is present. Tenth Segment. Four setae possibly “homotypic” with A, H, I, and J are located in a dorso-lateral position above the anal line. Four setae (undesignated) are located below the anus and above the anal proleg. Setae M, N, and P are present on the proleg. Two setae and a lenticle in addition are located anteriorly on the mesial surface of the proleg. A somewhat rounded rectang- ular anal plate makes its appearance on the dorsum. Comparison loith Other Lijcaenid Larvae: Setal maps of E. compyntas and Plebejiis (learieia) icarioides Bdv. were com- pared to see if there were any consistant differences. The pur-^ pose of this comparison was not only to discriminate the two species but possibly to gain some insight into generic differ- ences in the immatures. The taxonomy of the above species was originally derived by use of adult characters. The vestiture of the legs, prolegs, and prothoracic shield did not differ. The following is a summary of differences between the first instar larvae: Present in E. comyntas and absent in P. icarioides: Seta D on the sixth abdominal segment and lenticle 7 on the seventh abdominal segment. Structures present in P. icarioides and absent in E. comyntas: Setae K and F on the mesothorax; seta G on the seventh segment and lenticle S on abdominal segments 1, 2, 3, 4, and 5. With the limited material on hand (cast skins only), it was possible to make maps of only the first six abdominal segments of the first instar larva of Plebejus (Lycaeides) melissa Edw, There was nothing on the specimens examined to suggest that they were different from P. icarioides. A first instar larva of Strymon melinus Hub. differs from both Everes and Plebejus. Setae G and F are not clavate in the former species and dorsal lenticles S and T are very close together. A lenticle is present between K and L on abdominal segments 3, 4, 5, and 6, while no lenticles are found in these locations on 5 (2) -.61-96, 1966 IMMATURE EVERES 85 E. comxjntas, P. icarioides or P. melissa. The prothoracic shield of S. melinus differs in that it bears an extra pair of setae. The anal plate is longer than wide, roughly oval in shape (with the broader portion anterior) and has two punctures on the posterior half. Prothoracic Shield: The setae on the shield of the first instar have been described and figured (8C). The shield is defined on the second instar by a line, apparently a suture, enclosing a smooth green area. In contrast the surrounding area is granu- lated. The suture is less easily observed on the third and fourth instars, but the parts of it that are distinguishable enclose the area which, though green and therefore distinguishable on the living specimen, is granulated like the surrounding skin. Where- as the prothoracic shields of the first instar larvae of E. corny ntas and P. icarioides are relatively similar, later instars of the two species seem to differ more markedly. Other species, for example S. melinus, bear additional setae on this structure in the first instar. It is presumed that the prothoracic shield may be of some interspecific taxonomic importance. In spite of the great increase in the number of structures on the prothoracic shield in later instars (see Fig. 8D-8F), some setae can be traced ontogenetically. This can be done partly by position and setal type, but also by structures at the base of the setae. The most posterior-lateral setae on each side of the first instar shield are each located on a distinct sclerotized area, the pinaculum. The latter are slightly larger than the base of a typi- cal seta which elevation (tubercle) could be termed a papilla. The papillae are also present on these setae, marked ''a” in Fig- ure 8C to 8F. These very slender long setae represent a type not seen elsewhere on the larvae, even though it falls into a major setal category on the basis of its length. Another pair of setae, marked ‘T” in Figs. 8C to 8F, are more mesial on the shield, and are located on a prominence called a chalaza. The chalaza-borne setae “b” can be easily traced in the drawings. Mackay (1959) also found that the bases of the setae were useful in identifying particular setae in the Olethreutidae. The second instar prothoracic shield bears a pair of lenticles which are ‘‘homologous” to those on the first instar. They are in a somewhat similar location. These lenticles may or may not appear on the third or fourth instar, and their position may also vary. Clavate, blunt, or pointed pegged setae occur on the second, third, and fourth instar shields and are variable in number and position. 86 LAWRENCE AND DOWNEY /. Res. Lepid. Blackish areas may be present on the shield. Some first instar larvae had a pair of blackish spots along a longitudinal line pos- terior to each of the lenticles, others had a posterior pair of wedge-shaped blackish areas on each side of the mid-dorsal line. These areas were also noted in second and third instar larvae. On others the entire prothoracic shield was green. Anal Plate: The anal plate is located in the center of the dor- sum on the tenth abdominal segment in first instar larvae. It is square in shape with rounded corners. The sides of the plate are approximately equal to one third the width of the segment. The plate is devoid of setae and is not granulated, but on one specimen a minute lenticle (or puneture) was seen on each lateral edge of the plate. The plate did not occur on later instars. The anal plate of E, comyntas tended to approach a square in shape, while the anal plate of P. icarioides was hexagonal, being somewhat wider than long. This suggests that there may be some taxonomic usefulness in this structure. Legs: According to Fracker (1915) “No one has yet discovered characters of much value in the structure of the thoracic legs, so uniform are they throughout the entire order.” Comparison of the prothoracic leg of E. comyntas with the prothoracic leg of a larva of Cirphis unipuncta (Haworth) (drawing in Ripley, 1923) shows no significant difference in structure or setal pat- tern. Figures 5E to 5J, show the thoracic legs in the first instar larvae. No consistant difference between instars was noted. Prolegs: There are five pairs of prolegs. Those located on seg- ments 3, 4, 5 and 6 are referred to as the ventral prolegs and those on the last segment as the anal prolegs. Each proleg has a biordinal mesoseries of crochets interrupted in the middle by a fleshy spatulate lobe, see Figures 5A to 5D. This arrangement is diagnostie of the family Lycaenidae. There is generally the same number of crochets on each side of the lobe in each instar. Th average number of crochets on the proleg of the first instar is four (Fig. 5A); on the second, seven; on the third, fifteen; and on the fourth, twenty-three. The relationship between the fleshy lobe and the crochets in the fourth instar proleg is given in two views. Figure 5C and 5D. The number of crochets per proleg is constant on the first instar larva, there being four on each ventral proleg and three on each of the anal prolegs (Fig. 5B). In later instars the num- ber is subject to some variation between individuals of a given instar and even between members of a proleg pair. The average > number for an instar, however, appears to remain fairly constant. 5 (2) :61~96, 1966 IMMATURE EVERES 87 Lenticles and setae occur on the medial surface of the pro- legs as well as on the lateral surface. The medial surfaces of the anal prolegs and of the fourth abdominal proleg were studied on each instar. There is always one lenticle on the medial sur- face of each anal proleg of the first and second instar larvae. (Fig. 5B). Two were found in this position on a third instar and three on a fourth instar. No lenticles occur on the medial surface of the fourth abdominal proleg of the first instar, however, two are present on the second and third instars and only one was found on a fourth instar specimen. Setae on both surfaces of the prolegs become more numerous with each succeeding instar. Honey Gland and Protrusihle Tubercles: The honey gland and protrusible tubercles, see Fig. 8A, B, I, J, are found only in the later instars. Though the second instar larva does not have the tubercles, it does have a functional honey gland on the dor- sum of the seventh abdominal segment (Fig. 8b). On the third and fourth instars in addition to the honey gland a pair of pro- trusible sacs or tubercles (Fig. 8H) are located posterior and somewhat lateral to the eighth abdominal spiracles (Fig. 8A). A spiculate seta is located on each of the lateral margins of the gland on the second instar. Along the posterior margin of the glandular orifice is a cluster of lenticles with no definite arrange- ment. The anterior margin does not bear setae or lenticles. The gland on the third instar has the same vestiture except the len- ticles are more numerous and there are three spiculate setae at each of the lateral margins of the gland. Although there are more of these setae on the fourth instar honey gland, they are not arranged with an equal number on each side. For example, one specimen has four on one side and eight on the other. When the protrusible tubercles on the third and fourth instar are extended, they can be seen to bear spiculate setae (Fig. 8H) ! on their distal parts. These setae are unlike other spiculate setae in that the spicules are shorter and are proximal as well as distal I in position. When the tubercles are only part way protruded, the setae can be seen grouped in a vertical bundle in the center of 1 the tubercle. When fully protruded the setae point in all direc- tions. The tubercle is often fluttered up and down between these I two positions making the setae appear to move by their own 1 power. A count was made of the number of spiculate setae per j tubercle on fourth instar larvae: They vary in number from 29 j to 34 and may also be variable in numbers between members of i pairs on one individual. The third instar seta measured 76 mi- 88 LAWRENCE AND DOWNEY /. Res. Lepid. crons and the fourth instar about 112 microns. Other measure- ments of the gland and tubercles are given in Table 1. Colorationi Three basic colors, brown, green and black, make up the color pattern of the larva. Black color, which is probably due to pigments in areas underneath transparent epidermis is found only on the prothorax. On the first instar larva this color was seen just anterior to the prothoracic spiracle in a depression above the lateral ridge. The border of the prothoracic shield and certain spots on the shield also showed this color. When viewed from the side the larva appears to have at least twelve longitudinal alternating light green and brown areas. In gross examination these areas are not all obvious, so that the larvae may appear to have only a few anterior to posterior color stripes. When viewed under a microscope these appear as a linear arrangement of small pigmented patches. The larvae, however, are described in the following paragraphs as if the areas were not broken up into pigment patches. The fourth abdominal seg- ment of a first instar larva is shown in detail in Fig. 6B to illus- trate longitudinal areas of color. They have been numbered from one to twelve for ease in discussion. There is some variation in the color pattern, although its basic nature probably is fairly stable. For instance, the lighter stripe marked by number two, in a particular premolt fourth instar larva was nearly obliterated by the enlargement of stripes one and three; ordinarily the stripe is well defined in a freshly molted fourth instar larva. It is possible that some color changes are due to increasing pigmentation during a particular instar. Variations in which the larva has dark green areas instead of brown or red- dish-brown are about equally common and occur in siblings from the same female. First Instar Larvae: Mid-dorsal Line. — This line (marked 1 in Fig. 6B) is dark brown and extends from the dorsum of the mesothroax to the dorsum of the ninth abdominal segment where the line broadens to cover most of the dorsal area. The tenth segment bears five alternating light green and dark brown transverse bands, the most anterior being light green. Dorsolateral Area. — A broad light green stripe makes up this area extending from the prothroacic shield to the tenth ab- dominal segment. There is a hint of a fine dark brown stripe extending within the light green one, from abdominal segment 4, passing just dorsal to the lenticles and merging with the mid- dorsal line on abdominal segment 7. Lateral Area. ~ This area is basically made up of five alternat- 5 (2) :61-96, 1966 IMMATURE EVERES 89 TABLE 1 Measurements (in mm.) of the honey gland and eversible tubercle in larvae of Everes comyntas Godt . Instar No. Gland Orifice Length Gland Length Gland Width Tubercle Height Diameter of base Extended Tubercle Diameter of base Retracted Tubercle 4 5 256-372 370-549 204-256 255-362 155 154-179 3 5 154-256 282-384 128-204 155 77-128 2 5 102-154 256 102-128 ing reddish-brown and light green stripes extending the length of the body. The first of these lateral stripes (5) going in des- cending order is reddish-brown and starting just below the pro- thoracic shield it runs as far as the tenth abdominal segment where it merges with the last dark transverse band mentioned under the section on the mid-dorsal line. The second stripe (6) is light green and extends from the prothorax to the sixth ab- dominal segment. It is represented as haloes around the seventh and eighth abdominal spiracles and as small squarish patches on the ninth and tenth abdominal segments, the haloes and ! patches both being bordered by the merging first and third I reddish-brown stripes. The third stripe (7) is dark colored and ! starts as a fine line just on the posterior edge of the prothorax ! and extends back to the eighth abdominal segment where it 90 LAWRENCE AND DOWNEY /. Res. Lepid. merges with the first and fifth dark reddish-brown stripes. The fourth stripe ( 8 ) is a broad light green band extending from the prothorax, where it merges with the light green stripe on the lateral ridge, to the fourth abdominal segment. It is represented by haloes around the fifth and sixth abdominal spiracles. The fifth stripe (9) is reddish-brown and extends from the meso- thorax to the fourth abdominal segment where it becomes ob- scure. Lateral Ridge. — The entire lateral ridge is light green except on the underside of the last four segments where it is dark brown. Sublateral Ridge, Venter, And Prolegs. — These areas are en- tirely light green in color. Second Instar Larvae. The second instar has the same basic pattern as the first instar with the following exceptions: Stripe number three begins on the mesothorax and ends on abdominal segment 6, the honey gland being light green in color is in this area. An area between the prothoracic shield and lateral ridge is reddish-brown except for a small light green stripe in the mid- dle. Stripe number eleven extends from the second abdominal segment to the anus. Third Instar Larvae. The color pattern is the same as in the second instar. Fourth Instar Larvae. Same coloration as third instar only the dark stripes are broader and darker brown and the light stripes are greener. Other differences are as follows: Stripes number one and three nearly merge and the only remnant of stripe num- ber eight is found as haloes around the spiracles. Stripe number eleven under the lateral ridge merges with stripe number twelve on the sublateral ridge which is also brown. Together they ex- tend the length of the larva. In addition to the above the lateral sides of the ventral prolegs are reddish-brown, the remainder being green. The entire anal proleg is reddish-brown. The venter is green. THE PUPA The obtect pupa of Everes corny ntas is long and slender: average length of specimens examined was 6.7 mm.; average width at the widest portion was 2.5 mm. Thus the pupae are about three times as long as broad (not 4 to 1 as indicated by Scudder, 1889, vol. 3, p. 907). The ratio agrees with the length and breadth of the three European species of Everes figured by Lorkovic (1938). The average height at the highest point (3rd or 4th abdominal segment) was 2.4 mm. The small size of the pupa, its slender profile, and the num- erous long setae are fairly indicative of this species. These fea- 5 (2) ;61-96, 1966 IMMATURE EVERES 91 tures are shown on Figure 7 a, b, c. The living specimen is whit- ish-green in color, with the head and thorax tending to be slight- ly darker green than the abdomen. A conspicuous mid-dorsal black line, while variable between individuals, extends from the prothorax to the seventh abdominal segment, where it fades out. The lateral and dorsal surface is speckled with many small dark brown to black spots. A lateral row of larger spots is also present just dorsal and slightly posterior to each abdominal spir- acle. A larger spot occurs in a similar position on abdominal seg- ment one, a slightly smaller spot on the metathorax and a still smaller dark area occurs on the mesothorax just posterior to the spiracle. Reddish marks may be present in some individuals on both sides of the lateral lines on the abdomen. Setae, Hooks, and Papillae. The major setae are basically white in color, so that as one observes a pupa he notes first its greenish color and shape and is subsequently cognisant of num- erous long setae on the dorsal and lateral surface. These setae have been stippled on the drawing on Figure 7 in hopes that their unobtrusive nature may be imparted. In addition to the completely white setae, some are colored with one or more dark brown or black bands variously located at the base, tip or inter- mediate positions. Most of the setae on the pupae are major setae, but a few on the venter of the abdomen are small in comparison with the others. The size of the smallest on one specimen was 43 microns. Since there was a difference in number between groups of setae on one side and the corresponding groups on the other, and be- tween individuals, the taxonomic value of setae numbers is vague. Spiculate setae are found on the prothorax and sixth abdom- inal segment only. They are one third to one quarter the size of the major setae covering most of the pupa. Those on the pro- thorax were not constant in number although they do occupy the same general position on most specimens. Those on the sixth segment are more constant in number and position. Commonly there are two spiculate setae located dorsally and slightly pos- terior to the sixth abdominal spiracle although sometimes there may be one or none. A group of cremastral hooks is found near the anal opening on the tenth abdominal segment. Those measured range in size from 48.5 to 70.5 microns. They are variable in number and posi- tion but are prominently clustered in a “C” shaped band around the posterior part of the anal slit. 92 LAWRENCE AND DOWNEY /. Res. Lepid. Although the pigment spots in some individuals are suggestive of a roughened surface, no papillae were noted on the pupae which were not associated with setae. THE HEAD. Vertex: Perhaps the most difficult of the head structures to locate is the vertex, which is reduced to two small triangular sclerites. These lie on either side of the dorsal midline just anterior to the prothorax and just posterior to the bases of the antennae. The vertex lacks both setae and papillae. Its posi- tion and its small size preclude its being shown on Fig. 7A or 7B. Front, Clypeus, Labrumi There is no fronto-clypeal or clypeo- labral suture, which makes the boundaries of these particular areas obscure. Two pores, representing the anterior tentorial pits, are usually apparent between the eyes. Arms of the "‘U-shaped” suture demarking the limits of the labrum point to the tentorial pits. Since the pits are associated with the lateral margins of the clypeo-labral suture in other insects, we may assume the area between and cephalad of the pits to be the clypeus, which is not distinct from the frons to which the antennae are attached. Below the pits, the labrum is distinct. The region between clypeus | and glazed eye-piece is the gena. The pilifers, caudo-lateral projections of the labrum, are en- ^ larged and in most individuals, meet in the midline caudad of j the labrum proper. Labial Palps: In a few individuals, a small triangular portion of the labial palpi separates the lobes of the pilifers. The exter- nal appearance of the labial palps between the pilifers is thus variable in individuals of comyntas. The palpi also vary in ex- pression in other species. In Atlides halestis, for example, the pilifers touch and the labial palpi can only be seen between the proximal bases of the maxillae. Maxillae: The maxillae are very prominent just laterad of the mid-line between the pilifers and the central junction of the two antennae. They are only slightly longer than the length of the united distal parts of the antennae (see Fig. 7A). Eye-pieces: Between the labrum and the entennae are the prominent eye-pieces; a lunate mesal portion has a smooth glassy surface and lacks setae; the lateral sculptured eye-piece has the same texture as the gena. * Antennae: The distal one-third of the antennae are united in the midline. Externally the clavate ends of the antennae appear to stop at the fifth abdominal segment, as do the wings. How- j ever, there is a small oblique pocket beneath the fifth segment. 5 (2) :61-96, 1966 IMMATURE EVERES 93 into which the tips of the antennae ( usually less than 2.0 mm. ) fit comfortably. This is best observed in specimens preserved in fluid in which the intersegmental area might be slightly dis- tended due to swelling. THE THORAX. Thoracic segments are indicated in Figures 7B and 7C. They are similar to other lycaenid pupae in general shape. Proportionate lengths along mid-dorsal line are: prothor- ax, 2.5; mesothorax, 7.0; metathorax, 1. The mesothoracic spir- acle is level with the surface, is not associated with prominences or tubercles and appears plugged with a narrow, highly reticulate cover. A group of 6 to 8 small pores is found on the distal surface of the mesothoracic leg. Lesser numbers occur in the same posi- tion on the prothoracic leg. These openings may vary in number on each side of the body and between individuals, but they are always present. Other species of Lyaenidae also have these pores. They are of unknown function. THE ABDOMEN. Honey Gland Scar and Tubercles: The honey gland scar is located on the dorsum of the seventh ab- dominal segment in the same position as on the larva. It is more prominent on some specimens when it is pigmented, but it is very difficult to detect in others. Though in other Lycaenidae the pupal honey gland is functional, it is not functional in E. corny ntas. No markings or scars indicative of the protrusible tu- bercles could be found on segment eight. Stridulation: As most lycaenids, comyntas pupae are able to stridulate. The noises produced are barely audible and can be best described as slight chirping or squeaking sounds. It is best to place the pupae in a vial or other reflective container in order to hear the sounds. These organs were first reported in comyntas by Downey and Strawn (1963) and Strawn (1964). Downey (1966) subsequent- ly noted the stridulatory organs in pupae of the Palearctic E. argiades Pall. The sound-producing structures are located on the inter-seg- mental membrane between abdominal segments 5 and 6. To function, a posterior file containing numerous teeth is grated across an anterior stridulating plate. A recurved region of mem- branous folds lies between the grating surfaces. Stridulating plate: A narrow (0.05 mm.) band on the dorsal posterior membrane of segment five extending laterally to, or just lateral of, the spiracle. The surface is of grainy texture, with the main axis of the “grains” in a transverse row. Each “grain” 94 LAWRENCE AND DOWNEY /. Res. Lepid. has one or more small tubercles whose delineation makes more apparent the reigon of the stridulating plate. The latter is not sclerotized as it is in many other species so that the plate in comyntas is defined mostly by its roughened grainy surface. The tubercles range in height from 2 to 5 microns and vary slightly in number and position upon the plate. Inter segmental fold : This area of the intersegmental membrane is identified by its position and its lack of any sclerotized struc- ture. It is in this region that the membrane folds back on itself so that the anterior plate is in juxtaposition with the posterior file. Two relatively narrow longitudinal muscles are attached by means of short ligaments to the membrane. These attachments are in a dorso-lateral area, midway between the spiracular and the mid-dorsal line. Contraction of the muscles pulls the mem- brane and produces the grating action of the file against the plate. File-. This area is defined in comyntas solely on the presence of teeth. The latter are small, unsclerotized, indistinct protuber- ances which show a tendency for clustering. The same type of teeth, not quite as numerous, occur in other intersegmental mem- branes, and also on exposed portions of the pupal integument where they do not contact other surfaces. However, this situa- tion prevails in pupae of over 100 other lycaenid species ex- amined by Downey, and there seems to be little doubt that, in the region of the stridulating plate, the teeth serve as frictional devices in sound production. The file extends laterally a greater distance than the stridulat- ing plate. DEHISCENCE. Emergence of the adult is accomplished by a mid-dorsal split of the thoracic segments. As the adult exits through this longitudinal split, it forces the lips apart. This split- ting progresses posteriorly along the seam between the wings and the abdomen. Frequently ruptures also occur between the prothorax (to which the vertex usually remains attached) and the antennae, progressing for a variable distance along the an- tennal suture down the ventral face. The face covering, however, remains attached to the pupal skin. A gradual darkening of the pupae occurs about 48 hours prior to eclosion. SUMMARY AND CONCLUSIONS The external morphology of the immature stages of Everes comyntas is described and treated in some detail. This includes the egg, head capsules of the various instars, mouth-parts, setal pattern, honey gland, and characters of the pupa. 5 (2) .-61 -96, 1966 IMMATURE EVERES 95 Different types of setae not previously known for this species are drawn and described. Variation is found in number and position of the setae on the head and body between individuals of the same instar (except first instar larvae) and between individuals of different instars. Other characters showing variation include depth of the labral notch, segmental ratios of the antennae between instars, setal lengths, mo^rphological structure of the setae, number of crochets, and coloration. Individual variation is found in the following characters: setal pattern; presence or absence of spiculate setae; presence or absence of the honey gland scar; number size, and position of cremasteral hooks and coloration. Because of this extreme variation it is suggested that those involved in a comparative description of immature Lepidoptera make a study which is not only thorough in detail but which includes examination of a large number of specimens of all instars. The honey gland is described on the second, third, and fourth instar larvae, and the scar is noted on the pupa. Eversible tuber- cles on the third and fourth instar larvae and a pupal stridulating device are described. A complete setal map of the larva is plotted. Comparison of a first instar setal map of E. comyntas with one of P. icarioides shows that, though similar in many respects, there are decided differences in presence and absence of seta and lenticles. The usefulness of setal maps of butterfly larvae is demonstrated and should point the way to more work of this type. Some differences in the prothoracic and anal plates of several lycaenid species are pointed out which suggest that these struc- tures may be of importance in the taxonomy of the larvae of Lycaenidae. ACKNOWLEDGMENTS We are grateful to Mr. H. W. Capps, formerly of the U. S. National Museum for his useful suggestions early in the study and for the loan of specimens. Thanks are also given to Dr. Robert Mohlenbrock for identification of foodplants, to Dr. R. E. Blackwelder for his helpful criticisms of the manuscript, and to Mr. Gary Simmons for the illustrations. This is part of a larger study on the immature stages being supported by the National Science Foundation, GC 2423 and GB-4962X. Portions of this research were used by Mr. Lawrence for part of his Masters thesis at Southern Illinois University. 96 LAWRENCE AND DOWNEY J. Res. Lepid. LITERATURE CITED CLARK, G. C. and C. G. C. DICKSON. 1956. Proposed classification of South African Lycaenidae from the early stages. Jour. Ent. Soc. ' So. Africa, 19(2) : 195-215. CRUMB, S. E. 1929. Tobacco cutworms. U. S. Dept. Agr. Tech. Bull. 88, 180 pp. DETHIER, V. G. 1941. Antennae of lepidopterous larvae. Bull, Mm. Comp. ZooL Harvard, 87:45. DOWNEY, J. C. 1966. Sound production in pupae of Lycaenidae. Journ. Lepid. Soc., 20(3) :129-155. DOWNEY, J. C. and M. A. STRAWN. 1963. Sound producing mechan- isms in pupae of Lycaenidae. Bull. Ent. Soc. Amer., 9(3): 161 (Ab- stract ) . EDWARDS, W. H. 1876. The preparatory stages of Lycaena comyntm. Canad. Ent. 8( 11 ) :202-205. FERRIS, G. F. 1943. The basic materials of the insect cranium. Microen- tomology. 8(1): 8-24. FORBES W. T. M. 1911. A structural study of some caterpillars. Ann. Ent. Soc. Amer. 3:94-132. FRACKER, S. B. 1915. The classification of lepidopterous larvae. III. Biol Mono., 2(1):L169. HASENFUSS, IVAR. 1960. Die Larval systematik der Zunsler (Pyrali- dae ) . Abhandlungen zur Larval systematik der Insekten. Nr. 5. Akademie-Verlag-Berlin. 263 pp. HINTON, H. E. 1946. On the homology and nomenclature of setae of lepidopterous larvae, with some notes on the phylogeny of the Lepi- doptera. Trans. Roy. Ent. Soc., Lond., 97:1-37. 1947. The dorsal cranial area of caterpillars. Ann. Mag. Nat. Hist., 14:843-852. LORKOVIC, ZDRAVKO. 1938. Studien uber den Speziesbegriff. IL Artberechtigiing von Everes argiades Pall., E. alcetas Hffgg. imd E. decolorata Stgr, Mitt, der Munchner Ent. Gesellshaft, 28(2) :215-248, tables V-VII. MACKAY, M. R. 1963. Problems in naming the setae of lepidopterous larvae. Canad. Ent. 95:996-999. MUTUURA, A. 1956. On the homology of the body areas in the thorax and abdomen and new system of the setae on the lepidopterous larvae. Bull. Univ. Osaka Pref. 6(B): 93-122. PETERSON, A. 1951. Larvae of Insects. Part 1. Lepidoptera and plant infesting Hymenoptera. Edwards Bros. Inc. Ann Arbor, Mich., 315 pp. RIPLEY, L. E. 1923. The external morphology and postembryology of noctuid larvae. 111. Biol Mono., 8(4):1-102. SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada witli special reference to New England. Published by the auth- or, Cambridge, Mass., 3 vol. 40 + 1958 pp. 89 pi. SHORT, J. R. T. 1951. Some aspects of the morphology of the insect head as seen in the Lepidoptera. Proc. Roy. Soc., Lond., B, 26:77-88. STRAWN, M. A. 1964. Sound production of pupae of the butterfly fam- ily Lycaenidae. Unpub. Masters Thesis, Southern Illinois Univ. Journal of Research on the Lepidopter a 5 (2) :97“112, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A LITTLE-RECOGNISED SPECIES OF HELICONIUS BUTTERFLY (NYMPHALIDAE)'^ JOHN R. G. TURNER Genetics Laboratory, Department of Zoology, University of Oxford^. The Taxonomy of the South American butterfly genus Heliconius ( Nymphalidae ) Kluk is in confusion, not because the species are ‘"critical” as a result of inbreeding, apomixis or other evolutionary processes incompatible with the rigid species con- cepts inherited from the theory of special creation, but simply because the species are polymorphic, show remarkable geographi- cal variation, and mimic each other; thus as Fox (1956) has said of another South American group, the Ithomiidae, two apparently identical butterflies may belong to distantly related species, while two having hardly a single pattern in common may be conspecific. While studying geographical variation and mimicry in the genus ( See Turner 1963b, 1965 for preliminary summaries ) I find it necessary to give a definition of Heliconius elevatus Noldner, a species which has seldom been properly recognised, and to describe a new subspecies. The species H. elevatus shows strong geographical variation, each form resembling very closely a form of Heliconius mel- pomene ( Linnaeus ) , a highly variable species which shows both geographical variation and polymorphism ( Turner & Crane 1962; Sheppard 1963; Turner 1965); as a result the two species are usually confused in various ways. Neustetter (1929) came near to the truth in separating several forms of elevatus as a species different from melpomene, but spoiled the result by splitting melpomene itself into several species, and including two of the elevatus subspecies with melpomene. Eltringham (1917) was firm about the separation of elevatus from melpomene, although he regarded tumatumari Kaye, here listed as a subspecies of elevatus, as a species in its own right. Oberthiir (1916) correctly regarded tumatumari as a form of his own hari and separate from melpomene. In other works (e.g. Emsley 1964) and in most collec- tions, melpomene and elevatus are confused. The latest revision ^This work was supported by a grant from the Nature Conservancy. 2This paper is dedicated to the team working on the biology of Heliconiinae at the William Beebe Station for Tropical Research, Trinidad, whose papers appear in the New York Zoological Society’s journal Zoologica. 3Now at Department of Biology, University of York, U.K. 97 98 JOHN R. G. TURNER /. Res. Lepid. Tabl* I GONDIXrOHS FOR DYS-FSEDINO Dya Faading Start Collac of Time, . blend) LenKth.mo dav Time Dye Fed. days Pupation, Yield JSI- Punae Ooler Pueae 1 none - - - 23-31 ( 34 from green 65 lat-is- etar larvae) 2 ennrthcae Nila blue A (5) 22-24 20-21 1. 5-1.8 22-27 10/10 one blue (15 laa] root el, blue- green at most neutral red (5) 19-24 17-21^ 5 23-26 7/9 pink (esp. on abdomen) to deep red Nile blue 22 A (1) and neut. red (l) 21^ 3-5 24-26 5/5 green to green with blue ab- domen brill, ere- 18-20 >yl Us« (5) 21^ 4-5 25-26 3/3 green 3 pfailodice Mile blue A (5) 15-23 14 O.i-1,4 18-li 4/20 2 U.«e-green Nile blue A (1) 24-27 16 2.3-4. 5 20-22 2/8 greenish-bluei blue (1) neutral red (5) 15-19 14^ 4-5 18-19 7/10 dark- red (2) neutral red (5) 26-30 18 0.5 20-21 3/3 two daric-red) one green with red abdomen Nile blue 17-19 A (1) and neut. red (l) 15 0,5 19 4/5 el. blue tint brill, cre- eyl blue (§) 18-19 1 15^ 3-4 18-19 4/4 si, grayish green 4 Bhilodlba neutral red (i) ( 28-32 16-19^ one at 24) 0-3 19-21 32/32 red to dark-red 5 ^iQdlee Nile blue A(l) 27-30 16-17^ 2-3 18-20 4/10 blue-green ^fad dya to pupatlea. ^graao for controls. Fig. 1. Parallel variation of H. elevatus (left) and H. melpomene (right) Specimens of H. elevatus from the British Museum, (Natural History), of H . melpomene from the Hope Department of Entomology, Oxford (except ). Colour: ground, dark brown to black; pale marks, yellow; dark marks, red. Left: A. H. e. elevatus $ Sao Paulo de Olivenca, upp. Amazons, August 1907 (M. de Mathan); B. H. e. hari $ Essequibo R., Brit. Guiana; C, H. e. perchlorus $ , Mauchay, Beni R. viii 95 (Stuart); D. H. e. tumatumari $ . Tiger Creek, Potaro R., Br. Guiana, May ’07 ( C. B. Roberts); E. H. e. roraima $ , see type description; Right; F. H. m. aglaope $ , “Tonantins, Amazon”; G. H. m. thehiope $ , Para, L. Amazon, hi — vi, 1915 (A. M. Moss, ex W. J. Kaye coll.); H. H. m vicinus $ , no data, ex coll. J. A. Gibbs, Keble college; I. H. m. cybelel $ , Guyane francse. St. Jean de Maroni (coll, le Moult ) . 5 (2):97-112, 1966 SPECIES OF HELICONIUS 99 100 JOHN R. G. TURNER /. Res. Lepid. TABLE 2, DIFFERENCES BETWEEN H. ELEVATES AND H. MELPOMENE elevatus melpomene Male genital valves with a large strong hook and a marked txift of hairs at the tip. Male genital valves with weak hook near the tip, not accompanied 1 by a marked tu£t of hairs. Anterior margin of hindwing with a red stripe on the underside Anterior margin of hindwing with a yellow or red stripe on the \mderside. A yellow stripe a few millimetres behind the anterior margin of the hindwing, underside. A few millimetres behind the anterior margin of the hindwing' tinderside, either no stripe or a red stripe. No prominent red spots at the base of the hindwing, underside. Three to five deep red spots at the base of the hindwing, underside. Often a row of marginal white spots on the hindwing, underside. Seldom a row of marginal white spots on hindwing, underside. Rays on hindwing, upperside, tend to be thick. Rays on hindwing, upperside, tend to be thin. Apical spots on forewing often present. Yellow apical spots on forewing hardly ever present. Often a yellow fleck at inner angle of forewing. Never a yellow fleck at inner angle of forewing. Yellow mark between veins M3 and often with concavely indented distal border. Yellow mark between veins M3 and Cu]^ usually convex at its distal border. Spots on head and palpi often white. Spots on head and palpi often yellow. of Heliconius separates some forms of elevatus but places two of the subspecies as hybrids between H .melpomene and H.aoede (Hiibner); this revision (Emsley 1965) appeared while the present paper was in draft, and the evidence presented here is independent of Emsley 's study; the general agreement of the two studies testifies to their correctness. 5 (2):97-112, 1966 SPECIES OF HELICONIUS 101 TABLE 3, NUMBERS OF GENITALIA EXAMINED species infraspecies number melpomene aglaope 7 flavotenuiata 0 thelxiope 2 vicinus 7 meriana 0 others 6 total 22 elevatus elevatus 9 ps eudocupidineus 2° bari 2° + perchlorus 4 \ taracuanus r tumatumari 2 roraima 4e2 total 24 ® External only, without dissection + Including holotype 1 Holotype 2 Holotype and male paratypes No one looking at figure 1, which shows the main forms of elevatus and the parallel forms of melpomene, will be surprised at the confusion of the species ( the space in the lower right hand comer of the plate could also have been filled with an equivalent form of melpomene which is so rare that I was unable to obtain a specimen to photograph); the parallel variation is further summarised in table 1. The chief differences between melpomene and elevatus are summarised in table 2 and illustrated in figure 2, in many speci- mens of elevatus the hook on the genital valve can be seen without dissection; the characters used, the male genitalia and the basal markings of the hindwings, appear to be ‘"good” specific characters in the genus Heliconius, showing much less variation than the major wing markings, although they do vary between localities and a little between individuals; the range of variation of both characters in melpomene and elevatus is distinct and shows no overlap. Table 2 is based on the examination of thou- sands of H. melpomene, between fifty and a hundred of H.e.elevatus, perhaps two dozen of H.e.tumatumari, the five type specimens of the new subspecies, and not more than a dozen of 102 JOHN R. G. TURNER /. Res. Lepid. each of the other subspecies; table 3 shows the numbers of male genitalia examined. It could be argued that the apparent species are no more than genetic polymorphs. The basal markings are not known to be polymorphic in any other Heliconius. Poly- morphic genitalia are likewise not known in the genus, although not many specimens of each species have been examined; taxono- mists usually underestimate the variability of these organs (Ford 1955) despite the finding of Kerkis (1931) that the genital apodemes of a hemipteron have higher coefficients of variation than other parts of the body; a thorough survey of the African butterfly Papilio dardanus Brown showed strong variation and even polymorphism in the male genitalia (Turner 1963a). But if it can be established that the basal markings and genital char- acters are correlated, so that we do not find melpomene markings with elevatus genitalia or vice versa, then it becomes highly unlikely that the forms are merely polymorphs, as it would be too great a coincidence for two characters “good” in the rest of the genus to become in one species not simply polymorphic, but controlled by the same genetic switch mechanism. To check on this I selected from the Tring collection 14 males of H .melpomene (7 of the aglaope pattern and 7 of the vicinus pattern) and 8 of H. elevatus (5 of the elevatus pattern and 3 of the perchlorus pattern) all from the upper Amazon basin; the identification being made by means of the basal marks. The genitalia were prepared by Miss Susan May, an assistant in the Museum, and I then sorted them into melpomene and elevatus types, without knowing which butterfly they belonged to (all preparations were of course numbered). The result was: pattern: elev. melp. elev. 7 0 genitalia: melp. 3 11 A further elevatus had deformed valves. Of the three specimens placed in the “wrong” class, one was simply an error, having ordinary melpomene genitalia; the other two had the apical extension longer and more robust than is usual in melpomene, but on re-examining them I found that the extensions were still much weaker than those found in elevatus. The probability of finding such an association between the wing and genital char- acters in a sample if there was in fact no association in the total population is less than one in one hundred (Fisher’s exact test, two tails). 5 (2):97-112, 1966 SPECIES OF HELICONIUS 103 This is good evidence that elevatus is a species distinct from melpomene. Recently differences have been found between H.melpomene and H.e.elevatus in the distribution of the an- droconia (Emsley 1965). As an aside it is worth considering the possible function of the characters of the male genitalia. Experiments by Lorkovic (1953) have discredited the “lock and key” theory at least for some European butterflies subjected to forced mating (see also other work quoted by Dobzhansky ( 1951 ) . At least three authors (Lorkovic 1956, Turner 1962, Mayr 1963) are of the opinion that the valves are highly variable within species ( and therefore also between species) because natural selection has little effect on their precise shape: if they are gripping organs then it is not of great significance what shape they are, provided that they grip; the ridges in the human “finger print” are analogous. Emsley (1963) claims that on dissecting sundry unnamed Heliconids electrocuted while mating he found that the hooked tips of the genital valves were not touching the female, who was gripped by the median organs (uncus etc.) and possibly by the bases of the valves. It is very surprising that a hook on the end of a genital valve should be functionless. If the solution to the puzzle is not that the male’s muscles had been contracted by the electrocution, then it probably rests in the courtship behaviour of the butterflies; my own unpublished observations of the Heliconid species Dryadula phaetusa (Linnaeus), which has large appendages at the tips of the valves, are typical of most Heliconid courtships. When the female settles after the courtship flight, the male alights by her side, facing in the same direction, and by bending his abdomen in a semi-circle grips the tip of the female’s abdomen with his genitalia; this done he moves so that his abdomen and that of the female are in direct line, and the butterflies face in opposite directions; they remain in this position until they part. It is possible, and this could be checked by watching matings of males whose genitalia had been clipped, that the hooks at the tips of the valves grip the female during the first phase of copulation when the couple are facing in the same direction, and that on moving his position the male changes his grip on the female, holding her from then on with median organs and the bases of the valves. The small hooks of H .melpomene and some other species may be vestigial. 104 JOHN R. G. TURNER J. Res. Lepid. melpomene Fig. 2. Differences between H. elevatus and melpomene^ as shown by the forewing upperside, hindwing underside, and the tip of the male genital S (2);97~112,1966 SPECIES OF HELICONIUS 105 elevatus valve. Basal marks on the hindwing are shaded if unshaded if yellow; no otiber marks are shaded. 106 JOHN R. G. TURNER /. Res. Lepid. Appended is a synopsis of the infra-specific forms of H.elevatus, followed by a description of a new subspecies. A drawer in the main collection of the British Museum (Natural History) at Tring shows all the subspecies of H.elevatus. A distribution map, and a discussion of the interesting taxonomic and mimetic re- lations of elevatus with other Heliconius, will appear in a detailed study of the genus now in preparation. H.elevatus seems to be closely related to, or perhaps conspecific with H.luciana Lichy, a species with a superficially very different pattern resembling that of H.antiochus ( L. ) ( see Lichy 1960 ) . In the following list only the most important references are given; the short description refers to the characters differentiat- ing the subspecies: shape of the yellow band, presence or absence of apicals and presence or absence of hindwing rays (fig. 2). Subspecific epithets have, where necessary, been given mas- culine endings in accordance with the Code, although I am by no means convinced of the wisdom of this. The subgenus is that of Michener (1942). Heliconius (Heliconius) elevatus Noldner 1. H. elevatus elevatus Noldner Heliconius elevatus. Noldner, 1901, Berlin.ent.Zeit. 46, 5. Heliconius melpomene elevatus. Stichel & Riffarth, 1905, Tierreich 22, 120; Stichel, 1906, Gen.Ins. 37,25. Heliconius elevatus. Eltringham, 1917, Trans. R.ent.Soc.Lond. 1916, 134. Heliconius elevatus elevatus. Neustetter, 1929, Lep. Cat. 36, 52. Heliconius melpomene f. elevatus. Emsley, 1964, Zoologica 49, 262. Heliconius elevatus elevatus. Emsley, 1965, Zoologica 50, 210. Narrow band; no apicals; hindwing rays. Upper Amazon basin. The form griseoviridis (H.elevatus f. griseoviridis Neustetter, 1938, Ent. Rundsch. 55, 416) is a minor variation in which the band is extended posteriorly and proximally by an area of mixed black and yellow scales. Form noeldneri (H.elevatus f. noldneri Neustetter, 1938, Ent. Rundsch. 55, 415) has abnormally strong development of red marks, having red proximal to the band, a hammer-headed red band extending from the basal red along Cus to the margin of the forewing, two rows of red subapical spots on the forewing, and abnormally wide heads to the rays on the hindwing. It gives the impression of reverting toward the “Tiger” pattern of those Heliconii which mimic Ithomiids. The types of both these varieties, in the Naturhistorisches Mu- 5 (2). -97-1 12, 1966 SPECIES OF HELICONIUS 107 seum, Wien, come from Yiirimaguas, Peru; their relation with the following subspecies is not clear. 2. H.elevatus pseiidociipidineiis stat.nov. Heliconius elevatus f. pseudoctipidineus. Neustetter, 1931, Int.ent.Zeit., Guben 25, 169. As H.e. elevatus, but band very much narrower. Perhaps only just worthy of subspecific rank. Tarapoto and Yurimaguas, N.E. Peru. Types (one male, one female) in the Naturhistorisches Museum, Wien. I select the male as the lectotype of the new subspecies. Labels on the lectotype: (1) elevatus f. pseudo=/cupidineus Neust./ ^ Type. (2) Yurimaguas/Peru/0. Michael. (3) Coll./ Neustetter. ( 4 ) H.elevatus pseudocupidineus, /lectotype ( Turner ) . The form nigromacula {H.elevatus f. nigromacula Neustetter, 1932, Zeit.6sterr.Ent.Ver., Wien 17, 15) appears to be similar to form noeldneri ( but with a narrower band ) . I have not seen the type. 3. H.elevatus perchlorus Joicey & Kaye Heliconius elevatus perchlora. Joicey & Kaye, 1917, Ann.Mag. nat.Hist. (8) 20, 94; Neustetter, 1929, Lep. Cat. 36, 52. Heliconius melpomene f. perchlorus. Emsley, 1964, Zoologica 49, 262. Heliconius elevatus perchlorus. Emsley, 1965, Zoologica 50, 210. Broad band; no apicals; hind wing rays. Bolivia, valleys north- east of Lago Titicaca. Type in the British Museum (Natural History ) ; no locality data. 4. H.elevatus taracuanus Bryk. Comb.nov. et subspecies dubia H. melpomene taracuanus. Bryk, 1953, Ark.Zool. 5(1), 76. Broad band; no apicals; hindwing rays. Taracua, Rio Uaupes (a tributary of the Rio Negro), state of Amazonas (Brasil). Type in the Naturhistoriska Riksmuseet, Stockholm. This specimen is undoubtedly an elevatus, not a melpomene, and is very similar to perchlorus; individuals of this phenotype occur among the e.elevatus on the upper Amazonas, so it is not clear whether taracuanus is simply an intrapopulation variety or whether there are monomorphic populations of this phenotype in the basin of the Rio Negro which grade into the populations on the Amazonas. 5. H.elevatus bari ( Oberthiir). Comb.nov. Heliconia hari. Oberthiir,, 1902, Etudes dentomologie 21, 23. Heliconius melpomene bari. Stichel & Riffarth, 1905, Tierreich 22, 120; Stichel, 1906, Gen.lns. 37, 25. 108 JOHN R. G. TURNER J. Res. Lepid. Heliconius melpomene thelxiope forma aquilina. Neustetter, 1925, ZeitMsterr.Ent.Ver., Wien 10,12. Syn.nov, Heliconius elevatus schmmsmanni. Joicey & Talbot, 1925, AnnMag.nat.Hkt. (9) 16,647. Syn.nov. Heliconius melpomene melpomene forma bari. Neustetter, 1929, Lep. Cat. 36, 51. Heliconius elevatus aquilina. Neustetter, 1929, Lep. Cat. 36, 52. Heliconius elevatus aquilina forma schmassmanni. Neustetter, 1929, Lep.Cat. 36, 52. Heliconius melpomene X Heliconius aoede. Emsley, 1965, Zoologica 50, 210. Broken band; apicals present or absent; hindwing rays. Mato Grosso and north Bolivia (schmassmanni) , Rio Tapajos (state of Para) and Guianas (bari). Types of bari and schmassmanni in the British Museum (Natural History), the first in the Levick Collection; localities "Guyane fran^aise” and “River System, Cuyaba-Corumba, Mato Grosso, Brasil” respectively. Types of aquilina (one male, one female) in the Naturhistorisches Mu- seum, Wien; locality “Rio Machados, Mato Grosso-.” Clearly the male is the lectotype, and I have labelled it as such. The type of bari has apical spots, those of schmassmanni and of aquilina lack them, so one could regard bari and schmassmanni as separate subspecies, the one in the north and the other in the south, but the difference is small and based upon too few specimens. 6. H. elevatus tumatumari Kaye. Comb, et stat. nov. Heliconius tumatumari. Kaye, 1906, Entomologist 39, 53; Eltringham, 1917, Trans.ent.Soc.Lond. 1916, 134. Heliconius melpomene melpomene forma turrmtumari Neu- stetter, 1929, Lep.Cat. 36, 44. Heliconius melpomene X Heliconius aoede. Emsley, 1965, Zoologica 50, 212. Broken band; apicals; no hindwing rays. Potaro River, Guyana ( = British Guiana ) . 7. H. elevatus roraima subsp.nov. Heliconius melpomene, form near to .... . eulalia. Hall, 1939, Agric.] .Brit. Guiana 10, 39; 1940, Brit.Guiana Dept.Agric. Ent.Bull. 3, 15. Heliconius elevatus, form. Emsley, 1965, Zoologica 50, 212. Broad band; no apicals; no hindwing rays. Region of Mount Roraima, Guyana. Types in the British Museum (Natural History). Emsley (1965) reports a long series in the American Museum of Natural History. 5 (2):97-112, 1966 SPECIES OF HELICONIUS 109 Heliconim (Heliconius) elevatus romima subsp.nov. Holotype male (fig. 2, E). Upperside: Forewing black-brown, the proximal third extensively marked with red posteriorly from vein Sc to just anterior to the posterior margin; the red being traversed by black-brown along the veins, along a line running longitudinally down the centre of the cell, and a line which runs posterior to vein lA for 3 mms. from the proximal edge of the wings (venation is after Michener 1942), becoming coincident with the vein distally. At the base of this dark line, a yellow spot. The distal third of the cell, and portions of the wing between Sc, Ri, R2,Mi, Ms, Ms, Cui, Cu2, and posterior to Cus occupied by yellow areas giving the effect of a large yellow mark invaded by black along the veins. The yellow marks between Ms and Cui and between Cui and Cug have V-shaped indentations distally. Hindwing black-brown, a sub-triangular area about 1 cm. long and 2 mms. deep near the base being red, traversed by black-brown veins; posterior to the edge of this red mark three diffuse red markings lying between the veins. Anterior to the red triangle a silver-brown area extending right across the an- terior margin of the wing. Underside: forewing as upperside, except that the black line in the cell is broader, that all the wing posterior to the cell and Cui is silver-brown, without markings, and that there is a basal red mark 5 mms. long between the anterior margin and vein Sc. Hindwing black-brown, a red line 4 mms. long extending from the base along the anterior margin; 2 mms. posterior to this, and lying just posterior to vein Ri -f- Sc a yellow line 10 mms. long. A red line extending across the cell and proximally in a slight curve, following chiefly the position of the posterior edge of the red triangle on the upperside; posterior to this a row of four red marks, the three distal ones occupying the same position as the three red marks on the upperside. Antennae black-brown, slightly red-brown along one edge. Head black-brown, with white marks above and below the in- sertion of the antennae and on the ventral side of the palpae. Thorax with dorso-lateral yellow markings, but with too many scales missing for a detailed description. Abdomen black-brown, the first segment with two dorso-lateral spots. Genital valves (as seen in hand lens, without dissection) with strong hook at tip. 110 JOHN R. G. TURNER J. Res. Lepid. Labels: (1) Type/HT (2) Roraima,/B. Guiana. /H. Whitely. (3) ^ (4) Godman-Salvin/ Goll. 1913-2 (5) Heliconius elevatus/ roraima Turner 1967/HolO" type Dimensions — forewing: 3.8 cms.; antenna: 2 cms.; body from head to tip of abdomen: 2.6 cms. Paratype male, labelled as the holotype except that (1) and (5) read “Paratype”: similar except that some of the small un- derside marks on the hindwing are obscure or missing. Paratype male, labelled (in manuscript ink) Roraima. /Bt. Gui- ana/ (printed) Growley/Bequest./1901 — 78 and with paratype label as above. Similar to type except for obscurity of some small underside hindwing markings. Paratype male, labelled (in pencil)? Roraima/ (printed) Growley/Bequest/1901 — 78 and with paratype labels as above. Similar to the holotype, except that the row of red dots on the upper and underside of the hindwings is absent. Paratype female, labelled as the holotype, except that (3) is and (1) and (4) are paratype labels. Differs from the holotype in that the silver-brown areas of both wings are less extensive and less obviously differentiated in colour from the rest of the wing (a characteristic expression of sexual dimor- phism in the genus Heliconius); the diffuse red marks on the hindwing are missing on the upperside and obscure on the un- derside; on the forewing the yellow mark posterior to Gus is ab- sent; and the yellow mark between M3 and Gui has a slightly curved edge instead of the V indentation; and the tips of the antennae are obviously rufous. Holotype and paratypes in British Museum (Natural History). The “red” of the above descriptions refers to a pigment which changes colour over the years; it is now an orange red, but was probably a brilliant carmine red when fresh; similarly the “black-brown” was probably almost black. AGKNOWLEDGEMENTS I am much indebted to Professor E. B. Ford, F.R.S. for help- ing and encouraging my work on the taxonomy and distribution of Heliconids. I wish to thank also the authorities of the British Museum (Natural History) at South Kensington and Tring, of 5 (2);97-112, 1966 SPECIES OF HELICONIUS 111 the Hope Department of Entomology, Oxford, the Naturhis- torisches Museum, Wien, and the Natiirhistoriska Riksmuseet, Stockholm, particularly Professor G. C. Varley, and Messrs. T. G. Howarth, E. Taylor, and G. E. Tite, for their help and for the loan of specimens, and Miss S. May for making genitalia mounts. Professor P. M. Sheppard, F.R.S. and Mr. T. G. Howarth read the draft and I am indebted to them for their helpful sug- gestions. SUMMARY The South American butterfly Heliconius elevatus has seven subspecies, all resembling closely various forms of H. melpomene; the two species differ in the male genitalia and the detailed marks on the underside of the hindwings. Correlation between the characters of the wings and genitalia show that elevatus is a distinct species and not a form of melpomene, with which it is usually confused. The nomenclature of H. elevatus is sum- marised, and a new subspecies described. REFERENCES DOBZHANSKY, TH. 1951. Genetics and the origin of species. Colum- bia U.P., New York. ELTRINGHAM, H. 1917. On specific and mimetic relationships in the genus Heliconius, L. Trans, ent. Soc. Lond. 1916, 101-148. species. Zoologica (N.Y.) 49, 245-286. EMSLEY, M. G. 1963. A morphological study of imagine Heliconiinae (Lep.: Nymphalidae) with a consideration of the evolutionary relationships with the group. Zoologica (N.Y.) 48, 85-130. EMSLEY, M. G. 1964. The geographical distribution of the color-pat- tern components of Heliconius erato and Heliconius melpomene with genetical evidence for the systematic relationship between the two specis. Zoologica (N.Y.) 49, 245-286. EMSLEY, M. G. 1965. Speciation in Heliconius (Lep., Nymphalidae): morphology and geographic distribution. Zoologica (N.Y.) 50, 191-254. FORD. E. B. 1955, Moths. Collins, London. FOX, R, M. 1956. A monograph of the Ithomiidae ( Lepidoptera), Part 1. Bull. Amer. Mus. Nat. Hist. Ill (1), 1-76. KERKIS, J. 1931. Vergleichende Studien liber die Variabilitat der Merk- male des Geschlechts Apparats und der aiisseren Merkmale bei Eury- gaster intergriceps Put. Zool. Anz. 93, 129-143. LICHY, R, 1960. Documentos paraservir al estudio de los lepidopteres de Venezuela. IV. Una especie nueva del genero Heliconius Kluk ( Rhopalocera, Nymphalidae): Heliconius luciana sp.n. Rev. Fac. Ag- ron. (Maracay) 2 (3), 20-44. LORKOVIC, Z. 1953. L’accouplement artificiel chez les Lepidopteres et son application dans les reciierches siir la fonction de Pappareil geni- tal des insectes. Physiol. Comp. OEcol. 3, 313-320. 112 JOHN R. G. TURNER J. Res. Lepid, LORKOVIC, Z. 1956. Zavisnost varijabilnosti organa muskog genital- nog aparata kukaca a njihovoj funkcionalnoj vrijednosti. (Variability of the organs of the genital armature in insects due to their functional value.) Bioloski Glasnik 7, 234-235. MAYR, E. 1963, Animal species and evolution. Harvard U.P. Cambridge. MICHENER, C. D. 1942, A generic revision of the Heliconiinae (Lepi- doptera, Nymphalidae). Amer. Mus. Novit. 1197, pp. 8. NEUSTETTER, H. 1929. N3nnphalididae: subfam. Heliconiinae. Lepi~ dopterorum Catalogus pars 36. Junk, Berlin. OBERTHUR, C, 1916. Observations relatives aux planches CDIII, CDIV, CDV representant des Heliconia. Et. Lep. Comp. fasc. XII, 2e. partie, 30-38. Oberthiir, Rennes. SHEPPARD, P. M. 1963. Some genetic studies of Mullerian mimics in butterflies of the genus Heliconius. Zoologica (N.Y.) 48, 145-154. TURNER, J, R. G. 1962. A metliod for mass-producing mounts of Lepi- doptera genitalia, with a note on their evolutionary significance. Ento- mologist 95, 146-148. TURNER, J. R. G. 1963a. Geographical variation and evolution in the males of the butterfly Papilio dardanus Brown ( Lepidoptera: Papilioni- dae). Trans. R. ent. Soc. Lond. 115, 239-259. TURNER, J.R.G. 1963b. Mimetic multilocus polymorphism in South American butterflies {Heliconius spp. ) ( Lepidoptera, Nymphalidae). Proc. XI Int. Cong. Genet. Hague 1, 146. TURNER, J.R.G. 1965. Evolution of complex polymorphism and mimi- cry in distasteful South American butterflies. Proc. XII int. Congr. Ent. London 1964, 267. TURNER, J. R. G. & CRANE, J. 1962. The genetics of some polymor- phic forms of the butterflies Heliconius melpomene Linnaeus and H. erato Linnaeus. 1. Major genes. Zoologica (N.Y.) 47, 141-152. Journal of Research on the Lepidoptera 5 (2) :113»126, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A..91006 © Copyright 1966 COMPARATIVE SPECIATION IN TWO BUTTERFLY FAMILIES, PIERIDAE AND NYMPHALIDAE BJORN PETERSEN Departm,ent of Entomology, University of Lund, Sweden It is nowadays generally accepted that the main factors influencing evolution are mutation, recombination, selection, chance, and isolation. Naturally these five factors do not play the same role in all groups of animals and plants. Thus, for instance, the evolution of polyploid forms is of much greater importance in plants than in animals. Similarly the selective forces affecting the evolution of protozoans and mammals must be very different. The branch of biology dealing with such prob- lems we may call comparative evolution in analogy with com- parative anatomy. Comparative speciation, with which this paper is concerned, is a part of this science. An important method is here to compare the differences existing between the closest related species of different groups. Such differences must be the result of the evolution having taken place since the time of the common ancestral form. It is possible that the differences evolved are an integrating part of the mechanism isolating the two species. In other cases the differences may be the result of adaptations to different environmental conditions or be just a matter of chance. The two groups discussed here are the two butterfly families Pieridae and Nymphalidae. All sympatric closely related Euro- pean species of these two families are compared. To these species the two North American sibling species Colias eury theme and C. philodice have been added as they are especially well known predominantly from the investigations by Hovanitz ( 1943, 1948, 1949 a,b). Fam. Pieridae. 1. Pieris napi L. — P. bryonme Ochs. These two forms do not behave as different species in all areas where they meet. They do however in some areas which justifies a discussion of their differences. Male patterns rather similar. 113 114 PETERSON /. Res. Lepid. Female patterns more extended in bryoniae than in napi. Male wing color similar, female wing color white in napi, yellow in bryoniae. Genitalia similar ( Drosihn 1933 ) . Foodplants: napi: various cruciferous plants. bryoniae: Biscutella laevigata. 2. Pieris napi L. — P. ergane Hbn. The patterns of ergane are closer to those of P. rapae than to those of napi. The color of the females is sometimes yellow in ergane, less often in napi. The genitalia are similar (Drosihn 1933) as well as the scent of the males (Lorkovic, personal communication ) . Foodplants: napi: various cruciferous plants. ergane: various cruciferous plants (Forster and Wohlfahrt 1955, Verity 1947), Aethionema saxatile in the French Alps and Pyrenees (Descimon 1964, 1966), The very local oc- currence of this species makes a wide variety of foodplants rather unlikely. 3. P. rapae L. — F. manni Mayer. The patterns of the two species are rather similar, the back- ground color of the females less yellow in manni than in rapae. Genitalia similar. Foodplants: rapae: various cruciferous plants. manni: Diplofaxis temiifolia (Italy), D. eru~ coides, Lepidium graminifolium, Iberis spp., not on cultivated cruciferous plants (Verity 1947). 4. Colias hecla Lef. — C. nastes Bdv. The patterns are markedly different in the males, less so in the females. The color is in both sexes orange in hecla and yellow in nastes. Genitalia very similar. The number of teeth of the aedeagus on the average slightly higher in hecla. Only one specimen of 41 + 34 outside the range of variation of the other (Petersen 1963 b). Foodplants: hecla: Hedysarum americanum. Astragalus al- piniis. nastes: Astragalus deflexus and alpinus. 5. C. hyale L. — C. australis Ver. Slight differences are present in the patterns. The yellow color in australis is more “beautiful”, in hyale more greenish and “dirty” ( Reissinger 1960 ) . 5 (2): 113-126, 1966 COMPARATIVE SPECIATION 115 Genitalia slightly different (Warren 1951, Beuret 1951, Bret- schneider 1959, Schadewald 1959), Berger and Fontaine (1947), Reissinger (1960), and Petersen (1963 b) found no differences. Foodplants: hyale: various leguminose plants. australis: Hippocrepis comosa. Male scent: hyale: strong aromatic scent (Berger and Fon- taine 1947), rather peppery (Riley 1961). australis: absent or almost absent (Berger and Fontaine 1947), sweetish, honeylike (Riley 1961). 6. C. myrmidone Esp. — C. erate Esp. The patterns are markedly different in the males less so in the females. The color is orange in myrmidone, usually yellow in erate. Genitalia very similar ( Petersen 1963 b ) . Foodplants: myrmidone: Cytisus spp. erate: unknown. 7. C. eurytheme Bdv. -= C. philodice Gdt. As in two of the previous Colias species the pattern differences are greater in the males than in the females. Color is in eurytheme orange, in philodice yellow. Genitalia very similar ( Petersen 1963 b ) . Foodplants: eurytheme: Medicago. philodice: Trifolium. Male scent of eurytheme like sweet heliotrope. 8. C. aurorina H. Sch. — C. sagartia Led. Pattern differences about the same as in previous species. The color is in aurorina orange, in sagartia bluish green. Genitalia very similar ( Petersen 1963 b ) . Foodplants unknown. Only C. aurorina occurs in Europe. C. sagartia lives in S. W. Asia. 9. Gonepteryx rhamni L. ~ G. cleopatra L. There are hardly any patterns on the wings of the two species. The forewings of cleopatra males are orange except along the borders, and the yellow color is more saturated. The females of cleopatra have yellow borders on the wings. Genitalia: uncus much smaller in cleopatra, otherwise rather similar ( Verity 1947 ) . Foodplants: rhamni: Rhamnus spp., Vaccinium, Rosa, Prunus spinosa. cleopatra: mainly on other Rhamnus spp. than 116 PETERSON /. Res. Lepid. rhamni (Verity 1947). Male scent: rhamni: absent. Cleopatra: rich and powerful, freesia (strong) (Ford 1945). 10. Anthocaris cardamines L. — A. damone Boisd. and A. euphenoides Stgr. The two latter species are allopatric but not contiguous. The areas of distribution come fairly close together in Italy (cf. Verity 1947). Their great similarity suggests conspecificity. At least, they may both have arisen jointly from the cardamines stock and therefore cannot be treated here as two independent cases. The differences in pattern between cardamines and damone are slight compared with the geographical variations within the species. The ground colors of the males (yellow in damone and white in cardamines) are strikingly different. The differences be- tween the females are slight or none. Genitalia differences are only of a quantitative nature (Verity 1947). Foodplants. cardamines: various cruciferous plants. Reseda, damone: Isatis tinctoria and canescens. euphenoides: Biscutella spp. Fam. Nymphalidae. M edict a Billb. Higgins (1955) divides this genus into twelve species, seven of which occur in Europe. Three of these species, aurelia Nick. deione Geyer, and asteria Freyer together with the subsp. athalia celadussa Fruhst. do not revelop subuncal processes be- low the tegumen. Verity (1950) is of the opinion that M. deione and athalia celadussa are most similar and he connects on the other hand M. aurelia with M. asteria. It may very well be that one subspecies ( ) of one species is closest to one of the other species (B), while another subspecies (Ao) is closer to a third species (C). M. athalia with its marked intraspecific variation in the genitalia may very well be a species of type A. This, how- ever, is of lesser significance in the present investigation as the differences in the genitalia betweeen the forms mentioned are in all cases small but well distinct, colors similar, patterns similar or rather similar. 11. M. deione — M. athalia celadussa Golors and patterns similar but variable. 5 (2): 113-126, 1966 COMPARATIVE SPECIATION 117 Genitalia: small but distinct differences (Verity 1950). Foodplants. deione: Linaria spp., Antirrhinum spp. athalia celadussa: Melampyrum, Plantago. 12. M. aurelia — M. asteria. Colors and patterns similar. M. asteria much smaller. Genitalia, small but distinct differences (Verity 1950 p. 196). Foodplants: aurelia: Melampyrum, Veronica^ Digitalis^ Chry- santemum, Plantago. asteria: unknown. 13. Euphydryas cynthia Schiff. — E. intermedia Men. Color similar in the females, different in the males. Patterns more different in the males than in the females. Genitalia: slightly different (Higgins 1950). Foodplants. cynthia: Alchemilla, Plantago alpina and lance- olata, Viola, Vaccinium mijrtillus, Pedicularis. intermedia: Fraxinus, Populus, Salix, Alnus, Vi- hurnum, Melampyrum, Veronica, Scabiosa, Plant- ago, Viola, Prunus, Lonicera (Verity 1950). 14. Fahriciana adippe Rott. — F. niobe L. Color and patterns rather similar on the upper side; on the under side the colors are rather different. Genitalia: various parts shorter and thicker in niobe (Verity 1950). Foodplants: adippe: Viola spp. niobe: Viola spp., Plantago spp. 15. Brenthis ino Rott. — B. daphne Schiff. Patterns are slightly different in both sexes on both sides of the wings. Colors are similar. Genitalia: Verity (1950 p. 259) states that the differences found by Warren (1944) and dos Passos and Grey (1945) are individual variations present in both species. Foodplants: ino: Sangtiisorba, Spiraea, Rubus. daphne: Rubus, Viola. 16. Boloria pales Schiff. — B, olethea Hemming (arsilache) Esp. ) 1 Colors are similar in both sexes, patterns slightly different. I Genitalia: I was not able to confirm the differences found by Warren (1944). Foodplants: pales: Viola spp. ' alethea: Vaccinium oxy coccus, Polygonum spp., ■ Viola spp. i 17. Boloria frigga Thnbg. — B. improba Btl. 118 PETERSON /. Res. Lepid. B. improba is much darker and smaller than frigga. The patterns on the under side are rather similar. Genitalia: Slight differences are present (Bruun and von Schantz 1949). Foodplants: frigga: Rubus chamaemorus (Lingonblad 1946). improba: probably Salix herbacea (Bruun and von Schantz 1949 ) . In tables 1 and 2 the differences between the species have been summarized and indicated by value symbols. It is not al- ways easy to decide which symbol should be used but he tenden- cy is quite obvious: the differences in patterns are of the same magnitude in both families while differences in color are more marked in the Pieridae. Specific differentation without any or with very little change in the genitalia is the rule in the genera Pieris and C olios, but has probably also taken place in Brenthis and Boloria. When the colors of the Nymphalidae are different the differ- ences are usually restricted to the under side, except for in- trusion of melanistic scales on the upper side as in Boloria improba as compared with frigga. In the Pieridae the color differences are most striking on the upper side and are due to the intrusion of white, yellow, and orange pterins. In the C olios aurorina-group blue pigments are also present. In the genus C olios both sexes are different in color, in Pieris only the females, and in Gonepteryx and Anthocaris mainly the males. In the genera Pieris and C olios where the interspecific variation in female color is greatest, the differentiation in the male genitalia is less pronounced. TTiere are several reasons why closely related pierid species should be more divergent in color than Nymphalids. Pterins of white, yellow, and orange color are chemically related. The yellow color of P. bryoniae flavescens turns white when the pupae are exposed to pure oxygen during their development. Species of other families do not change color after similar treat- ment ( Deschka and Reichl 1964 ) . A mutation changing the wing color may happen more often in butterflies having pterins in their wing scales than in other Lepidoptera. The colors of butterflies have several functions: warning, concealment, sexual attraction and stimulation. In connection with isolation between species the sexual functions are of special interest. The colors of the females are known to attract the males though supraoptimal colors sometimes are present (Tin- bergen et al. 1952, Petersen et al. 1952). Thus the yellow color 5 (2): 113^126, 1966 COMPARATIVE SPECIATION 119 of P. hryoniae females is less attractive to their males than the white color (without ultraviolet) of napi females and males of both forms. Yellow color may have a higher selective value be- cause yellow females are less likely to hybridize. The yellow color may also be part of a protective coloration. The colors of the males have been supposed by Ford (1953, 1965) to have a sexually stimulating effect. As far as I know con- clusive experiments are still lacking. The presence of strikingly different colors in so many sibling species supports Ford's hypothesis and makes experimental evidence even more de- sirable. Differences in the foodplants of the larvae are very marked. In one case only, P. napi - ergane some authors mention "different cruciferous plants” for both species. Aetionema saxatile is at least in France the most important foodplant for ergane. In nine of the seventeen pairs of table 1 one of the species has a very wide distribution compared with the other. These species have been underlined in the table. As P. rmpi is present in two pairs there are together eight species of this kind. These eight species are all adapted to a temperate climate, some of them also to colder and warmer climates. Their larvae live in seven cases on plants of more than one genus while six of the species with restricted distributions live on only one plant genus, usually on a single species. Ecologically they are adapted to an Arctic or alpine climate or to the climate of the Mediterranean area. The wider ecological amplitude of the central species com- pared with the 'edge' species is in agreement with the ideas of Brown (1957). He points out that central species are more “potent' and therefore the source of higher categories. Contrary to this Mayr (1954) has stressed the importance of the edge populations, especially on islands. Such isolated populations can under unique environmental conditions form the real novelties of the animal world. It may be noted that the only novelty (not real novelty ) among the species treated here belongs to the edge species: Boloria improba. Living at a higher altitude than any other Scandinavian butterfly and with a concealing coloration well adapted to the dark rocks and stones at this it may be potentially equivalent to the many small Erehm and MelUcta spp. etc. of the higher parts of the Alps. However, the evolution of a new (sub) genus under such extremes is rendered less 120 PETERSON J. Res. Lepid. a •a c a “■5 0|0 in %o j1o1 p nJ £. u o :i T3 (ti 1) ^ - nj 1-1 ^ < P S a aa H 2 S| Table 1. Presence or absence of differences in some characters between sibling species of the families Pieridae and Nymphalidae. (+) = slight differences in color and pattern. Species in caps = widely distri- buted species, a= adapted to arctic or alpine climate; m = distribution mainly in the Mediterranean area. 5 (2): 113-126, 1966 COMPARATIVE SPECIATION 121 probable on account of the hazardous conditions for survival during the long time required for such a process. The more restricted ecological amplitude of the edge species may in some cases depend on competition from the central species. Sometimes , however, this is certainly not the explanation. P. bryoniae and A. euphenoides , two edge species, both live on Biscutella. This is the only plant genus of the family Cruciferae which is common at the altitude where the butterflies live. The close relationship larva — foodplant might from the beginning have been more or less a matter of coincidence following the adaptation to a certain climate and a plant family as source of food. However, as soon as the relationships have become established they become closer. When today P. bryoniae lives on Biscutella on the southern side of the Alps it is able to compete with P. napi down below 300 m ( Posavje, near Ljubljana, Yugoslavia ) . When both live on Arabia halleri, as on Monte Mottarone in the Italian Alps, the lower limit of bryoniae is at 700 m (Petersen 1955). By their morphology the edge species are exposed to a different selection than the central species. The polyphagy of the central species leads to an increase of the variation of their populations. The greater variation of the central species by the combination of geographic variation and dispersal (Mayr 1954) is in this way increased. patterns color genitalia larval foodplant over- + (+) - + {+) ” + lapping different Pieridae cf 4 5 0 6 1 3 2 1 7 $ 1 8 0 5 5 0 3(2) 5(6) total 5 13 0 11 6 3 2 1 7 Nymphalidae d* 1 2 4 1 2 1 4 5 2 0 1 ? 1 5 1 1 1 5 4 3 total 3 9 2 3 2 9 5 2 0 Table 2. Differences between sibling species of the families Heridae and Nymphalidae according to table 1. 122 PETERSON /. Res. Lepid. ISOLATING MECHANISMS - ! Ethological isolation. The differences between species leading ! to ethological isolation are thus different in the two families in- i vestigated. Pieridae often have different colors and in at least two cases different scents (G. hijale-australis and G. rhamni- cleopatra). Two species have separated from P, mpi without : developing recognizable scent differences. J The attractiveness of the color in P. napi is reduced to about ^ 1/10 by a very small amount of yellow pterins in the wings ! (Petersen 1963 a p. 224). Bees in training experiments are able :i to separate four colors only (Hertz 1937). Perhaps the reaction [ of the napi butterflies means that a limit between two of the colors in their color vision is situated between white (without [ ultraviolet) and the same white with a slight mixture of yellow. [ A similar slight difference is present in other pairs {napi - ergane, mpae - manni^ and cardamines - euphenoides) . It is possible that a + from the point of view of the butterflies is a : better expression of the differences than the ( + ) of table 1. The genitalia of many species of insects and other animals are i- often so characteristic that they enable the separation of closely i related species (cf. Dobzhansky 1941 p. 267). This has given I' rise to the lock-key theory. According to this theory the genitalia of males and females of a species match another so exactly that even a small deviation makes copulation physiologically im- j’ possible. Some experiments and observations tend to support to ' the lock-key theory but others show that the original formulation j was strongly exaggerated. Even after the removal of large parts [ of the genitalia except the aedeagus a male is able to copulate i| with a female of its own species ( Sengiin 1944, Lorkovic 1953 ) . ' These results led to the conclusion that a structural change | played no role in the isolation between species. No quantitative j: estimation of the isolation ( e. g. by choice experiments ) has ever '■ been made. Observations by Standfuss (1896), Federley (1932), j; and Sturtevant (1921) show that different genitalia probably . cause a certain isolation. I Differences between related species are more common where ; the genitalia have complicated structures as in the Nymphalidae, j In groups with less complicated genital apparatus (as in fam. j: pieridae) inter- and intraspecific variation is smaller. Dobzhan- sky, (1. c. ) suggests that a complicated structure is more often f changed in connection with the genetic revolution within a ] population. However, this does not explain the evolution of the | 5 (2): 113-126, 1966 COMPARATIVE SPECIATION 123 complicated structures which in many groups are more common in the genitalia than in any other part of the body. This may be explained by a greater efficiency of a complicated genital ap- paratus. The possibility of a greater chance of developing iso- lation between species as a factor promoting complication of structure cannot be excluded. A character which permits rapid speciation ( within reasonable limits ) must in the long run have a positive selective value. The ecological isolation is to a great extent dependent on the larval foodplant. The "typical” condition with a central species of wide distribution and a wide variety of foodplants and an edge species living on a single plant genus or species is present in only four (five) pairs (1, 3, 5, 10, perhaps also in 2), all of the family Pieridae. In another four pairs the foodplants are different, in seven (six) partly overlapping. No species have the same food- plants. In four pairs one of the members is a lowland species, the other a mountain species. As a consequence of this difference not only a spatial but also a temporal isolation exists as the mountain species are on the wing later in the season. Hybrids between sibling species have been found in several cases, though only within the family Pieridae. The hybrids be- tween C. eury theme and philodice show good though slightly lowered vitality (Hovanitz 1953). A similar result was obtained by Petersen and Tenow (1954) in crosses between sympatric bryoniae and napi from the northern Alps. In both cases the vitality of Fa -specimens was low. Fa -specimens of normal vitality where obtained by Bowden (1956) when crossing P. bryoniae from the Alps with English napi. In the genus C olios a number of different hybrids have been found, for instance sagartia x aurorina ( Lederer 1941 ) and nastes X hecla (Hovanitz 1963) among the pairs studied. Hovanitz (1. c. ) mentions another seven combinations of C olios hybrids described in the literature. In the genera Pieris and Anthocaris extensive hybridization experiments have been made by Lorkovic (1928, 1953). In crosses P. monni $ x rapae $ only males were obtained, a characteristic not uncommon among Lepidoptera. The reciprocal cross is in this respect normal. All hybrids are sterile. The ovaries of the females are rudimentary. Hybrids were obtained in the following crosses too, though the sexes are not mentioned. 124 PETERSON /. Res. Lepid. P. napi $ P. napi S P. napi S X ergane $ X manni 2 X rapae 2 A. cardamines S A. cardamines S X euphenoides 2 X ausonia 2 Adult Fi hybrids are thus obtained even between species less closely related than those counted as sibling species here. It seems, however, doubtful whether any of these hybrids exist in nature. The fact that so many more hybrids of Pieridae than of Nymphalidae have been found in nature can be explained in several ways : I 1. The Pieridae have in this respect been much more closely j studied. || 2. Hybrids are more easily detected in the Pieridae because of the clearcut difference in color and pattern. i 3. Sibling species in the Pieridae are genetically more similar because genetically similar populations can become isolated ! ethologically by color differences. Of the two families Pieridae and Nymphalidae the latter is the li most successful in so far as it has a higher number of species. The only factor among those investigated which could be j responsible for this is the more complicated structure of the j genitalia. Only the investigation of a greater number of families ; could give the answer whether this is an important factor in the ' evolution of species. , SUMMARY Ten pairs of sibling species, mainly European, of the family Pieridae and seven of the family Nymphalidae are compared. Differences in patterns, color, male genitalia, larval foodplants, and distribution have been investigated. The Pieridae are more often different in color, the Nymph- alidae in the male genitalia. The differences are of about the ! same size in the patterns of both families. The differences in color between Pierids are attributed to the greater chance in this family of a mutation markedly changing the color due to the presence in the wings of chemically related pterins of white, yellow, and orange color. These colors are ! supposed to be sexual attractants and stimulants as has been proved for the white color of P. napi, ! The greater differences in the genitalia between Nymphalid i 5 (2;.- 113^126, 1966 COMPARATIVE SPECIATION 125 species are connected with more complicated structures in this family. The foodplants of the larvae are markedly different in 8 of the 15 pairs where they are known. Species with a wide range usually live on a greater number of foodplants. In almost all pairs one of the species lives in the Mediter- ranean area or is adapted to Arctic or alpine conditions. Only under such circumstances geographical isolation leading to speciation was possible. LITERATURE CITED BERGER L. A. and M. FONTAINE, 1947, 1948. 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Res. Lepidoptera, 1:135 - 156. PETERSEN, B., O. TORNBLOM and N.-O. BODIN, 1952. Verhaltens- studien am Rapsweissling und Bergweissling ( Pieris napi L. und Pieris bryoniae Ochs.) Behaviour, IV:67 - 84. PETERSEN, B. and O, TENOW, 1954. Studien am Rapsweissling und Bergweissling ( Pieris napi L. und Pieris bryoniae O. ) Isolation und Paarungsbiologie. Zoo/. Bidr. Uppsala, 30:169- 198. REISSINGER, E., 1960. Die Unterscheidung von Colias hyale L. und Colias au.stralis Verity (Lep. Pieridae). Ent. Z. Stuttgart, 70:117- 131, 133 - 140, 148 - 156, 160 - 162. RILEY, N. D., 1961. The separation of Colias hyale L. and Colias australis Verity (Lep. Pieridae). Entomologist 94:296-210. SCHADEWALD, S., 1959. Colias australis calida Verity und hyale L. bei Jena in Thiiringen. Nachr.hl. Bayer. Ent. VIII: 49 - 52. SENGtiN, A., 1944. Experimente zur sexuell-mechanischen Isolation. Rev. fac. sci. univ. Istanbul, 9 B:239 - 253. STANDFUSS, M., 1896. Handbuch der palaarktischen Gross-Schmetterlinge fur Forscher und Sammler. Jena. STURTEVANT, A. H., 1921. The North American species of Drosophila. Carnegie Inst. Washington Puhl., 301:1 - 150. TINBERGEN, N., B. J. D. MEEUSE, L. K. BOEREMA, and W. W. WAROSSIEAU, 1942. Die Balz des Samtfalters, Eumenis {= Satyrus) semele ( L. ) . Z Tierpsych. 5:182- 226. VERITY, R., 1947, 1950. Le farfalle diurne dTtalia. Ill, IV. Firenze. WARREN, B. C. S., 1944. Review of the classification of the Argynnidi: with a systematic revision of the genus Boloria (Lepidoptera, Nymphalidae). Trans. Ent. Soc. London, 94:1 - 101. , 1951. Speciation in the genus Colias: with special reference to C. hyale and C. australis. Lamhill. 50 ( 1950) :90 - 98. Journal of Research on the Lepidoptera 5 (2) : 127-128, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE BUTTERFLY FAUNA OF A YELLOW PINE FOREST COMMUNITY IN THE SIERRA NEVADA, CALIFORNIA OAKLEY SHIELDS 5151 Alzeda Drive, La Mesa, California Below is a list of butterflies I collected in the vicinity of Carnegie Experimental Garden, 4600 feet, one mile east of Mather, Sierra Nevada Moutains, Tiioloiimne County, California. These were collected in an area of about one square mile during three summers: June 19 to September 6, 1964; June 14 to August 9, 1965; and June 20 to September 9, 1966. The area is a Transition Zone with a Yellow Pine Forest plant community; Pinus ponderosa, Lihocedrus decurrens, and Quer- cus Kelloggii are the predominant trees. Other indicator plants present are Abies concolor, Ribes Roezlii, and Ceanothus integerrimus. This plant community can be subdivided into micro-habitats of wet and dry meadows, stream banks, forest glades, and open and tree covered benches and slopes. PAPILIONIDAE Papilio zelicaon Papilio rutulus Papilio multicaudata Papilio eurymedon PIERIDAE Neophasia menapia Pieris protodice Pieris rapae Colias eunjtheme Anthocaris lanceolate Euchloe creiisa hyantis DANAIDAE Danaus plexippus SATYRIDAE Coenonympha tullia californica Cercyonis silvestris NYMPHALIDAE Limenitis lorquini Adelpha bredowii californica Vanessa atalanta Vanessa virginiensis Vanessa cardui Vanessa carye Junonia coenia Nymphalis californica Nymphalis antiopa Polygonia faunus rust ic us Chlosyne palla Phyciodes mylitta Phyciodes campestris Euphydryas chalcedona Euphydryas editha rubicunda ^^Speyeria zerene Speyeria calUppe inornata ^Speyeria hydaspe Sveyeria morwonia arge ’^Speyeria cybele leto LYCAENIDAE Habrodais gruntis Atlides halesus Mitoura spinetorum ^Mitoura johnsoni "^Mitoura nelsoni Incisalia augustinus iroidcs Incisalia eryphon Strymon melinus Satyrium californica Satyrium saepium Callophrys dumetorum Lycaena arota Lycaena xanthoides Lycaena editha Lycaena helloides Leptotes marina Plebejus saepiolus Plebejus icarioides Plebejus acmon ^Evres amyntula Philotes battoides intermedia Philotes enoptes Scolitantides piasus Glaucopsyche lygdamus behrii Celastrina argiolus echo HESPERIIDAE Amblyscirtes vialis Ochlodes sylvanoides Atalopedes campestris Polites sabuleti tecumseh Polites sonora "^Hesperia harpalus yosemite Hesperia juba Heliopetes ericetorum Pyrgus ruralis Pyrgus communis Erynnis persius Enjnnis lucilius afranius Erynnis propertius Thorybes pylades ^Thorybes diversus Epargyreus clarus 127 12S OAKLEY SHIELDS /. Res. Lepid. The species that Garth and Tilden (1963) consider restricted to the Transition Zone in Yosemite National Park are starred ( * ); only Colias occidentalis chrysomelas was not seen at Mather of their 10 species listed as indicators. 12 of 21 species they con- sider indicative of the Upper Sonoran Zone are also present at Mather. This may be partly explained by the fact that the Upper Sonoran Zone is about one air mile to the north so that some inflow of species might be expected. Also, some of these species breed in the Transition Zone as well, notably Incisalia augustinm iroides, Lycaena arota, T horybes pylades, and Epargyreus clarus. All of the 33 species listed by Garth and Tilden ( 1963) as occur- ring at Mather were duplicated in this study except Speyeria eg- leis. They do not list Speyeria mormonia urge for Mather, which is abundant in July in the meadows. The arge may have been mistaken for egleis since the two closely resemble each other. Some species were conspicuous by their absence at Mather: Farnassius clodius sol, Polygonia zephyrus, and Satyrium sylvinus. The food plants for all three were abundant. Earlier collecting in the year may produce such species as Anthocaris sara and Incisalia fotis windL Emmel and Emmel ( 1963 ) found 74 species in a six square mile area at Donner Pass, Placer County, California, between 6900 and 8300 feet. The zones included Transition, Canadian, and Hudsonian. It is interesting that 48 species (64.9%) are found both at Donner Pass and at Mather, and that the family com- position of both places is so similar: NO. OF SPECIES % OF TOTAL SPECIES FAMILY M^ D*^ M D PAPILIONIDAE 4 5 5.4 6.8 PIERIDAE 6 8 8.1 10.8 DANAIDAE 1 1 1.4 1.4 SATYRIDAE 2 2 2.7 2.7 NYMPHALIDAE 20 20 27.0 27.0 LYCAENIDAE 25 28 33.8 37.8 HESPERHDAE 16 10 21.6 13.5 totals 74 74 100.0% 100.0% ( *M rr Mather, D = Donner Pass) BIBLIOGRAPHY EMMEL, T. C., & J. F. EMMEL, 1963. Composition and relative abun- dance in a Temperate Zone butterfly fauna. /. Res. Lepid. 1: 97-108. GARTH, J. S., & J. W. TILDEN, 1963. Yosemite butterflies. /. Res. Lepid. 2: 1-96. MUNZ, P. A., & D. D. KECK, 1965. A California flora. University of California Press, Berkeley and Los Angeles, 1681 pp. THE JOUI^NJAL ©F RESEARCH ©Ml THE LEPIJ©©FTERA Volume 5 Number 2 June, 1966 IN THIS ISSUE Morphology of the Immature Stages of Everes Comyntas Godart Donald A. Lawrence and John C. Downey 61 A Little-Recognised Species of Heliconius Butterfly John R. G. Turner 97 Gomparative Speciation In Two Butterfly Families, Pieridae and Nymphalidae Bjdm Petersen 113 The Butterfly Fauna of a Yellow Pine Forest Gommunity Oakley Shields 127 ' September, 1966 Number 3 Volume 5 THE JOURNJAL ©F RESEARCH OH THE LEFIBOPTEKA a quarterly published at 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HOVANITZ THE PURPOSE OF THE JOURNAL is to combine in one source the work in this field for the aid of students of this group of insects in a way not at present available. THE JOURNAL will attempt to publish primarily only critical and complete papers of an analytical nature, though there will be a limited section devoted to shorter papers and notes. QUALITY WORK on any aspects of research on the Lepidoptera is invited. Analytical and well illustrated works are pre- ferred, with a minimum of long description. AUTHORS ARE REQUESTED to refer to the journal as an example of the form to be used in preparing their manuscripts. Illu- strations should be of the best quality black and white, or line draw- ings and should be pre-arranged by the author to fit a reduced size of 4” X 6V2.” Footnotes should be avoided; bibliography should be as indicated. Tables should be set-up for page size as indicated. Manuscripts in good form and requiring little work by the editor will naturally appear first. Authors, who wish drawings made for them, may submit rough sketches and will be billed for the cost, which will be very negligible. Work to be done on research grants should so specify. When possible, tabular matter should be typed on good paper with a carbon ribbon in a form suitable for a one-third reduction and in a size to fit 4” X 6V2'’ THE JOURNAL is not a suitable place for continued changes of nomenclature; unless the author is himself analytically studying a group from its biological point of view and finds a change necessary, the editor must ask authors to refrain from any changes from the McDunnough Check List unless superseded by a monograph published since that date. Popular books are not to be considered as giving scien- tific credence to any name. It is rare that name changes need be made and preference is given to old names unless in the editor s opinion sufficient evidence is given to warrant such change. SUBSCRIPTIONS should be sent to the above address. RATES are: $8.00 per volume, personal subscription (but see below) $12.00 per volume, institutional subscription. The personal subscription rate is included in the membership to the Lepidoptera Foundation indicated below. SPECIAL SERVICE TO FOREIGN ADDRESSES: THE JOURNAL will be maUed air mail or registered at cost to the subscriber, if so desired. Second Class Postage Paid at Arcadia, Calif. Journal of Research on the Lepidoptera 5(3) : 129-136, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 LABORATORY TECHNIQUES FOR MAINTAINING CULTURES OF THE MONARCH BUTTERFLY. F. A. URQUHART AND R. W. STEGNER Dept, of Zoology and Scarborough College, University of Toronto, Canada, and Dept, of Biological Sciences, University of Delaware INTRODUCTION The methods -described here represent the experiences of many years of work carried on at the University of Toronto and University of Delaware. Some of the procedures reported have been developed at both laboratories while others were developed at one particular laboratory. Our research has sometimes required mass rearing for migra- tion studies, and at other times the rearing of smaller numbers for studies on various research projects such as the study of scent receptors, scent organs, development rates, virus infec- tions, light periodicity, behaviour, and chemoreceptor stimulation by the food plant. For student laboratory exercises, the Monarch butterfly has proved to be an effective species for illustrating various aspects of invertebrate physiology such as, location of scent receptors, responses to light and temperature, effects of temperature on development rate, flight mechanisms, and receptor-oviposition responses. The Delaware Laboratory has supplied approxi- mately one hundred Elementary School Teachers in many States with viable eggs with which to carry out studies of the Monarch butterfly life cycle. The present paper outlines the culturing procedures we have followed in one or both laboratories, with a brief discussion of the use of artificial diets in maintaining living specimens of all stages during the winter months. ADULTS Flight Cages ( 60 cm. x 90 cm. x 90 cm. ) : These cages, of wood frame and cloth screen construction, have one side left open. The open end is fitted with a hinged, screened door or is covered with cheese cloth; the latter is pre- ferred since it allows easy access to the interior of the cage with- out liberating any of the actively flying butterflies, because the 129 130 URQUHART AND STEGNER J. Res. Lepid. cheese cloth can be pulled slightly aside allowing the specimen to be captured, at the same time keeping the cloth secured both on the cage and around one’s arm as one reaches into the cage. This open end also allows for introducing milkweed plants. By turning the cage so that the open end faces upward, it can also be used for mass rearing (Urquhart, 1960). Oviposition Cage ( 30 cm. x 30 cm. x 60 cm. ) : This is a smaller cage of the same construction as the flight cage, but it is open at the top and bottom. The cage is placed over a growing milkweed plant and the top covered with a piece of glass or a wood frame covered with plastic sheeting or cloth screen. The removable frame top or glass is pushed to one end thus allowing for removal of the eggs from the milk- weed plants without allowing the escape of the occupants. The smaller cage forces the gravid females to come into frequent contact with the milkweed plants, thus stimulating more rapid oviposition. Cylindrical Cage (90 cm. in height x 50 cm. in diam. ): This type of cage is constructed of wire mesh and equipped with two openings -- one near the top and one near the bottom. The openings are closed with wire mesh doors which swing open in or out on rings located at the tops of the doors. No latches are required. A wire mesh bottom keeps the butterflies from coming in contact with the honey solution that may have accumulated on the floor of the cage. With the bottom left open, a smaller form of this cage could be placed over a milkweed plant for oviposition. Cartons and Envelopes-. We have held adults alive for periods in excess of five months in small cardboard cartons (pint-sized freezer cartons). A small cotton pad, soaked in a four per cent honey solution, placed in the bottom of the carton, maintains a high R.H. and, at the same time, supplies the necessary nourishment. The carton is com- pletely sealed. The specimens are held at 7°C. and removed to 21 °C. for two hours once each week which allows sufficient ac- tivity for feeding and body movement. Specimens were also held in freezer cartons at 16“ C. continuously for four months, the temperature being low enough to decrease activity and still sufficiently high to allow for feeding and body movement. The low temperature decreased the incidence of mould formation and decreased the rate of fermentation of the honey solution -- the latter was renewed once a week. Specimens were held in cellophane envelopes ( 6 cm. x 12 cm. ) for periods in excess of one month. A small piece of cotton, 5 (3) a 29-136, 1966 MONARCH TECHNIQUES 131 soaked in honey solution, was placed in one comer of the en- velope and the butterfly, with wings folded together, was placed in the envelope with its head close to the cotton. The cotton was charged periodically by inserting the needle of a hypodermic syringe through the envelope and into the cotton. The envelope was sealed off by folding the free end and securing it with a paper clip. Specimens were held at 16“ C. Feeding: Monarch butterflies may be held throughout the winter on a diet of 10% honey-water solution. Among many dispensing methods, the following has proved practical for use with cylin- drical cage. Aluminum sheeting 1 mm. thick with 1 mm, holes spaced 1 mm. apart, is bent to fit the outside of the cage and held in place with hooks. The holes in the metal membrane serve as reservoirs for the honey solution. To feed the butterflies, the metal membrane is washed thoroughly and replaced on the cage with the water remaining in the small perforations. It is sometimes necessary to place the specimens in the vicinity of the m,etal membrane in order to instigate feeding. In doing so, care must be exercised in removing the butterflies from their hold on the cage wire as their tarsi may be broken, making it impossible for them to cling to the cage or to the milkweed plants. When the butterflies have had a drink of water, honey solu- tion is added to the membrane with a squeeze bottle. It may be necessary to "train” some of the butterflies to use the feeding membrane by first inducing feeding by unrolling the proboscis with the point of a pipette or squeeze bottle and giving the butterfly a small drop of honey solution. They can then be placed on the feeding membrane where they will continue to feed by themselves. Cheese cloth wicks, suspended inside the cages from one side of the cage to the other, and with the free ends secured by clip pins to the top of the cage, will hold the solution for a twenty- four hour period or more, depending upon the humidity of the environment. Such wicks can be charged with honey solution by means of a squeeze bottle, applying the solution to the free ends of the wick. Synthetic sponges, cut into thin strips, can be inserted into longitudinal slits (placed near the top of the cage) made in the cloth screen.- These can be recharged without opening the cage. Continuous feeders, designed after the principle of the intra- venous feeder, were found useful when the cages could not be 132 URQUHART RUD STEGNER J. Res. Lepid. attended to for a few days. This dispenser consists of a gallon jar fitted with a two-holed rubber stopper into which is in- serted a glass tube of sufficient length to reach the air space above the liquid when the jar is inverted, and a second glass tube, 10cm. longer than the stopper, protruding a sufficient length to allow for the attachment of a plastic tube approxi- mately two feet long. The jar containing the honey solution, is suspended above the cage in an inverted position. The rate of flow of the solution is controlled by a Hoffman Clamp (screw compressor type). The free end of the wick, suspended from the roof of the cage, passes into a gallon jar. Plastic vials have been used successfully by G. Grisdale of the Insect Pathology Research Institute, Sault Ste. Marie, On- tario. Shell vials (13 x 35mm) filled with honey solution, are taped together in bundles of six or more. These vial clusters are then suspended from the upper part of the cage. Humidity: Humidity is an important physical factor in the successful maintenance of adult butterflies over long periods of time. At the University of Toronto laboratory an R.H. in excess of 90% is maintained in the growth chambers. Adults, collected or reared in late August, have been kept alive until the following April at which time they were successfully mated with resulting viable eggs. Reproductive colonies are maintained the year round at the University of Delaware in a small laboratory with no special arrangements except the operation of a small humidifier and the maintenance of a temperature between 21° and 27° G. Adults have been kept alive for periods of four months under these conditions. Oviposit ion: Gravid females will readily oviposit on milkweed leaves either as growing plants or as individual leaves suspended in the cage. The plants are placed at the side of the cage that faces the most intense illumination. The butterflies congregate at the side of the cage where the light is most intense thus coming into more frequent contact with the milkweed plants. Scent receptors are located on the mesothoracic legs, (Urquhart, 1960) and con- tact with the milkweed leaves- triggers off the process of ovi- position. So long as the tarsi rest upon the Ifeaf, the egg is de- posited on any adjacent surface. Thus, eggs may be deposited on a piece of paper placed at the tip of the abdomen. This pro- cedure is especially useful when the eggs are to be shipped 5 (3) ; 129-136, 1966 MONARCH TECHNIQUES 133 by mail. The eggs develop normally on paper. Females will oviposit on thawed milkweed as well as on a paste of dehydrated milkweed. Females will sometimes oviposit on a cloth soaked in fermented honey solution. It is not unusual to find hundreds of eggs on the feeder wicks as a result of this peculiar reaction. Females will lay eggs even when held by the wings. Several eggs may be deposited in quick succession followed by a pause. If the specimen is allowed a brief period of flight, or to move its wings while being held, oviposition will take place more rapidly than when the specimen is not permitted such activity. Although when caged, the females may lay dozens of eggs on one plant, in their natural surroundings the eggs are widely scattered — rarely are two eggs found on a milkweed leaf. The leaves, bearing the eggs, may be placed in a petri dish or small jar until hatching has taken place. Hatching may be delayed by holding the eggs at a low temperature or accelerated by high temperature. Potted milkweed plants are desirable for oviposition since they can be left in the cage without deterioration and the larvae will have fresh food available when they hatch. Milkweed can be propagated from seeds or from root stalks, the latter being the quicker method. Ascelpias currasavica is convenient be- cause it sprouts new shoots readily after the tops have been cut off, and it is easily propagated from stem cuttings. Since larvae often devour the egg shell upon eclosion they may also devour adjacent eggs. To prevent this, the eggs are separated by cutting the leaves into small fragments, each frag- ment bearing an egg. Rearing Containers: Petri dishes are most suitable for early instars, and glass or plastic jars or tubes for more mature forms. There are many different kinds of containers that are satisfactory for holding large larvae but tight covers are necessary for the smaller ones. When large containers are used, the resulting pupae can be left until the adult butterfly matures. Or, a piece of paper or cardboard can be fastened to the top of the jar and in this way the pupae can be transferred to another jar or cage by simply removing the paper. When rearing large numbers for migration studies, eggs are placed in wide-mouthed gallon jars with screw tops. When the larvae reach the third instar, they are placed in flight cages. As many as three hundred larvae can be reared in one cage 134 URQUHART AND STEGNER J. Res. Lepid. (Urquhart, 1960). Pupae are formed on the wood frame where they remain until the emergence of the adults. Flight cages have also been used successfully for rearing large numbers by placing the open end of the cage over a stand of milkweed plants upon the leaves of which a number of larvae have been placed. When the plants have been consumed, the cage is placed over a fresh stand and the larvae transferred. Food: Although fresh milkweed leaves, grown in a green house during the winter months, are most effective, frozen leaves, at- tached to the stalks, which have been thawed, are used suc- cessfully. The leaves should be cut before the seed pods are formed — small, young plants being preferred. The thawed leaves may be supported on a wire-mesh platform or hung on the side of the rearing cage. Thawed leaves must be removed daily since deterioraiton is rapid. A stand of milkweed can be har\^ested several times each summer if only the tops of the plants are cut off each time. The old stalk will send out new shoots. If the mature milkweed plants in a field have no insects on them, such as milkweed bugs or beetles, there is a possibility that the field has been sprayed with an insecticide. At least, this possibility should be considered before starting a research project dependent on this source. To retain the highest nutritive quality in field-cut milkweed, freshly cut stalks, placed in plastic bags, are stored in a field cooler containing dry ice. The bags of milkweed are transferred to a deep freezer refrigerator as soon as possible. Larvae can be fed on a paste of dried milkweed powder re- constituted with distilled water. Apparently this food is not as stimulating to the appetite or as nutritious as fresh milkweed. The resulting adults, the larvae of which have been so fed, are small, but otherwise normal. Dale Grisdale (loc. cite) has used an artificial diet, reported by McMorran (1965), with considerable success. Grisdale re- ports that newly hatched larvae of the Monarch butterfly ac- cepted the food readily and almost one hundred per cent of the larvae established on the food. In order to start the larvae, the artificial food is mixed with powdered milkweed leaves. Without the addition of the powdered leaves, no feeding was evident and all the larvae died during the first instar. He also reports that the adult females were sterile. Nanavati, working at the University of Delaware, has reared Monarch larvae to adulthood on this medium both with and without the milkweed fraction. 5 (3) : 129-1 36, 1966 MONARCH TECHNIQUES 135 Lyon and Flake ( 1966 ) used a similar diet in connection with their studies of the Douglas-fir tussock moth larvae. E. Alan Cameron of the Division of Entomology, University of California, Berkeley, has used the following diet for rearing the tussock moth. This diet is based on that of Lyon and Flake. TABLE 1 Composition of Ingredients Ingredients Quantity A. Caesin-vitamin free 87.5 g Alphacel ..........12.5 g Salt mixture W .........25.0 g Sucrose 87.5 g Wheat embryo .75.0 g Choline Chloride 2.5 g Ascorbic acid ...........10.0 g Methyl parahydroxybenzoate 3.75 g Aureomycin 0.75 g Potassium sorbate 5.0 g B. Water, distilled 550.0 ml. 4 M Potassium hydroxide 12.5 ml. Vitamin solution 25.0 ml. Formaldehyde (36%) 1.25 mo C. Nutrient agar 62.5 g Water ....................1550 ml. Mix solution “A” and “B"’ in blender. Add “C” after melting. Pour into wax-paper lined shallow pans or teflon-coated pans. TABLE 2 Vitamin Solution Ingredients Quantity Water ....................................100 ml. Nicotinic acid (niacin) 100 mg. Calcium pentothenate 100 mg. Riboflavin 50 mg. Thiamine hydrochloride 25 mg. Pyridoxine hydrochloride 25 mg. Folic acid 25 mg. Biotin 2 mg. Vitamin B-12 0.2 mg. Disease and Diapause: Two factors may cause difficulty in rearing the Monarch butterfly successfully throughout the winter months. One is the presence of a virus disease and the other diapause females. During the past two years ,and commencing three years ago, the natural populations of the Monarch butterfly in North America have been greatly reduced as a result of a polyhedxosis 136 URQUHART AND STEGNER J. Res. Lepid. vims epizootic (Urquhart, 1966). Hence, a high mortality has been experienced in our laboratory cultures regardless of feed- ing methods used. Even careful sterilization methods have proved ineffective in preventing infection. It is possible to rear continuous generations of the Monarch butterfly during the winter, starting with eggs of gravid females collected in the field during the summer and fall. Monarch fe- males collected in late September at Cape May, New Jersey, readily laid eggs on milkweed when brought into the Delaware laboratory. No special treatment was necessary. However, at the University of Toronto, it was found that although females col- lected in the spring and early summer produced continuous generations throughout the winter months migrant females col- lected in late summer and early fall from over-night roosting sites, did not oviposit even when kept for periods of five months. Similarly, over- wintering specimens from Monterey, California collected from over- wintering sites during December and Janu- ary would not oviposit. Dissected females collected during Janu- ary and February from such over- wintering sites rarely contained eggs (Urquhart, 1960). In March, however, many females were gravid prior to their spring migration flight. Francis Munger, of Whittier, California, who has been work- ing with us at the University of Toronto for the past number of years, reared successive generation of Monarch butterflies throughout the winter months. Larval Monarch butterflies have been sent to us by Miss Martha Fussier of Orlando, Florida, in January. The possible factors involved in the production of a repro- ductive diapause in the migrating and over-wintering popula- tions are being studied at the laboratories of the University of Toronto. Note: Monarch eggs, with which to start a breeding colony, are usually available from Dr. Stegner, Reprints of this article may be obtained by writing to Dr. Stegner. LITERATURE CITED LYON, R. L. and FLAKE, H. W. 1966. Rearing Douglas-fir tussock moth larvae on synthetic media. J. Econ. Ent. 59 (3) : 696-698. McMORRAN, A. 1965. A synthetic diet for the spruce budworm, Chris- toneura fumiferana (Clem.) ( Lepidoptera : Tortricidae). Can. Ent. 97 (1) : 58-62. URQUHART, F. A. 1960. The Monarch butterfly. University of Toronto Press: 361 pp. 1966. Virus-caused epizootic as a factor in population fluctu- ations of the Monarch butterfly. J. Invert. Path. 8: 492-495. Journal of Research on the Lepidoptera 5(3) : 137-152, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 VITAL STAINING OF COLIAS PHILODICE AND C. EURYTHEME JOHN M. KOLYER 55 Chimney Ridge Drive, Convent, New Jersey 07961, U.S.A. In experiments with the larvae of Pieris rapae (Linnaeus) it was found that ingestion of the dyes neutral red or Nile blue A imparted conspicuous external color (red or blue) to all three stages, while certain other dyes, notably brilliant cresyl blue, gave colored pupae but not adults ( Kolyer, 1965 ) . In the present work the three dyes mentioned were tried on the interrelated species Colias philodice (Latreille) and C. eurytheme (Boisduval) to observe the extent of manifestation of dye color in strongly pig- mented butterflies ( as opposed to the white Pieris rapae ) , to gain an indication of the generality of the vital staining method, and to optimize the procedure for dye-feeding in the case of the Colias species. REARING Oviposition. — In all cases the eggs were obtained by confining the female(s) in a screen-covered jar (approximately a cylinder of 3.5 inches inside diameter and 5 inches depth ) with a layer of water in the bottom covered by a perforated wire screen through which sprigs of red clover {Trifolium pratense) were inserted. Oviposition was stimulated by direct sunlight or by electric light (suitably from a 100-watt bulb ten inches from the jar.) Oviposition began on zero day, and hatching started on the third day for broods 2 and 4 and the fourth day for the other broods (Tables 1 and 2). Temperature and Humidity. — The broods were reared during July - September, 1962 for brood 1 and July - October, 1966 for broods 2-5. For broods 2-5 the breeding room reached a high of 94 °F during the day and a low of 66 °F at night, while the rela- tive humidity was in the range 33-70% (usually 40-60%). 137 138 JOHN M. KOLYER /. Res. Lepid. n^ime (da^s) from katcking 5 (3) : 137-^1 52, 1966 VITAL STAINING 139 Methods for Supplying Leaves. ~ Both broods (1 and 2) of C olios eurytheme were reared on cut leaves of red clover in card- board boxes, e.g. shoe boxes as mentioned below, with gauze windows in the lids. The difficulty with this method was that rapid desiccation necessitated addition of fresh leaves up to five times a day. Cut leaves of white clover {Trifolium repens) were found to lose 50% of their original weight after 20 hours at 73- 83 °F and 34-45% relative humidity and 73% after 50 hours. To escape this difficulty, the following procedure was used for the three broods of C olios philodice. A cardboard shoe box, typically 10 inches long by 4 inches wide by 3.5 inches deep, with per- forated bottom and a lid with gauze window, was mounted atop a wide-mouth jar (3.5 inches inside diameter) nearly filled with water, into which white clover stems were inserted. Since the young larvae tended to drop from the leaves when disturbed, the clover leaves at first were situated at some distance from one large perforation by bending over a bundle of stems and taping them to the floor of the box. Later, when the larvae were in the fourth instar, the whole area of the box above the jar was per- forated with small holes, and individual stems were inserted to give a “carpet” of clover leaves. Figure 1 shows that the philodice larvae fed by the stems-in- water method developed more rapidly and attained larger size than the eurytheme larvae fed cut leaves. Klots (1951) lists a greater maximum expanse for eurytheme (2.4 inches) than for philodice ( 1.9 inches ) , and the forewing length for females col- lected in the general vicinity of Morristown, New Jersey, was greater for eurytheme (29 mm, mean of seven specimens) than for philodice (26 mm, also mean of seven specimens). Also, the larva of eurytheme has been described as attaining a slightly greater length than that of philodice (Comstock and Comstock, Fig. 1. Growth curves for larvae of broods 1-5. The curves are identified by brood no.; see Tables 1 and 2. The open circles mark the beginning of dye-feeding. In the case of broods 3 and 4 some readings were taken beyond this point, but these are not equivalent to the preceding values and are plotted only to show the greater length attained and more rapid development of these broods (vs. 1 and 2) even with the retarding effect of dyes. The solid circles mark the beginning of pupation. Maximum larval length was noted daily and plotted vs. time, and smooth curves were drawn through the points. In actuality there are short nO“growth steps, especially preceding the fourth molt. Approximate positions of the molts are indicated on the graph for brood 4 ( and the fourth molt of brood 5 ) . As a weight reference, two 25 mm larvae from brood 1 weighed 0.180 and 0,213 grams. 140 JOHN M. KOLYER J. Res. Lepid. ifcfel# l MMDCTIOIB FDR B«l-raiDlW %• Soll»§ (% of Ti*«, %• PmfKtl&Uf Ti*li Bmo4 lb SsMll Lwigtb«aiL d«r _ &im# 1 •MlTthia BOM - 23-11 (34 fro« gn«a iS ■tar larw#) Z swnrthmm Nil* U.u* A C5) 22-M 20-21 1.5-1.8 2S-W lO/lO •a# blu# (18 ■#), rmtst Bl* M«#- gr««n at momt asutral r*i C6) It- 24 i 2^26 f/9 piak (•«p, ea aMeiwB) t# d##p r#d Nil# bla# 22 A (1) ani •iw*. r«i (1) 9-5 24-26 S/S gr##a t» gr##a with M«# ab- d®M#a biin* er*- 18-20 •yl M.»* (5) 4-5 25-26 i/1 gmm a Bbllsdie# Nil# blu# A (5) 1&-2S 14 0,i-l*4 M-li 4/20 Nil# bla# A Cl) 24-2f 16 2.i-4,5 20-28 Z/% gr*#jd.afe-M«#i Ua# (1) »#atml rmA (5) 15-19 14^ 4-5 IS-lf 7/10 4ark-r#i (2) Bsutml r#d (5) 2i-30 18 0*5 20-a 3/3 tw» iart-r#4| oa# gr##a with rmi aMomm m# Ma# 17-1« A (1) 'tad B«ut. r#d (1) 15 0*5 If A/i ■1. bla# tlat brtU« cr#- 18-Ii •fl M.«# (5) I5I 9-4 M-lf 4/4 ■1. graylafa grmm 4 noutxml r#4 Ci| 2B-32 16-If^ Con# at 24) 0»3 it-a 3^32 r#i to tart»r#d i itdloil— Ml# Uu# A Cl) 2f-30 a.s 18-30 4/10 MM#-gr#«B 5(3) : 137-^1 52, 1966 VITAL STAINING 141 1943). Therefore, it seems reasonable that philodice is not inher- ently larger than eurytheme but, if anything, smaller, and that the improved rearing procedure was responsible for the more rapid growth rates and greater ultimate lengths of the philodice larvae as well as the greater expanse of the adults ( See forewing mea- surements in Table 2 ) . Mortality seemed reduced by the improved feeding method; for example, in brood 4, 39 eggs yielded 34 larvae which yielded 32 adults, while in brood 1 ( cut leaves ) 65 first-instar larvae ( from approximately 150 eggs) yielded only 34 pupae, and in brood 2 ( also cut leaves ) only 34 larvae remained of the original 54 ( from about 111 eggs) when the largest had reached 19 mm. No crowd- ing problem was evident in brood 4 when the newly-hatched larvae ( 34 ) were reared to . give 33 fifth-instar larvae ( 29 mm maximum length ) in a single box of dimensions specified above. Dye-Feeding Technique. — Dyes were fed by coating the leaves on both sides with dye-mica blend prepared by grinding and tumbling the dye with P-12 Davenite mica ( 325 mesh; Hay- den Mica Co., Wilmington, Mass.) as described in the earlier paper ( Kolyer, 1965 ) . However, the dye/leaf ratio in the present work was higher than with Pieris rapae because more dye blend was used (about 5 mg for a white clover leaf 1.5 cm wide) and because clover leaves are far less thick than the average cabbage leaf. The dyes (biological stains), all from National Aniline Divi- sion, Allied Chemical Corp., New York City, were brilliant cresyl blue (Colour Index No. 51010), neutral red (chloride, 70% mini- mum strength, 1.0% maximum water-insoluble content. Colour Index No, 50040), and Nile blue A (sulfate, 70% minimum strength. Colour Index No. 51180). Explanation of Table 1. — The days listed for inception of dye- feeding, pupation, and eclosion in the tables are all based on the start of oviposition, which continued no more than three days, at zero day. In the Yield Pupae column, 7/9, for example, means that 7 pupae were obtained from 9 larvae fed dye. The pupal color was noted when pupae were about two days old. Description of Female Parents. — Brood 1. — Eggs (approximately 150) were laid by one eury- theme, moderately marked with orange, forewing 23 mm, taken near Fall River, Mass., on July 22, 1962. Brood 2. — Eggs (approximately 111) were laid by some or all of five eurytheme, most by one moderately, marked with orange. 142 JOHN M. KOLYER /. Res. Lepid. Tabic Z Brood Dya (Ji Mo. of bland) QA7A ADULTS Color Inter- Fo rawing aagaantal Seloalon, Color JPu- Mo. of Adult a L«ngth,nB, Ground Sf* Abdonlnal dar oal Casa nala faqtale ao«ut (ransa) Color Color Maab|i^ 1 non© 29-38 pala yallow 15 14 uni a- 21(20-23} fanda- 22(20-24) omaga It. groan It . groan Z Milo blue A (5) 28-33 palo blua 6 4 iaala-19( 15-22) fcsala- 23(21- 24) oranga^ daap groan blue naut. rad (5) 28-32 pink 5^ 0 aala- 18(15-21) oraagai goldan- brown daap oranga for unaxpandad spaeinana pink Mila blue A (1) and naut. rad (l) 30-33 pala blua 1 4 ■ala-18 faml a- 20(18-23) oranga It. groan blua brill, cra- ■yl blua (5) 31-32 pala yallow 1^ 1 ■ala- 20 fsenda-BO oranga It. groan It. groan nona ( control ) 32-33 pala yallow 3 2 ■alo- 22(21-24) fawao-24(23,24) oruga It. groan It. groan 3 Nila blua A (5) 23-25 pala graaniah- yallow 2 2 mala-23( 22,23) faBMaa-27(26,28) faint groan groan tint (1 Mda) blue Mila blua A (1) 26-29 pala graanlah- yallow 1 1 ■ala- 21 f«wla-21 faint groan groan tint in nala blue (1) naut. rad (5) 23 pink 1* 0 - oranga rad daop-red (2) naut. rad (5) 25-27 flaah-eolor 1 2 and 0-23 famla- 26(25,26} al. daapar goldan- yallow to yallow to orango-taa daap-rad rad Nila blua A (1) and n«4t. rad (l) 24-25 palo yallow 0 4 fcMl*- 27(26-29) yellow It. groan to groan blua brill, oro- ayl blua (5) 23-25 pala graanish- yallow 1 3 ■alo- 20 f scud a- 22(21-23) yallow It. groan gray non© (control) 24-25 pala yallow 0 3 feeKla-29(28-29) yallow It, groan gray 4 naut. rod (5) 25-29 flash- color 12 UA® aal 0-23(20-25) fanla- 24(23-26) Y-aranga-tan Y-goldan rad A-pink to daap-rad irdaap-rad 5 Mila blua A (1) 24-25 pala blua or graanish-blua 2 1 ■alo- 21(21, 21) fsada-21 yallow groan groan (Hdaa), palo blua (albino fotaala) blua ^©nc al. gr©©alah iasld© ^rdsr} ©•© Flat© 1. *two failed to axpaad. ^aaothar aeloaad but not azpandad. 'not axpandad. ^ meana yalloof A moana albino. 5 (3) .-137-152, 1966 VITAL STAINING 143 forewing 26 mm, taken at Berkshire Valley, New Jersey, on July 3, 1966. Brood 3. — Eggs (81) were laid by three philodice, wings yellow with no trace of orange, forewings 24, 26, 27 mm (mean 26 mm) taken at Morristown, New Jersey, on July 12 and 13, 1966. Brood 4. — Eggs (39) were laid by three albinos, presumably philodice (as judged by width of the black borders of the fore- wings) with no trace of yellow or orange in the ground color, forewings 24,25,26 mm (mean 25mm), taken at Berkshire Valley, New Jersey, on September 3, 1966. Brood 5. — Eggs (11) were laid by an albino female with a touch of yellow-orange in the wings, forewing 25 mm, and/or a yellow philodice, no trace of orange, forewing 27 mm, both taken at Morristown, New Jersey, on September 10, 1966. RESULTS Color of Adults. — The situation is complicated by the fact that philodice and eurytheme are peculiarly interrelated and hy- bridize to some extent; see, for example, Gerould (1946) and Hovanitz ( 1949 ) . However, the adults reared in broods 1 and 2 were typical eurytheme, well marked with orange, while the un- dyed adults reared in brood 3 were all typical philodice, like the parent females, with no trace of orange. In brood 5 the female was an albino, and the males appeared to be typical philodice. Only in brood 4 among the philodice broods was there a trace of eurytheme; a male and female which pupated without feeding on the dye (neutral red) showed a faint orange suffusion in the basal area of the forewing, and this could be observed in some of the dyed specimens (except the albinos, which were evenly pink like dyed Pieris rapae and presumably would have been quite uncolored if not fed neutral red ) . The introduction of color solely by neutral red as opposed to eurytheme genes seems un- equivocal in brood 3, in which all 17 adults except the four fed neutral red (e.g., no j6, Plate 2) were quite yellow (with very faint greenish tint in some of the six fed Nile blue A). Approxi- mately half the females in brood 4 were albinos, which form is known to be controlled by a dominant allele (for example, Ho- vanitz, 1944). The color of the antenna club was altered by dye-feeding. This was tan in the controls (either species) and in specimens fed brilliant cresyl blue but was deep-green when Nile blue A had been fed and red-brown when neutral red had been fed ( orange 144 JOHN M. KOLYER /. Res. Lepid. VITAL STAINING 145 5(3) ;137-152, 1966 146 JOHN M. KOLYER /. Res. Lepid. PLATE 1. SPECIMENS STAINED WITH NILE BLUE A AND NEU- TRAL RED BY FEEDING DYES TO THE LARVAE. PLATE 2. SPECIMENS STAINED WITH NEUTRAL RED BY FEED- ING THE DYE TO THE LARVAE. 1 — male Colias eurytheme, forewing 15 mm, fed Nile blue A, brood 2, from blue pupa (length 15 mm vs. normal length of 20 mm); 2— -albino female C. philodice fed neutral red, brood 4. 3 — albino female C. philo- dice fed Nile blue A, brood 5. 4 — male C. philodice fed Nile blue A, brood 5. 5 — female C. philodice control, brood 3. 6 — female C. philodice fed neutral red, brood 3. 7 — male C. philodice fed neutral red, brood 4. 8 — female C. philodice fed neutral red, brood 4. The pupal case is shown with each specimen. 5 (3) : 137-152, 1966 VITAL STAINING 147 in the case of albino females). The liquid voided following eclosion was blue-green for adults fed Nile blue A and strongly red for those fed neutral red. Verbal description of color is vague at best (and photographic reproduction is hardly accurate), but the direction of vital stain- ing should be evident from Table 2 as well as Plates 1 and 2. Explanation of Table 2. — Forewing length is the distance from the base to the apex of the wing. The eye color was noted on living adults with the aid of a 16-power stereO' microscope. The color of the abdominal membrane was obsc'rved by flexing the abdomen of a living specimen under the microscope to separate the segments. This was a sensitive indicator of the presence of Nile blue A or neutral red. Concentration of Neutral Red in Adults. — Paper Chromatography on Wings. — Single forewings were extracted by grinding the wing in a mortar with 1.0 ml concen- trated hydrochloric acid (37-38% HCl). The liquid and undis- solved debris were transferred with about three ml distilled water to a watchglass and evaporated to dryness on a hot plate. The soluble part of the residue was taken up in 0.15 ml distilled water and chromatographed by the ascending method (for a general description see Lederer and Lederer, 1953). Whatman No. 1 filter paper (3 inches by 4.5 inches) was made hydropho- bic, in the manner of Ciglar, Kolsek, and Perpar, 1962, by dipping in 10% lauryl alcohol in 95% ethanol and allowing to dry. Spots (about 3 mm diameter) were applied with a capillary 5/8 inch from the shorter edge of the paper; three applications were made, with drying between, to reinforce each spot. The paper was dip- ped, spotted edge down, to a depth of 1/4 inch in a layer of sol- vent ( 2 volumes 95% ethanol : 2 volumes concentrated ammonium hydroxide (28-30% ammonia) : 1 volume distilled water) in a closed jar and left for one hour at room temperature (27°C). After drying in a draft of air the chromatogram was sprayed with 1.5% aqueous HCl to bring out the pink spot for neutral red. For a female philodice fed neutral red (brood 4, resembling no. 8, Plate 2) the tan spot, Rf 0.75, due to wing pigments, was well separated from the pink spot, Rf about 0.24, for neutral red; wings of undyed philodice or eurytheme gave only the tan spot, as expected. Several quantities of neutral red ( from same sample used to prepare the blend with mica) were evaporated with concentrated hydrochloric acid and chromatographed in the same way to allow a semiquantitative estimation of neutral red level in the wing by comparison of spot intensities. The result was very 148 JOHN M. KOLYER J. Re$. Lepid. approximately 0.01 mg neutral red, or, since the wing weighed 2.0 mg., 0.5% neutral red in the wing. The dye seems confined to the scales, incidentally; the membrane is colorless. Colorimetry, — A Bausch and Lomb Spectronic 20 Colorimeter (band pass 20 millimicrons) was used to take % transmittance readings (readily converted to optical density, which equals the negative logarithm, to the base 10, of the transmittance ) at every 10 millimicrons (in regions of absorption maxima) in the range 350-650 millimicrons ( 3500-6500 Angstoms ) . The % transmittance was readjusted to 100 with solvent at each wavelength. The sen- sitivity setting of the instrument was varied from nearly a maxi- mum level for, the wing extracts to lower levels for the other de- terminations. The spectra within each of the three sets were run at the same sensitivity and included standard neutral red solutions for calibration. Body. — The body of a dried specimen (wings removed) was ground in a mortar with 1.20 ml concentrated hydrochloric acid (37-38% HCl), and the mixture was partly neutralized (to pH about 0.3) by adding 3.75 ml ammonia solution prepared by diluting one volume concentrated ammonium hydroxide (28-38% ammonia) to five volumes with distilled water. Debris was re- moved by filtration. The spectra of dyed and undyed albino bodies (nos. 2 and 3) are shown in Figure 2 as well as the spec- trum of neutral red in the same solvent. The absorption maximum for neutral red was 530 millimicrons, as has been reported for acid solutions (Meyer and Treadwell, 1952), and a calibration curve (optical density vs. concentration) was obtained for this wavelength using several neutral red concentrations. By summing the optical densities of spectra 1 and 2, the predicted spectrum (neutral red superimposed on the undyed albino body extract) was obtained (no. 7 ) . This was a fair match for the actual spec- trum of the dyed albino ( no. 3 ) except for considerable deviation above 550 millimicrons. However, addition of the amount of neutral red (as a concentrated solution) predicted for the dyed albino body by means of the calibration curve to the undyed albino body solution gave spectrum 8, which is in rather good agreement with the spectrum of the dyed albino body over the whole wavelength range. Since the level of neutral red in the extract of the dyed albino body was 0,066 mg (after correcting for the difference between spectra 8 and 3) and the body weighed 26.6 mg, the neutral red extracted amounted to 0,25% of the body. This is about double the level for the living body, since this species should lose about half its original weight on 5 (3) : 137-1 52, 1966 VITAL STAINING 149 drying (Kolyer, 1963). Whole Adult. — The whole (dried) butterfly was ground in a mortar with about 5 ml acetone, transferred to a flask with about 25 ml acetone, and refluxed for 20 minutes with the in- tention of degreasing the body. Then the acetone was evaporated and replaced by about 10 ml distilled water. After refluxing for 30 minutes (neutral red is not affected by this treatment), the mixture was evaporated to about 4 ml, 0.035 ml concentrated hydrochloric acid was added, and the mixture was diluted to 7.0 ml with distilled water and filtered to remove debris. In Figure 2, spectra 4 and 5, respectively for undyed and dyed albinos, are generally similar to spectra 2 and 3 for the bodies. Addition of neutral red to the undyed albino extract gave spectrum 6, and the calibration curve indicated 0.022 mg in the extract or 0.06% neutral red in the butterfly. A dyed (orange) female (brood 4, resembling no. 8, Plate 2) and an undyed yellow female gave rather similar spectra to those for the albinos, and the neutral red level was the same (0.06% of the dyed butterfly, which weighed 37.0 mg). The low results suggest that this extraction method was less effectn^e than trituration with concentrated hydrochloric acid, which dissolves much of the specimen. Wing. — Single forewings were extracted in the same manner as the bodies (above). As in the case of the bodies, the calculated spectrum for the dyed albino (no. 10 in Figure 2) deviated from the observed spectrum (no. 11) above 550 millimicrons. By sub- tracting the optical density of spectrum 9 (undyed albino) at 530 millimicrons from than of spectrum 11 and converting to neutral red concentration by means of a calibration curve, a value of 0.0036 mg was found for the forewing of the dyed albino; this is 0.18% of the wing, which weighed 2.0 mg. Similarly, 0.0038 mg neutral red (0.19% of the wing) was found for the forewing of a dyed (orange) female from brood 4 resembling no. 8 in Plate 2. Incidentally, the spectrum of the extract of an undyed, yellow philodice wing showed 24% transmittance at 400 millimicrons and rose to a plateau of about 79% by 570 milli- microns, while an undyed eurytheme wing was less transparent in the lower ( blue ) region ( 12% transmittance at 400 millimicrons ) and more transparent ( e.g. 90% transmittance at 590 millimicrons ) in the yellow and red regions. This is consistent with the pres- ence of orange pigment. Concentration of Nile Blue A in Adults. — No assay was made on the three philodice specimens conspicuously colored by Nile blue A (brood 5). However, the intensity of this dye is com- 150 JOHN* M. KOLYER /. Res. Lepid. parable to that of neutral red; for solutions of equal weight con- centration neutral red gave optical density 0.27 at 530 milli- microns vs. 0.41 for Nile blue A at its maximum of about 640 millimicrons (Merrill and Spencer, 1948, report 634 millimicrons). Therefore, visual estimation suggests a concentration of on the order of 0.1% Nile blue A in the wings of the dyed albino from brood 5 ( no. 3 in Plate 1 ) . DISCUSSION Effect of Dyes on the C olios species vs. Pieris rapae. — The toxicity of Nile blue A was more pronounced with the C olios species than Pieris ropae when the level ingested was high enough to alter the wing color conspicuously. In brood 5, three well-dyed specimens were obtained ( two shown in Plate 2 ) only at the expense of 70% mortality. As with Pieris ropoe, Nile Blue A preferentially showed itself in Colios adults (abdominal mem- brane) when fed with neutral red at equal levels (Table 2). Neutral red gave strongly pink specimens with 30% mortality with Pieris ropoe (Kolyer, 1965), and good coloration with no mortality was achieved with Colios philodice (brood 4) when feeding was begun at the proper larval length. Brilliant cresyl blue gave strongly violet pupae when fed to Pieris ropoe larvae for 3-9 days, but when fed ( at a higher dye/ food ratio) to Colios philodice for 3-4 days (brood 3) there was only a subtle change in pupal color (Table 1). A feeding period of 4-5 days had no noticeable effect on the color of C. eury theme pupae ( brood 2 ) . It is interesting that this dye gave very different results for Pieris vs. Colios. Toxicity ond Growth-Retording Effect of Dyes. — Brilliant cresyl blue may have shown a slight retarding effect on the Colios species, while Nile blue A was toxic to the point of being very “tricky” to utilize and, besides killing larvae, produced some undersized specimens, the smallest being the dwarfed male shown in Plate 1. Neutral red retarded growth and had a toxic effect in C. eury theme when feeding was begun too early; a dwarfed male with 15 mm forewing was produced in brood 2, and only three properly-expanded adults were obtained from nine larvae. In this case the ground color was noticeably changed only in the un- expanded specimens, presumably because ft was difficult for neutral red to manifest itself against the orange pigment. With C. philodice, larval growth was retarded (maximum length at- tained was 27 mm) and undersized, deformed pupae resulted 5 (3) : 137-1 52, 1966 VITAL STAINING 151 when neutral red feeding was begun too soon (group(l) in brood 3). However, when feeding was begun (brood 4) with larvae that had attained 83-94% of the ultimate length for controls (34 mm), there were no deaths, and only about five specimens out of 31 were not well tinged with pink, A single specimen (brood 4) fed neutral red from, a length of 24 mm (71% of ulti- mate length) eclosed but failed to expand. It should be noted that staining with neutral red is reversible with C olios as with Pieris tapae and the silkworm (Edwards, 1921); loss of dye begins when the lan^a is transferred to undyed leaves. Tliis effect is avoided, of course, by feeding the dye to pupation. Optimum Dye-Feeding Procedure. — In agreement with the cited work with Pieris rapae, neutral red is judged particularly suitable for vital staining of the C olios species. The best pro- cedure ( at the level used in the experiments ) is to confine dye- feeding to the latter part of the final instar. Alternatively, it is possible that the dye might be fed earlier if used at a lower con- centration; a brief experiment with 1% neutral red in the blend showed that the larvae were visibly stained (as indicated con- veniently by change in color of the lateral stripe) after being fed dye for 12 hours. CONCLUSION Neutral red has been found to be an effective vital stain, with an acceptable degree of toxicity, in Pieris hrmsicae (linnaeus) (Roer, 1959), Pierk rapae (Kolyer, 1965), the Colias species (present paper), and also the silkworm (Edwards, 1921). Vital staining provides indelible marking, which might be of use in the rearing of experimental broods, but an area of more interest might be the effect of the unusual color on the behavior, e.g. mating, of adult butterflies. The concentration of dye in the adult eye might alter response to light, for example; this has been suggested as an area for experimentation with moths fed rhodamine B (Vail, Howland, and Henneberry, 1966 ) , SUMMARY Nile blue A and neutral red can be used to stain living larvae, pupae, and adults of Colias philodice and eurytheme by feeding leaves coated with dye extended with powdered mica. The use- fulness of Nile blue A is limited by its toxicity, but neutral red is effective with little or no mortality when fed to larvae within 152 JOHN M. KOLYER J, Res, Lepid. two or three days of pupation. The wings of yellow C olios philo- dice adults were stained by neutral red to a dull orange color, while dyed albinos were strongly pink. Paper chromatography and colorimetry on extracts of dried specimens indicated a level of on the order of 0.2-0.5% neutral red in the wing and 0.3% in the dried body. Brilliant cresyl blue, which is effective in stain- ing the pupa of Pieris rapae, had little efficacy with the C olios species. LITERATURE CITED CIGLAR, V. J. & J. KOLSEK, 1962. The paper chromatography of basic dyes. Chemiker Zeitung, 86: 41-43. COMSTOCK, J. H. & A. B. COMSTOCK, 1943. How to Know the Butter-^ flies: a Manual of Those Which Occur in the Eastern United States. Comstock Publishing Co., Ithaca, New York. (pp. 93,95) EDWARDS, W. F., 1921. Feeding dyestuffs to silkworms. Textile World, 60: 1111-1113. GEROULD, J. H., 1946. Hybridization and female albinism in Colias philodice and C. eurytheme. A New Hampshire survey in 1943 with subsequent data. Ann. Ent. Soc. America, 39(3): 383-396. HOVANITZ, W., 1944. Genetic data on the two races of Colias chrysotheme in North America and on a white form occurring in each. Genetics, 29(1): 1-30. 1949. Interspecific matings between Colias eurytheme and Colias philodice in wild populations. Evolution (New York), 3(2): 170-173. KLOTS, A. B., 1951. A Field Guide to the Butterflies of North America, East of the Great Plains. Houghton Mifflin Co., Boston, Mass, (pp. 183,185) KOLYER, J. M., 1963. Note on rates of drying and relaxation of specimens. Jour. Lepid. Soc., 17(3): 178-180. 1965. The feeding of coloring matters to Pieris rapae larvae. Jour. Res. Lepid., 4(3): 159-172. LEDERER, E. & M. LEDERER, 1953. Chromatography, a Review of Principles and Applications. Elsevier Publishing Co., New York. (p. 85) MERRILL, R. C. & R. W. SPENCER, 1948. Spectral changes of some dyes in soluble silicate solutions. Jour. Am. Chem. Soc., 70: 3683-3689. MEYER, H. W. & W. D. TREADWELL, 1952. Effect of acidity on the absorption spectra of some vat dyes and relationship to the redox potential. Helvetica Chimica Acta, 35: 1460-1470. ROER, H., 1959. Ueber Flug- und Wander Gewohnheiten von Pieris brassicae L. Zeit. Angew. Entomol., 44: 272-309. VAIL, P. F., A. F. HOWLAND & T. J. HENNEBERRY, 1966. Fluorescent dyes for mating and recovery studies with cabbage looper moths. Jour. Econ. Entom., 59(5): 1093-1097. Journal of Research on the Lepidoptera 5(3) : 153-176, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE NORTH AMERICAN SPECIES OF THE GENUS ZEIRAPHERA TREITSCHKE (OLETHREUTIDAE) A. MUTUURA and T. N. FREEMAN Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa, Ontario The North American species of Zeiraphera feed almost en- tirely on conifers of the tribe Abieteae. The only exception is Z. claypoleana (Riley) which feeds on Aesculus and is morphol- ogically remote from the others. Many of the, species occur abun- dantly enough to be of economic importance and because of this the Forest Insect and Disease Survey, Canada Department of Forestry and Rural Development, has been conducting investi- gations on them for several years. One result has been the ac- cumulation of about 1400 specimens reared from various coni- fers in many areas of Canada. The taxonomic study of these forms the basis of this paper. The specimens were divided in accordance with their respec- tive host trees and geographical areas. A close examination of the maculation and genitalia showed that seven conifer-feeding species are present. Some of these are very closely related but all can be recognized by differences in maculation and in the genitalia. These differences are not always apparent unless ex- amined in series. We have, therefore, usually illustratd the male genitalia of several specimens of each species to show the varia- tion within the species and the differences between them. A comparison of the North American species with the conifer- feeders of Europe showed that differences exist between them. It has therefore been concluded that the European species Z. diniana (Guenee) and Z. ratzeburgiana (Ratzeburg) do not oc- cur in North America. Z. improbana (Walker) is used for Z. diniana of American authors. The North American species for- merly considered as ratzeburgiana is described below as cana- densis new species. 153 154 MUTUURA AND FREEMAN J. Res. Lepid. GENITALIC DEFINITION OF ZEIRAPHERA TREITSCHKE Zeiraphera Treitschke, 1829, Schmett. Europa 7: 231. Type species: Tortrix corticana Hiibner, 1811, Europa Schmett. PI. 3, fig. 13 [= (Pyralis) Zeiraphera isertana (Fabricius) 1793, Ent. Syst. 3: 281; designated by Curtis, 1838, British Ent. 15: No. 711.] Male genitalia. Clasper sickle-shaped; ventral margin spined; apical region tongue-shaped; sacculus not spined; margin be- tween sacculus and apical region not strongly concave. Uncus rudimentary; with specialized scale-like hairs. Socii well devel- oped,, long, drooping, broad. Gnathos weakly developed. Aedea- gus short, moderately stout; cornuti a cluster of 3-60 elongate spines. Female genitalia. Eighth, ninth, and tenth abdominal segments very long and slender. Ovipositor retractile. Anterior and pos- terior apophyses long. Ostial plate arising from near bases of anterior apophyses, elongate, emarginate basally. Ostium some- what cup-shaped. Ductus bursae constricted; entirely sclero- tized, with sclerotized ring only, or membraneous. Bursa with one of two scobinate signa. Key to Species 1. Female bursa with one signum (Fig. 34). Male clasper without sub- apical constriction (Fig. 32) 2 Bursa with two signa (Fig. 37). Clasper with subapical constriction (Fig. 26) 6 2. Signum very small (Fig. 35). Ductus bursae membranous. Forewing with an oblique, rectangular, whitish grey patch near middle of dor- sum (Figs. 23-24). Hosts: Picea sitchensis, P. engelmanni 8 vancouverana McD Signum of moderate size (Fig. 40). Ductus bursae weakly sclerotized .. 3 3. Uncus well developed (Fig. 29). Forewing with indistinct maculation and dorsal patch (Figs. 7-11). Hosts: Larix laricina, L. occiclentalis 4. improhana ( Wlk. ) Uncus weakly developed (Fig. 30) 4 4. Frons and vertex whitish ochreous. Forewing with a distinct, white, median band not constricted near middle and containing a longitudin- al, shining, pale ochreous line (Figs. 12-14). Hosts: Picea glauca, P. engelmanni 5. fortunana (Kft. ) Frons and vertex greyish brown 5 5. Whitish median band of forewing constricted near middle or separated into a costal and a dorsal patch (Figs. 15-18). Hosts: Picea glauca, P. sitchensis, P. engelmanni, P. mariana, Ahies lasiocarpa, A. balsa- mea, A. amabilis 6. destitutana (Wlk.) Forewing with a distinct, whitish, semi-circular dorsal patch ( Figs. 19- 22). Hosts: Pseudotsuga menziesii 7. hesperiana n. sp. 6. One signum broader than the other 7 Signa almost equal in size 8 7. Neck of bursa with a slender sclerotized ring (Fig. 38). Ostial plate broad. Forewing with a pale brown, triangular, dorsal patch mixed with white scales (Figs. 3-4). Wingspread 12-15 mm. Hosts: Picea glauca, P. sitchensis 3. canadensis n. sp. Neck of bursa with a broad sclerotized ring (Fig. 36). Ostial plate slen- 5 (3) : 153-176, 1966 ZEIRAPHERA 155 der. Forewing with greenish, blackish, and whitish areas. Male with an indistinct, green dorsal patch; female with a large, blackish dorsal patch (Fig. 2). Wingspread 14-16 mm. Host: Aesculus glabra ...... 1. claypoleana ( Riley ) 8. Neck of bursa with a slender sclerotized ring. Forewing with a golden dorsal patch edged with white scales (Figs. 5-6). Wingspread 12-16 mm. Host: Picea sitchensis 2. pacifica Free. 1. Zeiraphera claypoleana (Riley) Sericorsis instrutana: Claypole, 1881, Proc. Am. Assn. Adv. Sci. 30: Riley, 1881, Am. Nat. 25: 1009. Proteoteras claypoleana Riley, 1882, Amer. Midland Nat. 10: 913; Forbes, 1923, Cornell Univ. Agric. Exp. Sta. Mem. 68: 440. Steganoptycha claypoleana (Riley), Claypole, 1882, Psyche 3: 364; Riley, 1883, Papilio 3: 191; Packard, 1890, Fifth Kept. U.S. Ent. Comm. 654; Lintner, 1897, Kept. N. Y. State Ent. 12: 214. Epinotia claypoleana (Riley), Fernald 1917, in Dyar List N. Am. Lep. No. 5232. Zeiraphera claypoleana (Riley), Heinrich, 1923, Proc. U. S. Nat. Mus. Bull. 123: 170. External characters. Frons fawn colored; roughly scaled. Ver- tex with long, light fuscous scales mixed with whitish ones. An- tenna with light brown and fuscous scales; scape whitish out- wardly. Palpus whitish inwardly, brownish fuscous with some whitish scales outwardly. Thorax whitish brown with two lateral, blackish-fuscous spots. Base of tegula blackish. Abdomen pale brown. Legs pale fuscous; fore- and mid-tarsi fuscous with paler bands. Male forewing (Fig. 1) with basal patch whitish on costal half, blackish brown at middle and greenish on dorsum; median band whitish, extending obliquely outward from costa to middle of wing, then acutely angling inward to dorsum, costal portions obscure; apical third whitish, with a brown subapical spot and three brown costal spots; fringe whitish fuscous, with three sub- apical white dashes. Hind wing pale fuscous; fringe whitish fus- cous. Female forewing (Fig. 2) with a large blackish-brown dor- sal patch; post-median area and subapical spot greenish. Wing- spread 14-16 mm. Male genitalia (Fig. 25). Uncus pointed at apex. Socii well developed, broadly sickle-shaped. Tegumen obtusely angled. Clasper arcuate, long, slightly expanded at apex, somewhat con- cave ventrally near base, densely spined. Sacculus weak. Aedea- gus short, stout, broader at base; cornuti a cluster of about 20 long spines. Female genitalia (Fig. 36). Apical region of anterior apophy- ses expanded. Ostial plate slender, strongly emarginate. Ostium small, cup-shaped. Ductus bursae with a small sclerotized plate. Neck of bursa with a broad sclerotized ring. Signa two, unequal in size, conical. 5 (3) : 153-1 76, 1966 ZEIRAPHERA 157 Flight period. June. Type locality. Ohio. Distribution. Ohio, Missouri, Texas. Host plant. Aesculus glabra Willd. Remarks. This species was placed in this genus provisionally by Heinrich ( 1923 ) on the basis of similarities in the male geni- talia. However, he indicated that the wing shape, pattern, color and habit were more characteristic of species of Proteoteras Riley than of the coniferous feeding species that comprise the genus Zeiraphera. Our studies of the female genitalia show a re- lationship to the coniferous feeding species and support Hein- richs arrangement. 2. Zeiraphera pacifica Freeman Zeiraphera pacifica Free. 1966, Can. Ent. 98: 588. External characters. Frons with smooth, silver scales. Vertex light golden brown, roughly scaled. Antenna light grey with darker annulations; scape blackish brown inwardly, pale out- wardly. Palpus whitish ochreous inwardly, blackish fuscous api- cally and outwardly. Thorax dark golden brown. Abdomen whit- ish grey. Fore- and mid-legs purplish black, with greyish bands. Hind leg purplish grey, with whitish bands. Forewing (Figs. 5, 6) with basal patch dark brown, whitish basally, narrowly margined with black apically, extending from just beyond basal fourth of costa obliquely outward to middle of wing, then at right angles inwardly to inner third of dorsum, costal region with Fig. 1. Z. claypoleana ^ , Cincinnati, Ohio; Fig. 2, Z. claypoleana $ , Cincinnati, Ohio; Fig. 3. Z. canadensis n. sp. $ , Ottawa, Ontario; Fig. 4. Z. canadensis n. sp. $ , McGillivary, B.C., Picea engelmanni; Fig. 5. Z. pacifica $ , paratype, Sandspit, B.C., Picea sitchensis; Fig. 6. Z. pacifica $ , paratype, Sandspit, B.C., Picea sitchensis; Fig. 7. Z. improbana, paratype of pseudotsugana (Kft. )Kaslo, B.C.; Fig, 8. Z. improbana 9 , Pagwal River, Ontario, larch; Fig. 9. Z. improbana $ , Tabor, Ontario, Larix laricina; Fig. 10. Z. improbana $ , Wawa, Ontario, Larix laricina; Fig. 11. Z. im- probana $ , Anarchist Mtn., B.C., Larix occidentalis; Fig. 12. Z. fortunana $ Paratype, Ottawa, Ontario, Fig. 13. Z. fortunana 9 , Black Sturgeon Lake, Ontario, Picea glauca; Fig. 14. Z. fortunana 9 , Yukon Territory, Picea glauca; Fig. 15. Z. destitutana 9 , Cape Breton, Nova Scotia; Fig. 16. Z. destitutana 9 , Black Sturgeon Lake, Ontario, Picea glauca; Fig. 17. Z. destitutana 9 , Black Sturgeon Lake, Ontario, Picea glauca; Fig. 18. Z. destitutana $ , B.C., Picea glauca; Fig. 19. Z. hesperiana n. sp. $ , Pavilion, B.C., Pseudotsuga menziesii; Fig. 20. Z. hesperiana n. sp. $ , Kelowna, B.C., Pseudotsuga menziessi; Fig. 21. Z. hesperiana n. sp. 9 , William Head, B.C., Pseudotsuga menziesii; Fig. 22. Z. hesperiana n. sp. 9 , Yale, B.C., Pseudotsuga menziesii; Fig. 23. Z. vancouverana $ , para- type, Ucluelet, Vancouver Island; Fig. 24. Z. vancouverana 9 , Vancouver Is., B.C., Picea sitchensis. 158 MUTUURA AND FREEMAN /. Res. Lepid. Male Genitalia: Fig. 25. Z. claypolana, Cincinnati, Ohio, Fig. 26. Z. pacifica, Sandspit, B.C., Picea sitchensis; Fig. 26a. Z. pacifica, Sandspit, B.C., Picea sitchensis; Fig. 27. Z. ratzburgiana, Germany; Fig. 28. Z. canadensis, n. sp. Ottawa, Ont.; Fig. 28a. Z. canadensis, n. sp. White Pt. Beach, Queens Co. N. S.; Fig. 28b. Z. canadensis, n. sp. Ottawa, Ont.; Fig. 28c. Z. canadensis, n. sp., Jacquet River, N. B.; Fig. 28d. Z. canadensis, n. sp., Zeballos R., B.C., Picea sp; Fig. 28e. Z. canadensis, n. sp., Zeballos R., B.C., Picea sp; Fig. 28f. Z. canadensis, n. .sp.. Prince Edward Island. 5 (3) :153-176, 1966 ZEIRAPHERA 159 three blackish dashes; median band golden brown, the posterior half more distinct, narrowly edged with white and forming a dorsal patch; post median band dark brown; apical third irreg- ularly mottled with golden, white and dark brown areas, the white area often appearing as a gradually broadening streak from tornus half way to costa; apical half of costa with evenly spaced, black dashes; fringe dark grey, with some white tipped scales, and with broken, darker basal line: Hind wing dark fus- cous; fringe light grey, with a darker basal line. Male Genitalia (Fig. 26). Uncus reduced, conical. Socii well developed, broadly sickle shaped. Tegumen broadly shouldered. Clasper well developed, long; heel broadly arcuate; apex broadly rounded. Aedeagus short, stout; apex truncate; cornuti a cluster of about sixteen elongate spines. Female genitalia (Fig. 37). Apex of anterior and posterior apophyses somewhat broad. Ostial plate broad, not emarginate. Ostium small, somewhat cup shaped. Ductus bursae membranous, with slender sclerotized ring at neck of bursa. Two similar, coni- cal signa present. Flight period. Late June. Type locality. Sandspit, Queen Charlotte Islands, British Co- lumbia. Distribution. Coastal British Columbia and Washington. Host plant. Picae sitchensis ( Bong. ) Remarks. This species is closely allied to the European Z. ratze- burgiana and to Z. canadensis. The head of pacifica is golden brown; that of canadensis is greyish. The dorsal patches of cana- densis and ratzeburgiana are pale brown and triangular; those of pacifica are golden brown and rectangular. 3. Zeiraphera canadensis n. sp. Steganoptycha ratzeburgiana: Fernald, 1884, Kept. U. S. Dept. Agr. p. 378; Packard, 1890, Fifth Report U. S. Ent. Com., p. 845. Epinotia ratzeburgiana: Fernald, 1903, in Dyar, List N. Amer. Lep., No. 5233. Enarmonia ratzeburgiana: Barnes and McDunnough, 1917, Check List Lep. Bor. Amer., No. 7157. Zeiraphera ratzeburgiana: Heinrich, 1923, Bull. 123, 17. S. Nat. Mus., p. 171; Forbes, 1923, Cornell Univ. Agr. Exp. Sta. Mem. 68, p. 446; Prentice, et al., 1965, Dept. Forestry Can., Publ. 1142, 4: 627. External characters. Frons smoothly scaled, silvery. Vertex brown, roughly scaled.^ Antenna light brown, with black scales; scape with blackish scales inwardly, whitish fawn outwardly. Palpus whitish-ochreous inwardly, blackish fuscous anteriorly and outwardly. Thorax blackish brown, with a sprinkling of 160 MUTUURA AND FREEMAN 5 (3) : 153-1 76, 1966 ZEIRAPHERA 161 pale ochreous scales. Abdomen pale fuscous; terminal segment with pale brown scales. Fore- and mid-legs blackish fuscous, with pale brown bands. Hind tarsus pale fuscous, with pale brown bands. Forewing (Figs. 3, 4) with basal patch brown, mottled with brownish black, extending from just beyond basal fourth of costa vertically for a short distance ,then obliquely an- gled outwardly to middle of wing, where it angles acutely in- ward to dorsum beyond basal third; costal area of basal patch with three or four small, blackish costal dashes; median band pale brown, divided into two large patches, one mottled with white scales on costal region, semi-circular, and containing a small costal dash, the other almost triangular with a broken, black, dorsal margin sometimes edged with white scales; post median band brown, becoming whitish on costal area where it contains a small costal dash; apical third irregularly mottled with whitish-grey, fuscous, and brown areas; near outer margin a brown, short, transverse band edged outwardly with white; fringe dark brown, with blackish areas and a broken, darker basal line. Hind wing dark fuscous; fringe light grey, with a darker basal line. Wingspread 12-15 mm. Male genitalia (Figs. 28-28f). Uncus triangular. Socii broad. Tegumen broadly shouldered. Clasper well developed, long, con- stricted before apex; heel broadly extended and arcuate; sacculus slender. Aedeagus short, stout; cornuti a cluster of 9-18 spines. Female genitalia (Figs. 38-38b). Apices of anterior and pos- terior apophyses somewhat widened. Ostial plate broad, not emarginate. Ostium small, cup-shaped. Ductus bursae membran- ous, with sclerotized slender ring near neck of bursa. One large and one small conical signum present. Type material. Holotype male, Aylmer, Quebec, 13 June, 1960 I (G. G. Lewis), reared from Ficea glauca. Rearing no. 60-107. Paratypes: three males, three females, same locality and host as holotype 13-15 June, 1960. One male, Aylmer, Quebec, 2 July, 19.35 (G. S. Walley). One male, Hull, Quebec, 26 June, 1946. F.I.S., reared from white spruce. Two males, Kingsmere (near Hull), Quebec, 14, 15 June, 1941. F.I.S., reared from white spruce. One male, Stittsville, Ontario, 25 June, 1948. F.I.S., reared from Male Gentialia Fig. 29. Z. improhana, n. sp., Vermilion Bay, Ont., Larix larcina; Fig. 29a. Z. improhana, Laniel, Que., Larix laricina; Fig. 29b. Z. improhana, Hopedale, Labr; Fig. 29c. Z, improhana, Priceville, Ont., larch; Fig. 29d. Z. improhana, Burgessville, Ont., larch; Fig. 29e. Z. im- prohana, Ottawa, Ont.; Fig. 29f. Z, improhana, Aweme, Man.; Fig. 29g. Z. improhana, Beaver R., B.C., Larix laricina; Fig. 29h. Z. improhana. Nelson, B.C.; Fig. 29i. Z. improhana, Yorkton, N.J; 162 MUTUURA AND FREEMAN /. Res. Lepid. Male Genitalia: Fig. 30. Z. fortunana, paratype, Ottawa, Ont.; Fig. 30a. Z. fortunana, S. Milford, N.S.; Fig. 30b. Z. fortunana, Baddeck, N.S.; Fig. 30c. Z. fortunana, Bobcaygeon, Ont; Fig. 30d. Z. fortunana, Black Sturgeon Lake, nt., Ficea glauca; Fig. 30e. Z. fortunana. Black Sturgeon Lake, Ont., Picea glauca; Fig. 30f. Z, fortunana. Black Sturgeon Lake, Ont., Ficea glauca; Fig. 30g. Z. fortunana. Black Sturgeon Lake, Ont., Ficea glauca Fig. 30h. Z. fortunana. Black Sturgeon Lake, Ont., Picea glauca; Fig. 30i. Z. fortunana, B.C., Picea glauca; Fig. 30j. Z. fortunana, B.C., Ficea glauca; Fig. 30k. Z. fortunana. Dragon Lake, B.C., Picea glauca; Fig. 301. Z. for- tunana, Hospital Cr., B.C., Picea engelmanni. 5 (3) : 153-1 76, 1966 ZEIRAPHERA 163 white spruce. One male, three females, Hazeldean, Ontario, 2, 4 July, 1947. F.I.S., reared from white spruce. Three males, one female. South March, Ontario, 21-24 June, 1949. F.I.S. , reared from white spruce. Four males, three females, Ottawa, Ontario, 27 June — 2 July, 1905 (C. H. Young). Two females, Ottawa, Ontario, 10, 12 July, 1906 (C. H. Young). One male, Ottawa, Ontario, 29 June, 1935 (C. H. Young). One female, Ottawa, On- tario, 28 June, 1931 (C. H. Young). Four females, Ottawa, On- tario, 26-29 June, 1948. F.I.S., reared from white spruce. One female, Ottawa, Ontario, 15 June, 1941. F.I.S., reared from white spruce. One female, Bell’s Corners, Ontario, 11 June, 1941 (J. McDunnough), reared from spruce. All type specimens No. 9485 in the Canadian National Collection, Ottawa, Ontario. Distribution. Transcontinental in Canada and the northern United States. Host plants. Picae glauca (Moench) Voss and P. sitchensis ( Bong. ) Carr. Remarks. This is the North American species that various authors have considered to be the European Z. ratzeburgiana. Our studies have shown that there are distinctive differences between them, mainly as follows: Z. canadensis. Outer margin of basal patch distinctly concave or indented on costal half. Subapical white streak prominent, without white dashes through base of fringe. Female with one large and one small signum in bursa. Z. ratzeburgiana. Outer margin of basal patch linear on costal half. Subapical white streak faint with prominent white dashes through base of fringe. Female with two equal sized signa in bursa. 4. Zeiraphera improbana (Walker) Sciaphila improbana Wik., 1863, Cat. Lep. Het. 28: 337. Zeiraphera improbana (Wlk. ), McDunnough, 1959, Amer. Mus. Novit. No. 1954: 2. Sciaphila indivisana Wlk., 1863, Cat. Lep. Het. 28: 344. (New synonomy). Paedisca diffinana Wlk., 1863, Cat. Lep. Het. 28: 378. Zeiraphera diffinana (Wlk.), McDunnough, 1959, Amer. Mus. Novit. No. 1954: 2. Cydia pseudotsugana Kft., 1904, Can. Ent. 36: 110. (New synonomy). Thiodia pseudotsugana (Kft.), Dyar, 1904, Proc. U. S. Nat. Mus. 27: 927. Epinotia pseudotsugana (Kft.), 1905, Can. Ent. 37: 89, 253. Enarmonia pseudotsugana (Kft.), Barnes and McDunnough, 1917, Check List Lep. Bor. Amer. No. 7143. Zeiraphera psuedotsugana (Kft.), Heinrich, 1923, U. S. Nat. Mus. Bull. 123: 171. Zeiraphera pseudotsugana (Kft.), Klots, 1942, Bull. Amer. Mus. Nat. Hist. 79: 408. 164 MUTUURA AND FREEMAN J. Res. Lepid. 5 (3) : 153-^176, 1966 ZEIRAPHERA 165 Zeiraphera diniana: Heinrich, 1923, U. S. Nat. Mas. Bull. 123: 171, Fig. 287; Forbes, 1923, Cornell Univ. Agt. Exp. Sta. Mem. 68: 446; Pren- tice, et ah, 1965, Dept. Forestry Can., Ptihl. 1142, 4: 629. Zeiraphera sp. probably diniana: MacKay, 1959, Can. Ent. Suppl. 10: 97. External characters. Frons roughly scaled, the scales greyish black, with pale fawn apices. Vertex rough, greyish black. Pal- pus fuscous inwardly, blackish outwardly. Antenna with alter- nate black and grey bands. Thorax blackish brown, the scales with pale fawn apices. Abdomen dark fuscous; terminal seg- ment brownish. Tibiae and tarsi of fore- and mid-legs blackish fuscous, with pale fawn bands. Hind tibia and tarsus pale fus- cous, with pale fawn bands. Forewing (Figs. 7-11) with basal patch, median band, and post median band variously obscured with a black or blackish grey ground color; basal patch extend- ing obliquely outward from just beyond inner fourth of costa to middle of wing, then acutely angled inward to dorsum; median band greyish, divided into a small costal patch and a large dorsal patch, the costal patch semi-circular and containing two costal dashes, the brownish dorsal patch quadrate and edged with whit- ish-grey scales; post median band blackish with some brown scales; apical third blackish grey and with three costal gemina- tions; submarginal line broken, black, with some golden brown scales; tornal spot greyish, obscure, and containing an irregular line of black and golden brown scales; fringe dark grey, whitish near tornus, and with broken black basal line. Hind wing dark grey; fringe fuscous, with dark grey basal line. Wingspread 10- 18 mm.. Male genitalia (Figs. 29-29i). Uncus broadly conical; apex rounded, sometimes shallowly notched. Socii well developed, long, widest near middle. Aedeagus short, stout; apex truncate; cornuti a cluster of 30-60 elongate spines. Female genitalia (Figs. 40-40c). Anterior and posterior apo- physes longer than in other species with one signum. Ostial plate Male Genitalia Fig. 31. Z. destitutana. Black Sturgeon Lake, Ont., Picea glauca; Fig. 31a. Z. destitutana, Parrsboro, N.S.; Fig. 31b. Z. destitutana, Baddeck, N.S.; Fig. 31c. Z. destitutana, Edmimdston, N.B., Abies balsamea; Fig. 31d. Z. destitutana, Mt. Lyall, Que; Fig. 31e. Z. destitutana, Upsala, Ont., Picea glauca; Fig. 31f. Z. destitutana, Finland, Ont., Picea glauca; Fig. 31g. Z. destitutana. Black Sturgeon Lake, Ont. Picea glauca; Fig. 31h. Z, destitutana, S. E. Corner, Yukon T., Picea glauca; Fig. 31i. Z. destitutana, Kamloops, B.C., Picea engelmanni; Fig. 31j. Z. destitutana. Big Bend Hwy., B.C., Picea engelmanni; Fig. 31k. Z. destitutana. Big Bend Hwy., B.C., Picea engelmanni; Fig. 311. Z, destitutana, Scully Ck., B.C,, Picea sitchensis; Fig. 31m. Z. destitutana. Nation River, B.C., Abies laciocarpa. 166 J. Res. Lepid. Male Genitalia Fig. 32. Z. hesperiana n. sp., Pavilion, B.C., Pseudotsuga menziesii; Fig. 32a. Z. hesperiana n. sp., Canal Flats, B.C., Pseudotsuga menziesii; Fig. 33. Z, vancouverana, paratype, Ucluelet, Vnacouver Is., B.C.; Fig. 33a. Z. vancouverana, paratype, Ucluelet, Vancouver Is., B.C. Fig. 34-42. Female genitalia; Fig. 34. Z. hesperiana n. sp. Powell River, B.C., Pseudotsuga menziesii; Fig. 35. Z. vancouverana, Vancouver Ls., B.C. 5 (3) : 153-1 76, 1966 ZEIRAPHERA 167 narrow, emarginate. Ostium small, cup shaped. Ductus bursae slightly sclerotized, without sclerotized ring at neck of bursa. One signum present, variable in size and shape. Flight period. Late June to early August. Type locality. St. Martin’s Falls, Albany River, Northern On- tario. Distribution. Transcontinental in southern Canada and the northern United States. Host plants. Larix laricina (Du Roi) K. Koch and L. occiden- talis Nutt. Remarks. This is the species that North American authors have usually associated with the name Z. diniana Gn. as previously noted in the introduction to this paper. The names improbana Wlk., and diffiniana Wlk. have been associated with various species during their taxonomic history. McDunnough (1959) with the help of lepidopterists at the British Museum (Natural History), London, England, established that the two Walker names were synonymous and applied to a Zeiraphera species. Subsequently Obraztsov (in litt. ) studied Walker’s type of in- divisana and realized that it also was synonymous with impro- bana. All three were described from St. Martin’s Falls, are vari- ants of a single species, and are indistinguishable from the North American larch feeding species. Kearfott (1904) described pseudotsugana from flown speci- mens taken by Dyar at Kaslo, Kokanee Mt., and iKtchener Gla- cier, British Columbia. Dyar also reared specimens from Pseudot- suga which he thought were the same species. The paratype at the American Museum of Natural History, New York City, N.Y., is a headless female from Kaslo (Fig. 7) and is undoubtedly the dark colored Larix feeder. The only other type material in exist- ence is a paratype in the Canadian National Collection, Ottawa, Ont., from Kitchener Glacier. It also is the Larix feeder. Both of these paratypes agree with the original description and it is apparent that Kearfott did not have specimens reared from Pseudotsuga. It follows therefore, that the oldest name improbana applies to this species. It may be recognized by the dark, indis- tinct maculation and its host preference of Larix. 168 MUTUURA AND FREEMAN /• Lepid. Female genitalia: Fig. 36. Z. claypoleana, Cincinnati, Ohio; Fig. 37. Z. paci- fied, paratype, Sandspit, B.C., Ficea sitchensis Fig. 38. Z. canadensis, n. sp., Belhs Corner, Ont., spruce; Fig. 38a. Z. canadensis, n. sp., Radium Hot Springs, B.C., Ficea sp; Fig. 38b. Z. canadensis, n. sp., Hazelton, B.C., Ficea sitchensis; Fig. 39. Z. ratzhurgiana, Germany, Ficea excelsa. 5 (3) ; 153-176, 1966 ZEIRAPHERA 169 5. Zeiraphera fortynana (Kearfott) Epinotia fortunana Kearfott, 1907, Can. Ent. 39:126. Enarmonia fortunana (Kft. ), 1917, Barnes and McDunnough, Check List Lep. Bor. Amer., No. 7168. Zeiraphera fortunana (Kft.), Heinrich, 1923, U. S. Nat. Mus. Bull. 123: 172, fig. 288; Forbes, 1923, Cornell Unw. Agric. Exp. Sta. Memoir 68:446; Klots, 1942, Bull. Amer. Mus. Nat. Hist. 79: 402; MacKay, 1959, Can. Ent. Suppl. 10: 96, fig. 85; Prentice, et. al., 1965, Dept. Forestry Can., Publ 1142,4: 630. External characters. Frons smoothly scaled, whitish grey. Vertex with long, whitish-grey scales. Palpus cream white, black- ish outwardly. Antenna pale fawn. Thorax dark brown, edged with pale fawn. Abdomen dark fuscous. Tibiae and tarsi of fore- and mid-legs blackish fuscous, with whitish-grey bands. Hind tibia and tarsus pale fuscous, with whitish- grey bands. Forewihg (Figs. 12-14) with basal patch cream-white, mottled with black- ish, and extending from beyond inner fourth of costa obliquely outward to middle of wing, then obtusely angled inward to dorsum, which it reaches just beyond inner third; median band creamy-white, angulated at middle of wing, and containing a shining, pale ochreous line and a blackish costal dash; post- median band blackish brown, irregular in outline, broader at trailing edge; apical third of wing irregularly mottled with whitish-brown and blackish-brown areas, with three costal dashes, and a broken submargioal line; fringe fuscous, with broken darker basal line. Hind wing fuscous; fringe pale fuscous, with fuscous basal line. Wingspread 12-1.5 mm. Male genitalia (Figs. 30-301). Uncus reduced, with apex trun- cate or slightly notched. Socii well developed, broadly sickle shaped. Tegumen not produced at the shoulder. Clasper short; apex narrow. Aedeagus short, stout; apex truncate; cornuti a cluster of 16-24 elongate spines. Female genitalia (Figs. 41-41c). Anterior and posterior apo- physes moderate in size. Ostial plate broad, emarginate. Ostium somewhat cup shaped. Ductus bursae weakly sclerotized. One signum present, somewhat conical, rather large. Flight period. Early June to late July. Type locality. Ottawa, Ontario. Distribution. Nova Scotia to British Columbia. Host plants. Picea glauca (Moench) Voss, F. engelmanni Parry. 170 MUTUURA AND FREEMAN /. Res. Lepid. Female genitalia: Fig. 40. Z. improbana, Hopedale, Labr., Que,; Fig. 40a. Z. improbana, Ottawa, Ontario; Fig. 40b. Z. improbana. Anarchist Mtn., B.C., Larix occidentalis; Fig, 40c. Z. improbana, Anarchist Mtn., B.C., Larix occidentalis. 5 (3) : 153-176, 1966 ZEIRAPHERA 171 Remarks. This species apparently is well distributed throughout the spruce regions of Canada. It is closely allied to Z. destitutana but fortunana is smaller in wingspread and in size of genitalia. The frons and vertex of fortunana are whitish ochreous; those of destitutana are greyish brown. The uncus of fortunana is some- what notched; that of destitutana is rounded at the apex. The ostial plate of fortunana is broader and shorter than that of destitutana. 6. Zeiraphera destitutana (Walker) Sciaphila destitutana Wlk., 1863, Cat. hep. Het. 28: 339. Zeiraphera destitutana (Wlk. ), McDunnough, 1959, Amer. Mus. Novit. No. 1954: 3. External characters. Frons smooth, white below, with over- lapping, long, greyish-brown scales laterally and dorsally. Vertex greyish brown. Palpus whitish grey inwardly, dark fuscous out- wardly and apically. Antenna fuscous, with darker fuscous scales; scape blackish fuscous inwardly, much paler outwardly. Thorax brownish fuscous. Abdomen pale fuscous. Tibiae and tarsi of fore- and mid-legs blackish fuscous, with greyish bands. Hind leg whitish. Forewing (Figs. 15-18) with basal patch whitish grey, mottled with blackish, margined outwardly with black, costal margin with three black costal dashes; basal patch extending from beyond basal fourth of costa obliquely outward to middle of wing, then almost at right angles inwardly to dorsum, which it reaches just beyond basal half; median band whitish, con- stricted at middle of wing to form a costal spot and a dorsal patch; an irregular blackish post median band extending across the wing; apical third irregularly mottled with black, white, whitish-grey, and brownish scales; costa with three blackish geminations on apical third; subapical spot black, confluent with the two outer costal geminations; tornal region whitish; subapical line broken into black spots; fringe with blackish and brownish scales, whitish near tornus and with a black and white basal line. Hind wing blackish fuscous; fringe blackish fuscous, with black basal line. The female is more distinctly marked with white than the male. Wingspread 14-17 mm. Male genitalia (Figs. 31-31m). Uncus poorly developed. Tegu- men not strongly shouldered. Clasper broad, broadly rounded apically. Cornuti a cluster of about 24-40 elongate spines. Female genitalia (Figs. 42-42c). Similar to those of Z. improb- ana Wlk. Flight period. July and early August. Type locality. St. Martins Falls, Albany River, Ontario. 172 MUTUURA AND FREEMAN /• «««• Lepid. Female genitalia; Fig. 41. Z. fortunana, paratype, Ottawa, Ont.; Fig. 41a. Z. fortunana. Black Sturgeon Lake Ont., Picea glauca; Fig. 41b. Z. fortunana. Black Sturgeon Lake, Ont., Picea glauca; Fig. 41c. Z. fortunana, Yukon Territory, Picea glauca; Fig. 42. Z. destitutana, Kazubazua, Que; Fig. 42a. Z. destitutana. Black Sturgeon Lake, Ont., Picea glauca; Fig. 42b. Z. des- titutana, Nelson, B. C., Picea glauca; Fig. 42c. Z. destitutana, Hudson-Hope, B.C., Picea glauca. ZEIRAPHERA 174 MUTUURA AND FREEMAN J. Res. Lepid. Distribution. Nova Scotia to British Columbia. Host plants. Picea glauca (Moench) Voss, P. mariana (Mill.) B.S.P., P. engelmanni Parry, P. sitchensis (Bong.) Carr., Abies balsamea (L. ) Mill., A. lasiocarpa (Hook.) Nutt, and A. amabilis (Dougl.) Forb. Remarks. This species has been confused with Z. fortunana (Kft. ). However, it is larger, and the forewing has a mottled appearance and a larger dorsal patch. It also has considerable variation from light contrasting markings to dark suffused ones. The apex of the male uncus is entire, not notched as in fortunana, and the female ostial plate of distitutana is longer and narrower, resembling that of improbana. 7. Zeiraphera hesperiana n. sp. External characters. Frons roughly scaled, dark fuscous. Vertex greyish brown, roughly scaled, with a large tuft of the scales between antenna and upper side of compound eye. Palpus whitish ochreous inwardly, darker apically and outwardly. Antenna fawn with whitish scales; inner side of scape blackish, outer side whitish. Thorax dark brown, sprinkled with pale scales. Abdomen pale fuscous. Tibiae and tarsi of fore- and mid-legs blackish fuscous, with greyish bands. Hind tibia pale fuscous; tarsus somewhat blackish fuscous with a greyish band at apex. Forewing (Figs. 19- 22 ) with basal patch grey, mottled with black spots, and extend- ing from just beyond inner fourth of costa obliquely outward to middle of wing, then acutely angled inwardly to just beyond inner third of dorsum; median band divided into a whitish grey costal area and a large, whitish, semi-circular patch on dorsum, the costal area usually with three blackish costal dashes, the dorsal patch extending from dorsum to middle of wing, with four obscure black dashes on dorsal edge, and with mottled black and light brown scales in the middle; post-median band irregular, blackish, extending across the wing, narrower at costa; apical third with four blackish costal geminations, and a black subapical spot; tornal spot golden brown or whitish, mottled with black; submarginal line broken, black and brown; fringe dark grey, with some white-tipped scales, and a broken, darker, basal line. Hind wing dark fuscous; fringe light grey, with a darker basal line. Wingspread 15.0-20.5 mm. Male genitalia (Figs. 32-32b). Uncus reduced; apex rounded. Socii well developed, broadly sickle shaped. Tegumen rounded at shoulder. Clasper large, long; apical region broad. Aedeagus short, stout; apex truncate; cornuti a cluster of about 23-34 elongate spines. 5 (3) d 53-1 76, 1966 ZEIRAPHERA 175 Female genitalia (Figs. 34). Ostial plate narrow, emarginate. Ostium small, cup-shaped. Ductus bursae slightly sclerotized, with a broad, scleritized ring at neck of bursa. One sub-conical, rather large signum present. Type material. Holotype male, Courtenay, B.C., 9 July, 1951, F.I.S., reared from Douglas fir. Paratypes: Two males, two fe- males, Cowichan Lake, 27 July, 1944 (M. L. Prebble). Two males, Bridesville (Osoyoos district), 17 July, 1952. One male, Kaleden (Similkameen district), 8 July, 1952. One male, one female, Eneas Creek (Okanagan district) 6, 5 July, 1954. One male, Kitchener, 20 July, 1950. One male, Shuswap Falls (Kamloops district), 24 July, 1951. One male, Tappen Valley (Kamloops district), 17 July, 1954. One male, Tlupana Inlet (Vancouver Island) 28 July, 1954. One female, Powell River (New West- minster district), 24 July, 1950. One male Canal Flats, 25 July, 1950. One female, Dutch Creek (Kootenay district), 26 July, 1949. One female, Fairmont Hot Springs (Kootenay district), 27 July, 1949. All the above paratypes are from British Columbia and reared from Douglas fir by the Canada Forest Insect Survey. All type specimens No. 9486 in the Canadian National Collection, Ottawa, Ontario. Distribution. Southern British Columbia. Host plant. Pseudotsuga menziesii (Mirble) Franco. Remarks. On the basis of similarities in the genitalia, this species is closely allied to Z. destitutana. 8. Zeiraphera vancouverana McDunnough Zeiraphera vancouverana McDunnough, 1925, Can. Ent. 57: 21. Eocternal characters. Frons and vertex roughly scaled, dark brown. Antenna blackish brown; scape blackish brown inwardly, pale brown outwardly. Palpus pale fuscous inwardly, dark brown outwardly. Thorax and tegula dark brown. Abdomen blackish brown; terminal segment paler. Fore- and mid-legs blackish brown, with pale whitish-brown bands. Hind leg pale whitish fuscous; tarsus somewhat fuscous, with pale whitish bands. Forewing (Figs. 23, 24) with basal patch fuscous, mottled with brown, margined outwardly with blackish on dorsal half, and extending from beyond inner fourth of costa obliquely outward to middle of wing, then acutely angled inwardly to dorsum just beyond inner third; median band divided into a pale brown costal portion and a whitish grey, sub-rectangular, oblique dorsal 176 MUTUURA AND FREEMAN /. Res. Lepid. patch; apical third mottled with whitish grey, brown, and dark fuscous areas; fringe dark brown, with a broken blackish-fuscous basal line. Hind wing dark brown; fringe paler, with a pale blaekish-fuscous basal line. Wingspread 11-14 mm. Male genitalia (Figs. 33, 33a). Resembling those of fortunana. Uncus round at apex, not notched. Socii well developed, broadly sickle shaped. Tegumen angled at the shoulder. Apical region of clasper only slightly narrowed. Aedeagus short, stout; apex truncate; cornuti a cluster of 3-6 elongate spines. Female genitalia (Fig. 35). Anterior and posterior apophyses short. Ostial plate short and broad, narrower basally. Ostium cup-shaped. Ductus bursae slender, not sclerotized. One small signum present. Flight period. July. Type loeality. Ucluelet, Vaneouver Island, British Columbia. Distribution. Known only from Vancouver Island, British Columbia. Host plants. Picea sitehensis ( Bong. ) Carr, and P. engelmanni Parry. Remarks. On the basis of the male genitalia, thi sspecies is elosely allied to Z. fortunana ( Kft. ) . It is the smallest species in the genus. Journal of Research on the Lepidoptera 5(3) : 177-180, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 NOTES ON EAGLES PENELOPE ( SATURNIIDAE ) BRIAN O. C. GARDINER 18, Chesterton Hall Crescent, Cambridge, England. The larva of the Neotropical moth Fades penelope (Cra- mer) has been figured and described previously, (Burmeister, 1879). Packard (1905) gives a copy of BurmeistePs plate. The coloring of the figure of the larva appears very dim and faded when compared against the living specimens which have very bright and vivid coloring. According to Packard it lives on Melastomeae and Guava (Psidium pomiferum) one of the Myr- taceae. As the result of being sent eggs of this species, I have been able to rear it, and make some further observations on its habits and life-history. On 30. IX. 1963 about 400 ova were received from Sr. Fritz Plaumann of Nova Teutonia, Santa Catarina, Brazil. Although these had been sent by air-mail they had been in transit for over two weeks; most had hatched en route; the few survivors ap- peared rather weak. They were at once put into a small glass- topped metal tin of 3 inches diameter, containing a wet filter paper to maintain them at a high humidity. They were offered a choice of Laburnum (Laburnum anagyroides) , Privet (Ligus- trum ovalifolium) , Walnut (Juglans regia), and Beech (Fagus sylvatica). All these leaves were nibbled, but the larvae finally settled on Beech. They were maintained on this until the end of the penultimate instar, when the supply failed. Evergreen oak (Quercus ilex) was offered and accepted throughout the final instar. Privet which was also offered was firmly refused. When fully grown the lan^ae left the food and started wandering. They were then removed to a large tin containing moist peat and they burrowed into this to form a pupa in a pressed out cell some nine inches down. Of about fifty larvae that were still alive on receipt, about fifteen actually started to feed, one or two died at each moult and only 3 pupae were obtained. It is con- 177 178 BRIAN O. C. GARDINER J, Res. Lepid. sidered that the larvae which failed to start feeding did so be- ; cause they were already weak from starvation rather than the | possible unsuitability of the food. The larvae were reared at j 20-25 °C and were subjected to British daylight conditions from [ September thro December, but had erratic artificial light during the evening while the stock was being attended to. Under these conditions the pupae entered diapause and the moths did not i eclose for six months. i The duration in days of the six instars is given below. To obtain the figure for the final instar an estimate of 6 days between j entering the soil and pupation has been allowed. This is a figure compatible with my experience of other underground pupating species that do not hibernate ( or diapause ) as pre-pupae. 1st - 11; 2nd - 10; 3rd --- 10; 4th 8; 5th ~ 15; 6th 31. Total — 85. Pupal duration 24 weeks. A total life-cycle of 9 months is clearly incompatible with normal feral conditions. Other species bred contemporaneously, such as Dirphiopsis eumedide ( Stoll ) and Automeris beckeri \ ( Herrich-Schaffer ) proved to have a six-month life-cycle. No I explanation can be offered for this, it being one of the few Neo- tropical species bred in England that has had an unusual life- cycle period. It may well be that this species has a facultative light-controlled diapause and is much more critical concerning j its requirements than are the species of various other Neotropical genera. One has only to study the enormous range of latitude over which both this species and E. imperialis (Druce) range (Packard, 1905; Draudt, 1930) to realise that they must have varying diapause requirements in differing parts of their range, j From the Atlantic States imperialis has a pupal period of nine i or ten months (Crotch, 1956) whereas from Panama, bred con- j temporary with penelope, the pupal period was two months, THE OVA — Like other Citheroninae, very large; in form, | eliptical, with a marked central, angular, ridge, flat on base, in- ^ dented inwards on top. Maximum diameter 3.9 mm; minimum, i 3.2 mm; height 2.0 mm. The larvae being mainly hatched on I arrival, the color was not observed, but in those unhatched, the larvae were clearly visible, curled up around the equatorial plane, j thro the translucent golden colored shells. •THE LARVA — Due to pressure of other work it was not possible to keep accurate details of all the larval stages. Fig. 1, | however gives a better idea of the fullgrown larva than can any ‘ description. Although only a few larvae were available it was | noted that they appeared to prefer a solitary life. Fig. 1. Full-grown larva of Eacles penelope. Fig. 2. Adult Eacles penelope; Top $ , Lower $ . 180 , BRIAN O. C. GARDINER J. Res. Lepid. THE IMAGO — The three obtained all eclosed about one hour after sunset, but did not become active in their cage until the following night. This is perhaps not surprising since during June in England there is only 2-3 hours of darkness — considerably less than would normally be experienced in Brazil. It is worth mentioning in this connection that E. imperialis, with ten hours of darkness, emerged at dusk and was paired within six hours. Particularly striking was the very pungent and all-pervading odor of the meconium from the newly emerged moth. To the author it resembled the odor of onions which have just started to go rotten and have been kept in a closed container for some days. It closely resembled, but was subtly different from, the odor given off by the meconium of Periphobia hircia (Cramer), but was not so persistent. The latter genus has previously been com- mented on by Blest (1960a), but the odor from E. penelope does not appear to have been recorded. The adults show a specialised procryptic coloration when at rest (Blest, 1960b) and it may be that the obnoxious odor serves as protection during the dangerous period while the wings are expanding and hardening. When disturbed shortly after this, the moths flicked themselves rapidly about the floor, from right way up to upside down, finally coming to rest with legs and antennae tightly tucked to body. ACKNOWLEDGEMENTS I should like to thank Dr. David Blest both for helpful criticism and for taking the photograph illustrating the larva, and Simon Frey of the Virus Research Unit for that illustrating the adults. The cost of obtaining the eggs was supported by the United States National Institutes of Health, Project No. GM-07109. LITERATURE CITED BLEST, A. D., 1960a., A Study of the Biology of Saturniid Moths in the Canal Zone Biological Area. Smithsonian Report for 1959. pp. 447-464. BLEST, A D., 1960b., The Evolution, Ontogeny and Quantitative Control of the Settling Movements of some New World Saturniid Moths, with some Comments on Distance Communication by Honey-bees. Be- haviour 16: 188-253. BURMEISTER, J., 1879., Atlas de la Description Physique de la Republique Argentine Contenant des vues Pittoresques et des Figures d’Histoire Naturelle, Lepidopteres 5. Buenos Aires. CROTCH, W. J. B., 1956., A Silkmoth Rearer’s Handbook .Amat. Ent. 12 : I-I65. DRAUDT, M. in SEITZ, A., 1930., The Macrolepidoptera of the World. Vol. 6. Stuttgart. PACKARD, A. P., 1905., Monograph of the Bombycine moths of North America. Part II. Memoirs of the National Academy of Sciences. Vol. IX. Journal of Research on the Lepidoptera 5(3) : 181-184, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 REMARKS ON THE GENUS ZERA EVANS IN MEXICO WITH A NEW RECORD H. A. FREEMAN^ 1605 Lewis Drive, Garland, Texas Four species of Zera have been previously recorded from Mexico, and in most cases the data was very incomplete. Hoff- mann lists three species from that country: scybis (Godman & Salvin), which is a synomym of nolckeni (Mabille); pelopea ( Godman & Salvin ) , which is a synomym of tetrastigma ( Sepp ) ; and hyacinthinus (Mabille), which he said he had not seen in Mexico. Evans lists three species contained in the British Museum from Mexico: nolckeni, hyacinthinus, and eboneus (Bell). During the process of making a careful study of the Hesperiidae of Mexico I have found that there are actually five species of in Mexico, one of which I am recording for the first time. Zera nolckeni (Mabille) 1891 Synomym.scybis Godman & Salvin 1894: Atoyac, Veracruz, Mexico. Type locality.— Bogota. General distribution.— Mexico, Panama, Colombia, Venezuela, Ecuador (Loja, Ambato), Peru (Cuzco, Chanchamayo, Huanca- bamba), and Bolivia (Cochabamba, Tanampaya, San Jacinthe). Mexican distribution.— Atoyac, Veracruz; Sierra Madre del Sur; Cuenca del Rio Balsas; and Colima. Remarks.— This species was first recorded from Mexico by Godman & Salvin as scybis based on a specimen from Atoyac, Veracruz. Hoffmann listed several locations which are mentioned above. In the large number of specimens of Hesperiidae that Dr. Tarsicio Escalante, Mexico, D. F. sent me for determination there were no examples of this species. During the eighteen trips that I have made to Mexico I have not as yet turned up this species. From the available data nolckeni is apparently rare in Mexico as well as in some of the Central American areas. 9 would like to express my thanks to the National Science Foundation for Research Frant GB-4122 which is making this study of the Hesperiidae of Mexico possible. 181 182 H. A. FREEMAN /. Res. Lepid. Zera hosta Evans 1953 Type locality. — Costa Rica. General distribution. — Mexico, Costa Rica, and Colombia. Mexican distribution. — Sta. Rosa, Comitan, Chiapas, 1 male May 1965. Remarks. — Evans described hosta as a subspecies of phila (Godman & Salvin) based on the general difference of the secondaries on the lower surface, hosta being bluish-white over the discal one half to two thirds of the wing, and phila being entirely tawny, with some black spots. The general range of the two overlaps thus bringing up the question as to whether they are separate subspecies or species. For the present I will con- sider them to be speciafically distinct. New record for Mexico. — the above mentioned male, which was obtained from Dr. Escalante, is the first record of this species from Mexico. Zera hyacinthinus (Mabille) 1877 ' Type locality. — ? General distribution. — Mexico, Guatemala, Honduras, Nicara- gua, Costa Rica, Panama, and Colombia. Mexican distribution. — Presidio, Veracruz, 1 male, August 1951 (A. Ramirez); Ajijic, Jalisco, 1 male, 27 September 1965, and 1 male, 5 October 1965 (Robert Wind); and 2 males and 1 female, Oaxaca, Oaxaca, 22 June 1966 (H. A. Freeman). Remarks. — Considerable confusion has centered around this species for many years. Godman & Salvin figured the male and genitalia of tetrastigma as hyacinthinus. Hoffmann stated that he had not seen any examples from Mexico indicating that possibly he was not familiar with the exact status of the species. Evans recorded four males and three females from Mexico with no other data. I have in my collection the above mentioned specimens listed under Mexican distribution of this species from Mexico. From the available data indications are that this species of is the predominant one in Mexico. Zera tetrastigma (Sepp) 1848 Synomyms. ™ menedemus Godman & Salvin 1894: Panama; pelopea Godman & Salvin 1894: Amazons. Type locality. — Surinam. General distribution. — Mexico, Guatemala, Nicaragua, Pana- ma, Colombia, Trinidad, British Guiana, French Guiana, Peru ( Chanchamayo, Chaquimayo), Bolivia ( Buenavista ) , Upper Amazons (Iquitos, Tonantins, S. Paulo de Olivenca), and Para. Mexican distribution. — Sierra Madre Occidental (Nayarit); 5 (3) : 18 1-184, 1966 GENUS ZERA 183 Catemaco, Veracruz, one male, November 1964, and one female, January 1952 (A. Ramirez). Specimens obtained from Dr, Escalante. Remarks. — The considerable variation of this species has resulted in two synomyms: menedemus, having the secondaries below with the dorsal half white; and pelopea, having the second- aries below entirely dark brown, with bands absent or vestigial. Typical tetrastigma is dark brown on the secondaries below with well-marked black central and submar ginal bands, and some bluish- white over the dorsal half of the wing. Zera eboneus (Bell) 1947 Type locality. — Ojo de Agua, Veracruz, Mexico. General distribution. ■— Mexico. Mexican distribution. — Ojo de Agua, Veracruz, 12 June 1941. Remarks. — From Bells original description and genitalia figure ehoneus is apparently a distinct species. Evans states that the genitalia are similar to those of belti ( Godman & Salvin ) , and that eboneus is probably an aberration or subspecies of belti. However as he never examined the type it seems most unlikely that the two are the same species since they are so different superficially. Key to the Mexican species of la. Primaries above with dark central area 2 lb. Primaries above plain black 5 2a. Primaries above with dark central area extending to base and lower end of cell; three apical spots ................................3 2b. Primaries above with dark central area not extending to base, which is broadly violaceous, nor to end of cell; 1-3 apical spots .4 3a. Primaries above with dark bar at end of cell; secondaries below with dorsal half bluish-white, but tawny colouring from costa runs narrowly along outer margin to vein lb nolckeni (Mab. ) 3b. Primaries above with no dark bar at end of cell; secondaries below with dorsal one half to two thirds bluish-white ........hosta Evans 4a. Primaries with tornus truncate and dorsum slightly concave; primaries below with tornal area broadly tawny, often reaching the costa and unmarked except for an inconspicious discal spot; usually 3 apical spots ....hyacinthinus (Mab.) 4b. Primaries not truncate at tornus and dorsum more or less straight; primaries below with tornal tawny area restricted and marked with a large dark central spot; usually only one apical spot tetrastigma (Sepp) 5. Primaries below brown, with faint spot in space 8; secondaries below with pale bar at end of cell eboneus ABellQ 184 H. A. FREEMAN /. Res. Lepid. LITERATURE CITED BELL, ERNEST L. 1947. New species and subspecies of Neotropical Hesperiidae. Amer. Mus. Novitates No. 1330, pp. 1-9, 1 pi. EVANS, W. H. 1953. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part III. Pyrginae. Sec. 1. London: British Museum. 178 pp., pis. 10-25. GODMAN, FREDERICK DUCANE, and OSBERT SALVIN. 1887-1901. Biologia centrali-Americana. Insecta. Lepidoptera-Rhopalocera, II, 244- 637; LII: pis. 112. HOFFMANN, C. C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros Mexicanos. Segunde parte-Hesperioidea. An. Inst. Biol. Mexico. 12: 237-294. Journal of Research on the Lepidoptera 5(3) : 185-195, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 STUDIES ON THE NE ARCTIC EUCHLOE Part 3. COMPLETE SYNONYMICAL TREATMENT PAUL A. OPLER Department of Entomology University of California Berkeley, California Full synonymies are presented here for Euchloe amonides, Euchloe creusa, Euchloe hyantis, and their named subspecific segregates. TTie author believed completeness to be essential so that future workers may refer to the material contained in past literature citations in light of the interpretations of these treat- ments. Synonymical treatments for Euchloe olympia and E. olympm rosa are not presented at this time. Euchloe (Euchloe) ausonides (Lucas) Anthocharis ausonides Lucas, 1852, Rev. ZooL, 4 (2nd ser. ): 340; Boisdu- val, 1852, Ann. Soc. Ent. Fr., 2;X;286; Realdrt [partim.], 1866, Proc. Ent. Soc. Phil., 6:135, Herrich-Schaffer, 1867, Corr.-Blatt. Zool. Min. Ver. Regnsb., 21: 144; Boisduval, 1869, Ann Soc. Ent. Belg., 12:40; Mead [partim.], 1875, in Wheeler, Rept. Geogr. ExpL, 5:747; Mosch- ler, 1878, Stett. Ent. Zeit., 39: 299, 309; Edwards, 1884, Trans. Amer. Ent. Sic., 9:261; Smith, 1891, List Lepid. Bor. Amer., p.l3; Skinner [partim.] [erroneously includes the name coloradensis as a synonym], 1898, Syn. Cat. N. Am^. Rhop., p. 64; Grote, 1900, Proc. Am. Phil, Soc., 39:40; Skinner, 1903, in Smith, Chklst. Lepid. Bor. Amera., p. 9; Franck, 1905, Ent. News 16:28. Anthocharis ausonides [lapsus calami]: Edwards [partim.], 1863, Proc. Ent. Soc. Phil., 2:81; Weidemeyer, 1863, Proc. Ent. Soc. Phil., 2:151; Edwards [partim.], 1897, Butt. N. Amer., 3:412. Euchloe ausonides: Kirby, 1871, Syn. Cat. Di. Lepid., p. 340; Beutenmuller, 1898, Bull. Amer. Mus. Nat. Hist, 10:240-241, pi. 14, fig. 1; Holland [partim.], 1898, Butt. Book, p. 283, pi. 32, figs. 24, 25, pi. 34, fig. 3; Butler [partim.], 1899, Can. Ent., 31:19; Wright [partim.], 1906, Butt. West Coast, p. 104, figs. 56, 56b, 56c; Coolidge [partim.], [er- roneously includes the name coloradensis as a synonym], 1908, Can. Ent, 40:298; Coolidge & Newcomes, 1908, Ent. News 19:204; Rober [partim.], 1909, in Seitz, Macrolepid.. World 5:95, pi. 28, fig. a’; Barnes & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3: 61-62; Barnes & McDunnough, 1917, Chklst Lepid. Bor. Amer., p. 3; Barnes & Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25:7; Com- stock [partim.], 1927, Butt. Calif., p. 37, pi. 10, figs. .6, 9; Holland, 1931, Butt Book, p. 284, pi. 32, figs.24, 25, pi. 34, fig 3; Martin & Ingham, 1931, Bull. So. Calif. Acad. Sci., 29: ; Davenport & Dethier, 1937, Ent Amer. 17: 179; McDunnough [partim.], 1938, Mem. So. 185 186 PAUL A. OPLER /, Res. Lepid. Calif. Acad. Sci., 1:7; Leighton [partim.], 1946, Univ. Wash. Pub. BioL, 9: ; Martin & Truxd, 1955, Los Angeles Co. Mus. Sci. Ser., 18(Zool.), 8:5; Bird, 1956, Lepid. News 10:107; Ehrlich & Ehrlich [partim.], 1961, How to know Butt., p. 75, fig. ; Ehrlich & Ehrlich, 1962, Microent., 25( 1 ) :4. Anthocharis ausonia var. ausonoides [lapsus calami]-. Strecker, 1878, Syn. Cat. Macrolep., p. 77. Anthocrais ausoniedes [lapsus calami]: Hy. Edwards, 1880, Pacific Coast Lepid., 22; Hy Edwards, 1881, Papilio 1:51; von Osten-Sacken, 1882, Papilio 2: 29. Anthocaris ausonides: Geddes, 1883, Can. Ent., 15:221. Euchloe “variety from Vancouver”, Butler [partim.], 1899, Can. Ent., 31:19. Synchloe ausonides: Dyar, 1902, Bull. U.S. Nat. Mus., 52:7; Anderson, 1904, Cat. Brit. Col. Lepid., p. 6. Synchloe aussonides [lapsus calami]: Elrod & Maley [partim.], 1906, Bull. Univ. Montana 30 ( Biol. ) : 32. Synchloe ausonides: Elrod & Maley [partim.], 1906, Bull. Univ. Montana 30 (Biol.): 32. Euchloe creusa {non Doubleday): Wright [partim.], 1906, Butt. West Coast, fig. 54b. Euchloe rosa (non Edwards): Wright [partim.], 1906, Butt. West Coast, fig. 54c. Euchloe coloradensis ( non Hy. Edwards ) : Coolidge, 1908, Can. Ent., 40:299 (synonymy); Storer & Usinger, 1963, Sierra Nevada Nat. Hist, Univ. Calif. Press, p. 195. Euchloe belia var. creusa {non Doubl^ay): Verity [partim.], 1908, Rhop. Pal., pi. 37, fig. 19. Euchloe belia var. creusa gen. aest. ausonides: Verity [partim.], 1908, Rhop. Pal., pi. 37, fig. 20 [type], figs. 21, 22, 23, 24 Euchloe ausonides $ var. flavidalis Comstock [partim.], 1924, Bull. So. Calif. Acad. Sci., 23: 5; Comstock [partim.], 1927, Butt. Calif., p. 37, pi. 10, fig. 10 [TYPE]. Euchloe ausonides var. semiflava Comstock [partim.], 1924, Bull. So. Calif, p 37, pi. 10, fig. 11 [TYPE]. Euchloe ausonides ab. semiflava: Barnes & Benjamin [partim.]. 1926, Bull. So. Calif. Acad. Sci., 25: 7; McDunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1(1): 7; Leighton [partim.], 1946, Univ. Wash. Pub. Biol., 9: ; dos Passos [partim.], 1964, Lepid. Soc. Mem., 1:49. Euchloe ausonides ab. flavidalis: Barnes & Benjamin [partim.], 1926, Bull. So. Calif. Acad. Sci., 25: 7; McDunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1(1): 7. Euchloe ausonides coloradensis {non Hy. Edwards): Comstock [partim.], 1927, Butt. Calif., p. 38; Brooks, 1942, Can. Ent., 74: 31; Martin & Truxal [partim.], 1955, Los Angeles Co. Mus. Sci. Ser., 18 (Zool. ): 8:5; Garth & Tilden [partim.], 1963, J. Res. Lepid., 2:50. Euchloe ausonides ab. boharti Doudoroff, 1930, Pan-Pac. Ent., 6: 143; McDunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1(1): 7; dos Passos [partim.], 1964, Lepid. Soc. Mem., 1:49. Euchloe ausonides ausonides: Talbot, 1932, Lepid. Catalogus 53:301; dos Passos [partim.], 1964, Lepid. Soc. Mem., 1:49. Euchloe ausonides race color^ensis tr. f. [sic!] hemiflava Field, 1936, J. Ent & Zool, 28:19. Euchloe ausonides coloradensis ab. hemiflava: McDunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1(1): 7; Martin & Truxal [partim.], 1955, Los Angeles Co. Mus. Sci. Ser., 18 (Zool), 8:5; dos Passos [partim.], 1964, Lepid. Soc. Mem,, 1:49. Euchloe creusa orientalides {non Verity): Martin & Truxal, 1955, Los Angeles Co. Mus. Sci. Ser., 18 (Zool.) 8:5. 5 (3) . -185-195, 1966 NEARCTIC EUCHLOE 187 Euchloe creusa lotta (non Beutenmuller) : Garth & Tilden [paHim.], 1963, J. Res. Lepid., pi. 58, fig. 1. Euchloe (Euchloe) ausonides coloradensis (Hy. Edwards) Anthocharis ausonides [lapsus calami]: Edwards [partim.], 1863, Proc. Ent Soc. Phil., 2:78; Edwards [partim.], 1897, Butt. N. Amer., 3:412. Anthocharis ausonides: Reakirt [partim.], 1886, Proc. Ent. Soc. Phil,, 6:135; Mead [partim.], 1875, in Wheeler, Rept. Geogr. ExpL, 5:747; Mead, 1877, Psyche 2:182; Skinner, 1902, Trans. Amer Ent. Soc., 29:37. Anthocharis coloradensis Hy. Edwards 1881, Papilio 1:50. Anthocharis ausonides coloradensis: Edwards, 1884, Trans. Amer. Ent. Soc., 9:261; Smith, 1891, List Lepid. Bor. Amer., p. 13; Skinner, 1903, Chklst. Lepid. Bor. Amer., p. 9. Anthocharis ausonides var. coloradensis: Gockerell, 1899, Ent. Amer., 5:33. Euchloe ausonides var. coloradensis: Beutenmuller [partim.], 1898, Bull. Amer. Mus. Nat. Hist., 10:241. Euchloe coloradensis: Butler, 1899, Can. Ent., 31:19. Synchloe ausonides coloradensis: Dyar, 1902, Bull. U.S. Nat. Mus., 52:7. Euchloe ausonides coloradensis: Wright, 1906, Butt. West Coast p. 104; Barnes & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:61; Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer., p. 3; Barnes & Benjamin, 1926, Bidl. So. Calif. Acad Sci., 25:7; Comstock [partim.], 1927, Butt. Calif., pL 10, figs. 13, 14; Klots, 1930, Bull. Brook. Ent. Soc., 25:154; Holland, 1931, Butt. Book p. 284; Talbot, 1932, Lepid. Cataiogus 53:302; Cross, 1937, Proc. Colo. Mus. Nat. Hist., 16:7; Mc- Dunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1(1):7; Brown, 1955, Proc. Denver Mus. Nat. Hist., 6:181; Martin & Truxal [partim.], Los Angeles Co. Mus. Sci. Ser., 18(Zool.), 8:5; Defoliart, 1956, Lepid. News 10:99; dos Passes [partim.] [erroneously includes the name hemiflava as a synonym], 1964, Lepid. Soc. Mem., 1:49. Euchloe helia var. belioides race montana Verity, 1908, Rhop. Pal., pi. 68, figs. 13, 14, 15. Euchloe ausonides montana: Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer., p. 3 [synonomyj; Barnes & Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25:7 [synonomyj; Holland, 1931, Butt. Book p. 284 [syn- onomyj; McDunnough, 1938, Mem. So. Calif. Acad. Sci., 1(1):7 [synonomyj; dos Passos, 1964, Lepid. Soc. Mem., 1:49 [synonomyj. Euchloe ausonides coloradensis ab. hemiflava (non Field): Martin & Truxal [partim.], 1955, Los Angeles Co. Mus. Sci. Ser., 18(Zool.), 8:5. Euchloe ausonides (non Lucas): Maeki & Remington, 1960, J. Lepid. Soc., 14: 42, 52; Ehrlich & Ehrlich [partim.], 1961, How to Know Butt., p. 75; Ehrlich, 1961 Microent, 24(4): 138. Euchloe ausonides race mayi Chermock & Chermock, 1940, Can. Ent., 72:82. Euchloe (Euchloe) ausonides mayi Chermock & Chermock Euchloe ausonides mayi: Brooks, 1942, Can. Ent., 74:32; dos Passos, 1964, Lepid. Soc. Mem., 1:49. Anthocharis creusa Doubleday, 1847, Gen. Di. Lepid., 1:56, pi. 7, fig. 1; Weidemeyer, 1863, Proc. Ent. Soc., Phil., 2:151; Herrich-Schaffer, 1867, Corr.-Blatt. Zook Min. Ver. Regnsb., 21:144; Smith, 1891, List Lepid. Bor. Amer., p. 13; Skinner, 1898, Syn. Cat. N. Amer. Rhop., p. 64; Skinner, 1903, in Smith, Chk. Lst. Lepid. Bor. Amer., p. 9. Euchloe creusa: Kirby, 1871, Syn. Cat. Di. Lepid., p. 340; Beutenmuller, 1897, J. N.Y. Ent. Soc., 5:208; Beutenmuller [partim.], 1898, Bull. Amer. Mus. Nat. Hist., 10:16; Buder, 1899, Can. Ent, 31:56; Coolidge, 1908, Can, Ent., 40:299; | ?] Rober, 1909, in Seitz, Macrolepid. World 5:95, pi. 28, fig. a”; Barnes & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:59; Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer,, p. 3; Barnes & Benjamin [partim.] [authors erroneously 188 PAUL A. OPLER J. Res. Lepid. include the name pumilio as a synonym], 1926^ Bull. So. Calif. Acad. Sci., 25:7; McDunnough [partim.] [erroneously includes the name pum- ilio as a synonym], 1938, Mem. So. Calif. Acad. Sci.,- 1:17; Martin & Truxal. 1955, Los Angeles Co. Mus. Sci. Ser., 18(ZdoL ), 8:5; Ehrlich & Ehrlich [partim.] [authors erroneously include Alberta in statement of distribution], 1961, How to Know Butt., p. 75; dos Passes [partim.] [incorrectly includes the name pumilio as a synonym], 1964, Lepid. Soc. Mem., 1:49. Anthocaris crensa [lapsus calami]: Edwards, 1881, PapiHo 1:51. Euchloe creusa var. elsa BeuterunuUer [partim.], 1898, Bull. Amer. Mus. Nat. Hist., 10:342, pi. 14, fig. 3; Beutenmuller [partim.], 1899, Can. Ent, 31:56. Euchloe elsa: Butler, 1899, Can. Ent.,31:19 [synonomy]; Coolidge, 1898, Can. Ent., 40:299 [synonomy]; Barnes & McDuimough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:59 [synonomy]; Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer., p. 3 [synonomy]; Barnes & Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25:7 [synonomy]; Holland [partim.], 1931, Butt. Book p. 285 [synonomy]; McDunnough [partim.], 1938, Mem. So. Calif. Acad. Sci., 1:7 [synonomy]; dos Passes [partim.], 1964, Lepid. Soc. Mem., 1:49 [synonomy]. Synchloe creusa: 1904, Anderson, Cat. Brit. Col. Lepid., p. 6. Euchloe belia var. orientalides Verity [partim.], 1908, Rhop. Pal., pi. 36, fig. 58. Euchloe belia var. creusa Verity [partim.], 1908, Rhop. Pal., pi. 68, figs. 11, 13 (“COTYPES”) [As mentioned in Barnes & McDunnough, 1916, verified through correspondence with Dr. N. D. Riley of the British Museum, 1964, these two figures represent the dorsal and ventral sur- faces of the single type specimen]. Euchloe creusa orientalides: Barnes & McDunnough [partim.], 1917, Chklst. Lepid. Bor. Amer., p. 3; Barnes & Benjamin [partim.[, 1926, Bull. So. Calif. Acad. Sci., 25:7; McDurmough ]partim.], 1938, Mem. So. Calif. Acad. Sci., 1:7; dos Passos [partim.], 1964, Lepid. Soc. Mem., 1:49. Euchloe (Euchloe) creusa: Klots, 1930, Bull. Brook. Ent. Soc., 25:87. Euchloe creusa creusa: Talbot [partim.], 1932, Lepid. Catalogus 53:302. Euchloe creusa creusa ab. elsa: Talbot [partim.], 1932, Lepid. Catalogus 53:302. Euchloe (Euchloe) hyantis hyantis (Edwards) Anthocharis hyantis Edwards, 1871, Trans. Amer. Ent. Soc., 3:205; Mead, 1877, Psyche, 2:182; Butler [partim.], 1882, J. Linn. Soc., 16:471; Edwards [partim.], 1884, Trans. Amer. Ent. Soc., 9:260; Smith, 1891, List Lepid. Bor, Amer., p. 13; Edwards, 1892, Can. Ent., 24:109; Skin- ner, 1903, in Smith, ChJdst. Lepid. Bor. Amer., p. 9. Anthocaris hyantis: Hy. Edwards [partim.] [erroneously treats the name creusa as a synonym], 1880, Pac. Coast Lepid., 22; Hy. Edwards, 1880, Pac. Coast Lepid., 28; Edwards, 1881, Papilio 1:51; von Osten Sacken, 1882, Papilio, 2:29. Anthocaris creusa: Hy. Edwards, 1880, Pac. Coast Lepid., no. 22 [synon- omy] . Euchloe hyantis: Butler [partim.], 1882, J, Linn. Soc., 16:471; Beutenmul- ler, 1897, J. N.Y. Ent. Soc., 5:208; Butler, 1899, Can. Ent, 31:19; Beutenmuller, 1899, Can. Ent, 31:56; Coolidge [partim.], 1908, Can. Ent, 40:298; [?] Rober, 1909, in Seitz, Macrolepid. World pi. 28, fig. a’”; Holland [partim.], 1931, Butt. Book, rev. ed., pi. 67, fig. 27 [“TYPE”]; Storer & Usinger, 1963, Sierra Nevada Nat Hist, pi, 67, fig. 72. Euchloe creusa: Beutenmuller [partim.], 1898, Bull. Amer. Mus. Nat. Hist, 10:241, pi. 14, fig. 2. 5 (3) ;185~19S, 19-66 NEARCTIC EUCHLOE 189 Anihqcharis ausonMes var, hy antis i SkimiCT, 1898, Syn. Cat. N. Amer, Rhop., p. 64. Antkocharis amonides hyantki ‘Franck, 1905, Ent, News, 16:28 [synonomy]. Anthocham ausofiides colorademis: Franck, 1905, Ent News, 16:29 [syn- onymy] . Euchhe crema vax. hyantisi Coolidge [partim.], 1908, Can. Ent,, 40:298. Euchloe belia var. creusa ( non DoubMa/) : Verity, 1908, Rhop. Pal., pi. 37, fig. 16. Euchlm cretisa cum. ab, [sic/] pumilio Strand [partim.]^ 1914, Archiv. f. Natiirgesck, 80(A 11). -153. Euchloe crema hyantis: Bames & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:60; Bames & McDunnough [partim.], 1917, Chklst Lepid, Bor. Amer., p. 3; Barn® & Benjamin [pariim.], 1926, Bull. So. CaLt. Acad. Sci, 25:7; Comstock [partim.], 1927, Butt. Calif., p. 38; Talbot, 1932, Lepid. Catalogus 53:302; McDunnough [partim.]^ 1938, Men. So. Calif. Acad. Sci, 1:7; Martin & Truxal [p-artim.], 1955, Los Angeles Go. Mus. Sci. Ser., 18(ZooL), 8:5; Garth & Thdm, 1963, J. Res. Lepid., 2:50, 87; dos Passes [partim.]^ 1964, Lepid, Soc, M«n., ^ 1:49. EtwMoe creusa pseudoausonidesi Barnes & McDunnough, 1916, Contr. Nat. Hist, Lepid. N. Amer., 3:61 [synonomy]; Barnes & McDunnough, 1917, Chklst. L^id. Bor. Ain»., p. 3 [synonomy]; Bames && Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25: [synonomy]; McDunnough, 1938, Mem. So. Cdif. Acad. Sci., 1:7 [synonomy]; dos Passos, 1964, Lepid. Soc. Man., 1:49 [synonomy], Euchloe creusa orientalidesi Bam« & McDumiough, 1916, Contr. Nat. Hist. Lepid, N. Amer., 3:61 [synonomy]; Bames & McDunnough [par- tim,], 1917, Chklst Lepid. Bor. Amo”., p. 3; Bames & Benjamin [partim.], 1926, Bull. So. Calif. Acad. Sci., 25:7; McDunnough [par- tim.] 1938, Mem. So. Calif, Acad. Sci., 1:7; dos Passos [p-artim.], 1964, Lepid. Soc. Mon., 1:49. Euchloe ausonides andrewd (non Martin): Powell, 1958, Lepid. News, 12:28. Euchloe (Euchloe) hyantis andrewsi (Martin) NEW COMBINATION Euchloe amonides race andrewsi Martin, 1936, Bull. So. Calif. Acad, Sci., ,35:94. Euchloe amonides andrewsi: Martin & Truxal, 1955, Los Angeles Co. Mus. Sci. Ser., 18(ZooL), 8:5; dos Passos, 1964, Lepid. Soc. Mem., 1:49. Euchioe (Euchloe) hyantis lotta (Beutenmuller), NEW COMBINATION Anthocharis creusa ( fWfi. Doubieday ) : Strecker, 1878, Syn. Cat. Macrolepid., p. 77. Anthocharis hyantis (non Edwards): Morrison, 1883, PapiliO' 3:9; Edwards [p-artim.], 1884, Trans. Amer. Ent. Soc., 9:260. Euchloe lotta BeutenmullCT, 1898, Bull. Amer. Mus. Nat Hist., 10:24, pi. 14, fig. 4; Butler, 1899, Gan. Ent., 31:56; Wright [partim.], 1906, Butt. West Coast, p. i04; Holland [partim.], 1931, Butt. Book, rev. ed., p. 285, pi 73, fig. 5 [“TYPICAL”, text], [“PARATYPE”, plate legend]. Euchloe crema (non Doubleday): Holland [purtim.], .1898, Butt Book, p. 283, pi 32, fig. 23, pi. 34, fig. 2; Wright [partim.], 1906, Butt West Coast, p. 194, figs. 54, 54a; Klots, 1930, Bull. Brook. Ent. Soc., 25: 153; Holland [partiwL], 1931, Butt. Book, rev. ed., pi 32, fig. 23, pi 34, fig. 2; Laghton [partim.], 1946, Univ. Wash.- Pub. Biol, 9: ; Bauer, 1953, Lepid. News 7:146; Brown, 1955, Proc. Denver Mus. Nat. Hist, 6:182; Ehrlich & Ehrlich [partim.], 1961, How to Know Butt., p. 75, ,fig. Euchloe amonides var. lotta: Bro'wning, 1901, Ent. News, 12(10):301, fig. 190 PAUL A. OPLER /. Res. Lepid. Synchloe creusa (non Doubleday); Dyar [partim.], 1902, Bull. U.S. Nat. Mus., 52:7. Synchloe lotta: Dyar [partim.], 1902, Bull. U.S. Nat. Mus. 52:7. Anthocharis lotta: Skinner, 1903, in Smith, Chklst. Lepid. Bor. Amer., p. 9. Euchloe hyantis (non Edwards): Wright [partim.], 1906, Butt. West Coast, p. 104; Strand [partim.], 1914, Axchiv. f. Naturgesch., 80(A 11):153; Holland [partim.], 1931, Butt. Book, rev. ed., p. 285. Euchloe rosa (non Edwards): Wright [partim.], 1906, Butt. West Coast, p. 104, figs. 56, 56b. Euchloe belia var. orientalides Verity [partim.], 1908, Rhop. Pal., pi. 36, fig. 58. Euchloe creusa var. lotta: Coolidge, 1908, Can. Ent., 40:299. Euchloe belia var. creusa (non Doubleday): Verity [partim.], 1908, Rhop. Pal., pi. 37, figs. 17, 18. Euchloe creusa cum. ab. [sic!] pumilio Strand [partim.], 1914, Archiv. f. Naturgesch., 80 (A 11): 153. Euchloe creusa lotta: Barnes & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:61; Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer., p. 3; Barnes & Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25:7; Comstock [partim.], 1927, Butt. Calif., p. 38, pi. 10, figs. 7, 8; Gunder, 1930, Bull. So. Calif. Acad. Sci., 29:9; Comstock & Dammers, 1932, Bull. So. Calif. Acad. Sci., 31:35; Talbot, 1932, Lepid. Catalogus 53: 302; Davenport & Dethier [partim.], 1937, Ent. Americana 17:179; McDunnough, 1938, Mem. So. Calif. Acad. Sci., 1(1):7; Martin & Truxal [partim.], 1955, Los Angeles Co. Mus. Sci. Ser., 18(Zool. ), 8:5; Garth & Tilden [partim.], 1963, J. Res. Lepid., pi. 5, fig. N; dos Passes, 1964, Lepid. Soc. em., 1:49. Euchloe belioides: Barnes & McDunnough, 1916, Contr. Nat. Hist. Lepid. N. Amer., 3:61 [synonomy]; Barnes & McDunnough, 1917, Chklst. Lepid. Bor. Amer., p. 3 [synonomy]; Barnes & Benjamin, 1926, Bull. So. Calif. Acad. Sci., 25:7 [synonomy]; Holland, 1931, Butt. Book, rev. ed., p. 285 [synonomy]; McDunnough, 1938, Mem. So. Calif. Acad. Sci., 1(1);7 [synonomy]; dos Passes, 1964, Lepid. Soc. Mem., 1:49 [synonomy]. Euchloe creusa hyantis (non Edwards): Coolidge, 1925, Ent. News, 36:65; Comstock [partim.], 1927, Butt. Calif., p. 38, pi. 10, figs. 4, 5; Daven- port & Dethier [partim.], 1937, Ent Americana 17:179; Leighton [partim.], 1946, Univ. Wash. Pub. Biol., 9: ; Martin & Truxal [par- tim.], 1955, Los Angeles Co. Mus. Sci. Ser., 18(Zool. ), 8:5. Euchloe creusa ab. pumilio: Strand, 1927, Bull. Soc. Zool. France 51:412. STUDIES ON THE NEARCTIC EUCHLOE Part 4. TYPE DATA AND TYPE LOCALITY RESTRICTIONS’ In any taxonomic revision, it is important for the revisor to know the biological identity and the type locality of the insect with which he is dealing. As a direct result of this precept, the information concerning the types of the named entities of Ne- arctic Euchloe, which are considered to be biologically applicable by this author, is presented in this paper. Also, the type localities 1 F. M. Brown, Colorado Springs, Colorado and J. A, Powell, University of California, Berkeley, California kindly read the manuscript of this paper and offered several helpful suggestions. 5 (3) : 185-195, 1966 NEARCTIC EUCHLOE 191 of three named taxa are restricted so as to provide a base for logical discussion of the distributional limits and the geographic variability of these units and their associated populations. For each name, the citation of the original description, a compendium of the available information concerning the type, and an interpretation of the type information are presented. Euchioe (Euchloe) ausonides (Lucas) Citation of original descriptions Lucas, 1852. Rev. Mag. Zool. 4 (2nd series ) : 324-345. Type data: The holotype is in the collection of the United States National Museum, Washington, D.C. The specimen is ex collection Barnes, ex collection Oberthur, and ex collection Bois- duval. Dr. Jerry A. Powell of the University of California at Berkeley examined the type specimen and obtained the data which are presented below. The specimen, which lacks the right forewing and both antennae, has a forewing costal measurement [base to apex] of 24.3 millimeters. The forewing discal spot is 1.2 millimeters in width at the middle and has fairly heavy whitish overscaling along the vein at the upper edge of the cell. The specimen bears five labels as follows: l./“Ex Musaeo Dris Boisduval” machine printed label with black border, 2./“Ober- thur Collection” machine printed label with red border, 3./“Fig- ure par R. Verity/Rhopal. Palaearctica PI XXXVII, fig. 20”, 4./‘Anthocharis Ausonides, Boisd., Lepid. Californie, II Partie, St. Entom. Belgiq. 1869 p. 40” handwritten label, 5./“Type ausonides Bdv. a/c Hofer 1925FHB” handwritten label with red border, different hand than 4. Interpretation of type data: That this is the type specimen described by Lucas is substantiated by the fact that the costal measurement of this unusually large male specimen is in close agreement with the measurement given by Lucas (1852). The specimen was part of the first lot of material sent to Boisduval by P. J. M. Lorquin who collected the material for that shipment in central California. Since Lucas, as well as Boisduval, stated nothing further than “California” as the type locality, since there are populations of this species occuring in California with significantly varying phenotypic expression, and since Lorquin could well have collected the specimen in the San Francisco Bay area, the type locality is hereby restricted to San Francisco, San Francisco County, California. Euchloe (Euchioe) ausonides coloradensis (Hy. Edwards) Citation of original description: Hy. Edwards, 1881. Papilio l(4):50-55. 192 PAUL A. OPLER /. Res. Lepid. Type data: The holotype is in the American Museum of Natural History, New York. The specimen bears the following labels: l./AMNH catalogue number “3577” on which the word “Colorado” is appended, 2./The original Henry Edwards label bears the catalogue number "14”. Interpretation of type data: The following information was supplied by F. M. Brown (personal communication). In his catalogue under entry “14”, Henry Edwards noted, ""ausonides Bdv. pastures Alameda Co. Calif, etc. HE 3-5, settles on flowers of Sinapis [=Brassica].” Brown states that the “14” on the type specimen means that Henry Edwards at first considered the specimen to be the same as the material he collected in California from March through May. Brown, who examined the type speci- men at the request of the author, sent the accompanying analysis which is quoted in part below. The specimen is without doubt one sent to Henry Edwards by W. H. Edwards from the material collected by T. L. Mead in 1871. In a letter from W.H.E. to Hy. E. dated July 8, 1871 about Mead specimens being sent to Hy. Edw., Edwards states 'And Anthocaris 1 is Auso- nides. The Anthocaris 2 is new if it is not ReakirtiV. The date of the letter places the capture in June, Mead (1875: 747-748) confirms this. From the itinerary of Mead pieced together by Brown (1955), it is seen that in June, 1871, Mead traveled from Denver to Fairplay via Morrison, Turkey Creek Junction, Bailey, and Kenosha House, whereupon he returned to Turkey Creek Junction by the same route. He then remained at Turkey Creek for a week (June 20-27, 1871) before moving elsewhere. Coolidge and Newcomer (1908) quote from a letter written by Mead to W. H. Edwards in which Mead told about finding the eggs and larvae of Euchloe ausonides coloradensis at Turkey Creek Junc- tion. From the information supplied by Brown, including a suggestion that Turkey Creek Junction would be a good type locality, and the fact that it is known that Mead found eggs and larvae of Euchloe ausonides coloradensis at Turkey Creek Junc- tion, the type locality of Euchloe ausonides coloradensis is hereby restricted to Turkey Creek Junction, Jefferson County, Colorado. Euchloe (Euchloe) creusa creusa (Doubleday) Citation of OEaGiNAL descriptions Doubleday, 1847, Gen. Di Lepid. 1:56, pi. 7, fig. 1. Type data: The original citation stated only that the type was collected in the Rocky Mountains of North America, there was no description, and the plate figure is a barely recognizable draw- S (3) : 185-195, 1966 NEARCTIC- EUCHLOE 193 ing. N, D. Riley of the British Museum of Natural History, m whose institution the holotype specimen is located, supplied the author with the necessary information concerning the type. The specimen bears two labels: l./“Rocky Mountains 45-135”, 2./ “Rhopalocera Palaearctica”, Interpbetation of type DATA: Riley (personal communica- tion) states that the first label which was mentioned above indicates that the specimen was presented in 1845 by the then Earl of Derby. F. M. Brown (personal communication) states that “the great majority of North American specimens collected by Lord Derby came from the vicinity of Banff, Alberta, I doubt that he got over to the B.C. side of the range and it is question- able that he got as far north as Kicking Horse Pass.” The second label shows that the specimen was figured by Verity (1908); the dorsal and the ventral surfaces of the type specimen are shown as figures 11 and 12 on plate 68. Taking into consideration the previously stated judgernen of Brown, as well as those of Barnes and McDunnough (1916) and McHenry (personal com- munication), and the fact that specimens readily assignable to the holotype have been collected near Banff, the type locality of Euchloe creusa Doubleday is hereby restricted to the vicinity of Banff, near 7,0(X)’ elevation, Alberta, Canada, Euchloe (Euchloe) hyantis hyantis (Edwards) Citation of original description: Edwards, 1871 Tram. Amer. Ent. Soc. 3:205-216. Type data: In the original citation it was stated that the type material was collected at Mendocino, California by R. H. Stretch. Specimens designated as types are located in the collection of the Carnegie Museum, Pittsburgh, Pennsylvania. H. K. Clench ( personal communication ) states that there is a specimen labeled as follows: '^Hyantis $ / type Gala” written in the hand of W, H. Edwards, the word “type” in red, the rest in black. A pin with an identical label except $ with a fragment of thorax on it is next to the supposed allotype. Interpbetahon of type data: “Mendocino, California” is now known as Mendocino City, Mendocino County, California. The significance of the red and the black ink on the type labels per- tains to the context of the time when they were written. Accord- ing to F. M. Brown the notations in black were probably written in 1889 or sometime later when Edwards shipped the specimens to Holland. The story of the word “type” written in red ink, according to Clench (personal communication), is that after Holland had purchased Edwards’ collection he removed the labels from the specimens that he considered to be types and 194 PAUL A. OPLER /. Res. Lepid. returned them to Edwards, who wrote the word “type” on them in red ink. He subsequently returned them to Holland who re- placed the labels on the appropriate specimens. It should also be noted that the original description stated “In the collection of Hy. Edwards”. Although this collection is under the care of the American Museum of Natural History, specimens designated as types of hyantis are not to be found in that institution. The above situation cannot be resolved at this time and a final decision of the proper types of hyantis must be held in abeyance. Euchloa (Euchloa) hyantis lotta Beutenmuller Citation of original description: Beutenmuller, 1898. BuU. Amer. Mus. Nat. Hist. 10(13): 243, pi. 14, fig. 4. Type data: Neither a type specimen nor a type locality was designated in the original paper. The distribution which was given as “Colorado, Arizona, Utah, Southern California”, and a drawing are sufficient to delineate, in the gross sense, the entity to which this name should apply. There is a specimen figured by Holland ( 1931 ) as figure 5 on plate 73 which is referred to by him in the text as “paratype” and on the plate legend as “typical”. The specimen is in the Carnegie Museum and bears the following labels: l./“P.— — — — , Arizona” (in pen, possibly in the hand of W. H. Edwards), 2./“lotta” (typewritten), 3./“Butterfly Book, Plate 73 Fig. 5” (the underlined numerals written in pen) ( Clench, personal communication ) . Interpretation of type data: The problem now arises whether the above specimen is a syntype loaned by Beutenmuller to Holland for illustration in his book as was done with “Theda ilavid" Beutenmuller (P. McHenry, personal communication), or if the specimen was selected by Holland from the W. H. Edwards material and was never seen by Beutenmuller. If the latter case were true the specimen is not an eligible candidate for a lecto- type. Thus, the situation of the absence of a suitable lectotype prevents one from restricting the type locality with any exactness. Euchfoe (Euchloe) hyantis andrewsi Martin Citation of original description: Martin, 1936. Bull. So. Calif. Acad. Sci. 35(2) :94“95. Type data: All of the type material was collected along the Crestline Highway, near Lake Arrowhead, San Bernardino Mountains, San Bernardino County, California between the ele- vations of 5000’ and 6000’ by R. H. Andrews- on the dates June 14-19, 1935 and June 8-18, 1936. The holotype, allotype, and 37 paratypes are located in the Los Angeles County Museum, Los Angeles, California. 5 (3) : 185-195, 1966 NEARCTIC EUCHLOE 195 Euchloe (Euchloe) ausonides mayi Chermock & Chermock Citation of original description: Chermock & Chermock, 1940. Can. Ent. 72(4):81-83. Type data: Holotype male, Riding Mountains, Manitoba, Canada, June 12, 1933, collected by Jack F. May. The specimen is in the collection of F.H. Chermock. The allotype female is located in the collection of the Carnegie Museum, Pittsburgh, Pennsylvania. The information of the accompanying labels of the allotype was noted for the author by H. K. Clench of the above institution: 1. /"Riding Mountains/ManitobaVII-31-33” locality in letterpress, date handwritten, 2./“ Euchloe ausonides/ var. mayi/ allotype / F. H. Chermock/ R. L. Chermock” the two authors names in script, the rest hand-lettered ,all in black ink on a hand bordered pink card. In the paper which included the original description the authors stated that 100 paratypes, all from the same locality but with various dates, were designated. Three paratypes are known to be located in the American Museum of Natural History, and one paratype is in the Carnegie Museum. Euchloe (Euchloe) olympia (Edwards) Citation of original description: Edwards, 1871. Trans. Ameri. Ent. Soc. 3:266-277. Type data: The holotype and allotype are located in the Carnegie Museum, Pittsburgh, Pennsylvania. The following in- formation was provided through the courtesy of H.K. Clench of the Carnegie Museum. Each specimen bears the label: "Olympia ^ [or $ ]/ type Kan ”. The word “type” is not in red ink. Interpretation of type data: The fact that the word "type” is not in red ink probably signifies that the word was placed on the labels at the time of the original description, and that these labels were not sent to Edwards by Holland at a much later date to have the word "type” appended in red ink (fide Clench, personal communication). The cryptic Kana refers to Kanawha according to Clench (personal communication), who states that "the name is born by a country, a river, and at least three com- munities or post offices have it as part of their name.” Klots ( 1951 ) cites the type locality as Coalburgh, West Virginia. F. M. Brown (personal communication) states that Edwards lived in Coalburgh on the Kanawha River and often collected there. Hence, to be precise and in agreement with the data on the type label, the type locality should be stated as "banks of the Kanawha River, near Coalburgh, West Virginia.” NEW INSECT NET FOLDS TO FIT IN POCKET OR BRIEF CASE An insect net which folds easily and quickly to a size which permits carrying in a coat pocket or brief case is offered by the BioQuip Divi- sion of Bio Metal Associates, P. O. Box 61, Santa Monica, California, 90406. Called the Collapsible Pocket Net, it is especially designed for inconspicuous use. The spring steel net ring folds to a 4" diameter, and flicks out quicklyto a IZ" Dacron aerial net. It may be refolded simply by twisting top of net ring and handle each 180® in opposite directions so that sections of ring fold into each other. The Pocket Net may be locked in closed position. Excellent for close-in collecting. The 5" aluminum handle may be extended by inserting a 5/8 " wooden dowel of desired length. Price of the Pocket Net is ^3. 50 each, ^ 35, 00 per dozen. Extra net bags are /l, 70 each, ^17. 00 per dozen. NEW STORAGE AND COLLECTION EQUIPMENT CATALOG ISSUED BY BIOQUIP Just off the press is the 1966-6? edition of their Specitenics and Field Equipment Catalog for the Natural Sciences, it was announced by BioQuip Division of Bio Metal Associates, Santa Monica, California. The 28-page illustrated catalog gives detailed descriptions and prices on the complete line offered by the firm, manufacturers since 1947 of quality storage and field equipment. Included is a variety of entomological instruments, as well as specially designed vials, trays, drawers, boxes and cabinets for housing and systematizing specimen collections. Collecting equipment covers nets, killing jars, pinning boards, forceps, etc. The catalog is divided into sections for easy reference: Botony, Entomology, Vertebrate Zoology, Geology- Paleontology, Ecology, Microscopy and Aquaria. There is no charge for the catalog. 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Stegner 129 Vital Staining of Colias philodice and C. eurytheme John M. Kolyer 137 The North American Species of the Genus Zeiraphera A. Mutuura and T, N. Freeman 153 Notes on Fades penelope (Saturniidae) Brian O. C. Gardiner 177 Remarks on The Genus Zera Evans in Mexico with a New Record, H. A. Freeman 181 Studies on the Nearctic Euchloe Part 3. Complete Synonymical Treatment 185 Part 4. Type Data and Type Locality Restrictions 190 Paul A. Opler THE JOURHA.L December, lb66 Volume 5 Number 4 THE J©URMJAL OF FESIAKCH OM THE LEFIOOPTERA a quarterly published at 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HOVANITZ THE PURPOSE OF THE JOURNAL is to combine in one source the work in this field for the aid of students of this group of inseas in a way not at present available. THE JOURNAL will attempt to publish primarily only critical and complete papers of an analytical nature, though there will be a limited section devoted to shorter papers and notes. 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Second Class Postage Paid at Arcadia, Calif. Journal of Research on the Lepkioptera 5 (4): 197-208, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A NEW SPECIES OF ONCOCNEMIS FROM THE WESTERN UNITED STATES ( NOCTUIDAE : CUCULLIIN AE ) JOHN S. BUCKETT AND WILLIAM R. BAUER University of California, Davis, and State Department of Agriculture, Sacramento For many years the authors have possessed a series of an Oncocnemis species that was believed to be new. Until recently, we were under the opinion that this new species was probably most closely related to O. hayesi Grote, as it superficially resem- bled that species more than it did any other species within the genus. Much to our surprise, when genitalic mounts of both sexes had been prepared and critically examined, it became ap- parent that the superficial resemblance to hayesi had been mis- leading. With further examination of other speices in this sec- tion of the genus, it was found that O. melantho Smith, described from specimens collected in the Yosemite Valley, California, was extremely closely related to O. sandaraca Buckett and Bauer, herein described as new. Both O. melantho and O. sandaraca occur sympatriacally at various localities in the central Sierra Nevada, and the adults rely, in part at least, on the same species of Rabbit brush ( Chry- sothamnus spp.) as a food source. Both species occur where there is floral influence of the Great Basin type. It is also in- teresting to note that while both species occur diurnally on flowers, only sandaraca has been taken noctumally. The fre- quency of sandaraca occurring during diurnal hours is much less than that for melantho. Oncoenemis sandaraca Buckett and Bauer, new species Male: Ground color of primaries dorsally yellowish-red (thus the name sandaraca, from Latin), transverse lines and marks conspicuous; secondaries dorsally a pale yellow with black ex- terior band. Head with vertex clothed in bicolorous elongate flattened dentate hairs, basally tan, apically brown; between vertex and frons a dark brown transverse band is evident; frons clothed in tan simple hairs; palpi exterolaterally clothed pre- 197 198 BUCKETT AND BAUER J. Res. Lepid. Fig. 1. Distribution map of Oncocnemis sandaraca Biickett and Bauei showing known distribution. 5 (4): 197-208, 1966 NEW ONCOCNEMIS 199 Fig. 2. Holotype male, O. sandaraca, dorsal view. Johnsville, Plumas ® September 1959, ex. fluorescent black light ( W. R. Bauer & J. S. Buckett ) . Fig. 3. Allotype female, O. sandaraca, dorsal view. Same locality and collectors as in fig. 2, 8 September 1960. 200 BUCKETT AND BAUER /. Res. Lepid. Fig. 4. Male, O. melantho Smith, dorsal view. Road to Spencer Lakes, 8 miles southwest of Johnsville, Plumas County, California, 10 September 1961 (W. R. Bauer & J. S. Buckett). Fig. 5. Female, O. melantho, dorsal view. 13 miles northeast of Garden Valley, Boise County, Idaho, 27 August 1965 (W. R. Bauer, J. S. Buckett & M. R. Gardner). 5 (4): 197-208, 1966 NEW ONCOCNEMIS 201 Fig. 6. Holotype male, O. sandaraca, ventral view. Data same as for fig. 2. Fig. 7. Female, O. melantho, ventral view. Data same as for fig. 5. 202 BUCKETT AND BAUER J. Res. Lepid. Fi^. 8. .Paratype male, O. sandaraca, genitalia minus adeagiis. Johnsville, Plumas County, California, 10 Oetober 1966 (H. J. Pini), Bauer- Buckett Slide No. 66L19-3. Fig. 9. Paratype male, O. sandaraca, aedeagus. Data same as for fig. 8. 5(4): 197-208, 1966 NEW ONCOCNEMIS 203 dominently in tan simple scales, but a few brown scales present also, ventrally clothed in elongate tan and brown simple hairs; terminal segment of palpi short, clothed in brown scales; com- pound eyes weakly fringed dorsally and posteriorly with tan tipped brown simple hairs or “lashes”; antennae with scape and pedicle clothed in tan scales; flagellomeres brown, dorsally clothed weakly in tan-brown scales, ventrally weakly ciliate, ciliations becoming more pronounced apically. Thorax with collar tricolor, composed of elongate scales, basally and apically yellowish-red, medially brown so as to form a transverse band; tegulae clothed in white, tan, brown, and dark brown simple hairs and elongate scales, appearing most lightly colored inward- ly (border near disc); as in tegulae except possessing a greater percentage of simple hairs vs. elongate scales; posterior tuft composed of tan colored elongate scales; ventrally clothed in light tan simple, elongate hairs; legs with femora dorsally clothed in tan and brown scales and hairs, ventrally clothed in elongate ochreous and tan simple hairs; tibiae with tibial claw pronounced; meso and meta tibiae dorso-basally clothed with a bunch of elongate tan simple hairs, dorso-apically clothed in mixture of tan and dark brown scales; tarsi predominently clothed in black scales, but each segment possessing an apical annulus of tan scales; primaries dorsally of ground color; costal subcostal and radial veins clothed in ochreous except where transverse lines intersect costa; basal one-half line geminate on costa, basally brown, apically ochreous; basal and transverse anterior areas contiguous, clothed in reddish-yellow scales; trans- verse anterior line geminate, basally ochreous, apically brown, strongly marked costally, thence undulating to inner margin; median area with conspicuous brown transverse shade, strongly marked costally, also dark brown scaling present on Cua and 2dA veins; orbicular round, outlined in dark brown, thence ochreous, centrally overlain with brownish scales; reniform trapazoidal, broadest apically, narrowest basally; transverse posterior line geminate on costa, dark brown, centrally filled with ochreous, thence a single brown undulating line separating dark median area from yellowish-red portion of subterminal area; subterminal area basally more ochreous in area of radial and medial veins and more yellowish-red in area of cubital veins, apically brown to subterminal line, veins outlined in dark brown; subterminal line an ochreous transverse shade from costa to Mi, thence in- terrupted on veins to inner margin; terminal aera checkered yellowish-red and dark brown, the darker color appearing on veins; terminal line brown, faint; fringes tricolor, basally ochre- 204 BUCKETT AND BAUER J. Res. Lepid. f Fig. 10. Male, O. melantho, genitalia minus aedeagus. Road to Spencer f Lakes, 8 miles southwest of Johnsville, Plumas County, California, 7 September 1960 (W. R. Bauer & J . S. Bucket! ), Bauer-Buckett i; Slide No. 66L16-3. I Fig. 11. Male, O. melantho, aedeagus. Data same as for fig. 10. j 5(4): 197-208, 1966 NEW ONCOCNEMIS 205 ous, medially dark brown, apically light brown; ventral surface ochreous basally to just preceding subterminal line, clothed in ochreous simple, elongate hairs; transverse posterior line rep- resented costally as dark brown dash, thence only a suggestion of it appearing on medial veins; exterior border dark brown; fringes appearing tricolor, basally ochreous, somewhat darker between veins, medially dark brown, apically tan; secondaries dorsally with basal two-thirds ochreous, basal one-third scantily irrorated with brown scales, this irroration appearing concealed due to additional clothing of elongate simple ochreous hairs; faint brown scalation present just preceding exterior band on Cui, Cua, Ml, and Mg; apical one-third of wing a dark brown band, the veins contained therein outlined in near black scales; fringes tricolor, basally ochreous, medially dark brown, apically creamy- white; ventral surface ochreous for basal three-fourths, apically with a broad brown band (otherwise marked as in dorsal sur- face); discal lunule faint; fringes lighter than dorsal surface, mostly ochreous, but with brown medially. Abdomen dorsally clothed in brown and ochreous scales, intermixed with sparsity of light tan simple hairs; apically clothed in admixture of brown and ochreous elongate flattened and simple hairs; ventrally clothed in tan simple scales and simple hairs. Greatest expanse of forewing 16 mm. Genitalia as in figures 8 and 9. Female: More somber in coloration than in male, lacks as much reddish coloration as in male; primaries with more brown coloration; basal one-third of secondaries darker than in male, as though a dirty brownish coloration; ventral surface of pri- maries with brown in the tan basal three-fourths of surface, both median shade and transverse posterior line represented in dark brown costally, otherwise surface as in male; secondaries ven- trally as in male, but with less yellow coloration, so as to appear “washed out.” Greatest expanse of forewing 16 mm. Genitalia as in fig. 12. Holotype in collection of Entomology type collection. University of California, Davis, California. Specimens examined Holotype male: Johnsville, Plumas County, California, 6 September 1959, ex. 15 watt fluorescent black light (W. R. Bauer and J. S. Bucket! ). Para- types: 64 males and 33 females; 1 female (designated Allotype) same locality and collectors as for Holotype, 8 September 1960; 1 male, same locality as holotype, 21 September 1962 (H. J. Pini), Bauer-Buckett Slide No. 66L16-1; 1 male, same locality as preceding, 10 October 1966 (H. J. P. ), Bauer-Buckett Slide No. 66L19-3; 1 male, same locality as preceding, 3 October 1966 (H. J. P. ), Bauer-Buckett Slide No. 66L19-1; 1 female, same locality as holotype, 15 September 1962 (H. J. P. ), Bauer-Buckett I Slide No. 66L16-2; 1 female, same locality as preceding, 8 October 1966 i (H. J. P. ), Bauer-Buckett Slide No, 66L19-2; 1 female, same locality as I preceding, 3 October 1966 (H. J. P. ), Bauer-Buckett Slide No. 66L19-4; I i I 206 BUCKETT AND BAUER J. Res. Lepid. Fig. 12. Paratype female genitalia, O. sandaraca. Johnsville, Plumas County, f California, 15 September 1962 (H. T. Pini), Bauer-Buckett Slide jl I No. 66L16-2. ■ li 5 (4): 197-208, 1966 NEW ONCOCNEMIS 207 Fig, 13. Female genitalia, O. melantho. Mt. Ingalls, Plumas County, Cali- fornia, 12 September 1961 (W. R. Bauer & J. S. Bucket! ), Bauer- Bucket! Slide No. 66L16-4, 208 BUCKETT AND BAUER J. Res. Lepid. 45 males, 23 females, Johnsville, 2 September - 5 November 1959 - 1966 (H. J. P., W. R. B., J. S. B., and M. R. Gardner); 1 male, Road to Spen- cer Lakes, 8 miles s.w. Johnsville, Plumas County, California, 24 August 1961 (W. R. B. & J„ S. B.); 5 males, 2 females, same locality as pre- ceding, 10 September 1961 (W. R. B. & J. S. B.); 1 male, same locality as preceding, 9 September 1966 (J. S. B., M. R., R. C., J. L. & B. W. Gardner); 1 male, 1 female, same locality as preceding, 26 September 1959 (H. J. P. ); 1 male. Nelson Creek, Plumas County, California, 17 September 1940 (W. R. B.); 1 male, same locality and collector as pre- ceding, 2 September 1940; 1 male, 1 female. Monitor Pass, Alpine County, California, 8 September 1964 (W. R. B. & J. S. B.); 1 male, same locality and collectors as preceding, 9 September 1964; 3 males, Woodfords, Al- pine County, California, 8 October - 21 October 1962; 1 male, Grizzly Meadows, Trinity County, California, 10 September 1964 (G .E. Buxton) 1 female, 6 miles n.w. Cedarville, Modoc County, California, 8 September 1963 (W. R. B. & J. S. B.); 1 female, Tulelake Inspection Station, Modoc County, California, 22 September 1965; 1 male, 2 females, 3 miles n. Toll- gate, Umatilla Co., Oregon, 6 September 1963 (W. R. B. & J. S. B.). Oncocnemis sandaraca can readily be distinguished from O. melantho by the great differences in color and lesser differences in maculation. O. sandaraca possesses yellowish-red to ochreous- brown primaries dorsally; whereas, melantho possesses greyish to greyish-olive primaries dorsally. The ventral surface of the pri- maries in sandaraca is as described, but in melantho, the lighter portion is off white and not at all ochreous, or possessing any j yellowish color; also melantho may have a black transverse med- ial band (corresponding to the median shade of the dorsal sur- ' face) either strongly or weakly expressed; the exterior band in melantho is considerably broader than that of sandaraca. The secondaries of each speices are distinctly different interspecifi- cally. O. sandaraca possesses secondaries as described; whereas, dorsally melantho possesses a dark basal one-third, a white, or brown medial one-third and a broad black exterior band (as in figures 4 and 5); the fringes are noticebly more whitish than , in sandaraca. The ventral surface of the secondaries in melantho are also quite different than for sandaraca: dorsally they are with basal and medial transverse areas white, or irrorated with black scales; two areas are separated by a black band; just pre- ceding broad exterior band there is another faint balck line which parallels the inner margin of this broad black band. This exterior ’ band in melantho is much broader than the exterior band in sandaraca, and melantho possesses much whiter fringes in com^ parison to sandaraca. Nothing is yet known concerning the immature stages of either species, but it would not be all too surprising to discovel that larvae of both species feed on the same plant species. The ' genitalic slides were prepared using lignin pink stain and balsam as the mounting media. , Journal of Research on the Lepidoptera 5 (4): 209--214, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 SPECIATION IN THE AGATHYMUS ( Megathymidae ) H. A. FREEMAN' 1605 Lewis Drive, Garland, Texas I CAN REMEMBER BACK in the past when we used to collect but- terflies with a net and killing jar and then there were fifteen recognized species in two genera in the family Megathymidae. Today we have 47 species in five genera in that family and the net has long since been retired after we discovered that it was much more efficient to collect these “critters” with a fox hole pick, drain spade or sharp knife. Once when we had available a few hundred specimens of these insects the taxonomy seemed comparatively simple, and now that we have several thousand specimens available for study complications present themselves. In making a study of speciation in the Agathymus we must recognize a number of species complexes that exist in that genus. They are the neumoegeni, chisosensis, hoffmanni, evansi, aryxna, baueri, alliae, stephensi, polingi, mariae, remingtoni, rethon, and indecisa complexes. Of these the following are very compli- cated and indicate a considerable amount of evolution is in progress: neumoegeni, hoffmanni, aryxna, mariae, and reming- toni, and it is with this group that I would like to go into with some detail. Agathymus speciation particularly in the mariae complex has iptrigued me for a number of reasons, one of which has been the method by which new species apparently developed during the past ages from a mariae-like prototype. In making a de- tailed study of this species complex in the United States a num- ber of very interesting observations were encountered some of which I would like to discuss briefly. One of the first things that was noticed about mariae was its close association with its lar- val food plant Agave Lecheguilla Torr. During the number of ^All my research on the Megathymidae during the past six years was made possible by the National Science Foundation through research grants G-9900 and GB-398 for which I am deeply grateful. 209 210 H. A. FREEMAN }. Rc.s. Lepid. years that I have collected mariae and other related species I have never found a specimen in any other species of Agave other than members of the lechegiiilla complex, with the possible ex- ception of a Mexican species that feeds in Agave falcata Engelm. Indications are that at one time lechegiiilla occurred extensively from the Edwards Plateau in Texas over into New Mexico and well down into Mexico. Fossil seeds have been found in areas of Texas now completely devoid of these plants. In this vast expanse a prototype of our modern species was well established with a gene pool common to all areas. Conditions arose result- ing in the eradication of lechegiiilla plants in many sections of this great area thus disrupting the gene pool. During the many years that followed mutations occurred in the restricted popu- lations resulting in the production of individuals that differed from the original species. These isolated populations gradually changed both biologically and morphologically into five separate species in the Texas area. Mariae was the first described in this complex and it was found at El Paso, in the Franklin Moun- tains, and for many years was considered to be the only species present all over southwestern Texas. After carefully studying specimens from 32 locations in Texas and New Mexico it was found that there was actually five different species going under the name of mariae. Chinatiensis Freeman, in the Chinati Moun- tains and near vicinity; lajitaemis Freeman, in the Fajita region of the Big Bend; rindgei Freeman, and gilherti Freeman, in the general vicinity of Del Rio; and mariae (B. & B.), from the Big Bend and McCamey regions westward and north to Carlsbad, New Mexico and El Paso, Texas. In no area do more than two of these species occur together and when this happens it was found that they ha\'e changed somewhat their larval feeding habits. In the Boquillas Canyon, Dryden, and Langtry areas where gilberti and mariae fly together, gilberfi larvae penetrate into the caudex of the plant to a depth of 20 mm, whereas mariae does not penetrate more than 5 mm into the caudex, but does most of its feeding in the lower portion of the leaves. In the Del Rio, Juno, and Bracketville areas where gilberti and rindgei fly together gilberti penetrates sometimes to a depth of 30 mm into the caudex, whereas rindgei seldom penetrates below 20 mm. It is not unusual to find these two species and estelleae (Stall- ings & Turner) feeding in the same plant, each with a somewhat different feeding level, with estelleae feeding only in the leaves, rindgei the base of the leaves and upper caudex, and gilberti in the lower base of the leaves and deeper into the caudex. A rea- 5 (4): 209-214, 1966 SPECIATION IN AGATHYMUS 211 sonable explanation for this is the rapidly declining population of lecheguilla plants in this general area, thus resulting in greater competition for food among these insects. The pH of the soil has considerable bearing on the occurrence of these species in a given habitat. Mariae is associated with distinctly alkaline soil, often from 7.7-8 pH; chinatiensis and lajitaensis, with soil somewhat less alkaline, around pH 7.2-7.4; and gilberti and rind- gei, in soil that is neutral or only slightly alkaline, pH 7-7.1. An interesting observation was made concerning the feeding habits of the larvae of Aegiale hesperiaris (Walker) and Aga- thymus hoffmanni (Freeman) in Agave americana L. in Mexico. In areas where hoffmanni is fairly abundant, especially in the valley of Mexico, the larvae feed in a small species of a parryi- like Agave, and overlook the larger americana plants, in which the larvae of hesperiaris are found. The hoffmanni trap doors are located on the lower surface and usually near the center of the leaf. In areas where both species are found in the same plant, some member of the americana complex, hoffmanni pre- fer the hip of the leaf for their feeding area leaving the center portion for the much larger hesperiaris larvae, or else the hes- periaris forced them into that position. Specimens of hoffmanni from these areas are atypical, indicating that they have evolved or else are in the process of evolving into a separate species. Both biological and morphological work on these specimens is difficult due to the scarcity of the individuals. When sufficient material is present for study it may then be possible to determine the exact status of these hoffmanni-\ike specimens. In Arizona some parallel evolution appears to be present in various populations of Agathijmus aryxna (Dyar). In the gen- eral area of the Chiricahua Mountains southward through Ram- sey Canyon or into northern Mexico is found one population with similar characteristics. Another population begins near Globe and extends through Santa Catalina, Rincon, and Santa Rita Mountains southward through the Patagonia Mountains into Sonora, Mexico. The third is a rather variable population that occurs in the Baboquivari Mountains and possibly on south- ward into Mexico. Indications are that at one time this entire area was covered with a palmeri-like Agave in which fed an aryxna-like prototype of our modern species. Environmental con- ditions plus man made factors eliminated the Agave in large areas thus producing desert and mountain islands where the plants are still undisturbed. Individuals from each population may be recognized by the width of the spots on both wings. 212 H. A. FREEMAN /. Res. Lepid. Starting with the eastern population, the spots are generally separate and fairly small. In the central population, the spots are somewhat larger and tend to be closer together. While in the western population, the spots are the largest, tending to fuse together in some cases. To me there seems no doubt that we are seeing evolved three separate subspecies that eventually will assume specific status within a number of years. In the neumoegeni comolex we have a number of so called species that exhibit characteristics indicating a common ancestor. Possibly some of these have evolved farther than others and are specific, however others may still be in the process of changing from distinct subspecies into full species. The test of breeding has just started to be worked upon by Stallings, Turner & Stall- ings and the results will likely be very conclusive in forming our conception of this complex. I have collected larvae of wild hybrids of a cross between male mariae and female judithae (Stallings & Turner), as well as the same in a cross between male mariae and female diabloensis Freeman and the resulting Fi did not resemble each other as much as the pure judithae | and diabloensis do, especially the females. In all of the wild hybrids that I have seen to date involving mariae with some species of the neumoegeni complex it has always been the males of mariae that have resulted in the hybrids. This is based on the presence of the larvae in species of Agave other than mem- bers of the lecheguilla complex. From all evidence presented these Fi are sterile. Two members of the neumoegeni complex are associated with juvenile plants, neumoegeni (Edwards) and florenceae (Stall- ings & Turner). All other species are associated with mature plants running from small to large in size. Florenceae and neu- moegeni morphologically resemble each other more closely than do any of the other species in this group even though there is a great distance separating them in their distribution. Genetically 1 speaking the entire complex is closely related. In the remingtoni complex we have three described species, remingtoni (Stallings & Turner), estelleae (Stallings & Turner) and comstocki (Harbison), and several undescribed species. Genetically they are related, biologically they are very different. Remingtoni is found in the mountains in very rugged terrain, often on cliffs, with the pH on the acid side, and feeding upon a member of the lecheguilla complex. Estelleae is associated with alkaline or near neutral soil, usually in the plains country or else 5 (4): 209-214, 1966 SPECIATION IN AGATHYMUS 213 where it is slightly hilly, and feeding upon another member of the lecheguilla complex. Comstocki is found in hilly, desert, areas of Baja California (pH has not been checked), feeding upon Agave shawii Engelm., which is not at all related to leche- guilla. The general range of remingtoni is Victoria, Tamps., Mexico southward to the Jacala, Hidalgo, area, while estelleae occurs north and west of Victoria. All indications point to a common ancestor of these two species that occurred in an area where the gene pool was available for all regions of the range. Due to environmental changes, both man made and natural, the gene pool was disrupted, eventually resulting in the formation of separate subspecies and eventually species. In conclusion I would say that in the genus Agathymus we have had three major prototype complexes, the neumoegeni, aryxna, and mariae, and from these three all of our present day species evolved. REFERENCES COMSTOCK, JOHN A., 1957. Notes on the metamorphosis of an Agave boring butterfly from Baja California, Mexico. Trans. San Diego Soc. Nat. Hist. 12: 263-276, pi. 22. EDWARDS, W. H., 1882. Description of species of butterflies taken in Arizona by Jacob Doll, 1881. Papilio 2: 19-29. FREEMAN, H. A., 1950. Notes on Megathymus, with the description of a new species. Field & Lah. 18: 144-146. 1951a. Notes on the Agave feeders of the genus Megathymus. Field & Lah. 19: 26-32. 1951b. Ecological and systematic study of the Hesperioidea of Texas. So. Methodist Univ. Studies, no. 6:1-64. 1952. Two new species of Megathymus. Amer. Mus. Novi- tates, no. 1593:1-9, figs. 1-13. 1955. Four new species of Megathymus. Amer. Mus. Novi- tates, no. 1711: 1-20, figs. 1-34. 1958. A revision of the genera of the Megathymidae, with the description of three new genera. Lepid. News. 12: 81-92, 1 pi. 1960. Notes on Agathymus in Texas, and the description of a new species from Mexico (Megathymidae). Journ. Lepid. Soc. 14: 58-62. 1962. A new species of Agathymus from Texas. Amer. Mus. Novitates, no. 2097: 1-7, figs. 1-6. 1963. Type localities of the Megathymidae. J. Res. Lep. 2 (2): 137-141. 214 H. A. FREEMAN J. Res. Lepid. 1964a. The effects of pH on the distribution of the Megathy- midae. /, Res. Lep. 3(1): 1-4. 1964b. Four new species of Agathymus from Texas. Journ. Lepid. Soc. 18: 171-185, 4 pis. 1964c. Larval habits of Agathymus mariae ( B. & B. ) . /. Res. Lep. 3 (3): 145-147. HARBISON, C. F., 1957. A new species of Megathymus from Baja Cali- fornia, Mexico. Trans. San Diego Soc. Nat. Hist., 12: 231-262, pis. 18-21. 1963. A second new species of megathymid from Baja Cali- fornia, Mexico. Trans. San Diego Soc. Nat. Hist., 13: 61-71, 4 pis. STALLINGS, DON B. & J. R. TURNER, 1954. Notes on Megathymus neumoegeni with description of a new species. Lepid. News. 8: 77-87. 1957. Four new species of Megathymus. Ent. News. 68: 1-17. 1958. A revision of the Megathymidae of Mexico, with a synopsis of the classification of the family. Lepid. News. 11; 113-137, 8 pis. 1960. A new species of Agathymus and a new subspecies of Megathymus. Ent. News. 71: 109-115 STALLINGS, DON B., J. R. TURNER & VIOLA N. STALLINGS, 1961. A new subspecies of Agathymus mariae from Mexico. Journ. Lepid. Soc. 15: 19-22. Journal of Research on the Lepkloptera 5 (4): 215-219, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE EGGS AND FIRST INSTAR LARVAE OF THREE GALIFORNIA MOTHS JOHN ADAMS COMSTOCK Del Mar, California Research Associate In Entomology of the Los Angeles County Museum of Natural History In rearing lepidoptera from captured females of identified species it occurs not infrequently that foodplants and early stages have not previously been recorded, and the rearing ends with the newly emerged laiA^ae because of their refusal to accept experimentally offered plants. It is justifiable to publish such incomplete records for the purpose of encouraging other entomologists, located in areas of varying plant associations to carry on the experiments with plants not available to the original recorder. Three examples of such incomplete records are here included: Scotogramma defessa Grote Eggs were secured from a gravid female during the second week in April, 1962, at Del Mar, California. They hatched April 20, ’62. EGG: (Figure 1, No. 1), hemispherical, the base flattened and the top evenly rounded. It is topped by a minute round micro- pyle with a raised center and a surrounding depressed circlet. The ground color is a light straw. The basal diameter is approxi- mately 0.65 mm. and the height 0.4 mm. The surface is covered by 48 vertical ribs, many of which coalesce as they approach the curved top. From 12 to 15 of these extend across the de- pressed circlet and about on the micropyle. The raised vertical ribs are topped by minute glistening pearly- white nodules. Higher magnification discloses low horizontal ridges or lines running across the troughs between the vertical ridges. The illustration shows the egg on lateral aspect, tipped slightly forward to show the micropyle. 215 216 J. A. COMSTOCK /. Res. Lepid. FIGURE 1 No. 1. Egg of Scotogramma defessa, lateral aspect, X 50. No. 2. First instar larva of S. defessa, X 45. No. 3. Egg of Camptogramma neomexicana X approximately 70. No. 4. First instar larva of C. neomexicana X approx. 38. 5(4): 215-219, 1966 LARVAE OF MOTHS 217 FIRST INSTAR LARVA: (Figure 1, No. 2.), Body, cylindical, translucent and white. Length, 2. to 2.25 mm. The head is yellow, and wider than the first segment. Numerous setae are scattered over the body. These are black, and arise from black papillae. Those on the thoracic segments are in line transversely across each segment. On the typical abdominal segments they alternate in a zig-zag pattern, giving the impression of two transverse rows running along each segment. There are two pairs of prolegs in addition to the anal pair. No food plant notes were available. I tried lettuce, clover, parsley, sage, yarrow, oak, willow, Ceanothus sp., Eriogonum sp., Adenostoma sp., Artemisia sp., Oenothera sp., Mimulus sp., and Rhus of three separate species, without avail. Campto gramma neomexicana Hulst A captive female laid numerous eggs April 24, 1962. They were deposited singly on their sides. EGG: (Figure 1, No. 3.), oval; ground color a delicate pink, but appearing nearly white because of the reticulation of raised white walls outlining hexagonal cells. Length of egg, 0.75 mm. Width, 0.4 mm. Just before hatching the eggs lose some of their pink tinge and become spotted with light chrome. Hatching oc- curred May 3, 1962. FIRST INSTAR LARVA: (Figure 1, No. 4.), Length, 2.5 mm. Head width approximately 4.35 mm., broader than first segment and somewhat flattened; dull yellow with a darker tinge on the outer edges of cheeks and many bulging prominent black ocelli. Apparently there are six on each cheek, but they are somewhat confused with numerous black spots in the near vicinity. These spots occur along the frontal and adfrontal sutures and on the sides of the cheeks. The body color is yellow. From the 4th to about the 10th seg- ment there is a longitudinal middorsal stripe, discontinuous in some examples and more conspicuous in others. This line has a darker shading on the segmental junctures. There are raised black spots or papillae running in line longitudinally on the spiracular area, one on each segmental juncture. The legs are yellow as are also the anal prolegs and single pair of prolegs. I The setae are predominantly white. The young larvae were tested on honeysuckle, oak, willow, bur clover, chamise, sage, toyon, pine, Cyprus, and Rhus of three species, all without success. 1 218 J. A. COMSTOCK /. Res. Lepid. FIGURE 2 No. 1. Egg of Sphinx vashti X 45. No. 2. First instar larva of S. vashti X 15. ! il / 5 (4): 215-219, 1966 LARVAE OF MOTHS 219 THE EGG AND FIRST LARVAL INSTAR OF Sphinx vashti Strecker Eggs of the rare hawk moth, Sphinx vashti, were sent me September 20, 1955 by Frank Sala. They were obtained Septem- ber 13, '55 from a confined female collected at Wheaton Springs, Mescal Range, San Bernardino County, California. EGG: (Figure 2, No. 1), length 1.8 mm., by 1.2 mm. tall. Color, a delicate green. The surface is smooth, and there is apparently no reticulated pattern or network of cell impressions. The eggs hatched September 21, 1955. FIRST INSTAR LARVA: (Figure 2, No. 2), length, 5 mm., not including the caudal horn. The latter measures 2.55 mm. Head: larger than first segment; translucent light green in- cluding the ocelli and all appendages. Later the ocelli and tips of the mandibles show a tinge of gray. Body: cylindrical, tapering gradually toward the cauda, but slightly expanded at the 11th segment where there is a large caudal horn. This is recurved caudally, and ends in a bifurcated tip. It is brownish-black in color. The legs and prolegs are concolorous with the light green body. I was unable to find a food plant that the larvae would accept, and they aU died shortly after the accompanying notes and il- lustrations were made. AN ADDITIONAL FOOD PLANT RECORD FOR Papilio thoas autocles R. & J, JOHN ADAMS COMSTOCK Del Mar, California In Holland’s revised edition of the “Butterfly Book”, 1931, p. 317, he lists the genera Ptelea, Xanthoxylon, and the various species of Citrus which have been introduced into America, as the food plants of Papilio thoas autocles. As a result of rearing this butterfly in Puerto Vallarta, Mex- ico, during the summer of 1957, as recorded in “Estudios de los Ciclos Biologicos en Lepidopteros Mexicanos”, authored in co- operation with Dr. Leonila Vazquez Garcia, and run in the Anales del Institute de Biologia, Vol. 31, 1961, we were able to add the genus Piper to the list of host plants. An additional food plant for this butterfly was collected at the time, but proved puzzling to the cooperating botanists. This month (August, 1963) I received word from Dr. Reid Moran, botanist of the San Diego Museum of Natural History that a determination of the unknown plant has finally been made. He states that “it is Monnieria trifolia L., very rarely collected in Mexico and overlooked in the North American Flora; also omitted from Trees and Shrubs of Mexico, though perhaps be- cause not considered a shrub. Accordingly, we can now list Monnieria trifolia L., as another food plant of Papilio thoas autocles. 220 Journal of Research on the Lepidoptera 5 (4): 221-228, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A NEW SPECIES OF FOLIA OCHSENHEIMER FROM CALIFORNIA AND NOTES ON FOLIA DISCALIS (GROTE) ( NOCTUIDAE : HADENINAE ) JOHN S. BUCKETT and WILLIAM R. BAUER University of California, Davis and State Dept, of Agriculture, Sacramento, California PoLiA piNiAE Buckett AND Bauer, new species, has long been masquerading under the name P. discalis ( Grote ) , and until recently this error has gone unnoticed. While working over this section of the genus Folia, the authors recognized a confused situation, and with concentrated effort and further research, it was agreed that there were at least two species under the name discalis. We also have a large series of P. discalis from the eastern Sierra Nevada of California which is perhaps of subspecific rank, but at the present time it is felt best to retain this series as merely a light colored form of discalis. POLIA PINIAE Buckett and Bauer, new species Male: Ground color of primaries a whitish grey, transverse lines hardly dis- cernable. Head with vertex and frons clothed predominently in whitish flattened hairs, but with some dark brown flattened hairs intermingled; frons smooth, broadly truncately rounded; palpi clothed exterolaterally with whitish flattened hairs, but predominently with brown and dark brown : elongate, flattened hairs; antennae appearing dorso-ventrally bicolor under lOX power, with scape and pedicle clothed in brown and white intermixed elongate scales; flagellomeres with small fasciculate hairs ventrally, dor- sally clothed in whitish and brownish scales; compound eyes moderately to heavily haired; latrad of eyes, a buch of black fine hairs forms a dense cluster. Thorax with collar composed of flattened hairs and elongate scales, basally whitish, medially black ( so as to form a conspicuous transverse bar ) , apically white tipped; dorsally clothed in silvery-whitish elongate scales, ) some brown elongate scales intermixed also; anterior tuft composed of predominently dark brown flattened, dentate scales, some whitish scales of same type intermixed; posteriorly from anterior tuft a thin brown line I of flattened hairs leads to suggestion of diminutive divided posterior tuft; 1 patagia exterolaterally composed of dark brown fine silken hairs; legs with I pro and meso femora dorsally clothed in brown, metafemora clothed dor- ! sally in dirty whitish colored scales; all legs with tibiae clothed in mixture : of whitish and brown scales; tarsi clothed dorsobasally in dark brown 221 222 J. S. BUCKETT /. Res. Lepid. Fig. 1. Distribution map indicating the presently known distribution of Polia piniae Buckett and Bauer. 5(4): 221-228, 1966 NEW FOLIA 223 scales, dorso-apically and ventrally clothed in whitish scales; ungues mod- erately interolaterally bifid; primaries dorsally clothed in ground color, sparsely irrorated with fuscous scales (which gives a light ashey grey, or greyish white appearance); basal line present as thin black apically pointed “V” on costa, thence wanting, or hardly discernable; basal and transverse anterior areas of ground color, contiguous; transverse anterior line present on costa as is basal line, but broader and geminate costally, thence dimin- ishing into a faint transverse line to inner margin, hardly discernable; median area with very slightly darker shade than ground color; orbicular very broad, subquadrangular, open on costal margin, outlined faintly in black, filled with ground color; area between orbicular and reniform ir- rorated with blackish scales, less than width of orbicular; reniform large, color as in orbicular, nearly contiguous with orbicular on Cuj; transverse posterior line represented as dark shade on costa, thence almost entirely obliterated; subterminal space of ground color; subterminal line strongly represented costally, opposite of discal area and in tornus area (as in figure 2); tornus area with blackish bilobed mark, lobes terminally directed; terminal space of ground color; terminal line faint, represented by blackish lunules between veins; fringes basally whitish, thence of intermixed brown and whitish spatulate scales; ventral surface with costal edge and outer margin silvery-grey; central portion of wing fuscous; veins outlined in deep smokey fuscous; transverse posterior line represented costally as dark dash; secondaries dorsally light ochreous brown, basally and ter- minally darker than lighter median band; veins outlined in smokey; discal lunule faint, dark brown; terminal line dark brown; fringes tricolor, basally ochreous, medially brown, apically white tipped; ventral surface whitish, veins faintly outlined in fuscous; discal lunule faint; remainder of surface as in dorsal surface. Abdomen dorsally whitish; mid dorsal hairs blackish, almost appearing tuft-like; posteriorly whitish hairs and scales intermixed with dark brown; ventrally clothed in whitish hairs. Greatest expanse of forewing 25 mm. Genitalia as in figures 6 and 9. Female: As in male except antennae lacking fasciculations as in male, but rather possessing fine ciliations; primaries and secondaries slightly darker than in male. Abdomen as in male except slightly darker dorsally. Greatest expanse of forewing 26 mm. Genitalia as in figure 4. SPECIMENS EXAMINED HOLOTYPE male: Johnsville, Plumas Gounty, California, 5 July 1962 (Helena J. Pini). Paratypes; all specimens from California unless otherwise stated; 175 males, 39 females; 1 female (designated Allotype), same locality as Holotype, 3 July 1959 (W. R. Bauer and J. S. Bucket! ); 1 male, Johns- : mile, Plumas Co., 24 July 1955 (W. R. B. & J. S. B.), Bauer-Buckett slide ' No. 66L2-3; 1 male, 6 July 1962 (H. J. Pini), Bauer-Buckett slide No. j 64D26-4; 1 female, 5 August 1962 (H. J. P. ), Bauer-Buckett slide No. j 66L6-5; 1 female, 10 August 1962 (H. J. P. ), Bauer-Buckett slide No. i 66L7-2; four males, 1 female, 5 August 1962 (H. J. P.); 1 female, 20 ; August 1962 (H. J. P. ); 1 female, 1 August 1962 (H. J. P. ); 14 males, 8 females, 7 August 1962 (H. J. P. ); 6 males, 2 females, 9 August 1964 I (H. J. P.); 1 female, 11 August 1965 (H. J. P.); 1 male, 21 July 1965 1 ( W . R. B. & J. S. B. ); 3 males, 6 July 1965 (H. J. P. ); 10 males, 1 female, 24 July 1965 (H. J. P.); 1 male, 27 July 1964 (H. J. P.); 1 male, 21 i July 1964 (H. J. P.); 1 male, 13 July 1964 (H. J. P.); 1 male, 16 ; July 1964 (H. J. P.); 2 males, 24 July 1964 (H .J. P.); 2 males 17 July 1964 (H. J. P.); 1 male, 12 July 1964 (H. J. P.; 1 female, 26 ij July 1965 (H. J. P. ); 13 males, 2 females, 17 July 1964 ( H. J. P. ); 3 |1 males, 16 July 1964 (H. J. P.); 3 males, 13 July 1964 (H. J. P.); 4 males. 224 J. S. BUCKETT /. Res. Lepid. Fig. 2. Holotype male, Folia piniae. Johnsville, Plumas Co., California, 5 July 1962 (Helena J. Pini). Fig. 3. Allotype female. Folia piniae. Johnsville, Plumas Co., California. 3 July 1959 ( W. R. Bauer and J. S. Buckett). 5 (4): 221-228, 1966 NEW FOLIA 225 5 Fig. 4. Paratype female. Genitalia of Folia pinkie. Johnsville, Plumas Co., 10 August 1962 (H. J. Pini), Bauer-Buckett slide No. 66L7-2. Fig. 5. Female genitalia of Folia discalis (Grote). Rabbit Ears Mountains, 10 miles southeast of Steamboat Springs, Routt Co., Colorado, elevation 8200’, 28 July 1962 (J. S. Bucket! and G. M. Trenam), Bauer-Buckett slide No. 66L7-1. L 226 J. S. BUCKETT /. Res. Lepid. Fig. 6. Paratype male. Genitalia minus aedeagus of Folia pmiae. Johns ville, Plumas Co., 24 July 1955 (W. R. Bauer and J. S. Buckett), Bauer- Buckett slide No. 66L2-3. Fig. 7. Male genitalia minus aedeagus of Folia discalis. Data same as for figure 5, except for slide No. which is: Bauer-Buckett slide No. 66L2-1. Fig. 8. Aedeagus of Folia discalis. Data same as that for fig. 7. Fig. 9. Aedeagus for Folia piniae. Data same as that for fig. 6. 5 (4): 221-228, 1966 NEW FOLIA 227 7 July 1965 (H. J. P. ); 1 male, 9 September 1963 (H. J. P. ); 2 males, 6 July 1965 (H. J. P. ); 1 male, 1 female, 21 July 1964 (H. J. P. ); 12 males, 2 females, 19-23 July 1964 (H. J. P.); 6 males, 2 females, 27 July 1964 (H. J. P.); 1 male, 17 July 1962 (H. J. P.); 3 males, 1 female, 11 July 1962 (H. J. P.); 21 males, 1 July 1965 (H. J. P.); 3 males, Mohawk, Plumas Co., 4-7 July 1946 (W. R. Bauer); 1 female, 12 July 1946 (W. R. B.); 3 males, Mt. Ingalls, Plumas Co., 11 July 1964 (W. R. B., J. S. B., M R Gardner); 4 males, 1 female, Pinecrest, Tuolumne Co., 25 July 1965 (Robert Mason); 14 m^es, 1 female, 9 July 1965 (R. M.); 11 males, 16 July 1965 (R. M.); 1 male, 2 July 1965 (R. M.); 1 male, 2 miles west, Sonora Pass, Tuolumne Co., 1 July 1966 (G. M. Buxton); 3 males, Went- worth Springs, El Dorado Co., 8 July 1961 (W. E. Simonds); 1 male, Donner Lake, Nevada Co., 20 July 1962 (G. M. B.); 3 males, Carnelian Bay, Placer Co., 12 July 1965 (F. D. Parker); 3 males, 4 females, Man- zanita Lake, Shasta Co., 13 August 1963 (G. M. Buxton); 1 male, Hat Creek, Shasta Co., 26 July 1964 (R. R. Finger); 1 male, Cedar Pass, 6 miles northwest of Cedarville, Modoc Co., 4 July 1962 (W. R. B., J. S. B., G. M. Trenam); 1 female, Lassen Creek, north of Davis Creek, Modoc Co., 28 July 1957 (W. R. B. & J. S. B.); 1 male, Mineral King, Tulare Co., 19 July 1963 (W. E. S.); 1 male, 1 female, Zeypher Cove, Douglas Co., Nevada, 20 August 1963 (Bobbie Ellis); 1 male, 27 August 1963 (B. E.). The Holotype is deposited in the Entomology Type Collec- tion, Department of Entomology, University of California, Davis, California. The female Allotype and paratypes are deposited in the private collection of the authors. Other paratypes will be deposited in the following institutions and collections: American Museum of Natural History, New York; California Academy of Sciences, San Francisco; California State Department of Agri- culture, Sacramento; Canadian National Collection, Ottawa, Ontario; J. G. Franclemont collection, Ithaca, New York; Los Angeles County Museum of Natural History; United States Na- tional Museum, Washington, D. C.; University of California, Berkeley and Davis. PoUa piniae can be readily distinguished from P. discalis by the former possessing sparsely fuscous irrorated primaries, there- fore appearing ashey-grey, or silvery-grey; whereas, discalis possesses primaries that are thickly and evenly irrorated with fuscous, therefore appearing darker or bluish grey. The space between the orbicular and reniform is less than the width of , the orbicular in piniae; whereas, in discalis this same space is as wide as the orbicular or wider in some specimens. Both the male and the female genitalia of piniae are specifically distinct from the genitalia of discalis also (as can be seen in figures 4-9). P. piniae is apparently a mid to late summer flier, predom- I inently at moderate elevations in the Sierra Nevada of California I and elsewhere it occurs. The light form of discalis occurs on the I i 228 J. S. BUCKETT /. Res. Lepid. eastern edge of the Sierras where there is considerable influence from Great Basin type flora; whereas, typical discalis occurs at higher elevations in the Rocky Mountainous regions of the central United States, and more specifically in Colorado. To date nothing is known concerning the immature sages of piniae. We take great pleasure in naming this species in honor of the ardent collector and naturalist, Mrs. Helena J. Pini of Johns- ville, California. The genitalic illustrations were prepared by the first author. 229 KNOW YOUR AUTHOR BRIAN OLIVER CORDERY GARDINER Born: Ealing, England, March 10, 1923. St. Bartholomews Hospital Medical School, first M, B. Married in 1951, 3 boys. Positions: Left St. Bartholomews due to war, after which worked for a short time in the Entomology Department, British Museirm (Natural History). Transferred to present position on Agricultural Research Council. Interests: The lepidoptera of Cambridgeshire, especially the fenland area; Pieris brassicae, its varieties, races and virus disease; breeding the Saturniidae of the world, and breeding lepidoptera out- of- season and on artificial diets. Publications: About 75 notes and articles on lepidoptera, mostly in British journals. Journal of Research on the Lepidoptera 5(4): 230, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1966 A GYNANDROMORPH OF LYCAENA GORGON PAUL A. OPLER 1131 E. 10th St. Albany, California Fig. 1. A gynandromorph of Lycaena gorgon collection data: Castle Rock Park, Mt. Diablo, Contra Costa County, California, May 10, 1953. Collected by Paul A. Opler. ' 230 Journal of Research on the Lepidoptera 5 (4): 231-242, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1966 THE DISTRIBUTION AND BIONOMICS OF ARCTIC-ALPINE LYCAENA PHLAEAS SUBSPECIES IN NORTH AMERICA OAKLEY SHIELDS 5151 Alzeda Drive, La Mesa, California and JOHNSON C. MONTGOMERY 3660 Altamont Way, Redwood City, California INTRODUCTION Ford (1923) defines the range of Lycaena phlaeas (Lin- naeus) and its subspecies as “throughout the greater part of the Northern Hemisphere,” including most of the Palaearctic and Nearctic Regions and part of the Oriental and Ethiopian Regions. The species is subject to remarkable seasonal, geographical, and individual variation within this range (Ford, 1923). In spite of this variation, the haploid chromosome number for three sub- species of L. phlaeas from Japan, Finnland, and the United States is 24 (Maeki & Remington, 1960), strongly indicating a stable chromosome number for the species throughout its range. Lees ( 1963 ) has shown that a changed environment can radically alter the phenotype of phlaeas. TYPE LOCALITIES Linnaeus (1761) described L. phlaeas from “in pratis West- manniae.” Westmannia is located in Sweden (Tite, 1957). Bois- duval (1852) described L. p. hypophlaeas from “Nord de la Californie. II se retrouve dans tout le nord des Etats-Unis.” This translates, “North of California. It is found in all the northern United States” (Dod, 1907). Thus the type locality is not “Cali- fornia” as listed by Klots (1951) and Comstock & Huntington (1960) and alluded to by Forbes (1960) and Garth & Tilden (1963). We do not know of a precise locality for hypophlaeas nor where the type specimen ( s ) is located. ( California material 231 232 SHIELDS AND MONTGOMERY J. Res. Lepid. Figs. 1 and 2. Lycaena phlaeas feildeni, I, 2; phlaeas ssp., 3-10; p, “hypophlaeas,"’ 11, 12; phlaeas, 13, 14; p. americana, 15-18; p. americana f. fasciata, 19; p. arethusa, 20. 1. & 2. Clyde Inlet, Baffin Id., N.W.T. (BMNH), $, ^ . 3. & 4. McKinley Park, Alaska (AMNH), $, ^.5. & 6. !4 mi. W. Haffmoon Park, Crazy Mtns., Sweet Grass Co., Montana, $ , ^ . 7. & 8. Bear Tooth Mtns., Carbon Co., Montana (AMNH), $ , $ . 9. Amphitheater Lake, 10,000’, Grand Tetons, Wyoming (CU), $ . 10. Beartooth Lake, Wyoming (CU), 11. & 12. N. slope Mt. Dana, 11,000-12,000’, Mono Co., California, $ , <5 . 13. & 14. Uddevalla, Swe- den (BMNH), $ , $ . 15. & 16. Near Prairie Village, Johnson Co., Kan- sas, 2 } $ • 17., 18., & 19. Camp Lucerne, Waushara Co., Wisconsin, $ , , $ . 20. Plateau Mt. 8200’, Alberta, $ . 5 (4)231-242, 1966 LYCAENA PHLAEAS 233 234 SHIELDS AND MONTGOMERY /. Res. Lepid. Fig. 3. Map of the North American Lycaena phlaeas subspecies distribu- tion (excluding americana). Localities indicated by black dots. Shaded portion is the distribution of Oxyria digyna as figured by Mooney & Billings ( 1961 ) and Billings ( in litt. ) . 5(4) 231-242, 1966 LYCAENA PHLAEAS 235 4 Fig. 4. Variation of “hypophlaeas” series from N. slope Mt. Dana, Mono Co., California. Left half females, right half males. I i ! 236 SHIELDS AND MONTGOMERY J. Res. Lepid. Fig. 5. Map showing the distribution of ‘"hypophlaeas” in the Sierra Nevada Mtns., California. Localities indicated by black dots. Map adapted from Starr ( 1956 ) . 5(4) 231-242, 1966 LYCAENA PHLAEAS 237 will be referred to as “hypophlaeas” in this paper as a matter of convenience.) MXachlan (1878) described L. p. feildeni from two males and one female from “Lat. 81° 45’.” The British Museum of Natural History contains these three specimens which bear the label, “Grinnell Land, west side of Smith Sound, Arctic America. 78-83 Lat. (81-45) Capt. Feilden R. N. 77-101.” (Tite, in litt. ). These were collected in 1875 or 1876 (Wolff, 1964). Dod (1907) described arethusa from five males and eight females: one male from ca. 35 mi. SW Calgary, Alberta (Lineham’s lower log camp, S. Fork Sheep Creek), and the rest from ca. 25 or 20 mi. SW Calgary (“near the spruce woods”), July 5 to 20 (no years given). The holotype and allotype are in the United States National Museum, and six paratypes are in the Canadian National Museum. FOOD PLANTS Langer ( in litt. ) mentions that European texts list Rumex and Polygonum as larval foodplants for phlaeas forms. Yokoyama (1955) lists “daikon” (a type of garden radish) and various grasses as larval foodplants for Lycaena phlaeas daimio Seitz in Japan. This was the only reference we found that listed food- plants other than members of Polygonaceae for phlaeas sub- species. Rumex species are gh en as foodplants for L. p. ameri- cana Harris in various U. S. texts (see Davenport & Dethier, 1937; Klots, 1951). The only reference we found to a foodplant of the three subspecies studied was that of M’Lachlan (1878) to Oxyria. He suggested that Oxyria digyna ( L. ) , then known as O. reniformis, probably serves as the foodplant for feildeni since no Rumex was found “at all the stations” while Oxyria was. Wyatt (in litt.) found feildeni at Coppermine, N. W. T., always in association with “a very Rumex~\i\ie plant” with reddish seeds and 6-8 inches high. The foodplant for “hypophlaeas"’ in the Sierra Nevada Moun- tains of California is almost certainly Oxyria digyna, Mountain Sorrel, although evidence at present is circumstantial. No speci- mens of “hypophlaeas” were found by us more than a quarter mile from O. digyna. Both of us independently have seen fe- males slowly flutter over digyna plants (one was seen doing so at 12:35 P.S.T., July 28, 1966, Mt. Dana, Mono County) and repeatedly alight on the flower heads without feeding. One fe- male on August 4, 1964, Mt. Dana, walked extensively on a digyna plant. No such activity by females was seen directed toward other plants. However, no oviposition or abdominal 238 SHIELDS AND MONTGOMERY /. Res. Lepid. probing by these females was seen. At the Mt. Dana locality, no Rumex was found growing on the slopes where "‘hypophlaeas"' flies. Flying, feeding, and sunning of both sexes were confined to the extensive digyna colony there. One of us (JCM) found "‘hypophlaeas” in four or five localities where digyna was found. The distribution of O. digyna is characterized by Mooney & Billings (1961) as “arctic-alpine circumpolar, with disjunct lo- cations far to the south in the mountains of Europe, Asia, and North America.” The North American range of digyna, as fig- ured by them, encompasses the known localities for arethusa, feildeni, and “hypophlaeas'' (see fig. 3). Mooney & Billings (1961) found that O. digyna in North America can be classed into two primary morphological groups. Based on stamen number, inflorescence branch number, and presence or absence of rhizomes, one group includes “all the populations from southern Alberta southward in an area largely to the south of maximum Pleistocene continental glaciation. The other group includes all of the northern populations (p. 27).” Presuming that digyna is the foodplant for the subspecies con- sidered here, arethusa and "‘hypophlaeas” would correspond to the southern digyna populations, and feildeni would correspond to the northern populations. (We do not rule out the possibility that these populations may have other larval foodplants.) Klots ( 1951 ) lists Rumex acetosella L. and “perhaps” R. acetosa L. and R. crispus L. as larval foodplants for L. p. americana. These are all introduced weeds from Europe and Asia (Fernald, 1950; Munz & Keck, 1965). In California, R. acetosella is found in cismontane areas, and R. crispus is found in low areas (Munz & Keck, 1965). The California "‘hypophlaeas^' apparently has not extended its range into the habitats of these two weeds. HABITAT One of us (JCM) found adult feeding confined to a small i yellow composite at four localities for ‘"hypophlaeas."' The other (OS) found feeding by both sexes at the Mt. Dana locality oc- i curring on a number of small alpine flowers and a large yellow composite. At all localities we checked, the males often flew I rapidly over the steep talus slopes and alighted to sun themselves on rocks; most of the females were collected while feeding on flowers. Both sexes perched on rocks and sunned either toward or away from the sun. One male on July 28, 1966, Mt. Dana, 11:20 P.S.T., lit on an O. digyna flower head momentarily. [ MacNeill (in litt. ) says that ‘‘hypophlaeas^^ at Mono Pass, Mono Co., California, is partial to the rocky “nunatak”"like plateaus j 1 il 5(4) 231-242, 1966 LYCAENA PHLAEAS 239 of gentle relief and the slopes of these near their base in the small canyons and chutes. One of us (JCM) twice took “hypophlaeas"' in conjunction with Lycaena cupreus (Edwards) and L. editha (Mead). Neither of these species flew directly with ‘*hypopMaeas” at the Mt. Dana locality (see fig. 6), although both did fly some 500 feet lower in elevation to the west. Legge (in litt. ) says that atethusa at Plateau Mt. in Alberta is found in small grassy meadows, while L. cupreus snowi (Ed- wards, will fly o\^er the talus as near as few hundred feet away. Scott ( in litt. ) says that phlaeas ssp. at Haifmoon Park, Sweet Grass Co., Montana, flew in the Hudsonian zone; these were taken on a rocky jeep road in the trees below a barren rockslide. ADULT MORPHOLOGY The British Museum of National History contains one male and three females of a phlaeas form labelled “California, Felder Colin.” These specimens are very similar to Eastern U. S. ameri- carm except that the upper forewing spots are elongated inward. An aberration of americana named fasciata (Strecker, 1878) has these elongated spots (see no. 18, fig. 1). Brower & Brower (1954) obtained fasciata individuals under uncontrolled rearing conditions and speculated that the condition is genetically de- termined. Lees (1963), however, produced this form environ- mentally. He reared ten L. phlaeas from Ilkley, England, at 35° C., a temperature much higher than the insect normally en- counters. His description and illustration of the female adults correspond closely to the B.M.N.H. California specimens. No such ruddy coloration or spot formation was noted in 104 "%ypo- phlaeas'' specimens we have examined from the Sierra Nevada Mountains. Perhaps the four Felder specimens came from a warm habitat or were reared under heated conditions. Figures 1 and 2 illustrate geographical variation in North American L. phlaeas. Certainly long series from many places coupled with experiments to determine how much of the varia- tion may be due to environmental influences will be necessary to establish the status of the names arethusa, feildeni, and hypo- phlaeas. Figure 4 illustrates the variation in one population of ''hypophlaeas' collected on three different years . 240 SHIELDS AND MONTGOMERY /. Res. Lepid. Fig. 6. Rocky slope habitat of “hijpophlaeas” at N. slope Mt. Dana, Mono Co., California. 5(4) 231-242,1966 LYCAENA PHLAEAS 241 ACKNOWLEDGMENTS We wish to thank the following people for loaning specimens or for contributing records and information to this study: W. D. Billings, F. H. Chermock, J. D, Eff, J. G. Franclemont of Cor- nell University (CU), T. N, Freeman of the Canadian National Museum (CNM), J. S. Garth, J. A. Justice, T. W. Danger, J. A. Legge, C. D. MacNeill, L. M, Martin of the Los Angeles County Museum (LACM), C. W. Nelson, F. H. Rindge of the American Museum of Natural History (AMNH), J. A. Scott, G. E. Tite of the British Museum of Natural History (BMNH), T. P. Webster, and C. W. Wyatt. Our thanks also go to P. McHenry for sup- plying some original descriptions, to R. Brock for translation of a passage in Japanese and to M. Evans for the color prints. BIBLIOGRAPHY ANONYMOUS, 1962. Season summary. News Lepid. Soc., no, 3, 14 pp. (p. 4). ANONYMOUS, 1965. Season summary. News Lepid. Soc., no. 3, 16 pp. (p. 12). ANONYMOUS, 1967. Season summary. News Lepid. Soc., no. 3, 17 pp. (p. 15). BOISDUVAL, J. B. A. D., 1852. Ann. Ent. France, 2nd Ser. 10 (2): 291, no. 2^ BROWER, L. P & J. V. BROWER, 1954. The heredity of some spot aberrations in Lycaena phlaeas and L. hypophlaeas. Lepid. News 8: 125-129. COMSTOCK, W. P., & E. I. HUNTINGTON, 1960. An annotated list of of the Lycaenidae ( Lepidoptera, Rhopalocera) of the Western Hemis- phere. J. New York Ent. Soc. 68: 176-186. .DAVENPORT, D., & V. G. DETHIER, 1937. Bibliography of the de- scribed life-histories of the Rhopalocera of America north of Mexico, 1889-1937. Ent. Amer. 17: 155-196. DOD, F. H. W., 1907. Notes on Chrysophanus hypophlaeas and its allies, with description of a new species. Canad. Ent. 39: 169-171. FERNALD, M. L., 1950. Grays Manual of Botany. American Book Co., pp. 570-571. FORBES, W. T. M., I960'. Lepidoptera of New York and neighboring states. IV. Agaristidae through Nymphalidae including butterflies. Cornell Univ. Agr. Exper. Sta. Memoir 371: 1-188. FORD, E. B., 1923. The geographical races of Heodes phlaeas L. Trans. Ent. 'Soc. London 71: 692-743. GARTH, J. S.. 1935. Butterflies of Yosemite National Park. Bull. So. Calif. Acad. Sci. 34: 37-75. GARTH, J. S., & J. W. TILDEN, 1963. Yosemite butterflies. J. Res. Lepid. 2: 1-96. KLOTS, A. B,, 1951. A Field Guide to the Butterflies. Houghton Mifflin Co., Boston, 349 pp. LEES, E., 1963. Experimentally induced sexual dimorphism in Lycaena phlaeas (Lycaenidae). /. Lepid. Soc. 17: 105-106. 242 SHIELDS AND MONTGOMERY /. Res. Lepid. LEGGE, A. H., 1965. A collecting trip in Yukon and Alaska. /. Lepid. Soc. 19 : 57-62. LINNAEUS, C., 1761. Fauna Svecica, p. 285, no. 1078. MAEKI, K., & C. L. REMINGTON, 1960. Studies of the chromosomes of North American Rhopalocera, 3. Lycaenidae, Danainae, Satyrinae, Morphinae. /. Lepid. Soc. 14: 127-147. M’LACHLAN, R., 1878. /. Linnean Soc. Zoology 14: 111. MOONEY, H. A., & W. D. BILLINGS, 1961. Comparative physiological ecology of arctic and alpine populations of Oxyria digyna. Ecol. Mono. 31: 1-29. MUNZ, P. A., & D. D. KECK, 1965. A California Flora. Univeristy of Calif. Press, Berkelely and Los Angeles, 1681 pp. NABOKOV, V., 1950. Remarks on F. Martin Brown’s “measurements and Lepidoptera.” Lepid. News 4: 75-76. STARR, W. A., JR., 1956. Guide to the John Muir Trail and the High Sierra Region. James J. Gillick & Co., San Francisco, 131 pp., 1 map. STRECKER, F. H. H., 1878. Butterflies and Moths of North America, a Complete Synonymical Catalog. Reading, Pennsylvania, p. 101. TITE, G. E., 1957. On the typical subspecies of Lycaena phlaeas L., and the Scandinavian distribution of the species ( Lepidoptera, Lycaenidae ) . The Entomol. 90: 37-38. WOLFF, N. L., 1964. The Lepidoptera of Greenland. C. A. Reitzels For- lag, Kobenhaven, 74 pp., 21 pis. WYATT, C,, 1957. Collecting on the Mackenzie and in the Western Arc- tic. Lepid. News 11: 47-53. YOKOYAMA, M., 1955. Coloured Illustrations of the Butterflies of Japan. Hoikusha; Osaki, Japan, 136 pp. Continued on Page 265 Journal of Research on the Lepidoptera 5 (4): 243-247, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 LIFE HISTORIES OF THREE WESTERN SPECIES OF POLITES E. J. NEWCOMER 1509 Summitview, Yakima, Washington The early stages of three western species of Polites, one of which is very common, have apparently not been studied thor- oughly. This paper discusses the life histories of P. sabuleti (Bdv. ), P. sonora (Scud.) and P. mardon (Edw. ). METHODS OF REARING It was found that females would oviposit readily in the small glass containers that season salt, dill salt and various other seasonings come in. These are about V/z x 4 inches in size and they have perforated plastic caps. The holes in the tops prevent too much moisture from accumulating but do not allow the skippers to escape. A small piece of absorbent material saturated with a sugar solution can be placed inside, held at the top of the bottle by the cap, on which the females will feed. Oviposi- tion occurs either on grass placed in the bottle or on the sides of the bottle. Females will live for three of four days under these conditions and may deposit as many as 25 eggs during this time. Newly hatched larvae were put into salve boxes with pieces of grass which was renewed daily. A damp piece of paper towel- ling in the bottom kept the grass from drying out rapidly. Lar- vae of these skippers develop slowly and when they are fairly large, they may be transferred to the same type of bottles used for oviposition, where they will conclude their development and pupate. They usually spin a loose cocoon among the pieces of grass in which pupation takes place. POLITES SABULETI This species occurs from Washington to Arizona and east to Colorado. In the Yakima Valley, Washington, there are two broods, one flying in May and June and the other in August and September. Sabuleti seems almost to have become domesticated 243 244 E. J. NEWCOMER /. Res. Lepid. in that it has adopted the lawns in cities and towns as its prin- cipal habitat. In the Yakima Valley, at least, it is seldom seen elsewhere. In this habitat the larvae feed on lawn grass, which is usually a mixture of Kentucky bluegrass and other species, and the adults feed on the blossoms of wild morning glory, dandelion and other garden flowers. Comstock (1927) states that the larvae “feed on Car ex filifolia and Trifolium monan- thum.” The former is a sedge occurring from British Columbia and Saskatchewan to California and Texas, and the latter is a clover occurring only in the Sierra Nevada Mountains of Cali- fornia and Nevada. I tried feeding larvae of sabuleti on the sedge and also on various clovers, but they would not eat any of these plants. Since the normal food of Polites is grass, it is reasonable to expect this to be true of sabuleti. Duration of the egg stage was 7 days at room temperature. ; The larval period lasted 35 to 50 days and the pupal period 10 , to 13 days. Eggs deposited about May 10 produced adults | July 19 to 26, and eggs deposited May 21 produced adults July ; 21 to 26. Hibernation of the second brood is in the pupal stage. I DESCRIPTION Polites sabuleti \ EGG. — Basal diameter 1.0 mm, height 0.8 mm. Color light : bluish green, darker at micropyle. Dome-shaped, base flattened. Finely reticulate. LARVA. — FIRST INSTAR. - Head width 0.5 mm, shiny black. Body length 2.0 mm, light cream; cervical shield narrow, I dark brown, a few setae. SECOND INSTAR. — Head width 0.8 mm, shiny black with | numerous fine setae. Body length 4 mm, greenish with many small brown dots, each bearing a seta. THIRD INSTAR. — Head width 1.1 mm, color and setae as ■ before. Body length 7.0 mm increasing to almost 10 mm, grayish green with many brown dots and brown setae of various lengths; dark median line and a distinct dorso-lateral line; cervical > shield black. FOURTH INSTAR. — Head width 1.6 mm, color as before but with two whitish streaks anteriorly and a shorter one laterally. * Body length 10.0 mm growing to 15.0 mm, grayish brown with dark median line, lighter ventrally; suranal plate becoming pro- i nounced, light colored; thoracic legs black; spiracles on first ' segment large, black, others smaller, brown. 5 (4): 243-247, 1966 SPECIES OF POLITES 245 FIFTH INSTAR. Head width 2.5 mm, color as before, punctate and with fine setae. Body length 18.0 to 22.0 mm, ground color light gray with numerous small brownish patches giving it a griseous appearance, lighter ventrally; median line brown but not sharply defined; numerous fine setae; cervical shield black; suranal shield white with a dorsal and two lateral black streaks; spiracles dark shiny brown; thoracic legs black; prolegs gray. PUPA. ™ Length 16 mm, width at thorax 4 mm; color green- ish with some brown on head, tips of legs and last two abdominal segments; numerous fine setae dorsally and a few laterally and ventrally on the abdomen. Cremaster short, not emarginate, with a single cluster of hooks. POLITES SONORA This is a common species occurring from British Columbia (Llewellyn Jones, 1951) to California and east to Montana, Wyoming and Colorado. In the Northwest, I have taken it on the prairies south of Olympia, Washington, and in Oregon along the upper Rogue River in Douglas County and at Camp Sher- man in Jefferson County. The larval foodplant has not been determined, but Festuca idahoensis Elmer^ is very common on the Washington prairies, it occurs throughout the range of sonom, and it is probably at least one of the foodplants. The larvae were easily reared on lawn grass and they also fed on F. idaho- ensis. The adults feed on various flowers growing in the habitat. Duration of the egg stage was 8 days at room temperature. Larvae hatching at the end of June were carried through until the end of July, when they had reached the third instar. No larvae were reared beyond this stage. ^DDetermined hij Professor Marion Ownhey, Washington State Univer- sity, Pullman. DESCRIPTION Polites sonora EGG. •— Basal diameter 1.0 mm, height 0.7 mm. Color very light green. Spherical with small flattened base, not flanged. Finely reticulate. LARVA. — FIRST INSTAR. — Head width 0.6 mm, shiny black. Body length 1.75 mm, creamy white, a few setae on last two segments; cendcal shield black. SECOND INSTAR. Head width 0.75 mm, black. Body length 3-5 mm, greenish, covered with numerous minute brown dots; cervical shield black. 246 E. J. NEWCOMER /. Res. Lepid. THIRD INSTAR. — Head width 1.0 mm, solid black, punc- tate. Body length 5 mm, grayish green with many fine black setae and a few longer ones on posterior segment. ROUTES MAROON PoUtes mardon was described in 1881 by W. H. Edwards (Edwards, 1881) from three males and three females “taken at Mt. Hood” in Oregon by H. K. Morrison. It has been scarce, but has since been taken near Tenino, Thurston County, Wash- ington, by D. L. Bauer, at Grand Mound, also in Thurston County, by J. F. Gates Clarke, on the south slope of Mt. Adams, Yakima County, Washington, at about 6500 feet elevation by Stanley G. Jewett, and on the open grassy slopes of Signal Peak, Yakima County, Washington, at 4800 to 5000 feet by the writer. It has also been reported from Seattle, A search of the area about Tenino and Grand Mound on June 6 and 20, 1966, by the writer did not turn up any mardon, although sonora was found there at that time. It is possible that mardon flies earlier. The Mount Adams and Signal Peak locations are about 55 miles north and 70 miles northeast of the type locality, respectively. i: i, i I I I Festuca ovina L.- is abundant on Signal Peak, as is Bromus carinatus Hook, h Arn.,^ and one or both of these grasses may be the native foodplant. The larvae feed readily on lawn grass, i however. The adults feed on the blossoms of dandelion and wall- j flower (Erysimum capitatum (Dougl. )). Males are often seen resting on rocks or bare patches of soil. ! Duration of the egg stage was 6-7 days at room temperature, j The larval period yasted about three months. Hibernation is in the pupal stage. ■ DESCRIPTION Polites mardon EGG. — Basal diameter 1.0 mm, height 0.8 mm. Color cream [ becoming yellow-orange. Spherical, base flattened, not flanged. Very finely reticulate. LARVA. FIRST INSTAR. Head width 0.5 mm, light brown. Body length 3.0 mm, very light brown with transverse rows of darker brown dots and a few setae; cervical shield black. ; 5 (4): 243-247, 1966 SPECIES OF POLITES 247 SECOND INSTAR. — Head width 0.75 mm, color as before. Body length 5.0 mm, color as before. THIRD INSTAR. — Head width 1.0 mm, dark brown with a darker narrow dorsal stripe. Body length 6.5 mm, color as before. FOURTH INSTAR. - Head width 1.5 mm, color as before. Body length 8.0 mm, color as in third instar but with a darker dorsal stripe. ^Determined by Professor Marion Ownbey. FIFTH INSTAR. — Head width 2.0 mm, black with two lighter dorsal stripes, surface covered with small pits. Body length 16.0 mm, tapering anteriorly; color light gray, sprinkled with numerous dark brown dots of irregular shape and varying size; a black median stripe; cervical shield black, suranal shield with dark margin and three dark spots just behind anterior edge; ventral surface same color as dorsal; spiracles and thor- acic legs black. PUPA. — Length 15.0 mm, width at thorax 4.0 mm. Smooth without protuberances. Color ashy gray with some light brown areas on abdominal segments and “shoulders” of wings; dor- sum of thorax darker gray, darker spots of various sizes scattered over thorax, abdomen and wings; eyes rich brown, darker pos- teriorly; many fine, light brown setae on abdomen and several tufts of them about the eyes and elsewhere on the head. Cremas- ter acute, laterally emarginate, hooks many, in dense cluster, scorpioid. LITERATURE CITED COMSTOCK, JOHN ADAMS, 1927. Butterflies of California. Publ. by the author, 334 pp. EDWARDS, W. H., 1881. Description of new species of butterflies. Papilio 1(4): 43-48. LLEWELLYN JONES, J. R. J., 1951. An annotated check list of the Macrolepidoptera of British Columbia. Soc. Brit. Columbia, Occ. paper no. 1. KNOW YOUR AUTHOR BRYANT MATHER Born: Baltimore, Maryland, December 27, 1916. Johns Hopkins University, A. A. , 1936 Johns Hopkins University, graduate work American University Positions: Curator (Mineralogy), Field Museum of Natural History (1939-1941). Geologist, Corps of Engineers, U. S. Army, West Point and Mt. Vernon, New York. Research Engineer, Corps of Engineers, Jackson, Miss- issippi, 1946 to present. Began , jointly with K. Mather in 1946, a survey of lepidoptera of Mississippi, now extended to include neuroplera and trichoptera. Author of papers on lepidoptera in Jour, Miss. Acad. Sci. , Maryland Naturalist, Lepid, , Jour. Lep. Soc, , and co-author (with K, Mather) of "Butter- flies of Mississippi]' Tulane Stud, Zool. , 6:63-109, 1958. Interested in the occurrences of lepidoptera, neuroptera, and trichoptera in Mississippi. Offers specimens of lepidoptera, neuroptera, and trichoptera, mostly from Mississippi for determination and study. Member: Am. Inst. Biol, Sciences, Lep, Foundation, Lep, Soc, , Ent Soc Canada, Nat. Hist. Soc. Md. , Mis s. Acad, Sci. , Am. Assn, Adv. Sci. (Fellow), Phi Beta Kappa, American Concrete Institute (President, 1963-64), Reference: Euphyes dukesi, J, Res, Lepid. , 2:161-169, 1963. 248 Journal of Research on the Lepkloptera 5 (4): 249-252, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 OVERCOMING DIFFICULTIES WITH THE PUPAE OF EUPROSERPINUS PHAETON MOJAVE (SPHINGIDAE) NOEL McFarland South Australian Museum Adelaide, South Australia On April 18, 1957, I collected over 30 last instar larvae of Euproserpinus phaeton mojave Comst., in a small area near Llano, Los Angeles County, California (at 3200 feet elevation on the western Mojave Desert). Around 10:00 A.M., many of the larvae were feeding on certain species of the abundant small annuals that cover the desert after a winter of ample rain- fall. Those observed feeding were eating Coreopsis bigelovii (Gray) Hall, and two small, annual Oenothera spp. Those not feeding were either crawling over the ground to other plants, or quietly sunning themselves on the warm sand. Although the larvae are marked with black, pink, and white on a soft green ground-color, they do not stand out conspicuously when in the natural habitat, because during the short time they are present as nearly fullgrown larvae, the sand around them is covered (to a varying degree) with brightly-colored petals, both fresh and dry, which have fallen from the many ephemeral spring flowers. These fallen petals (which are mostly pink, white, yellow, or blue) blow over the sand and collect in small depres- sions, or underneath plants where the larvae are feeding and resting. These collected larvae were transported to the Santa Monica Mountains near the coast (where I was living), and they were oflFered the leaves of the large, biennial Oenothera hookeri T. & G., which was accepted casually. Other possible foodplant sub- stitutes were not available, but as most of the larvae were about to pupate, this was not a problem. In order to approximate con- ditions in the natural habitat, the larvae were placed outdoors, in a screen-covered 15-gallon aquarium, in the bottom of which 249 250 NOEL McFarland J. Res. Lepid. Fig. 1. Euproserpinus niojave (adult, 3 views of Larvae). 5 (4): 249-252, 1966 PUPAE OF EUPROSERPINUS 251 was about 3 inches of sand from the habitat. They showed an interest in active feeding only after they had had a sufficient sunning each morning. Soon, they burrowed into the sand to make underground cells, in which they all pupated successfully. The pupae were dug up about two weeks later, and were stored in a suitable container, indoors, for the diapause period (i.e. — from May, 1957 until the following March ) . During the diapause period, the pupae were kept nearly dry, but for an occasional sprinkling, and they were overwintered outdoors in Albuquerque, New Mexico, where I spent the school year of 1957-1958. (Winter temperatures in that locality approximate those on the high Mojave Desert ) . They remained in excellent condition the whole time, responding with abdominal movement when handled. How- ever, when it came time for them to emerge (in March, 1958), problems developed. By late March, maculation on the fore- wings of the developing moths was easily visible through the pupal shells, so it was assumed that warmer temperatures, plus dampened soil, would bring about emergence in short order. The pupae were in a non-toxic, clear plastic box, on top of about one inch of the original sand from the habitat. The sand was then kept lightly damp. In most of the pupae, the moths were developed to the full extent, appearing ready to emerge at any moment, and at last they reached the final stage (just prior to emergence) where there is expansion between the abdominal segments of the pupa. But nothing happened thereafter! The pupae would not emerge after reaching that point of readiness; if they were opened by hand, the moths scrambled out in ap- parent good health, but the wings always failed to expand. If the pupae were left alone, the moths finally died within them, never emerging. When all but one of the pupae had been lost in one way or the other, something different was tried with the one remaining healthy pupa, which was ready to emerge in mid April. On April 18, 1958, the plastic box containing this pupa (lying upon damp sand ), was placed in the sun at about noon. The box “steamed” inside. Shortly the moth emerged of its own accord! Then another detail was observed: The newly-hatched moth did not seek a place where it could climb up, in order to let its wings hang down as they expanded and dried (which is the normal procedure with most freshly-emerged moths). Instead, the moth continued to sit on the damp sand, in the sun, and its wings rapidly “mushroomed” straight upward as they ex- panded. Then they were held erect, over the thorax. Full ex- 252 NOEL McFarland J, Res. Lepid. pansion of the wings took less than one minute, once started. When the box was taken out of the sun for a moment, the moth showed immediate distress and began running about; its limp, newly-expanded wings fell down and dragged on the ground as the moth ran. Upon being replaced in the sun, the moth stopped running and the wings again went straight up, and were held that way (together) until dry. A few minutes later, the fully-developed moth was flying around inside the box. It was subsequently spread for the collection. (See the photograph of this reared specimen, and three unhatched pupae). This account illustrates one of the difficulties sometimes en- countered by those who would rear Lepidoptera — the problem of breaking pupal diapause, and/or causing emergence of the imago; this problem is very commonly encoutered when at- tempting to rear species from arid regions. Although the treat- ment described above was successful with the diurnal, sun-loving Euproserpinus , that procedure would not necessarily succeed in causing emergence in very many other cases, except, possibly, with other diurnal desert moths. If such a technique is tried, the pupae should not be sun-warmed until the day they appear completely ready to emerge; then, the sun must not be too hot, the soil should be damp, and conditions in the container holding the pupae should be watched closely, to avoid over-heating. REFERENCES COMSTOCK, J. A. ( 1938). A new race of Euproserpinus phaeton from the Mojave Desert (Lepidoptera: Sphingidae), Bull. S. alif. Acade. Sci. 37 (1), pp. 33-42. COMSTOCK, J. A. ( 1938) ~ Bull., S. Calif. Acad. Sci., 38(1), pp. MUNZ, P. A. and D. D. KECK (1959). A California flora. Berkeley, Univ. of Calif. Press. 1681 pp. Journal of Research on the Lepidoptera 5 ( 4): 253-254, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 SPEYERIA CYBELE IN MISSISSIPPI ^ ^ (ARGYNNINAE: ARGYNNIS) BRYANT MATHER Box 2131, Jackson, Miss. 39205 Mather and Mather (1958) listed Speyeria cybele cybele (Fabricius) among the butterflies that were not then known to ha\’e been found in but were of probable occurrence in Mississippi. They noted that Lambremont ( 1954 ) had reported one male taken at Lafayette, La. on 2 October 1931, in the col- lection at Southwestern Louisiana Institute; that H. A. Freeman (1951) had reported it as usually rather scarce in Arkansas; and that Roever had taken it in southwestern Tennessee. They noted that dos Passes and Grey (1947) had listed it for Tennessee, Illinois, Arkansas, and Oklahoma; but not for Mississippi or Louisiana. In October 1960 in a butterfly collection displayed at the Mississippi State Fair by Miss Lynn Schabilion of Rolling Fork, Miss, there was a specimen of S, cybele. Upon investigation it was determined that this specimen had been collected in Arkan- sas. Through the courtesy of Dr. Gordon Gunter, Director, Gulf Goast Research Laboratory, Ocean Springs, Mississippi, we were put in touch with Dr. E. Avery Richmond of Moorestown, New Jersey, who was compiling data on the flora and fauna of Horn Island. This work has now been published ( Richmond, 1962 ) , and includes Speyeria cybele cybele (Fabricius). Dr. Richmond has informed me ( in litt. ) that his records indicate that this specimen was taken on 23 September 1944; they do not indicate the determiner of the specimen or the disposition of the speci- men. Most of the determinations were made at the U. S. National Museum. In January 1962 a search was made of the collections ‘ In this note the name “Speyeria cybele cybele (Fabricius)” is used to conform with previous usage in current relevant literature ( Mather and Mather 1958; Klots, 1951; Ehrlich and Ehrlich, 1961; and dos Passos, 1964). For a discussion of an alternative usage, see Hovanitz (1962). The butterfly with which this note is concerned has been given the common name “Great Spangled Fritillary.” ^ Editorial Note: In the Journal of Research on the Lepidoptera, name usage is the personal prerogative of the author, and is not an editorial matter. The editor prefers that authors use names of general significance, rather than of local preference. However, as is the practice in all entomologi- cal journals, when the name of a smaller taxonomic group is used in a title, a larger group of which the smaller is part, is shown in parenthesis for the orientation of the reader. This should aid bibiographers looking for various Argynnids. 253 254 BRYANT MATHER J. Res. Lepid. there and it was established that no specimen of S. cybele from Mississippi could be found. It was thus concluded that while this constituted a probable occurrence it could not be regarded as a confirmed record. On 10 June 1963 at Oxford, Lafayette Co., Miss., Mr. John L. Daniel took by hand, in the afternoon, a male S. cybele which is now before me. The specimen has a forewing of 40 mm. It was taken while feeding on flowers of cultivated marigolds. On the basis of this record Speyeria cybele cybele (Fabricius) is added to the list of butterflies known to have been taken in the state of Mississippi. The range of this species, which has been given as “ ... to North Carolina and Georgia (mountains), Tenn- essee, Arkansas, and Oklahoma” (Klots, 1951) and as “ ... to North Carolina, Georgia, Arkansas, Oklahoma ...” (Ehrlich and Ehrlich, 1961), can now be given “ ... to Georgia, Tennessee, Mississippi, Louisiana, Oklahoma ...” REFERENCES EHRLICH, PAUL R. AND ANNE H. EHRLICH 1961. How to Know the Butterflies. Wm. C. Brown Co., Dubuque, Iowa. 262 pp. FREEMAN, H. A. 1951. Southeast — Florida to Louisana, north to Arkan- sas and Maryland (compiled by Ralph L. Chermock). Lepid. News. 5 : 101-102. HOVANITZ, WILLIAM 1962. Argynnis and Speyeria. Jour. Res. Lepid. 1 : 95-96. KLOTS, ALEXANDER B. 1951. A Field Guide to the Butterflies. Houghton Mifflin Co. Boston, 349 pp. LAMBREMONT, EDWARD N. 1954. The butterflies and skippers of Louisana. Tulane Stud. Zoo/., 1 ; 125-164. MATHER, BRYANT AND KATHARINE MATHER 1958. The butterflies of Mississippi. Tulane Stud. Zoo/., 6 : 63-109. DOS PASSOS, CYRIL F. AND L. P. GREY 1947. Systematic catalogue of Speyeria ( Lepidoptera, Nymphalidae) with designations of types and fixation of type localities. Amer. Mus. Novitates No. 1370: 1-30. DOS PASSOS, CYRIL F. 1964. A Synonymic- List of the Nearctic Rhopalocera. Lepid. Soc. Memoir No. 1, 145 pp. RICHMOND, E. AVERY 1962. The flora and fauna of Horn Island, Mississippi. Gulf Research Reports, 1 : 59-106. Euphyes dukesi — Additional Record Bryant Mather, Bex 2131, Jackson, Mississippi 39205 Following publication of the review of knowledge of this species, (Mather, 1963), Mr. H. A. Freeman wrote (in litt. ) that he had recently found among his specimens a 9 E. dukesi that had been collected by him at Hope Hill Farm, Faulkner Co., Arkansas on 29 June 1947. This locality, somewhat north of Little Rock in central Arkansas, represents a westward extension of the known range and adds an eighth state to the list of those from which it is now known. REFERENCE MATHER, BRYANT. 1963. Euphyes dukesi. J. Res. Lepid. 2(2) : 161-169. Journal of Research on the Lepidoptera 5 (4): 255-261, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE LITTLE KNOWN MOTH EUXOA SCULPTILIS (HARVEY) IN ARIZONA, WITH DESCRIPTIONS, ILLUSTRATIONS, AND NOTES ON EUXOA VIOLARIS (GROTE AND ROBINSON ( NOCTUIDAE-AGROTIINAE ) J. S. BUCKETT University of California, Davis, California Euxoa SCULPTILIS (Harvey) WAS DESCRIBED from Specimens collected in Texas, and until recently has been poorly known in collections. This species is in the adult stage during the autumn months, and because of its occurrence during this late time of year, most collectors have not taken species. E. sculptilis has been collected from the first part of October into early Decem- ber, and without the aid of a collector-in-residence, the large series at hand would probably not have come before the author. Nothing is yet known concerning the immature stages of this species. Since the original description of sculptilis in 1874, the species has been shifted from Agrotis to Ammaconia to Richia to Car- neades to Paragrotis and finally to Euxoa in 1903 by Hampson, where is it presently placed by most authors. Strangely enough, Euxoa violaris (Grote and Robinson), the most closely related species to sculptilis, was not even placed in the genus Euxoa until McDunnough (1928) in his generic revision of the North American agrotid moths, placed it next to sculptilis. E. violaris, as in the case with sculptilis is poorly represented, or entirely lacking in most collections. “Carneades” xyliniformis Smith was described in 1890, and Smith stated “This species is unique in appearance and has a striking habitual resemblance to Litholomia napae” He further stated “The species is unique wherever placed, and is readily recognizable.” “C.” xyliniformis is nonspecific with E. sculptilis, and therefore was placed into synonymy by Smith (1893). E. sculptilis is quite distinctive (as can be seen in figures 1 and 2) 255 256 J. S. BUCKET /. Res. Lepid. Fig. 1. Euxoa sculptilis (Harvey), male. Madera Canyon, Santa Rita Moun- j tains, Santa Cruz County, Arizona, ele. 4880’, 17 November 1966 |i (D. N. Harrington). j Fig. 2. E. sculptilis, female. Same data as preceding. j: and does remind one of Litholomia napae in general appear- |' ance. Hampsons colored illustration of sculptilis is quite good, !jj and will readily aid in identification of this species; whereas, 5 (4): 255-261, 1966 EUXOA SCULPTILIS 257 the colored illustration by Draudt (in Seitz, 1923) is altogether off color and the shape of the forewing is not correct either. EUXOA SCULPTILIS (Harvey) Agrotis sculptilis Harvey, 1874. Bull. Buffalo Soc. Nat. Sci 2:271. MALE: Ground color of primaries transversely bicolor, basal one-half silvery grey, terminal one-half blackish grey; secondar- ies dirty whitish. Head with vertex clothed in elongate, blackish flattened hairs, white tipped, hairs porrect so as to form eyelash- like tufts over base of antennae; between antennae a transverse black bar is present; frons also with eyelash-like divided tufts dorsally, composed of dark brown white-tipped flattened hairs and white scales; beneath eyelash-like tufts, and between com- pound eyes a coal black broad transverse band is present, com- posed of simple and flattened elongate hairs; beneath this band, a tuft of brownish, white-tipped elongate flattened hairs, one each exterolateral of frons; central portion of frons of brown short hairs; frons roughened, centrally with ovate protrusion, pointed end directed dorsally, palpi with apparent basal seg- ment exterolaterally black, ventrally clothed in brownish; second segment exterolaterally composed of brownish scales, ventrally clothed in elongate black, greyish brown and whitish hairs; third segment short, less than one-half length of second seg- ment, clothed in greyish and blackish scales; antennae with scape, pedicle clothed in brown, reddish-brown and dark brown elongate hairs, elongate, dentate scales; flagellomeres bi-serrate, bipectinate, pectinations and serrations becoming reduced to ciliations and filamentous apically. Thorax with collar composed of reddish brown elongate hairs and elongate dentate scales, with two bilobed transverse bands, one basally, another med- ially; scales and hairs between bands white-tipped; slight an- terior tuft composed of bicolor, basally brown, apically dark brown, white-tipped elongate hairs; disc composed as is an- terior tuft; tegulae conspicuously -of black elongate dentate scales and hairs, with inner margin possessing a band of brown- ish, white-tipped, elongate dentate scales; posterior tuft basally brown, preapically dark brown, apically white-tipped; posterior- ly with elongate simple, silken hairs; at base of forewing, and beneath it, a bunch of deeply cleft elongate scales forms a yel- lowish-red cluster; primaries dorsally transversely bicolor; basal one-half of surface heavily overlain with white scales, therefore appearing grey; basal half line geminate, black, centrally grey 258 J. S. BUCKETT /. Res. Lepid. Fig. 3. Euxoa violaris (Grote and Robinson), male. Lakehurst, New Jer- sey, 21 September 1910 (F. Lemmer). Fig. 4. E. violaris, female. Long Island, New York, 29 September 1900. filled, very distinct; basal and transverse anterior area contiguous on inner margin; transverse anterior area grey, centrally with white dot; transverse anterior line black, % ery distinct, geminate. 5 (4): 255-261, 1966 EUXOA SCULPTILIS 259 centrally filled with grey, from where transverse anterior line intersects 2dA vein to where it reaches inner margin, it bows apically, therefore appearing like a geminate subclaviform; med- ian area to median shade grey; claviform distinct, neatly out- lined in black; orbicular large, more triangular than round, neat- ly outlined in black, thence white, centrally filled with grey; median shade black, contrasting markedly with grey portion of median area; remainder of median area dark brown, costally with light brown scales; transverse posterior line represented on costa just above reniform, appearing as geminate black dash, thence undulating out around reniform and very faintly on to inner margin; subterminal area dark, but with more grey than ad- joining median area; subterminal line represented costally as dark wedge, thence hardly discernable; terminal line represented as black lunules between veins; fringes basally light brown, re- mainder smokey; ventral surface bordered in grey, medially brownish; transverse posterior line represented as black costally, thence a very faint line on to inner margin; secondaries dorsally whitish; inner margin with brown scales and hairs; veins out- lined in brown; terminal line dark brown; fringes tricolor, basally light brown, medially brown, apically white-tipped; ventral sur- face whitish, costal margin irrorated with black so as to form faint dark band; exterior line thin, black; fringes whitish, be- coming darker toward apex; ventrally, thorax clothed in brown- ish, white-tipped simple hairs; legs with profemora dorsally black, ventrally composed of elongate white-tipped, brown sim- ple hairs; protibiae heavily spined; tarsi dark brown, a sugges- tion of a lighter apical annulus on each tarsomere; ungues weak- ly bifid. Abdomen dorsally clothed in brown and white simple elongate hairs and elongate dentate scales; ventrally clothed in elongate, brown, white scales. Greatest expanse of forewing 16mm to 19mm. Genitalia as in figures 6 and 7. FEMALE: For all practical purposes, as in male, except anten- nae simple, ciliate, thorax lacking basal bilobed black band of collar; primaries slightly darker; secondaries dorsally fuscous. Greatest expanse of forewing as in male, may average very slightly larger. Genitalia as in figure 5. Specimens examined One male, Madera Canyon, Santa Rita Mountains, Santa Cruz County, Arizona, ele. 5880’, 19 October 1959 (J. G. Franclemont), Bauer-Buckett slide No. NY63L11-15; 1 female, same locality and collector as preceeding 7 October 1959, Bauer-Buckett slide 260 J. S. BUCKETT /. Res. Lepid. Fig. 5. E. scluptilis, female genitalia. Madera Canyon, Santa Rita Moun- tains, Santa Cruz County, Arizona, 7 October 1959, ele. 5800’ j (J. G. Franclemont), Bauer-Buckett slide No. NY63L11-16. Fig. 6. E. sculptilis, male genitalia minus aedeagus. Locality and collector ! same as preceding, 19 October 1959, Bauer-Buckett slide No. | NY63L11-15. I Fig. 7. E. sctdptilis, aedeagus of male genitalia. Data same as for fig. 4. 5 (4): 255-261, 1966 EUXOA SCULPTILIS 261 No. NY63L11-16; remainder of specimens all Madera Canyon unless otherwise stated; 2 males, 2 females, 8 October 1963 (V. L. Vesterby); 1 male, 5 November 1965 (D. N. Harrington), Bauer-Buckett slide No. 66C31-1; 1 female, 5 November 1965 (D. N. H.), Bauer-Buckett slide No. 66C31-2; 8 males, 9 fe~ males, 2 November 1966 (D. N. H. ); 1 male, 7 October 1965 (D. N. H.); 20 males, 25 females, 2-5 November 1965 (D. N. H. ); 15 males, 8 females, 21-28 October 1965 (D. N. H.); 4 males, 14 October 1965 (D. N. H.); 3 males, 6 December 1965 (D. N. H. ); 16 males, 21 females, 27 November 1965 (D. N. H. ); 15 males, 27 females, 14 November 1965 (D. N. H. ); 1 female, Kerr- ville, Texas, with label “det. & pres, by Wm. Barnes.” Over 200 specimens of sculptilis were examined from the southwestern United States, the majority of these being from Madera Canyon, Santa Rita Mountains, Santa Cruz County, Arizona, elevation 4880'. Specimens were also collected at 5800' elevation in Madera Canyon. The genitalic slides were prepared while the author was studying under Dr. John G. Franclemont in Ithica, New York, in 1963. Merthiolate instead of ligin pink was used as the stain. Thanks are due Dr. -Franclemont for his encouragement and instruction during that most fruitful ex- perience. At this time I would like to express my appjreciation to Master Danny Harrington of Madera Canyon, Arizona for his splendid efforts in collecting moths during the past few years. Without his effort, this presentation would not have been possible. REFERENCES BARNES, W., and J. H. McDUNNOUGH, 1917. Checklist of the Lepi- doptera of Boreal America. The Herald Press, Decatur, 111., viii -}- 392 pp, DRAUDT, M. (In: Seitz, A. A.), 1923. The Macrolepidoptera of the World. Alfred Kernen Press, Stuttgart, 396 pp. + 64 plates. DYAR, H. G., 1903 (1902). A list of North American Lepidoptera and key to the literature of this order of insects. Bull. U.S. Natl. Mus., No. 52, xix -f- 721 pp. GROTE, A. R., 1887. On the genus Richia. Canad. Entom. 19(3) :44. HAMPSON, G. F., 1903. Catalogue of the Noctuidae in the collection of the British Museum. Longmans & Co. Press, London, xx + 689 pp. McDUNNOUGH, J. H., 1928. A generic revision of North American Ag- rotid Moths. Bulletin No. 55, Biological Series, No. 16, Canadian De- partment of Mines, 78 pp. SMITH, J. B., 1890. Contributions toward a monograph of the insects of the Lepidopterous family Noctuidae of temperate North America. Revision of the species of the genus Agrotis. Bull. U.S. Natl. Mus., No. 38, 237 pp. , J. B., 1893. Catalogue of the Lepidopterous superfamily Noctuidae found in Boreal America. Bull. U.S. Natl., Mus., No. 44, 424 pp. Journal of Research on the Lepidoptera 5 (4): 262-264, 1966 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 DISCOVERY OF A LARVAL HOSTPLANT FOR ANNAPHILA LITHOSINA WITH NOTES ON THE SPECIES (NOCTUIDAE: AMPHIPYRINAE) JOHN S. BUCKETT University of California Davis, California On May 13, 1967, it was the author’s pleasure of discovering larvae and eggs on Mimulus guttatus DC. which were thought to be deposited by specimens of Annaphila lithosina Henry Edwards. By observation of the Mimulus through late morning and early afternoon, oviposition by lithosina was observed. This phenomenon was studied quite closely, and occurred between 1:00 P.M. and 1:30 P.M. (Pacific Daylight Saving Time). The female would hover over small clumps of Mimulus for a few seconds, alight on new growth and walk over the area, ap- parently examining the new shoots for an oviposition sight, and perhaps also in some way attempting to detect the presence of previously deposited eggs by members of the same species. The value of female lithosina being able to detect the presence of eggs of her own species would be of paramount importance in exhibiting awarness of food supply versus population density for the immatures to emerge in due time. Whether this “ability” is in reality present in lithosina is doubtful, and could only be positively ascertained through timely intricate experimentation; but, when females alighted on new growth that already contained what seemed to the observer to be an overadequacy of eggs in relation to the amount of plant material available, they would not oviposit, but would seek another clump of new growth. When a female located the “desired” clump, she would re- main in one position for a few seconds, and while slowly waving her wings up and down, would thrust the abdomen downward between new leaves or sepals, and would either singly deposit 262 5 (4): 262-264, 1966 HOSTPLANT OF LITHOSINA 263 an egg, or would lay up to 10 or 12 in a row. Eggs were also deposited on stems and some of the older growth of the plant. She would then move to another site on the same clump and repeat the same process until a number of eggs had been deposited. On a single stalk of new growth, as many as over 75 eggs were counted. If gusts of wind would appear in sufficient force to shake the plant vigorously, the female would either return to her hovering above the plant, or would fly away to another clump. If she returned to her hovering position, generally a few inches to nearly a foot above the plant, she would return soon after the gust of wind subsided and continue oviposition. Examination of Mimiiliis flowers yielded one second instar larva, and some damage to petals was evident; however, the majority of flowers examined showed only a few to have been fed on . . . presumably by immature lithosbia. The adult lithosina abounded in this study area which is located 3 miles east of Auburn, in El Dorado County, California, and a number of specimens were collected with minimal effort. Adults were also collected 15 miles northeast of Auburn, Placer County, again in the close proximity with Mimulus guttatus. In previous years the author and associates have collected lithosina near the F’eather River at Elephant Butte, Plumas County, California and Mimtdiis giiftatiis was present, but prior to the present time, this species was not a suspect larval host- plant, and therefore was not examined for eggs or larvae. This year adult lithosina were collected in mid morning during which time they were seen to alight on green foliage while exhibiting the typical “wing-waving” pattern so noteworthy of members within the genus. As noon and early to mid afternoon came about, adults were observed feeding on the flowers of Montia parviflora (Moq. ) Greene, Nemophila heterophylla F. and M.; one specimen was taken while in association with Rannunculus calif ornicus Benth., but it was not ascertained whether or not the specimen was actually feeding. Another foodplant adult lithosina were collected on is Plagiobothrys nothofiilvus Gray (personal communication with Mr. Terry A. Sears), but the author was unable to locate lithosina feeding on this plant species. At no time were any adult specimens of lithosina seen feeding on the flowers of Mimtilus guttatus. As is usual with the large, succulent form of M. guttatus, the specimens concerned were growing at the edge of a stream at J. Res. Lepid. 264 J. S. BUCKETT ± 2,000' elevation where moisture is available well through the flowering season. The surrounding environment is intermixed deciduous and coniferous woodland with occasional meadows to be found. It is possible that lithosina, like other members of the genus is host specific on M. guttatm, as time may bear out. To my knowledge M. guttatus is abundant throughout the known range of lithosina. It is rather interesting to note a peculair quotation referring to the immature stages of lithosina as quoted by Rindge and Smith (1952). It reads as follows: “Larvae feeding in numbers on larvae of wasps. Sacramento, Calif. Harry S. Smith Febr. 16, 1915.” It is highly unlikely that this observation (?) is repre- sentative of the normal feeding habits of immature lithosina, and it is possible that this claim has no foundation in reality. Some lepidopterous larvae will turn cannibalistic when under an ecological stress such as food shortage, but in most instances this habit should not be considered usual. There is no actual proof that the specimen bearing the aforementioned label was one of these wasp consumers, but Rindge and Smith (op. cit. ) do state of the specimen “The wings of this specimen, a male, are rudimentary, but a genitalic preparation shows that it is properly referred to this species.” At present, the author is attempting to rear specimens of lithosina, and should success occur, detailed descriptions, il- lustrations and setal maps will be presented in an additional paper. Comparisons are being made between natural populations and laboratory populations in relation to duration from egg to adult. With the large natural population at hand, perhaps the observation of parasites and predators will be made. A. lithosina is in the species group of the genus comprised of lithosina, miona Smith, and casta Henry Edwards. For a colored photograph of the members of this species groups, see “A reevaluation of Annaphila casta” (Ruckett, 1966). To my knowl- edge, no larval hostplant records have been published concerning either miona or casta, but I believe Mr. Chris Henne (personal communcation ) has worked these out and will publish the results in the near future. LITERATURE CITED BUCKETT, J. S., 1966 (1965). A reevaluation of Annaphila casta (Noc- tuidae)./. R. Lepidoptera 4(3):199 - 204. RINDGE, F. H., and C. I. SMITH, 1952, A revision of the genus Annaphila Grote (Lepidoptera, Phalaenidae) . Btdl. American Mus. Natur. Hist. 98(3)191 - 256. TABLE 1. LOCALITY DATA 265 RECORDS FOR NORTH AMERICAN LYCAENA PHLAEAS SUBSPECIES (EXCEPT AMERICANA) CANADA. ALBERTA. Banff, Vn-23-25, "1" (O. Bryant, AMNH). Near Billings lumber mill, Calgary, Vn-19-03, "1" (F. H. W. Dod, AMNH). Brobokton Creek, halfway between Lake Louise and Jasper, VII- 8, 12, 13, 16-07 (Mrs. Nicholl, BMNH and CNM). Near the spruce woods, 20 or 25 mi. SW Calgary, VII- 5 to 20- ?, 4 cf 89 (F. H. W. Dod, in Dod, 1907, 6 arethusa paratypes in CNM, cf and 9 "types" in USNM). Crow's Nest Pass, S, Alberta (G. Geddes, in Dod, 1907). Fallentimber Creek, 51° 45' N. , 114° 39' W. , 5500' (J. Legge). Foothills, Lineham's lower log camp, S. Fork Sheep Creek, ca, 35 mi. SW Calgary, 1 cf (F.. H. W. Dod, in Dod, 1907). Peyto Lake, Banff Nat. Park, 51° 43' N. , 116° 31' W., (fide J. Scott). Plateau Mt. , 8200', VIII-2-62, "3" (J. and A. Legge). BRITISH COLUMBIA. Atlin, 4000', Vni-2-55 (CNM). Moosehorn Lake, 132° 07', 58° 10', 4500' (CNM). Western slopes, Mt. Sidney Williams, N. C. B. Columbia, 54° N. , 125° W. , ca. 5500', Vn-31-53, I cf (R. J. Gilpin, BNMH), Spray Lake, "2" (in Anonymous, 1962). NORTH WEST TERRITORIES. Arctic Bay, Baffin Land, Vin-2-42, "1" (AMNH), Baker Lake, VI- 30 to VII- 7- 51 (CNM); "2" (F. H. Chermock). Bathurst Inlet, VIII-1-51 (CNM); "1" (C, Wyatt, in Wyatt, 1957). Bernard Harbour, 68° 45' N. , 114° 45' W. , Vni-6-15 (CNM). Caribou Valley, head of Clyde Inlet, Baffin Island, VII- 31- 50, 1 cf (V, Wynne-Edwards, BMNH). Head of Clyde Inlet, Baffin Island, VII-29-50, Z cf 1 9 (V, C. Wynne-Edwards. BMNH); Chesterfield Inlet, Vn-?-66, "1" (in Anonymous, 1967^ VIII- 7- 58 (CNM). Coppermine, VIII-2-51 (CNM); late VII- 55, "series" (C. Wyatt, in Wyatt, 1957). Cockburn Point, Coronation Coast, 68° 52' N. , 115° O' W. , IX-1 to 2-14 (CNM). S. slopes of Dyke Mt. , head of Clyde Inlet, Baffin Island, Vn-26-50, 1 cf (V. C. Wynne- Edwards, BMNH). Eskimo Point, "2" (F. H. Chermock). Falcon Hollow, head of Clyde Inlet, Baffin Island, Vn-27-50, 1 cf (V. C. Wynne-Edwards, BMNH). Frobisher Bay, Baffin Island, VII- 6 to 20-48 (CNM); VIII- ?-64, Id* (J. P. Paton, BMNH). Grinnell Land, W. side of Smith Sound, Lat. 81° 45' (Capt. Feilden; holotype cf, allotype 9, and a paratype cf in BMNH, in M'Lacklan, 1878, feildeni TL). Hazen Camp, Ellesmere Island, 81° 49' N. , 71° 18' W.”7”vn^ 29 and 30-66; VI- 23 to Vn-31-61 (Both CNM), Holman, Victoria Island, VII-5-52 (CNM). Lake Harbour, Baffin Island, Vn-7-35 (CNM). Mellville Penn., 56° 51' N, [probably 66° 51' N. ], 84° 51' W., Vn-19~48 (CNMf. Pelly Bay, Vn-19-52 (CNM). Penny High- land Area, Cumberland Penn. , Baffin Island, VIII- 5- 53, 1 cf (A„ Watson, BMNH). Repulse Bay, VII- 28 to VIII- 9- 50 (CNM). Southhampton Island, VII- 5- 30, "1" (G. M. Sutton, AMNH); "2" (F. H. Chermock, CM). Wager River, NW Hudson Bay,. 65° 26' N. , 88°40'W. , "1" (F. H. Chermock). YUKON. Dawson (CNM). Dry Creek, on Alaska Hwy, near Alaska border, VII 23-48 (CNM). Near Haines Junction, ?-?-66. Yukon Terr., Lat. 64° 31', Long. 138° 30', ?-?-l6 (CNM). GREENIAND. Qanaq, MacCormick Fjord, 77° 41', Vni-26-41, 1 9 (C. Vibe, in Wolff, 1964). UNITED STATES OF AMERICA ALASKA. Arctic Village, VII-15 and 20-64 (in Anonymous, 1965). Cape Thompson, Vn-26- to 29-61 (CNM). Driftwood, 68° 49' N. , 161° 09' W., VIII-6-52 (CNM). Highway Pass, McKinley Nat. Park (A. H. Legge, in Legge, 1965). McKinley Nat. Park, VIII-4-31, Id* (F. Morand, LACM); VII- 20 to VIII- 1- 31, "20" (AMNH). Toklat River, McbSnley Nat. Park (A. H. Legge, in Legge, 1965). CALIFORNIA. Fresno County: near Baboon Lake, VIII-1-60, "sighting" (J. C. Montgomery). Ridge S. of Golden Trout Lake, VIII-4-60, 4 d (J. C. Montgomery). N. side of pead (12,492'), Wahoo Lake, VIII- 3- 60, 5 cf 3 9 (J. C. Montgomery). S. side of peak (12,492'), Wahoo Lake, VIII-5-60, 2 d (J. C. Montgomery). Fresno- Inyo County line: Bishop Pass, 12, 000', Vn-31-60, I9 (J. C. Montgomery). Mono Pass (C. D. MacNeill). Madera County: W. slope above Emerald Lake, 10, 000', 1/4 mi. S. Thousand Island Lake, VIII-14-62, "1" (T. P. Webster). Mono County: W. above Barney Lake, est. 10, 200', VIII-4-35, 1 cf (L. M. Martin, LACM). Mammoth Mountain, est. 11,000', VII-27-34, Id (LACM). N. slope Mt. Dana, NE of Tioga Pass, 11, 000-12, 000', Vni-8-33, "1" (J. S. Garth); VIII-6-60, 13 d 49; VIII-7-60, 14 d 69; VIII- 8-60, 4d 69; VIII-9-60, 4d 29; Vni-13-60, 5d 29 (all O. Shields, in Garth and Tilden, 1963); VIII-4-64, 10 d 39; VII-28-66, 5 d 1 9; VIII- 6- 66, 6 d 49 (all O. Shields). Tuolumne County: Bert Lake, below Mt. Maclure, est. 11, 700', Vni-6-33, Id (J. S. Garth, LACM, in Garth, 1935). Cathedral Lake, est. 9000-9500', VII-16-29, "1" (G. E. Bohart, AMNH). Kuna Crest, Yosemite Nat. Park, 11, 000', VIII-5-33, Id (J. S. Garth, LACM). MONTANA. Carbon County: Bear Tooth Mtns. , Vni-9 to 12-41, "5" (G. H. and J. L. Sperry, AMNH); VII-15-42, "1" (AMNH). Sweet Grass County: Halfmoon Park, Crazy Mtns. , ca. 15 to 20 mi7~NW Big Timber, VII- 3 to 4-66, 2d 1 9 (J. Scott). WYOMING. County undetermined: Amphitheater Lake, Grand Tetons, 10, 000', VIII- ?-49, 1 9 (V. Nabokov, CU, in Nabokov, 1950). Beartooth Lake, VIII-21-49, Id (V, Nabokov, CU). Teton Glacier, Teton Mtns. , VII-19-31, "2" (AMNH). 266 TABLE 1, VARLA.TION OF H, MELPOMENE AND H, ELEVATUS cut out this table and insert it in place of the table as printed on page 9^8 of this volume hi O I s 1 ^ © ^ g O ^ © m S m ^ s fiS .S p o ^ o ffl o © ft © ft TO rt d U © g o © © © \J ®Hi 0 d u ft S d u ft r-i & 0) -M d © ft o ®H > © g © f— 1 © u o g B +j © u © rH o @ e e ft (ti 'd o ft ft ft ft o rH ft d d CD m m TO 0 m c © © Cfi m m TO ft to A X I &0 , o p. u m p ’-d d A © •d 50 Z © d ®s=l 'd 4J U © 1-3 m d u TO d rt ®r=l u fti h o d m u h d d g d 0 O 0 rH § d > ■d d o o (ti +3 • f4 £0 © ® fH u ?H £ N TO © fd +3 3 0 © ft ft +J U d o ®H m © ch u © P. P. d © I o m •U cd d ’d fti g d N u fd ^ g W d cd p. m & ^ ^ © fl © fi rd < see Turner, Ji R„ , 1967, J, Linn^ Soc« , Lond, (Zool, ) 46, Z63 NOTICES CHANGE OF ADDRESS: When sending a change of address, please include both old and new addresses. This v^^ill greatly fac ilitate /our changeover. ZIP CODE: If your zip code is not present, or is incorrect, on your address plate, please inform us as the Post Office requires it on all domestic mailings. BUTTERFLIES OF THE DELAWARE VALLEY, by Arthur M. Shapiro. Special Publication of the American Entomological Society, Philadel- phia, Pa. i-vi, 1-79, 13 b. and w. plates, keys and check list. ^1. 50 from publisher at 1900 Race St. , Philadelphia, Pa. AUTOMERIS AURANTIACA, living ova or cocoons urgently required to complete research project, contact Brian O. C. Gardiner, 18, Chesterton Hall Crescent, Cambridge, England. LEPIDOPTERA OF FLORIDA, by C. P. Kimball. 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Bauer 197 Speciation in the Amthumus (Megathymidae) H. A. Freeman 209 The Eggs and First Instar Lar\^ae of Three California Moths John Adams Comstock 215 An Additional Food Plant Record for Papilio thoas autocles R. & J. John Adams Comstock 220 A New Species of Folia Ochsenheimer from California and Notes on Folia discalis (Grote) ( Noctuidae : Hadeninae ) John S. Buckett and William R. Bauer 221 Know Your Author — Brian O. C. Gardiner 229 A Gynandromorph of Lycaena gorgon Paul A. Opler 230 The Distribution and Bionomics of Arctic- Alpine Lycaena phlaeas Subspecies in North America Oakley Shields and Johnson C. Montgomery 231 Three Western Species of P elites E. J. Newcomer 243 Know Your Author — Bryant Mather 248 Overcoming Difficulties With The Pupae of Euproserpinus phaeton mojave Noel McFarland 249 • Speyeria cybele In Mississippi. Argynninae; (Argynnis) Bryant Mather 253 Euphyes dukesi — Additional Record Bryant Mather 254 The Little Known Moth Euxoa sculptilis (Harvey) in Arizona, with Descriptions, Illustrations, and Notes on Euxoa violaris ( Grote and Robinson ) ( Noctuidae- Agrotiinae) J. S. Buckett 255 Discovery of A Larval Hostplant for Annaphila lithosina With Notes on the Species (Noctuidae Amphipyrinae ) John S. Buckett 262 Appendix to Distribution of Lycaena phlaeas Oakley Shields and Johnson C. Montgomery 265 Correction to “A Little-recognized Species of Heliconuis Butterfly” John R. G. Turner 267 Established in 1962 Edited by WILLIAM HOVANIT/ Volume 6 1967 Published at 1160 W. Orange Grove., Arcadia, California, U.S.A. THE JOURNAL OF RESEARCH ON THE LEPIDOFTERA CONTENTS Volume 6 Number 1 March, 1967 Spring Moths of a Natural Area in Northeastern Kansas Noel McFarland 1 A New Species of Nepticula on Bur Oak in Ontario T. N. Freeman 19 Fixation of the Type Locality of Lycaena phlaeas hypophlaeas and a Foodplant Correction Oakley Shields 22 Description of a New Species of Xylomiges from California John S. Buckett 23 The Life-Histories of South African Colotis erone, C. ione, C. vesta and Leptosia alcesta (Pieridae) Cowan C. Clark and C. G. C. Dickson 31 A New Species of Feralia John S. Buckett 43 Homonymy of the “New Genus” Petaluma and Proposal of the Name Petalumaria J. S. Buckett and W. R. Bauer 52 Rearing Euleucophaeus ruhridorsa and E. lex Brian O. C. Gardiner 53 The Status of Some Hesperiidae from Mexico H. A. Freeman 59 The Generic, Specific and Lower Category Names of the Nearctic Butterflies Paddy McHenry 65 Volume 6 Number 2 June, 1967 HILLTOPPING AN ECOLOCICAL STUDY OF SUMMIT CONGREGATION BEHAVIOR OF BUTTERFLIES ON A SOUTHERN CALIFORNIA HILL OAKLEY SHIELDS Volume 6 Number 3 September, 1967 Origin of Autumnal “False Broods” in Common Pierid Butterflies A. M. Shapiro 181 The Lepidoptera Research Foundation, Inc. 194 Polyctor Polyctor in Mexico II. A. Freeman 195 Ecological Color Variation in Some Argynnis of the Western United States W. Hovanitz 197 Natural Habitats W. Hovanitz 199 Estimating the Density of An Animal Population William R. Hanson 203 Volume 6 Number 4 December, 1967 Life History Studies on the Lithosina-Miom-Casta complex of the genus Annaphila Christopher Henne 249 Studies in the Life Histories of North American Lepidoptera. California Annaphila III. J. A. Comstock and C. Henne 257 The Generic, Specific and Lower Category Names of the Nearctic butterflies. Pt. 6, The Genus Dryas. Paddy McHenry 263 A Melanie form of Pieris rapae J. P. Donahue 266 Man-made Habitat for Colias Eunjtheme W. Hovanitz 267 A New Species of Army worm — Genus Faronta J. S. Buckett 268 Early Stages of Lygomorpha regulus. J. A. Comstock and C. Henne 275 A Previously Unrecognized Subspecies of Philotes speciosa. J. W. Tilden 281 -A.'* V'?- ^ Mi*' March, 1967 Volume 6 Number 1 THE J0URNJAL ©F RESEARCH ON THE LEPIDOPTERA a quarterly published at 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HOVANITZ THE PURPOSE OF THE JOURNAL is to combine in one source the work in this field for the aid of students of this group of inseas in a way not at present available. THE JOURNAL will attempt to publish primarily only critical and complete papers of an analytical nature, though there will be a limited section devoted to shorter papers and notes. QUALITY WORK on any aspects of research on the Lepidoptera is invited. Analytical and well illustrated works are pre- ferred, with a minimum of long description. AUTHORS ARE REQUESTED to refer to the journal as an example of the form to be used in preparing their manuscripts. Illu- strations should be of the best quality black and white, or line draw- ings and should be pre-arranged by the author to fit a reduced size of 4” X 6V2." 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Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 SPRING MOTHS (MACROHETEROCERA) of a NATURAL AREA IN NORTHEASTERN KANSAS NOEL McFarland South Australian Museum, Adelaide, South Australia While attending the University of Kansas, between 1958 and 1961, I had the opportunity for considerable (but sporadic) moth-collecting during the spring semesters of 1959, 1960, and 1961. The results are presented in the annotated list that follows. Most of the collecting was restricted to the University of Kansas Natural History Reservation, 7 miles NE of Lawrence, Douglas County, Kansas, at approximately 900 feet elevation. A limited amount of collecting took place in a garden, at one address ( 1212 Louisiana St. ) in Lawrence, Kansas. BIOTIC DESCRIPTION OF THE RESERVATION The following is quoted from a paper by Henry S. Fitch (1957) on “The Univ. of Kansas Natural History Reservation — Objec- tives and Uses”: “Natural areas preserving samples of the original flora and fauna in an undisturbed state are almost com- pletely lacking in Kansas; therefore, this reservation created in 1948, will have increasing usefulness and significance as it reverts to an approximation of its original condition, and as other areas that are not protected are further altered by agricultural and industrial uses.” Prior to 1948, the area had been used chiefly for grazing. The Natural History Reservation, excluding the Rockefeller Tract, covers almost one square mile (590 acres), at elevations between 900 and 1100 feet. Most of the slopes, ravines, and lower areas are heavily wooded, with American elm, oaks, and hickory predominating. Some of the relatively flat hilltops are covered by grassland, and are thought to be remanants of blue- stem prairie, which once dominated larger areas. The habitats represented on the Univ. of Kansas Natural History Reservation have been described in some detail by Fitch (1952, 1958), and by Fitch and McGregor (1956). These papers include not only 1 2 McFarland /. Res. Lepid. descriptions of the Reservation as a whole, but also some de- tailed descriptions of specific small areas, and a discussion of the past history of the area in general. For the purposes of this paper, it would be of interest to list (alphabetically) the woody plants growing on the Reservation, as they typify the undisturbed woodland habitat of the area. Of course herbaceous plants (both forbs and grasses) are abund- antly represented by numerous species, particularly in the open areas, along roadsides and trails, in disturbed places, around the pond, and in the forest understory. For nearly complete coverage of the higher herbaceous plants (except grasses) of eastern Kansas, see Stevens ( 1948). Woody Plants Growing on the Natural History Reservation (Common species starred; dominant species double-starred) Acer negundo Pyrus ioensis * ^ Ailanthus altissima (scarce) Quercus macrocarpa ( scarce ) Amorpha fruticosa Q. marilandica Asimina triloba Q. muehlenbergii Canja ovata Q. prinoides * Celastrus scandens * Q. rubra (scarce) Celtis occidentalis * Q. stellata (scarce) Cercis canadensis ** Q. velutina * Cornus asperifolia ** Rhus spp. * Crataegus mollis ** Ribes sp. Euonymus sp. Rosa sp. Fraxinus americana * Salix nigra ^ ( at the pond. Gleditsia triacanthos ^ * near Reservation head- Gymnoeladus dioica * quarters, along with three Juglans nigra * other Salix spp. ) Juniperus virginiana ( the Sambucus canadensis only coniferous plant present ) Smilax spp. * Lonicera sp. Symphoricarpos orbiculatus"^ Madura pomifera ** Tilia americana (scarce) Morus rubra * Ulmus americanus * * ( the PaHhenocissus quinquefolia * commonest tree, but now in Pkitanus occidentalis ( scarce ) the process of dying off ) Populus deltoides U. rubra Primus americana Vitis vulpina * Prunus serotina Zanthoxylum americanum Fitch & McGregor (1956) discuss 29 of the above-listed woody plants, with reference to their ecology and occurrence on the Reservation. 6(1): 1-18, 1967 SPRING MOTHS 3 TOPOGRAPPIY The collecting station was near the head of a small valley at about 920 feet elevation. Another somewhat larger valley is in the eastern part of the Reservation. The two valleys are separated by a relatively flat hilltop 1040 to 1060 feet in elevation; the hilh top has shallow soil with an underlying layer of hard rock of Pennsylvanian age, the Oread Limestone, and for the most part has a grassland type of vegetation. The slopes between the valleys and hilltop are steep and wooded, with thorny thickets dominated by honey locust, osage orange, and other xeroseral vegetation on the south-facing slopes, but with more mesic woodland including oaks, hickory, and ash, on the north-facing slopes. Much of the woodland is of recent origin, having encroached onto former grassland areas since prairie fires were eliminated, METHODS OF COLLECTING Except for the diurnal moths, and those species listed for Lawrence, Kansas, all were collected around the Reservation headquarters buildings, where they were attracted to several 15"Watt black lights (ultraviolet tubes — F15T8/BL, without filters). These lights were placed against white walls, or white sheets, about six feet above the ground. No traps were used. Three of the lights faced rather dense woodland edge and streamside habitats, at the south and east edges of a relatively clear area where the buildings stand. Two other lights faced north and northeast toward a rather large, open, grassy field, bordered by woodland. A small pond, with characteristic vege- tation, was approximately 200 yards to the northeast. The lights were turned on sporadically, whenever time was available, during suitable weather from the first mild nights in late March up to the end of May each spring, from 1959 to 1961. On no occasion were the hours between 1 to 5 A.M. sampled. Rarely was it possible to collect on more than three nights per week, and it was not possible to continue this study beyond the end of May each year. It should be emphasized that collecting was restricted to one location on the Natural History Reservation. Even though the survey was restricted to this extent, some 226 species of moths were turned up, representing 12 families of the “Macrolepidoptera,” and considerable information was gained on the relative abundance of these moths during the spring months. Collecting in the city of Lawrence turned up a few species that were never encountered at the Reservation; these species are 4 McFarland J. Res. Lepid. included in the list, but with the comment “to be looked for at the Natural History Reservation/' As might be expected, there were numerous species taken at the Reservation, with its varied native flora, that were never seen in Lawrence. Extensive daytime collecting at the Reservation was also car- ried out, during April and May, in favorable weather. INTRODUCTION TO THE ANNOTATED LIST OF SPECIES The families follow the arrangement in Part I of the Check List of Lepidoptera of Canada and the United States of America by J. McDunnough (1938). The genera and species are arranged adphabetically , within each family, for ease of locating the species in this list. ( The McDunnough number are included how- ever, at the right markin of the page, opposite the name of the moth). If a question-mark precedes a specific name, this indi- cates that there is some doubt as to the correct identity of the species; where there is considerable doubt ,no specific name is included. Occasional subspecific names are included. In most cases, no mention is made of mere color forms. Following (1), is given the flight-period of the adult, as re- corded within the limits of this spring survey. “Early” (preceding the month) indicates the period from the first through the 10th of the month; “mid” indicates the period from the 11th through the 20th; “late” indicates the period from the 21st through the 31st. The word “diurnal” is included, in the few cases where it applies. The last item after (1) is the abundance-rating which I have given to the moth, based on its occurrence at the Univ. of Kansas Natural History Reservation during the period of March through May, in 1959, 1960, and 1961. The relative abundance of the moth is expressed by the letter A, B, or C, as follows: A = abundant. B+ B = moderate abundance (neither notably common or rare). B- C = scarce (6 or fewer records for all three years combined). Cl = only one record during the entire survey; complete date given. “B-f-” (= approaching “A”) and “B“” (= approaching “C”) are intended to show an inclination either towards abundance or scarcity; but the species is still better placed within the “B” category. Little use is made of these “B-f-” and “B— ” ratings, as 6(1): 1-18, 1967 SPRING MOTHS 5 they imply rather fine differences in relative abundance, which can hardly be ascertained during a short period of time, with only sporadic and limited collecting. Some of the species placed in the “C” category may actually be commoner, but it would take more (and consistent) collecting to indicate this, if it is the case. Other “C— rated” species are scarce in the spring, as they are just beginning to emerge in late May; but in some cases, these moths become increasingly abundant during June, and perhaps later in the summer. However, in this paper, their abundance- ratings are derived strictly, from their occurence during March, April, and May, and do not imply abundance for the year as a whole. Of course, some of the “C — rated” species are, indeed, truly scarce! But the possible complications should be kept in mind. There is little likelihood of error with those placed in the “A” category, as this degree of abundance is quite apparent to the collector within a two or three year period at the same locality. Some of the very earliest species to appear ( i.e. — late March — April) are single-brooded only, and will not fly again until the following spring. Thus, their flight-periods are completely cover- ed by this survey, and abundance-ratings are probably more accurate for these early species. Following (2), occasional foodplants are listed, but only when they are known to be the plants eaten by the larvae of these moths at the Univ. of Kansas Natural History Reservation. Gen- eral information on the foodplants of many of these moths may be found in the writings of Forbes (1920 - 1960), Jones (1951), Crumb (1956), Holland (1905), Peterson (1959), and J. H. Com- stock (1950). Follotving (3), are miscellaneous remarks, which may refer to the adult or to the early stages ( i.e. — variability; pecularities in behavior; short notes on the early stages; anything distinctive or worth noting). Under many species, (2) and/or (3) are not included. MISCELLANEOUS REMARKS The specimens collected during this spring survey of the Univ. of Kansas Natural History Reservation are chiefly in the authors collection, and in the Snow Entomological Collection, at the University of Kansas. Some are in the Los Angeles County Museum, and in the collection of W. R. Bauer and J. S. Buckett (Davis, California). 6 McFarland J. Res. Lepid. The following is a listing of the total numbers of species collected per family: 1. Sphingidae .... 10 7. Noctuidae ......121 2. Citheroniidae ......... .... 1 8. Notodontidae 18 3. Saturniidae .... 2 9. Lasiocampidae .... 2 4. Amatidae .... 1 10. Drepanidae ...... 1 5. Arctiidae .... 13 11. Geometridae ........ ....... 55 6. Agaristidae .... 1 12. Epiplemidae ........ 1 (TOTAL species. 226) More extensive and consistent collecting, over a period of several consecutive years (covering all months of the year), and sampling the various habitats on the Reservation, would perhaps bring the total to somewhere between 300 and 400 species of “Macroheterocera.” Inclusion of the “Microlepidoptera” would probably add another 200 (or more) species, and inclusion of the butterflies and skippers would probably add another 70 - 80 species to the total for the order Lepidoptera occuring on the Reservation. It is of interest to note the preponderance of scarce, uncommon, or only moderately common moths in this list. The Reservation is characterized by a moth fauna that is rich in number of species, but relatively few of these species are particularly abund- ant. This condition is usually seen in undisturbed woodland habitats, where successional changes (in the plant associations) are slow, and the plant species are many; it should become in- creasingly the case at the Reservation, as it continues to be pro- tected from disturbance. Woodland and forest associations ( other than those of mostly coniferous trees) usually have a rich moth fauna, with relatively few of the moths reaching great abundance; this is even more clearly demonstrated in cases where the trees and shrubs are of numerous species, none of which is overwhelm- ingly common. ( Tropical rain forests are an outstanding example of this condition, having great numbers of species — both plants and animals — but individuals of most of the species being un- common or widely-scattered ) . The following species (known to occur on the Reservation) have been preserved in the author’s collection of preserved early stages of Lepidoptera, and are catalogued under the code- numbers given in parentheses. SPHINGIDAE: Celerio lineata (Sp. 4), Cressonia juglandis (Sp. 13), Deidamia inscriptum (Sp. 1), Hemaris diffinis (Sp. 2) Jsogramma hageni (Sp. 5), Smerin- thus jamaicensis (Sp. 6); CITHERONIIDAE: Adelocephala 6(1): 1-18, 1967 SPRING MOTHS 7 bicolor (Ci. 1); SATURNDIIAE : Actias luna (St 1), Telea poly- phemus (St2); AMATIDAE: Scepsis fulmcollis (Am. 1); LITHOSIIDAE: Hypoprepia fucosa (Li. 1); ARCTIIDAE: Diacfisia virginica (Ar. 11), Ecpantheria deflorata (Ar. 8), Estigmene acmea (Ar. 2), Euchaetias egle (Ar. 7), Haploa lecontei ( Ar. 1 ) , Ma Isabella ( Ar. 10 ) ; AGARISTIDAE : Fsycho- morpha epimemis epimenis (AS. 1); NOCTUIDAE ;Acron!/cte oblinita (N. 20), Caenurgina crassiuscula (N. 18), Catahena Uneolata (N. 11), Heliothis zea (N. 4), Ipimorpha sp. (N. 27), Mamthyssa inficita (N. 30), Orthosia garmani (N. 26), O. ? alurina (N. 2), Palthis angulalis ? (N. 57), Rhodophora gaurae (N. 16), Schinia trifascia (N.15), Simyra henrici (N. 22), NO- TO'DONTIDAE: Cerura sp. (Nd. 11), Heterocarapa umhrata (Nd. 3), Melalopha inclusa (Nd. 2), Nadata gihhosa (Nd. 9), Schizura P ipomome (Nd. 13); LASIOCAMPIDAE; Epicnaptera americana (La. 4); GEOMETRIDAE: Ahhotana clemataria (G. 2), Anavitrinella pampinaria (G.24), Biston cognataria (G. 25), Calothysanis armturaria (G. 15), Catopyrrha coloraria (G. 13), Palaeacrita vernata (G. 1), Plagodis phlogosaria (G. 33). Many (but not all) of the above preserved inmatures were collected on or near the Natural History Reservation. In most cases, at least the last instar larva was preserved, and for some the complete life history was preserved ( all stages and all larval instars ) . ACKNOWLEDGEMENTS I wish to thank the following, whose assistance has been of great value: Dr. George W. Byers, of the Entomology Depart- ment, University of Kansas, for advice and assistance on various aspects of this paper, for reading the original manuscripts, and for taking the photographs of the two larvae illustrated in this paper. Dr. and Mrs. Henry S. Fitch, of the Zoology Department, University of Kansas, and residents at the Natural History Reservation, for permitting me to collect around their dwelling at all hours of the night, and for reading the original manuscript. Dr. John G. Franclemont, of the Entomology Department, Cornell University, for determination of over 90% of the 226 species collected. Dorothy Flenne, of Pearblossom, California, for typing the manuscript. Mr. Carl W. Kirkwood, of Summerland, California, for determination of several of the geometrids. Dr. Frederick H. Rindge, of the American Museum of Natural History, for determination or verification of 17 of the geometrids. 8 McFarland J. Res. Lepid. The National Science Foundation (Undergraduate Research Par- ticipation Fund), for financial assistance during this project. REFERENCES COMSTOCK, J. A. and N. McFARLAND (1961) -- Notes on the life history of Palthis angulalis Hubner ( Lepidoptera; Herminiinae ) . Bull. S. Calif. Acad. Sci. 60 (2): 112 — 116. COMSTOCK, J. H. (1950) — An introduction to entomology (9th ed. ). Ithaca, N. Y., Comstock Pub. Co. 1064 pp. CRUMB, S. E. (1956) — The larvae of the Phalaenidae. Washington, D. C. U. S. Dept, of Agric. Tech. Bull. no. 1135. 356 pp. FIELD, W. D. (1938) — A manual of the butterflies and skippers of Kansas, Bull., Univ. Kans. Biological Series, 39(10). 328 pp. FITCH, H. S. (1952) — The Univ. of Kansas Natural History Reservation. Univ. Kans. Mus. Nat. Hist., Misc. Publ., 4:1-38, 4 pis., 3 figs. FITCH, H. S. (1958) — Home ranges, territories, and seasonal movements of vertebrates of the Natural History Reservation. Univ. Kans. Publ. Mus. Nat. Hist., 11(3): 63-326. FITCH, H. S. and R. L. McGREGOR (1956) — The forest habitat of the Univ. of Kansas Natural History Reservation. Univ. Kans. Publ. Mus. Nat. Hist., 10(3) : 77-127. FLORA, S. D. (1948) — Glimate of Kansas. Kept. Kans. State Board ' Agric. 67, no. 285, pp. xii -j- 320. FORBES, W. T. M. (1920 - 1960) — The Lepidoptera of New York and neighboring states (4 vols.). Ithaca, N. Y., Gornell University. FRANGLEMONT, J. G. (1957) — The genus Euclidia, with the descrip- tion of a new species (Lepid., Noctuidae, Gatocalinae). Bull. Brooklyn Ent. Soc., LH (1): 5-15. HOLLAND, W. J. (1905) — The moth book. N. Y., Doubleday, Page, and Go. 479 pp., 48 pis. JONES, J. R. J. L. (1951) — An annotated check list of the Macro- lepidoptera of British Golumbia. Occas. Paper No. 1, Ent. Soc. B. C. 148 pp. McDUNNOUGH, J. (1938) — Gheck list of the Lepidoptera of Ganada and the United States of America, Part 1. Los Angeles, S. Calif. Acad. Sciences. 275 pp. PETERSON, A. (1959) — Larvae of insects, Part I (4th ed.). Ann Arbor, Mich., Edwards Bros. 315 pp. RINDGE, F. H. (1954) — A revision of the genus Toronos Morrison (Lepid., Geometridae ) . Bull. Amer. Mus. Nat. Hist. 104 (Art. 2): 181-236. RINDGE, F, H. (1956) — A revision of the American species of Deilinia (Lepid., Geometridae). Amer. Mus. Nov. No. 1810: 1-31. RINDGE, F. H. (1964) — ■ A revision of tlie genera Melanolophia, \ Pherotesia, and Melanotesia (Lepid., Geometridae). Bull. Amer. Mus. Nat. Hist. 126 ( Art. 3 ) : 241-434, 7 pis. STEVENS, W. G. (1948) — Kansas wild flowers. Lawrence, Univ. Kans. i Press. 463 pp. 6(1) : 1-18, 1967 SPRING MOTHS 9 THE ANNOTATED LIST OF SPECIES I. Family SPHINGIDAE Ampeloeca myron Cram. (1) Late Apr. - May; B (779) Celerio lineata Fabr. (799) (1) Late Apr. - May; B- Ceratomia sp. ( — -) (1) May 13, 1961; Cj Cressonia juglandis A. and H. (744) (1) Mid-late May; C (2) Jug Ians nigra (3) Seems to be more abundant in the city of Lawrence than on the Natural History Reservation. Deidamia inscriptum Harr. (782) (1) Late Apr. - early May; B (2) Vitis, Parthenocissus (3) This is the earliest and smallest nocturnal sphingid. Most specimens came in at dusk, or just after. The maximum number taken in one evening (at light) was 20 (Apr. 28, 1959). Only three were seen in I960 (May 2), and none in 1961. The period of flight is short, lasting only about two weeks (at the most), and the moth is only single-brooded. It appears to be quite cyclic from year to year. The larvae were abundant in May 1959, nearly stripping some of the Virginia Creeper vines that climb on tree trunks in the woods. Under these conditions, the larvae did not attempt to hide themselves. They were also common on wild grape, but harder to find by casual searching. A peculiar habit which the larva has (especially when young), is to throw its head clear back, thus pointing the true legs upward; at the same time it usually spits a droplet of fluid. This odd posture is assumed as a reaction to disturbance, and is quite the opposite of the usual sphingiform position. Hemaris diffinis Bdv. (770) (1) Apr. - May; diurnal only; A (2) Symphoricarpos orbiculatus, on the Reservation. (3) This is the earliest sphingid. The moth is sometimes attracted to redbud (Cercis) blossoms. The larva bears a caudal horn, like many other sphingids, and also has the general appearance of a "typical" sphingid larva. ( photograph of a last instar larva). It has a distinctive mode of locomotion, best described as hesitant, and "forward-inching", which allows it to move inconspicuously along, without drawing much attention to itself. Most of the time, it is quiet, and blends in rather well with the foodplant. The eas- iest way to obtain Hemaris larvae is to collect the eggs, which are laid singly on the \indersides of leaves, as the female moth hovers over the plant. Only an instant is needed for oviposition. Although the foodplant is abundant on the Reservation, the moths show a preference for ovipositing on plants which are in certain types of situations, as follows: (a) single plants, growing in open sunny places, or along roadsides; (b) groups or short rows of plants, growing in sxinny clearings in the woods, or at the bases of limestone outcroppings (in the sun ). The moths seem to avoid ovipositing on plants that form the shaded understory, where Symphoricarpos grows abundantly beneath trees. Isogramma hageni Grt. (705) (1) May; A (2) Madura pomifera (3) The larvae can be foxind on Osage orange, in the late summer. They were more common on small trees, out in open grassy fields near the Reservation headquarters. An unusual feature of the larval behavior is that they readily let go of the plant and drop if disturbed; most sphingid larvae hang on tenaciously when disturbed. 10 McFarland J. Res. Lepid. Paonias myops A. and S. (742) (1) Early - mid May; B- Smerinthus jamaicensis Dry. (739) (1) May 20, 1961; Cl (2) Ulmus americana (3) The single specimen was reared from a larva collected in September, I960, by Mrs. Fitch, near the Reservation headquarters. Sphinx ? drupiferarum A. and S. (7 30) (1) May; B- II. Family CITHERONIIDAE Adelocephala bicolor Harr. (849) (1) Late Apr. - mid May; B + (2) Gleditsia triacanthos (3) As is the case with quite a few of the moths in this list, a summer brood follows. The first indi- viduals (in spring) have deep gray forewings, whereas most of the summer brood moths have tan (or light golden-brown) fore- wings; the difference in general appearance is striking. III. Family SATURNIIDAE Actias luna L. (811) (1) Late Apr. - early May; B+ (2) Juglans nigra (3) Fifteen individuals came to light on Apr. 28, 1959; this was a maximum count for any single night. Telea polyphemus Cram. (812) (1) Late Apr. - May; B (2) Quer- cus, etc. IV. Family AMATIDAE Scepsis fulvicollis Hbn. (871) (1) May; diurnal and nocturnal; B- (2) Bromus and other grasses (3) Although not common in the spring, it is abundant in Sept. , on Solidago blossoms; it is also more common at light in late summer. A larva was collected on Bromus, in Sept. , I960. The hairy larva could be mistaken for an arctiid, but there are no strictly grass-feeding arctiids in this locality. V. Family ARCTHDAE Apantesis arge Dru. (1038) (1) Late Apr. ; C^ Apantesis nais Dru. (1058) (1) May; B Apantesis phalaerata Harr. (1061) (1) May; A (3) No females were ever taken at the lights. Color and maculation are quite variable on the hindwings, and to a lesser degree on the forewings. Cycnia ? inopinatus Hy. Edw. (992) (1) May; C Cycnia tenera Hbn. (991) (1) May; B Diacrisia virginica Fabr. (=Spilosoma) (1065) (1) Late Apr. May; A (3) This is the first arctiid to appear in the spring. Estigmene acraea Dru. (1070) (1) May; C Euchaetias egle Dru. (1001) (1) Late May; larvae were collected in Lawrence (in Sept. ) and the adults emerged the following May; to be looked for at the Natural History Reservation (2) on Gono- lobus, in Lawrence. Euchaetias oregonensis Stretch (1005) (1) May; C Halysidota tessellaris A. and S. (984) (1) May (one specimen collected inLpwrence; to be looked for at the Natural History Reservation). 6(1) : 1-18, 1967 SPRING MOTHS 11 Haploa lecontei Guer„ (1103) (1) Late May; diurnal and nocturnal; B+ (3) The moths are highly variable^ from immaculate white to white heavily marked with chocolate-brown on the forewings. The nearly mature larvae are common in early May, especially west of the Rockefeller Tract, where they climb into Ceanothus Ovatus , Symphoricarpos, small box elders^ and other plants, to feed and sit in the sun, Hyphantria cunea Dru. (1075) (1) May; A (3) This species is highly variable in maculation, from immaculate white to heavily- spotted with black. It is the smallest white arctiid in this locality, Isia Isabella A. and S. (1069) (1) May; C (3) This species is more abundant in open fields and roadside habitats, where the familar hairy larvae (banded in black, rust-brown, and black) are seen crawling over the ground in Sept. - Oct. After overwintering as mature larvae, they spin cocoons in the early spring and emerge in May and June. VI. Family AGARISTIDAE Psychomorpha epimenis Dru, (2864) (1) Apr.; diurnal only; B (2) Vitis, Parthenocissus (3) This moth is on the wing just as the buds are beginning to swell and open, in early spring. It is fairly common on mild, sunny days. Although not often inclined to alight, it sometimes comes down to drink on damp or wet ground. Numerous eggs were obtained from a confined female, by feeding her daily and giving her some filtered sunlight each morning, along with sprigs of foodplant. A preference was shown for ovipositing on or under the stipules of the leaves. VII. Family NOCTUIDAE Acontia aprica Hbn. (3203) (1) May; C Acronycta connecta Grt. (1167) (1) Early May; C (3) A melanic specimen was captured. Acronycta ? hastulifera A, and S. (1149) (1) Early May; C Acronycta impleta Wlk. (1201) (1) Late Apr. - early May; A (3) As in some other Acronycta spp. , melanic individuals are common. Acronycta longa Gn. (1207) (1) May; B- Acronycta morula G. and R. (1181) (1) Early May; C (3) This species is much more common in the town of Lawrence. Acronycta oblinita A. and S. (1215) (1) Mid Apr. - early May; B (3) The bristly, colorful black and yellow larvae are sometimes common on Polygonum (in roadside ditches, etc. ) in Sept. Acronycta pauper cula Grt. (1169) (1) Late Apr. - early May; B (3) This species is more common in Lawrence. Agriopodes teratophora H. -S. (2582) (1) mid - late May; C Agrotis ypsilon Rott. (1435) (1) Late Apr. ; B Amathes c-nigr\im Linn. (1511) (1) May; C Anagrapha falcigera Kby. (3252) (1) Apr. - mid May; B Anepia capsular is Gn. (1804) (1) Early - mid May; C Anorthodes tarda Gn. (2651) (1) May; B- Argyrostrotis anilis Dru. (3458) (1) May; C Autographa biloba Steph. (3279) (1) May; B (3) This species is more common in Lawrence. 12 McFarland J. Res. Lepid. Baileya australis ■ Grt. (3240) (1) Early - mid May; A Baileya dormitans Gn. (3241) (1) Early - mid May; C Baileya ? ophthalmica Gn. (3239) (1) May 2^ 1959; Balsa malana Fitch (2669) (1) May; B+ (3) More common in 1959 than in i960 or 1961, Bendis detrahens Wlk. (3527) (1) Early - mid May; C (3) This species is more common in Lawrence. Bleptina caradrinalis Gn. (3797) (1) May; B (3) This species is more common in Lawrence. Bomolocha albalienalis Wlk. (3691) (1) Mid May; C Bomolocha bijugalis Wlk. (3689) (1) Mid - late May; C (3) Sexual dimorphism is noticeable in this species, Bomolocha sordidula Grt. (3694) (1) Mid May; C Caenurgina crassiuscula Haw. (3430) (1) Apr. - early May; diurnal and nocturnal; A Caenurgina erechtea Cram. (3431) (1) Late Apr, - May; diurnal and nocturnal; B- Catabena lineolata Wlk. (2737) (1) Apr. - early May; C (2) Verbena sp. __ Cerastis tenebrifera Wlk. (1495) (1) Early - mid Apr.; B Cerma cora Hbn. (2559) (1) One collected inLawrence^ May 12, 1961; to be looked for at the Natural History Reservation. Chamyris cerintha Treit. (3131) (1) One collected in Lawrence, May, I960; to be looked for at the Natural History Reservation. Chorizagrotis auxiliaris Grt. (1387) (1) Late Apr. - early May; B Chytolita morbidalis Gn. (3766) (1) Mid May; C Cissusa spadix Cram. (3539) (1) Mid-late Apr. ; C Crambodes talidiformis Gn. (2661) (1) Late Apr. - early May; B Crocigrapha normani Grt. (1950) (1) Apr. - early May; B+ (3) The peak of flight lasts only a week or less. Delta ramosula Gn. (2589) (1) Mid - late Apr.; C Diptergia scabriuscula L. (2587) (1) May 12, 1959; Cl Elaphria festivoides Gn. (2647) (1) May; B Elaphria grata Hbn. (2650) (1) Late Apr. - May; A Erastria bellicula Hbn. (3117) (1) May; C Erastria carneola Gn. (3124) (1) May; B (3) This species is more common in Lawrence, Erastria muscosula Gn. (3118) (1) Mid - late May; C Erastria musta G. and R. (3123) (1) May 18, 1959; Cj Erastria synochitis G. and R. (3120) (1) May; B Epizeuxis americalis Gn. (3734) (1) Mid May; C (3) May be more common than indicated, as members of this genus are not readily attracted to lights , and they often leave after coming in. Epizeuxis lubricalis Geyer (3746) (1) Late May; C (3) Adults vary considerably in size. Euagrotis illapsa Elk. (1483) (1) May; C Euclidia cuspidea Hbn. (3426) (1) May 5, I960; diurnal only; C^ (3) At first glance, the diurnal adult (when on the wing) could be mistaken for a skipper of the genus Srynnis (dusky- wings ), but the skippers have a more erratic, darting flight. Euparthenos nubilis Hbn. (3414) (1) May 11, 1961; C^^ Eupsilia morrisoni Grt, (2305) (1) Mid Feb. - early Apr. ; B- Eutolype electilis form depilis Grt. (— ) (1) Mid - late Apr. ; B+ (3) This species was fairly common in 1961, but not in 1959 or 1960. 6(1): 1-18, 1967 SPRING MOTHS 13 Eutolype grandis Sm. (2194) (1) Late Mr. - early Apr.; B (3) The period of flight lasts only a few nights, but the moth is not uncommon during the limited period that it is on the wing, Faronta diffusa Wlk. (1962) (1) Late Apr. - early May; B Galgula partita Gn. (2666) (1) Apr. - May; B (3) This species is more common inLawrence. Its color is quite variable, from pale tan to rich, dark brown. Heliothis zea Boddie (corn-ear worm) (2932) (1) May; C Himella intractata Morr. (1840) (1) Early - mid Apr. ; B Homohadena infixa Wlk. (2153) (1) May 28, 1959; Cj Homorthodes lindseyi Benj. (1890b) (1) May 18, 1959; Hypena humuli Harr. (3706) (1) Early - mid Apr. ; C Hypsorophora hormos Hbn, (3611) (1) One collected in Lawrence, May 26, 1959; to be looked for at the Natural History Reservation. Isogona tenuis Grt. (3643) (1) Mid - late May; B- Lacinipolia anguina Grt. (1716) (1) Mid May; C Lacinipolia renigera Steph. (1738) (1) May; B (3) Although common in 1959, it was scarce in I960 and 1961. Lascoria ambigualis Wlk. (3805) (1) May; C (3) Sexual dimorphism is notable; the male has a cleft forewing, but the female does not. Legna perditalis Wlk. (3685) (1) One collected in Lawrence, May 25, 1959; to be looked for at the Natural History Reservation. Leucania phragmitidicola Gn. (1978) (1) May, especially early May; Leucania ursula Fbs. (1974) (1) May; B Leuconycta diphteroides form obliterata Grt. (2576) (1) Mid - late May; C Lithophane antennata Wlk. (2242) (1) Apr. 6, I960; Marathyssa inficita Wlk. (3223) (1) May 13, I960; (2) Rhus trilobata Matigramma pulverilinea Grt. (3506) (1) Mid May; C Melipotis indomita Wlk. (3549) (1) May 23, I960; Cj Melipotis jucunda Hbn. (3551) (1) One collected in Lawrence, mid May, I960; to be looked for at the Natural History Reservation. Mocis texana Morj-. (3435) (1) May; B Morrisoriia confusa Hbn. (1904) (1) Apr. - early May; C Morrisonia distincta Hbn. (1671) (1) Late Apr. - early May; B + Neoerastria apicosa Haw. (3126) (1) May 18, 1959; Cj Ogdoconta cinereola Gn, (2773) (1) May; C Orthodes crenulata Butl. (1871) (1) May 18, 1959; Cj Orthosia alurina Sm, (1941) (1) Apr. ; A Orthosia garmani Grt. (1938) (1) Early - mid Apr. ; A (3) The flight period is extremely short, with the moths at a "peak" for only one or two nights, at which time they are abundant. Largest numbers were seen on April 4, 5, and 6, 1959 and April 11, I960. Larvae, reared from eggs, proved to be general feeders on various woody plants, including Ulmus and Cornus. Orthosia hibisci Gn. (1943) (1) Apr.; A Orthosia rubescens Wlk. (1937) (1) Apr.; B- Palthis angulalis Hbn, (3807) (1) May; B (3) This species is more common in Lawrence. At the Natural History Reservation, larvae (Sept. - Oct. ) were feeding on the dead, dried, papery seed cap- sules of Lobelia syphilitica, which they closely match in colora- tion. The larvae are quite inactive, and slow in their movements, resting quietly on the stem below the empty seed capsule, or on 14 McFarland J. Res. Lepid. the capsule itself. They are multiple- hr ooded^ and have been re- corded from many other foodplants. See Comstock and McFarland (1961). Panopoda carneicosta Gn, (3535) (1) May 28, 1959; Panopoda refimargo Hbn. (3534) (1) May 21, I960; Ci (3) As is the case with quite a few of the moths listed for late May only, P. r ufimargo is barely represented as its flight is just starting, but it becomes more abundant in June. Fhrallelia bistriaris Hbn. (3422) (1) One collected in Lawrence, mid-May, 1959; to be looked for at the Natural History Reser- vation. Peridroma margaritosa Haw. (1946) (1) Late Apr. - early May; B + Perigea xanthioides Gn. (2610) (1) Mid May; C Phalaenophana pyramusalis Wlk. (3750) (1) May; C Phalaenostola larentioides Grt. (3769) (1) May 28, 1959; Phiprosopus callitrichoides Grt. (3639) (1) May 24, 1961; Cj Phoberia atomaris Hbn, (3545) (1) Late Mar. - mid Apr.; A Phosphila miselioides Gn. (2549) (1) Mid - late May; C Plathypena scabra Fabr, (3705) (1) Mid May; B Platysenta vecors Gn. (2620) (1) Late Apr. - early May; B Platysenta videns Gn. (2613) (1) Apr. - May; B + Plusia aerea Hbn. (3296) (1) One collected in Lawrence, May 25, 1959; to be looked for at the Natural History Reservation. Plusiodonta compressipalpis Gn. (3612) (1) Late Apr. - early May; B (3) This moth is more common in Lawrence, Polia adjuncta Bdv. (1691) (1) May 23, I960; Prodenia ornithogalli Gn. (2678) (1) Late Apr. - early May; C Proxenus miranda Grt, (2662) (1) Mid - late May; B (3) The forewings have a distinct glossy sheen. Psaphida grotei Morr. (2191) (1) Early - mid Apr.; B Pseudaletia unipuncta Haw. (1994) (1) Late Apr, - May; A Psychomorpha epimenis (see AGARISTIDAE) Pyrrhia umbra Hufn. (2715) (1) A few collected in Lawrence, late May; to be looked for at the Natural History Reservation. Raphia abrupta Grt. (1140) (1) Late Apr. - mid May; B + Salia interpuncta Grt. (3731) (1) May 11, 1961; Sericaglaea signata French (2298) (1) Mid - late Apr.; C Simyra henrici Grt. (1222) (1) Late Apr. , C (3) One of the color- ful, bristly larvae was collected on Sorghum, in a roadside ditch (early Oct. ). Spiloloma lunilinea Grt. (3595) (1) Mid - late May; B- Stiriodes obtusa H. -Sch. (2832) (1) May 28, 1959; Cj Tarachidia candefacta Hbn, (3176) (1) Mid May; C Tarachidia erastrioides Gn. (3172) (1) Mid - late May; B Trachea delicata Grt. (2547) (1) May 11, 1961; C^ (3) The fore- wings are marked with olive green, which often discolors in the killing jar. Trichoplusia ni brassicae Hbn. (3269) (1) Late Apr. - early May; C Ulolonche culea Gn. (1831) (1) May; C Xanthoptera nigrofimbria Gn. (3113) (1) Mid - late May; B- (3) As is the case with some of the other species listed, this moth is abundant later in the year, although uncommon in its spring brood. Zale galbanata Morr. (3478) (1) Apr. - May; C Zale ? minerea Gn. (3484) (1) Late May; C Zanclognatha cruralis Gn. (3762) (1) Collected in Lawrence, mid- late May; to be looked for at the Natural History Reservation. 6(1):1~18, 1967 SPRING MOTHS 15 Vni. Family NOTODONTIDAE Dasylophia anguina a. and S. (3880) (1) May 13, I960; Cj Gluphisia septentrionalis Wlk. (3939) (1) May; B- Heterocampa bilineata Pack. (3908) (1) May; C Heterocampa guttivitta Wlk. (3907) (1) Early - mid May; C Heterocampa sp. (- — — ) (1) May 23, 1959; Heterocampa umbrata Wlk. (3902) (1) May 29, I960; (3) This single record is from a pupa, collected under a log (Oct. ), about 5 mi. WSW of the Natural History Reservation. A female emerged on the date given above. Hyperaesehra georgica H. -S. (3845) (1) May 14, 1959; Ci Lophodonta angulosa A. and S. (3854) (1) One collected in Lawrence, late May; to be looked for at the Natural History Reservation. Melalopha albosigma Fitch (=Ichthyura a. ) (3827) (1) Mid Apr. early May; B Melalopha apicalis Wlk. (3822) (1) May 4, 1959; Melalopha inclusa Hbn. (3825) (1) Early - mid May; B (3) A number of larvae were collected on Populus (in Sept. ), near the Reservation headquarters; these emerged in May, 1961. The lar- vae are gregarious, in a web-nest of two or more leaves. Misogada unicolor Pack. (3909) (1) Mid May; C Nadata gibbosa A; and S. (3857) (1) May 11, 1961; Cj (3) This moth was often seen in Lawrence. Nerice bidentata Wlk. (3858) (1) Late Apr. - early May; B + (3) The period of flight is rather short. Oligocentria lignicolor Wlk, (3912) (1) May; C Schizura apicalis G, and R. (3926) (1) Mid - late May; C Schizura ipomoeae Dbldy. (3920) (1) May 21, I960; Cj Schizura unicornis A, and S. (3924) (1) Late Apr. - mid May; C IX. Family LASIOCAMPIDAE Epicnaptera americana Harr. (3999) (1) Apr. - early May; B Heteropacha rileyana Harv. (3998) (1) Apr.; B+ (3) The peak of abundance lasts only a few nights. Only two or three females came to the lights during the entire survey. X. Family DREPANIDAE Eudeilinea herminiata Gn. (4017) (1) May; C (3) There are pure white geometrids for which this moth could be mistaken, XL Family GEOMETRIDAE Abbotana clemataria A. and S. (5199) (1) Apr. - early May; B+ (2) Fraxinus, etc, ( 2, photograph of a last instar larva). Anacamptodes defectaria Gn. (4915) (1) Mid - late Apr.; C Anavitrinella pampinaria Gn. (4908) (1) Late Apr. - May; A Apicia confusaria Hbn. (5184) (1) Fairly common during mid - late May, in Lawrence; to be looked for at the Natural History Reservation. Bapta vestaliata Gn. (4606) (1) Late Apr. - May; duirnal and noc- turnal; B+ (3) As this small geometrid is pure, shining white, it is very conspicuous when flying in the daytime. 16 McFarland J. Res. Lepid. Biston cognataria Gn. (4968) (1) May 24, 1961; Cj Calothysanis amaturaria Wlk. (4205) (1) Mid - late May; B- (2) Rumex crispus (3) The very unusual dark brown larvae are considerably swollen in the thoracic region. They make no attempt to hide on their foodplant, and are easily collected by searching. Catopyrrha coloraria Fabr. (4642) (1) Late Apr. - mid May; diurnal and perhaps nocturnal; B+ (2) Ceanothus ovatus (3) This moth was encountered only in a grassland-field, directly NW of the Natural History Reservation, in places where Ceanothus was growing amongst the grasses. In this restricted habitat, for a short period, it was rather common. The moth rests on the ground, with its wings up over its back. It flies up when approached, but lands again, a short distance away. The behavior is reminiscent of Sericosema spp. A large number of larvae were reared from eggs in 1961. Although offered many different plants from the habitat, they accepted only Ceanothus. Chlor ochlamys chloroleucaria Gn. (4095) (1) Late Apr. - mid May; _ Cleora sublunaria Gn. (4876) (1) Apr. - mid May; A Coryphista meadi Pack. (4248) (1) Two records for early May, 1959, in Lawrence; the moth might occasionally turn up at the Natural History Reservation, although its only recorded foodplant is Berberis ( = Mahonia). Deilinia quadrifasciaria Pack. (4611) (1) May 28, 1959; Dichorda iridaria Gn. (4082) (1) May; C Dyspteris abortivaria H. -S. (4234) (1) May; B- Ectropis cr epuscularia Schiff, (4946) (1) Mid Apr. - early May; B Enconista dislocaria Pack. (= "Semiothisa" d. ) (4703) (1) Late Apr. - early May; B Euacidalia sericeata Pack. (4109) (1) Mid - late Apr.; C (3) This is the smallest spring geometrid collected on the Natural History Reservation. Euchlaena pectinaria D. and S. (5003) (1) Late Apr. - mid May; C Eugonobapta nivosaria Gn. (4991) (1) May; C (3) This moth is of the same size and color as Eudeilinea herminiata Gn, , a dre- panid, which also occurs on the Natural History Reservation. Eumacaria latifer rugata Wlk. (=Itame 1. ) (4774) (1) Mid Apr. mid May; C Euphyia centr ostrigaria Woll. (4559) (1) Late Apr. - May; B Eupithecia herefordaria C. and S. (4310) (1) Apr. - May; B + Eupithecia swetti Grossb. (4354) (1) Apr. - May; B Haematopis grataria Fabr. (4204) (1) Collected during May, in Lawrence; to be looked for at the Natural History Reservation. (3) Remarks under Xanthoptera nigrofimbria (Noctuidae) apply here, as well as to a number of the other geometrids listed. Heter ophleps refusata Wlk. (4231) (1) May; C Heterophleps triguttaria H. -S. (4233) (1) Mid - late May; C Horisme intestinata Gn. (439 3) (1) Abundant in Lawrence, in mid to late May; to be looked for at the Natural History Reservation, Hydriomena pluviata Gn. (4477) (1) One collected in Lawrence, Apr. 26, I960; to be looked for at the Natural History Reservation. Hyperetis amicaria H. -Sch. (5043) (1) Late Apr. - May; B- Lobocleta os s ularia Hbn. (4115) (1) One collected in Lawrence, in mid May; to be looked for at the Natural History Reservation. 6(1): 1-18, 1967 SPRING MOTHS 17 Lycia ypsilon S. A. Forbes (4952) (1) Apr.; A (3) The moth ““r^aches'a peak of considerable abundance in one or two nights, and then rapidly disappears until the following April. On Apr. 6, 1959, over 200 individuals came to one light. No females were ever taken at the lights. MelanolopMa signataria signataria Wlk. (4857) (1) Apr. - early “‘■■'■Ma^rT^yT^'FeTnacuIaH^ variable. Mellilla xanthometata Wlk. (4657) (1) Apr. ^ May; sometimes on "“The^ingTnAhe^aytim although primarily nocturnal; A Metanema inatomaria Gn. (5054) (1) Mid - late May; C Metarranthis duaria Gn. (5050) (1) Early - mid May; C (3) This moth may be more common than indicated, as individuals have a tendency to leave the immediate vicinity of the light after once flying in. Metarranthis ? homuraria Grt. (5046b) (1) Mid May; C Palaeacrita merriccata Dyar (4961) (1) Early Apr. ; C Palaeacrita vernata Peck (4960) (1) Jan. - Mar. - Apr. - mid May; A+ (2) Ulmus americanus, U. rubra, etc. (3) The males occasionally come to lights (on mild nights) in Jan, and Feb. , and they become extremely abundant in March and April. Periodically, the larvae of this moth defoliate the elms, shortly after they leaf out, but the trees usually recover (with a secondary crop of leaves) by late' May, at which time the larvae have dropped to the ground to pupate. Percnoptilota obstipata Fabr. (4535) (1) Mid Apr. - May; B- (3) Sexual dimorphism is notable. Pero ? honestarius Wlk. (5072) (1) Apr. - early May; B Phigalia olivacearia Morr. (4956) (1) Apr. (particularly early Phigalia titea Cram. (4958) (1) Apr. (particularly early Apr. ); A Philobia aemulataria Wlk. (4665) (1) Late Apr. - early May; B Plagodis phlogosaria Gn. (a spring form) (5040) (1) Late Apr. - mid May; C Scopula inductata Gn. (4158) (1) Mid - late May; C Scopula limboundata Haw — (1) Mid - late May; B (3) The maculation is variable. Semiothisa continuata Wlk. (4710) (1) May; C Semiothisa gnophosaria Gn. (47 38) (1) Mid - late May; C Semiothisa ocellinata Gn. (4723) (1) May; C Synchlora aerata Fabr. (407 0) (1) Two specimens collected in Lawrence, in late May; to be looked for at the Natural History Reservation. T etracis crocallata Gn. (5197) (1) Mid - late May; B T ornos abjectarius calcasiata C. and S. (— -) (1) Mid Apr. - mid May; B- Tornos scolopacinarius scolapacinarius Gn. (4812) (1) Mid Apr. mid May; B Xanthotype ? sospeta Dru. (5007) (1) Mid - late May; B= (3) This moth is more common in Lawrence, although not abundant. Xystrota rubromarginaria Pack. (4134) (1) Apr, - mid May; diurnal and nocturnal; A (3) This is a common diurnal spring geometrid on the Reservation, but it also comes to lights after dark. Xn. Family EPIPLEMIDAE Callizzia amorata Pack. (5223) (1) May; C 18 McFarland /. Res. Lepid. Fig. 1 — Hemaris diffinis (last instar, alive and feeding) on foodplant, Sym- phoricarpos orhiculatus. Univ. Kansas Nat. Hist. Res., 7 mi. N.E. Lawrence, Douglas Co., Kansas. May, 1960. Reared from egg. Fig. 2 — Abhotana clemataria ( Geometridae), last larval instar, alive. Rear- ed from egg on Fraxinus at Lawrence, Douglas Co., Kansas. A general feeder on many trees and shrubs. Journal of Research on the Lepidoptera 6(1) : 19-21 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 A NEW SPECIES OF NEPTICULA ON BUR OAK IN ONTARIO (NEPTICULIDAE) T. N. FREEMAN Entomology Research Institute, Canada Department of Agriculture, Ottawa During the past few years some officers of the Forest Insect Survey of Canada, Department of Forestry and Rural Develop- ment, have been studying an undescribed species of Nepticula V. Heyd. which is abundant on bur oak in the Lake Simcoe District of Ontario. The description is presented here to assist those officers in publishing the results of their investigations. Nepticula macmcarpae sp. n. General. Face black. Vertex creamy yellow. Antenna black- ish at base, gradually becoming ochreous toward apex. Eye-cap, thorax, base of fore wing, and median wing fascia creamy yellow. Median fascia broad, occupying about one-third of wing just before middle, and slightly wider on coastal and trailing mar- gins. Remainder of wing brownish-black; apical fringe whitish; trailing fringe fuscous. Hind wing and fringe fuscous. Abdo- men dull steel-grey dorsally, creamy yellow ventrally. Legs creamy yellow. Wing spread 4 mm. There are two generations each year. The moths occur in early June, and again in late July and early August. Male genitalia (Fig. 1). Uncus heavily sclerotized, bilobed. Gnathos heavily sclerotized, M-shaped. Vinculum broadly arcu- ate laterally, truncate ventrally. Clasper broad, with apical tooth. Transtilla bifid, each arm subtriangular. Aedeagus broad, short, tubular; cornutus broad, arcuate. 19 20 T. N. FREEMAN /. Res. Lepid. Female genitalia (Fig. 2). Ovipositor lobes short, subconical. Apophyses short, thick. Ductus bursae membranous. Bursa with two bands of short stout teeth. Mine. Upper side serpentine, about 2 cm. long and with a central frass line. Holotype. Male, Orillia, Ontario, 12 August 1965. Forest Insect Survey No. S65-4100-01. No 9539 in the Canadian National Collection, Ottawa. Paratypes. Seven males, thirteen females, same locality as holotype, August 12, 15, 30, 1965. All type material reared from Quercus macroearpa Michx. Distribution. Known only from Orillia, Ontario, but probably occurs elsewhere within the distribution of the host tree. Remarks. This species will key to latifasciella Chambers in Braun’s key in Forbes (1923, Cornell Univ. Agr. Exp. Sta. Mem. 68: 84). 1 have examined Chambers’ type in the Museum of Comparative Zoology, Cambridge, Mass. It is a female securely glued to a paper point, and except for the thorax, is in good condition. There are several differences between it and macro- earpae. Firstly, Chambers collected his specimen in Kentucky from the bark of a sweet chestnut tree {Castanea dentata (Marsh.) Borkh. ), the leaves of which contained many empty Nepticula mines. Secondly, there are significant colour differ- ences as outlined below. N. latifasciella. Face rusty-yellow. Vertex dark brown. An- tenna silvery. Abdomen fuscous dorsally, pale greyish fuscous ventrally. Legs dark fuscous. N. macrocarpae. Face black. Vertex creamy yellow. Antenna blackish basally, ochreous apically. Abdomen steel-grey dorsally, creamy yellow ventrally. Legs creamy yellow. 6(1) : 19-21, 1967 NEPTICULA 21 Figs. 1, 2.— Genitalia of Nepticula macrocarpae sp. n. 1, male. 2, female. Journal of Research on the Lepidoptera 6(1) : 22, 1967 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 FIXATION OF THE TYPE LOCALITY OF LYCAENA PHLAEAS HYPOPHLAEAS (BOISDUVAL) AND A FOODPLANT CORRECTION OAKLEY SHIELDS 5151 Alzeda Drive, La Mesa, California C. F. DOS Passos ( IN LiTT. ) POINTED OUT that the type locality of Lycaena phlaeas hypophlaeas listed as “north of California” (J. Res. Lep. 5: 231) should instead be translated “Northern California.” The type(s) was collected by P. J. M. Lorquin in 1852. Lorquin's California explorations appear in a mimeograph- ed pamphlet, “Pierre Joseph Michel Lorquin — Naturalist-voy- ageur,” by Estelle H. Lorquin, published by the Lorquin Service in San Francisco in 1938. L. M. Martin (per. communication) of the Los Angeles County Museum determined where Lorquin collected during 1952-1853 in relation to the pamphlet text. During that time, Lorquin “explored at first all the neighborhood of San Francisco, then the banks of the Sacramento and Plumas rivers; made trips in the Sierra Nevada chain, and ventured as far as the great forests of the interior.” In view of what is known about the California distribution of hypophlaeas, the probable type locality is the Sierra Nevada Mountains, California. F .M. Brown ( in litt. ) says that the type locality could well have been the gold region in Tuolumne County. T. Iwase pointed out (in litt.) that the record of daikon by Yokoyama as a foodplant for Lycaena phlaeas daimio in Japan was a misprint. He says there are five foodplants recorded in Japan: Riimex japonicus, R. acetosa, R. acetosella, R. obtusi- folius, and R. domesticns. Thus the known foodplants for L. phlaeas subspecies are all in Polygonaceae—daikon (Raphanus) previously being the only exception. 22 Journal of Research on the Lepidoptera 6(1) : 23^30 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 DESCRIPTION OF A NEW SPECIES OF XYLOMIGES FROM CALIFORNIA WITH NOTES AND ILLUSTRATIONS (LEPIDOPTERA: NOCTUIDAE: HADENINAE) JOHN S. BUCKETT Systematic Entomologist, Bureau of Entomology, California Department of Agriculture, Sacramento, Calif. It is my pleasure at this time to present a description of an undescribed noctuid species of the genus Xylomiges Guenee. This species has been in the Bauer-Buckett collection for well over ten years now, and in the interim many specimens have accumulated. This species has a peculiar flight habit, and be- cause of this characteristic, males of the species were never taken until relatively recent times. The females come out in the evenings, usually between 7:30 P.M. and 8:00 P.M., and can be secured until 10:00 P.M. on normal nights. The males have never been taken before 10:00 P.M., and become abund- ant about 11:00 P.M. and later on normal occasions. The tem- peratures along the coast are cool in the early part of the year (January and February), and the humidity is very high, and therefore, collectors seldom would stay out until 10:00 P.M. collecting. Because of this factor, males were never taken until recently when colleagues and I remained out all night collecting. The type locality is a typical coastal Pine belt vegetation, the dominant tree being Bishop Pine (Finns muricata Don.). Ma- drono (Arbusus menziesii Pursh. ), Tanbark (Lithocarpus densi- florus (H. & A.)), California Laurel (Umbellularia calif ornica (H. & A.)), and Coast Live Oak (Quercus agrifolia (Nee.)), are other common trees of the area. Common shrubs of the area are various species of Ceanothus, Coyote bush (Baccharis pilularis DC.), Manzanita (Arctostaphylos spp.). Huckleberry 23 24 J. S. BUCKETT J. Res. Lepid. Fig. 1 — Map depicting the distribution of Xylomiges baueri Buckett and X. variabilis (Smith). Note the restricted distribution of baueri. 6(1) : 23-30, 1967 XYLOMIGES 25 ( Vaccinium spp. ) , and Salmon-berry ( Rubus spectabilis Pursh. ) . Many other smaller herbaceous plants abound also. While in the company of Mr. Bill Bauer and Mr. Chris Henne, search for the eggs and larvae was done. The only larva we found that resembled one of the Xylomiges species was one found in the tip of staminate cone of a young Bishop Pine. The larva was in an intermediate instar, and as yet has not emerged (aceord- ing to Mr. Chris Henne who is rearing the specimen). It seems probable that Bishop Pine will prove to be the host plant of this new species. Xylomiges variabilis Smith, the closest relative to Xylomiges baueri n. sp., is a pine feeder, and thus, this is the reason for suspecting Bishop Pine in this case. XYLOMIGES BAUERI n. sp. Male: Ground color of primaries dorsally maroon, secondaries dorsally with pink tint. Head with vertex clothed in tricolor, elongate hairs, basally brown, subapically maroon, apically white-tipped; frons clothed as in vertex, for most part, but with more maroon elongate hairs; maxillary palpi exterolaterally brownish, ventrally possessing elongate white-tipped maroon hairs; proboscis normal; eyes very hairy; antennae with scape and pedicel clothed in whitish and maroon scales and hairs; flagellomeres bipectinate-fasciculate, pectinations becoming shorter apically, terminally ciliate. Thorax with collar maroon, composed of elongate white-tipped hairs; disc and tegulae cloth- ed in brown, maroon, white-tipped dentate hairs; ventrally clothed in maroon and whitish elongate hairs; tarsi black and white banded; ungues very weakly bifid; primaries dorsally maroon; basal line bicolor, basally black, apically irrorated with white scales so as to appear grey, or blue-grey; transverse an- terior line uneven in course, bicolor, basally blue-grey, apically black; median area blackish on inner margin, remainder maroon; orbicular squarish, grey, centrally blackish, outlined with light colored scales, anastomising with reniform; reniform centrally maroon, outlined in creamy scales; transverse posterior line undulating, represented as line of demarcation between median and subterminal areas; subterminal area bicolor, basally grey, apically cream yellowish; subterminal line jagged, represented as line of demarcation between cream yellow coloration and grey terminal area; terminal area with transverse black dashes on veins; fringes composed of spatulate white-tipped scales, alternating maroon and brown from apex to point of inner margin, or tornus; inner margin basally and medially with 26 J. S. BUCKETT J. Res. Lepid. Fig. 2-— Holotype male, Xylomiges baueri. Inverness, Marin County, Cali- fornia, 6 February 1965 ( J. S, Buckett). Fig. 3 — -Allotype female, X. baueri. Same locality as holotype, 2 February 1962 (J. S. Buckett and G. M. Trenam). 6(1) ; 23-30, 1967 XYLOMIGES 27 fringe of elongate maroon hairs; ventral surface dark, dusky, costally and apically, basally dirty white, overlain with tint of maroon; dark spot present in area of reniform; secondaries dor- sally with pink tint; discal dot dusky, prominent; terminal line dusky; veins slightly outlined in fuscous; fringes a pinkish- maroon; ventral surface pinkish basally; remainder of surface a dirty white, overlain with hue of rose-colored scales; discal dot prominent; suggestion of exterior line, fuscous; a short, faint band of fuscous scales running parallel to exterior line inner- marginally; fringes pinkish. Abdomen clothed both dorsally and ventrally in brown and rose colored elongate hairs and paucity of scales. Greatest expanse of forewing ± 16mm. Geni- talia as in figures 4 and 5. Female: As in male, very little variation, except somewhat larg- er; antennae ciliate, rather than as in male. Greatest expanse of forewing ±: 18 mm. Genitalia as in figure 6. SPEGIMENS EXAMINED CALIFORNIA: Holotype male, Inverness, Marin County, 6 February 1965 (J. S. Buckett). Paratypes: 1 female, (designated allotype) , Inverness, Marin County, 2 February 1962 (J.S.B. & G. M. Trenam); remainder of paratypes all from Inverness, Marin County, unless otherwise stated; 1 female, 14 February 1950 (W .R. Bauer); 1 female, 14 February 1955 (W.R.B. & J.S.B. ); 1 female 31 January 1956 (W.R.B. & J.S.B.); 1 fe- male, 16 February 1960 (W.R.B. & J.S.B.); 1 female, 31 January 1961 (J.S.B.); 1 female, 1 February 1961 (J.S.B.); 2 females, 4 February 1961 (W.R.B. & J.S.B.); 3 males, 3 females, 1^ February 1962 (J.S.B. & G.M.T. ); 10 males, 6 females 2 February 1962 (J.S.B. & G.M.T.); 4 fe- males, Tomales Bay State Park, Marin County, 6 February 1960 (W.R.B. & J.S.B.); 1 male, Mendocino, Mendocino County, 28 November 1959 (J. R. Heifer). Holotype deposited in the Type Collection, Department of Entomology, University of California, Davis. Allotype female deposited in the Bauer-Buckett Collection, Davis, California. Remainder of paratypes deposited in the following institutions and collections: American Museum of Natural History, New York; Bauer-Buckett Collection, Davis; California Academy of Sciences, San Francisco; California Department of Agriculture, Sacramento; Franclemont Collection, Ithaca, New York; Los Angeles County Museum of Natural History; United States National Museum, Washington, D.C.; University of California, Berkeley and Davis. There is also a large series of Xylomiges haueri contained in the Franclemont Collection, Cornell University, Ithaca, New York, which was unavailable for study at this time. The speci- mens of this large series were collected at the same localities as these listed in the “specimens examined” section of the present 28 J. S. BUCKETT /. Res. Lepid. Fig. 4 — Paratype, Xijlomiges baueri. Male genitalia minus aedeagus. In- | verness, Marin County, California, 3 February 1962 (J. S. Buckett and ; G. M. Trenam), Bauer-Buckett Slide No. 65D2-4. ji Fig. 5 — Paratype, Xijlomiges baueri. Male aedeagus. Data same as in j fig. 4. jl 6(1). -23-30, 1967 XYLOMIGES 29 Fig. e—Faiatype Xylomiges baueri. Female genitalia. Locality and col- lectors same as in fig. 4, 11 February 1962, Bauer-Buckett Slide No. 65D1-2. 30 J. S. BUCKETT /. Res. Lepid. work (primarily at Inverness, Marin County), and were collected by the author and colleagues. Xylomiges baueri is most closely related to X. variabilis, as is evidenced by both the male and female genitalia, as well as by maculation of the primaries. In distribution, baueri is thus far known to occur in the Central California coastal area, particular^ ly immediately adjacent to the ocean. In the past, baueri has not been collected over one air mile from the ocean. This re- stricted distribution is, no doubt, due to the confined range of its probable host plant, Bishop Pine. Xylomiges variabilis is not particularly restricted in distribu- tion, as is hauerg and it occurs throughout much of the western United States, and at higher elevations (see distribution map). In the Bauer-Buckett collection specimens of variabilis are repre- sented from the following localities: Arizona: Madera Canyon, Santa Cruz County. California: Alturas, Modoc County; Benton Station, Mono County; Bishop, Inyo ounty; Cobb Mountain, 8 miles northwest of Middleton, Lake County; Fish Camp, Madera County; Johns ville, Plumas County; Markleville, Alpine County; Mohawk, Plumas County; Pinyon Crest, Riverside County; Valyermo, Los Angeles County. Utah: Dividend, Utah County; Eureka, Juab County; Provo, Utah County. Due to the distribu- tion of both species, it seems more probable that variabilis was an earlier occurring species evolutionarily speaking, and that it could have even given rise to baueri. Mr. William R. Bauer of this Bureau was the first person known to discover this undescribed entitity, and therefore, I take great pleasure in naming it in his honor. REFERENCES DYAR, H. G., 1903 (1902). A list of North American Lepidoptera and key to the literature of this order of insects. Bull. United States Natl. Mus. No. 52. MUNZ, P. A. and D. D. KECK, 1959. A California Flora. University of California Press, Berkeley and Los Angeles, 1681 pp. SMITH, J. B., 1891. Notes on some Noctuidae, with descriptions of new genera and species. Trans. American Entomol. Soc. 18:103 - 135 plus 1 plate. Journal of Research on the Lepidoptera 6(1) : 31-42, 1967 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE LIFE-HISTORIES OF SOUTH AFRICAN COLOTIS ERONE, C. lONE, C. VESTA AND LEPTOSIA ALCESTA (PIERIDAE) The late Gowan C. Clark and C. G. C. Dickson^ Cape Town, South Africa The early stages of two of these very beautiful Pierids were recorded in full, comparatively recently, apparently for the first time, and they are now being described and illustrated in colour in the present paper. The first species, Colotis erone (Angus), was reared from eggs and larvae procured by the second writer at Umhlanga Rocks, Natal, in 1955 and 1956, and the other species, Colotis tone (Godart), from eggs laid by a captive female caught by Mr. T. W. Schofield at the Shongweni Dam, Natal, in 1958; while the more detailed observations and paintings were made from this living material by the first writer in Port Elizabeth, Cape Province. Colotis erone (Angus) Egg. Eggs are laid singly, generally on a young shoot. They are pale yellow, darkening slightly in time and developing salmon spots, 0.6 mm. in diameter by 0.9 mm. in height and with 14-15 longitudinal ribs, only half of which reach the micropile. These ribs are cross-braced by some 25-28 lesser ribs. Eggs hatch after 4-7 days. Larva. The larva eats its way out of the top of the egg and devours the discarded shell. It is 1.75 mm. in length on hatching and of a pale yellow colour, and soon develops brown longi- tudinal stripes. It feeds on the surface of a leaf and grows to 3 mm. in 6-12 days. In the second instar larvae are pale green with brown mottled stripes and with the ventral parts dirty yellow. They grow to 5.5 mm. in 4 days. ^C. G. C. Dickson, “Blencathra,” Cambridge Avenue, St. Michael’s Estate, Cape Town, South Africa. 31 COLOTIS KRONE (ANGUS) for legend see page 41 GOWAN C. CLARK, del ad. nat. CLARK AND DICKSON /• B.es. Lepid. 6(1): 31-42, 1967 LIFE HISTORIES 33 In the third instar larvae are of a darker green with firmer brown stripes and with the ventral portions yellow. They grow to 9 mm. in 3-4 days. In the fourth instar larvae are of a still deeper green and the stripes are of a faint greenish-brown colour, the thin spiracular one of the two former instars still being present. There is a broad white dorsal line ( which shows faint development in the previous instar). Larvae grow to 14 mm. in 4-5 days. In the final instar larvae are green throughout with a thin white spiracular line. They feed on the edge of a leaf, filling the eaten-out gap with their bodies. They grow to 31 mm. in 7 days. Pupa. The pupa is some 22 mm. in length and is mainly of green shades, to match its surroundings. It is secured in an up- right position by cremastral hooks and a girdle. Emergence takes place after some 18 days. A succession of broods occurs throughout the year, with a distinct seasonal difference both in markings and size. This butterfly has a quick, direct flight only a little distance above the ground, which has been rather aptly described as “jet-propelled” — in contrast to that of so many other Pierids. It is found in the open spaces in the neighborhood of the sub- tropical coastal forests or in glades and clearings in these forests, but it is also known to occur a short distance from the coast (of Natal) in places showing some approach to savannah country (e.g., at Shongweni Dam) and which, on the whole , are more characteristic as habitats of the following species, C. ione. Food-plant : Maerua racemulosa (D.C.) Gilg. and Benn. ( Capparidaceae ) — alternatiae generic name: Niehuhia^ as re- corded originally by Platt (1921) for Colotis erone, i.e., N. pedunculosa Hochst. Distribution. The only reliable records for this species appear to be those from or close to the coast of Natal and Pondoland — with a possibly dubious record from Swaziland. Most experienced workers, such as Mr. K. M. Pennington and Dr. van Son, do not now accept the earlier supposed locality records from other territories in Southern Africa, which may have resulted from confusion with C. ione. Colotis ione (Godart) Egg. Eggs are laid singly on a leaf and are white at first but later assume a pale salmon tint with salmon spots. They are 0.45 mm. in diameter and 0.8 mm. in height and have 14-15 34 CLARK AND DICKSON J. Res. Lepid. COLOTIS lONE (GODART) for legend see page 41 GOWAN C. CLARK, del ad. nat. li LEPTOSIA ALCESTA (STOLL.) for legend see page 41 GOWAN C. CLARK, del ad. nat. 36 CLARK AND DICKSON /. Res. Lepid. longitudinal ribs, only 7-8 of which reach the micropile. The ribs are cross -braced by some 24 finer ribs. The larva emerges from the egg after 9 days. Larva. The larva eats its way out near the top and the dis- carded shell is devoured. It is 1.25 mm. in length and of a pale yellow colour, later developing brown irregular lines in the vicinity of the spiracles. It feeds on a leafs surface and grows to 2.75 mm. in 11 days. In the second instar larvae are greyish with a whitish dorsal stripe. They grow to 4 mm. in 6 days. In the third instar larvae are lighter grey, inclining to green, with a thin white spiracular line. They grow to 7 mm. in 6 days. In the fourth instar larvae are green with white dorsal and pale green spiracular lines, while the anal segments are in- clined to brown. They grow to 13.5 — 14 mm. ( depending upon the sex) in 8 days. In the final instar larvae are green with a thin pale-green dorsal stripe and a white spiracllar stripe. The primary moles are yellow and very noticeable. Larvae grow to 23-24 mm. in 11 days. (The foregoing measorements applied to specimens which were below average size. ) Pupa. The pupa is about 22 mm. in length and is whitish or green in colour. It is secured by cremastral hooks and a girdle, in an upright position. The butterfly emerges after some 18 days. A succession of broods occurs throughout the year and, as in the previous species, there is noticeable seasonal difference in development of marking, colouring of the underside and size. The flight is like that of C. erone. This species generally inhabits country of savannah types or areas transitional to it, but the butterfly occurs to some extent, anyway, on the Natal coast itself. In the writer’s experience, it is decidedly scarce today, in comparison with C. erone, on the coast near Durban. Food-plant : Maerua racemulosa (D.C.) Gilg. and Benn. ( Cappiaridaceae ) . Distribution in Southern Africa. Pondoland (Port St. Johns — on the first writers authority); Natal (from the coast to some distance inland in suitable areas) and portions of Zululand; S.W. Africa; Bechuanaland; N. and N.E. Transvaal; Rhodesia; Mozambique. Presumably, also Swaziland. LIFE HISTORIES 37 6(1): 31-42, 1967 COLOTIS VESTA (REICHE) for legend see page 41 GOWAN C. CLARK, del ad. rmt. 38 CLARK AND DICKSON /. Res. Lepid. j The complete life-cycles of the two species remaining do not appear to have been recorded previously or figures given of all the immature stages and these are therefore now being published, with all the stages represented in colour. Material for these studies was procured in Natal: that for Leptosia alcesta on the coast, at Umhlanga Rocks, by the second author in 1955, and for Colot is vesta at Keat’s Hill, on the Greytown-Dundee road, by Mr. T. W. Schofield, of Pietermaritz- burg, in 1958, while the actual breeding and recording were done at Port Elizabeth, Cape Province. Leptosia alcesta (Stoll.) Egg. Eggs are laid singly on young shoots. They are pure white, 1.5 mm. high by 0.4 mm. in diameter and have 10 longi- tudinal ribs, only 6 of which reach the micropile. These ribs are cross-braced by 28-30 fine ribs. Eggs hatch after 4-6 days. Larva. The emerging larva eats its way out of the top of the egg and devours the discarded shell. It is 1.5 mm. long and is transparent water-white, the transparency, until it feeds on the leaves, making it difficult to see. It feeds on the edge of a leaf and grows to 3 mm. before moulting, after 6 days. In the second instar larvae are greenish above and whitish below. They keep to one resting place, though they are not gregarious, and crawl away from it to feed. They grow to 6 mm. in 3 days. In the third instar larvae are green throughout; they have the same habits as before and grow to 8 mm. in 3 days. In the fourth instarf larvae are of the same green colour but they have more setae. They grow to 12.5 mm. in 3 days. In the final instar larvae are perhaps a shade darker in colour and there is a faint whitish sub-spiracular stripe. They grow to 22 mm. for male, and 23 mm. for female specimens in 10 days before pupating. Fupa. The pupa is 15.5-16 mm. in length and is pale green. It is secured to a twig by cremastral hooks and is supported by a girdle. Emergence occurs after 10 days. There is a succession of broods throughout the year, at any rate in the warm coastal portions of its range. This delicate species has a very feeble flight and habitually 6(1): 31-42, 1967 LIFE HISTORIES 39 frequents the tropical or sub-tropical forests and their under- growth, or at least localities in which the vegetation is sufficient to provide the shade in which it is usually found flitting about in its leisurely and rather restless manner. Food-plant Capparis zeyheri Tiircz. ( Capparideae ) . Distribution in South Africa. Natal (the coastal, or more coast- al, zone); N. E, Transvaal; portions of Rhodesia; Swaziland; Mozambique. Colotis vesta (Reiche). Egg. Eggs are laid singly on a leaf. They are white when laid but change to a pale yellow, are 0.45 mm. in diameter by 0.75 m.m. in height and there are 16 longitudinal ribs, only 9 of which reach the micropile. The ribs are cross-braced by some 24 small, fine ribs. Some eggs are faintly blotched with pale salmon. Eggs hatch 4 days after being laid. Larva. The larva, on emergence, ‘is 1.5 mm. long and is pale yellow throughout. It feeds on the younger leaves and grows to 3 mm. in 3-4 days. In the second instar larvae are pale green with a light dorsal stripe. They grow to 5 mm. in 3 days. In the third instar larvae are green with a yellow-bordered, whitish dorsal stripe. They feed on the edge of a leaf, occupying the part eaten out, with the dorsal stripe matching the missing edge. Larvae grow to 8 mm. in 3 days. In the fourth instar larvae are green with a solid yellow dorsal line and a faint, whitish-green spiracular line has developed in this instar. They grow to 12-13 mm. in 3 days. In the final instar larvae are green, with the dorsal line broken into alternate white and yellow portions, and each white portion thinly edged with brown. The spiracular line has widened into a white patch round the spiracle on segment VII. Larvae will now devour a whole leaf. They grow to 20-23 mm. in 4-5 days. Pupa. The pupa is some 13.5 mm. in length and there is individual variation in colour from light green to shades of light brown. There is a well defined lateral stripe and the extended wing-case has a diagonal stripe. The pupa is secured by cremas- tral hooks and a girdle. The butterby emerges after 6-11 days. 40 CLARK AND DICKSON ]. Res. Lepid. The broods are continuous throughout the year, with seasonal variation noticable in specimens. This species is characteristic of certain tracts of country of more or less savannah type with a prevalence of native Acacia trees, sometimes interspersed with scattered but not very high bush. The flight is not as rapid as that of some of the other members of its genus. Food-plant: at least one species of Capparideae. Distribution in Southern Africa. Natal (excluding, from avail- able records, the southern portion); S.W. Africa; Bechuanaland; N. and N.E. Transvaal; Rhodesia; Mozambique. Presumably, also Swaziland. Much information pertaining to the two foregoing species, as regards the imagines, may be obtained from the works listed below. REFERENCES DICKSON, C. G. C., 1965. “Recently observed food-plants of some South African lepidopteroiis larvae”, J. Ent. Soc. S.Afr., 28: 11-20, 1965. PINNEY, E. C. G., 1949. Butterflies of Rhodesia, Salisbury, Rhodesia. 1965. Butterflies of Southern Africa, Johannesburg. PLATT, E. E., 1921. List of food-plants of some South African Lepidopter- ous larvae”, S. Afr. Journ. Nat. Hist., 3: 65-138, 1921. SEITZ, ADELBERT, 1925. The Macrolepidoptera of the World, Vol. XIII. Stuttgart. SWANEPOEL, D. A., 1953. Butterflies of South Africa, Cape Town. TRIMEN, ROLAND and BOWKER, JAMES HENRY, 1889. South African Butterflies, Vol. III. London. VAN SON, G., 1949. The Butterflies of Southern Africa, Part I. Papilianidae and Pienidae, Pretoria. Soutli Africa. 6(1): 31-42, 1967 LIFE HISTORES 41 LEGEND TO THE FIGURES COLOTIS KRONE 1. ImagOj natural size 2. Egg 27 3. Larva on hatching 4. 7th Segment, 1st instar — x 60 5. Head, 1st instar — x 30 6. Larva, 2nd instar 7. 7th Segment, 2nd instar — x 30 8. Larva, 3rd instar 9. 7th Segment, 3rd instar ~ x 18 10. Larva, 4th instar 11. 7th Segment, 4tli instar — x 10 12. Larva, final instar 13. 7th Segment, final instar x 5 14. Spiracle enlarged 15. Head, final instar — x 5 16. Seta, much enlarged 17. Pupa, natural size 18. Cremastral hooks, much enlarged COLOTIS lONE 1. Imago natural size 2. Egg™x35 3. Larva on hatching 4. 7th Segment, 1st instar — x 65 5= Head, 1st instar — x 35 6. Larva, 2nd instar 7. 7th Segment, 2nd instar — ■ x 60 8. Larva, 3rd instar 9. 7th Segment, 3rd instar — x 25 10. Larva, 4th instar 11. 7th Segment, 4tli instar — x 15 12. Larva, final instar 13. 7th Segment, final instar — x 10 14. Spiracle, enlarged 15. Head, final instar — x 8 16. Seta, enlarged 17. Pupa, natural size 18. Cremastral hook, much enlarged LEPTOSIA ALCESTA 1. Imago, natural size 2. Egg=^.x20 3. Larva on hatching — x 30 4. 7th Segment, Lst instar — x 50 5. Head, lst instar 6. Larva, 2nd instar 7. 7th Segment, 2nd instar x 25 8. Larva, 3rd instar 9. 7th Segment, 3rd instar — x 16 10. Larva, 4th instar 11. 7tli Segment, 4th instar — x 12 12. Larva, final instar 13. 7th Segment, final instar — x 9 14. Spiracle, enlarged 42 CLARK AND DICKSON /. Res. Lepid. 15. Head, final instar 16. Seta, enlarged 17. ditto 18. Pupa — X 2 19. Cremastral hooks, much enlarged COLOTIS VESTA 1. Imago, natural size 2. Egg — X 16 3. Larva on hatching — x 30 4. 7th Segment, 1st instar — x 60 5. Head, 1st instar — x 25 6. Larva, 2nd instar 7. 7th Segment, 2nd instar — x 30 8. Larva, 3rd instar 9. 7th Segment, 3rd instar — x 20 10. Larva, 4th instar 11. 7th Segment, 4th instar — x 15 12. Larva, final instar 13. 7th Segment, final instar — x 10 14. Spiracle, enlarged 15. Head, final instar — x 9 16. Setae, enlarged 17. Pupa, natural size 18. Cremastral hooks, much enlarged Journal of Research on the Lepidoptera 6(1) i 43=^51, 1967 1160 W. Orange Grove Ave., Arcadia, California, US. A. 91006 © Copyright 1966 A NEW SPECIES OF FERALIA FROM SANTA CATALINA ISLAND OF CALIFORNIA, WITH NOTES ON THE IMMATURE STAGES OF FERALIA FEBRULIS GROTE ( NOCTUIDAE:CUCULLIINAE ) JOHN S, BUCKETT University of California Davis, California While in the United States National Museum, Washington, D. C., in January, 1964, it was the authors pleasure to discover an apparently new species, or subspecies of Femlia that was closely allied to februalis Grote. Upon further studies, in particu- lar genitalia mounts, it was found that meadowsi Buckett, new species was distinct anatomically as well as by maculation from februalis. The specimens of this new species were all collected during the short interval between 1930 through 1933 by the amateur Lepidopterist, Schoolteacher, Mr. Don C. Meadows. The Los Angeles Times Sunday Magazine for 26 February 1933, pg, 16, states, “For five years Master of Science Don C, Meadows, moth-man of Catalina Island, has been studying ‘fly- by-nights,' and he has captured 400 different species of moths, mapy of which are newer to entomology than the neutron is to physics.” Even though the specimens are labeled “Avalon,” the article further states “Prof. Meadows sets the trap in canyons back of Avalon . . This statements leads one to infer that it is quite possible that meadowsi may have been collected in mountainous canyons of the island. Judging by its affinity to februalis, one wouldn't expect meadowsi to be an unusually rare moth during the early part of the year. Its scarcity in collections is probably due to the lack of collecting on the Channel Islands. F. februalis is not a species which flies great distances, and most ilkely meadowsi would ex- hibit a similar habit. Santa Catalina is over 20 miles from the mainland, and it is highly unlikely that februalis and meadowsi have come in contact with one another for sometime by natural means, yet the two species are obviously closely related, I take great pleasure in naming this species after the ardent “moth- man of Catalina Island,” Mr. Don C. Meadows. 43 44 J. S. BUCKETT /. Res. Lepid. Fig. 1. Holotype male, Feralia meadowsi J. S. Buckett. Avalon Club House, Santa Catalina Island, California, 23 February 1933 (D. C. Meadows). Fig. 2. Allotype female, F. meadowsi. Same locality and collector as holotype, 10 February 1932, Bauer-Buckett slide No. 67D17-53 (in U.S.N.M.). Feralia meadowsi J. S. Buckett, new species Male: Ground color of primaries dorsally pale green (Speci- men compared to Reinhold Color Atlas, pi. 27 A3); secondaries dorsally with black dots at anal angle. Head with vertex clothed in whitish and pale green dentate scales and hairs; frons clothed dorsally in short black simple scales, medially clothed as in 6(1): 43-51, 1967 FERALIA 45 Fig. 3. Feralia februalis Grote, male. Santa Rosa, Sonoma County, Cali- fornia, 14 December 1954 (J. S. Buckett). This is the typical februalis which expresses very little transverse blackish coloration of the primaries. Fig, 4. F. februalis, male, illustrating slight transverse blackish coloration. Inverness, Marin County, California, 31 January 1964 (J. S. Buckett & M. R. Gardner). vertex, cuticula externally with minute reticulations, otherwise smoothly rounded; maxillary palpi exterolaterally black, ventrally clothed in whitish and brown flattened hairs, terminal segment greatly reduced; antennae bipectinate, each bipectination bear- ing ventrally directed ciliations, becoming setose apically; com- pound eyes moderately lashed with elongate brownish hairs. 46 J. S. BUCKETT /. Res. Lepid. Fig. 5. F. februalis, male, illustrating an extreme amount of transverse li blackish coloration. Middletown, Lake County, California, 12 [ February 1955 (W. R. Bauer & J. S. Bucket! ). j Fig. 6. F. februalis, female, illustrating extreme transverse blackish colora- I tion, as well as an olive ground color of primaries. Cobb Mountain, i; Lake County, California, 7 March 1959 (W. R. Bauer & J. S. Bucket! ) . Thorax dorsally with divided collar basally, with weak black j dash, medially pale green, apically white tipped; tegulae with ' anterior-most portion clothed in jet-black dentate scales; inner, posterior and exterior margins of tegulae bound subterminally [ in black, terminally white; disc clothed in pale ^een, white and | black dentate scales; ventrally clothed in blackish simple hairs ji 6(1): 43-51, 1967 FERALIA 47 Fig. 7. F. meadowsi, male genitalia minus aedeagus. Avalon, Santa Cata- lina Island, California, 12 November 1931 (D. C. Meadows), Bauer-Buckett slide No. 67C31-21 (in U.S.N.M.). Fig. 8. F. meadowsi, aedeagus. Data same as for fig. 7. 48 J. S. BUCKETT /. Res. Lepid. Fig. 9. F. februalis, male genitalia minus aedeagus. Inverness, Marin County, California, 31 January 1964 (J. S. Bucket! & M. R. Gardner), Bauer-Buckett slide No. 67C31-23. Fig. 10. F. februalis, aedeagus. Data same as for fig. 9. anteriorly, remainder clothed in white simple hairs; legs with femora clothed dorsally in blackish scales, ventrally clothed in elongate whitish hairs; tibiae and tarsi banded, black and white; primaries dorsally with ground color pale green; basal line gem- inate, terminally white; basal and transverse anterior areas of ground color; transverse anterior line represented costally as 6(1): 43-51, 1967 FERALIA 49 Fig. 12. Mesal view of right mandible, fifth instar larva. Fig. 13. Frontal view of head, fifth instar larva. Fig. 14, Lateral view of head, fifth instar larva. Fig. 15. Setal maps showing chaetotaxy of fith instar larva, lateral view. Tl, T2 = Thoracic segments 1 and 2, respectively; A3, A7, A8, A9, AlO = Abdominal segments 3,7,8,9,10 respectively. black wedge, weak from subcosta to cubutis, thence ' geminate, basally black, apically whitish; median area of ground color for most part, orbicular squared off, weakly outlined in black, thence white, centrally of ground color; median line irregular, black; reniform an open “figure 8,” colored as in orbicular; transverse 50 J. S. BUCKETT /. Res. Lepid. I: posterior line very jagged, geminate, basally white, apically ; black, commencing on costa just basally of subterminal line, j thence meandering onto lower portion of reniform, thence pro- i ceeding to inner margin where it intersects closer to tornus ' than to median line; tornus area with conspicuous black dash; i subterminal line a black wedge on costa, thence diminishing and i fading out in the vicinity of Mi; combined subterminal and ter- , minal areas washed with whitish, therefore appearing very pale, !! or “washed out;” terminal line represented as black undulating line, dipping basally between veins, fringes checkered, black ' and pale green; ventral surface whitish, with slight trace of pale green; basal, transverse anterior, median, and subterminal lines represented costally in black, thence wanting; veins faintly out- lined in brownish; terminal line and fringes as in dorsal surface, but paler; secondaries dorsally whitish, with slight suggestion of black exterior line; inner margin clothed in tan elongate hairs; terminal line as in primaries; fringes pale green; ventral surface possessing two faint transverse black lines; fringes as in dorsal surface. Abdomen dorsally clothed in brownish and tannish hairs, * for most part, some very stout; terminally clothed in whitish and blackish simple and flattened hairs; laterally clothed in blackish hairs and scales; ventrally clothed in whitish simple hairs. Greatest expanse of forewing 16 mm. Genitalia as in figures 7 ’ and 8. Female: As in male, except antennae weakly fasciculate instead of being bipectinate. Judging by the single female at hand, this | sex seems to be of lesser forewing expanse than in the male ( this is not necessarily the case in fehrualis, though). i Specimens examined Holotype male, Avalon Club House, Santa Catalina Island, ! California, 23 February 1933 (Don C. Meadows), U.S.N.M. type no 64645. Paratypes: 1 female, 13 males; 1 female (designated Allotype), same locality and collector as holotype, i 10 February 1932; 13 males, same locality and collector as holo- type, 12 November through 19 February, 1930 - 1933; all in ■: United States National Museum except for a single male dated 8 January 1932, in Bauer-Buckett Collection, Davis, California. i Judging by the series of 15 specimens before me, the Santa I Catalina Island species is more constant in color and in macula- tion than the mainland species, fehrualis, its closest relative. F. meadowsi differs from F. fehrualis superficially by its con- stant, drab, pale green coloration; pronounced black lunule, or [ dash of the tornus area on dorsal surface of primaries; constant ; 6(1): 43-51, 1967 FERALIA 51 conspicuous black spot at anal angle of secondaries, dorsally. The male genitalia of meadoivsi are distinctly different from februalis in that meadoivsi possesses a broader cucullus, much broader valvae, and broader ventral process of the terminal por- tion of the uncus. The aedeagus possesses considerably fewer spines on the vesical sac also. The female genitalia of meadoivsi differ from februalis by the former possessing virtually no large setae on the ovipositor lobes, whereas februalis possesses many; meadoivsi has a less sclerotized ductus bursae, and the genitalia on the whole is of lesser size than is februalis. The single female before me probably is representative of meadoivsi, but it could be a teneral individual. The immature stages of Feralia februalis have been briefly discussed ( at least the fifth instar ) by Crumb ( 1956 ) ; however, to my knowledge, the chaetotaxy of the fifth instar larva has not been described or illustrated. Crumb (op. cit. ) states in his key to known speices of Feralia larvae that februalis “Feeds on broad-leaved trees.” Under the description of februalis he cites as food plant ''Cer cocarpus betuloides and oak (Quercus doug- lasi)” The author has been successful in rearing this species on California Buckeye, Aesculus californicus (Spach) Nuttall, from eggs obtained from an adult female, and McFarland (Master’s Thesis) cites Sambucus, Cercocarpus and Quercus as host plants for februalis in southern California. Crumb described the general habitus of the fifth instar larva and therefore it will not be necessary to do so here. In the key as well as in the description, he stresses the presence of a “very decided subconical dorsal hump on 8.” The eighth abdominal segment on my three preserved fifth instar larvae lacks this “subconical hump;” however, on a second instar larva there is a suggestion of this hump. Chaetotaxy of the pertinent thoracic and abdominal segments may be seen in fig. 15. Chaetotaxy of the head may be seen in both front and lateral views (as in figs. 13 and 14, respectively). At this time, I would like to express my appreciation to Mr. Michael R. Gardner for preparing the illustrations of the larva of februalis. The genitalia slides were made using lignin pink stain and balsam as the mounting media. LITERATURE CITED CRUMB, S. E., 1956. The larvae of the Phalaenidae. U. S. Government Press, Technical Bull. No. 1135, Washington, D. C., 356 pp. McFarland, a. N., 1963 (Master’s thesis, not published). The Mac- roheterocera ( Lepidoptera ) of a mixed forest in western Oregon. KORNERUP, A. and J. H. Wanscher, 1961. Reinhold Color Atlas. Rein- hold Publishing Corporation, New York, 224 pp. Journal of Research on the Lepidoptera 6(1) : 52, 1967 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 HOMONYMY OF THE “NEW GENUS” PETALUMA BUCKETT AND BAUER (NOCTUIDAE) UNDER PETALUMA HULST (PYRALIDAE) AND PROPOSAL OF THE NAME PETALUMARIA FOR “PETALUMA’’ CALIFORNICA. J. S. BUCKETT and W. R. BAUER University of California, Davis, California and California State Department of Agriculture Sacramento, California In 1964 (J. Res. Lepidoptera) the authors named a new noc- tuid moth Petaluma californica, new genus and species, the generic name being preoccupied. The name Petaluma Bkt. and Br. is then a primary homonym of Petaluma Hulst. This error stemmed from not having referred to “Nomenclator Zoologicus” by Neave. Only recently the first author was checking “Nomen- clator Zoologicus” in connection with another new generic name to be proposed in a manuscript in preparation, and while doing this, he thumbed into the “P” section and looked to see if Peta- luma was listed. There, appearing in bold print, was ‘Petaluma Hulst 1888, Ent. Amer., 4, 116 — Lep.” The irony of the situa- tion is worthy of note, as the name was used in the Lepidoptera, Phycitidae. Petaluma illibella Hulst, the type species of the genus is now placed under the genus C oenochroa, in the subfamily Anerasti- inae of the Pyralidae. We herein propose Petalumaria, new generic term, for the species “Petaluma” californica Bkt. and Br. The type species of Petalumaria is P. californica ( Bkt. and Br. ) . REFERENCES BUCKETT, J. S. and W. R. BAUER, 1964. Petaluma, a new genus, with the description of a New Species. J. Res. Lepidoptera 3(3) :193-196. HULST, G. D., 1888. New Genera and Species of Epipaschiae and Phy- citidae. Entomologica Americana 4(6) :113-118. McDUNNOUGH, J. H., 1939. Check list of the Lepidoptera of Canada and the United States of America, par t2, Microlepidoptera. Memoirs of the Southern California Academy of Sciences 2(1):1-171. NEAVE, S. A., 1940. Nomenclator Zoologicus. A list of the names of genera and subgenera in Zoology from the tenth editon of Linnaeus 1758 to the end of 1935. Richard Clay and Co., Ltd., Bum gay, Suf- folk, vol 3, pp. 1-1065. 52 Journal of Research on the LepidotAera 6(1) : 53-58, 1967 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 REARING EULEUCOPHAEUS RUBRIDORSA AND E. LEX (SATURNIIDAE) BRIAN O. C. GARDINER 18 Chesterton Hall Crescent, Cambridge, England. INTRODUCTION Final and penultimate instar larvae of Euleucophaeus rubridorsa Felder and E. lex Druce were collected in the -vicinity of the Great Pyramid, Teotihuacan, near Mexico City about the middle of September 1961 by Dr. A. D. Blest and Dr. T. S. Collett who kept them in plastic boxes and delivered them to me in England on 26. IX. 61. Upon receipt the larvae, which were as yet unidentified, were sorted into two species and reared separately. When the larvae arrived both were feeding on rather dried leaves of some species of Leguminosae which it was not possible to identify, but from the leaf form and bits of stem appeared to be a kind of prickly Mimosa. This was the plant from which the wild larvae had been collected. They were offered various European trees and shrubs and the rearing was carried out at 20-25 °C. under natural conditions of British daylight. EULEUCOPHAEUS LEX On receipt the larvae were offered Robinia pseudo-acacia to which they transferred after a few days. As the leaves of this tree shortly began to fall Fagus sylvatica was later offered and this too was accepted by all the larvae. The larvae were very slow in their movements and showed no sign of being gregarious. When disturbed there was a tendency for them to drop and curl up on the ground although this reaction is by no means so intense as I have observed in, for instance, Automeris vinosus Conte, and is about equal to the intensity in the final instar larvae of the rather better known Hemileuca maia Druce. 53 54 BRIAN O. C. GARDINER /, Res. Lepid. About 80 larvae were received. Of these 25 per cent died for unknown reasons and a further 25 per cent produced parasites of the genus Apanteles. It has not to date been possible to determine the exact species). From 12-24 specimens of Apanteles emerged through the skin of each lex larva. They then proceeded to spin their oval white cocoons amongst the spines of the larva. The adult parasites emerged about a week after pupation. It is interesting to speculate what they then do in nature. The first adult parasites appeared only 16 days after receipt of the larvae and there seems no doubt that they would be emerging simult- aneously in Mexico. What then does the next generation live on? The lex larvae are now all final instar. Do the Apanteles (which normally parasites the early instars of Lepidopterous larvae) have an alternative host? Here is an opportunity for some field research in Mexico. Even more curious is the failure of the Apanteles to kill the lex larvae immediately. Although a few remained motionless after the parasites had emerged, most of them continued to walk slowly about, without feeding, for up to 14 days before dying. The adult parasites were thus emerging before the death of their host. The remaining 50 per cent of the larvae successfully pupated. Pupation took place in a flimsy cocoon, generally in the corners of the rearing cage, a few amongst the leaves of the foodplant, where it would probably normally occur in nature. The pupa itself is a bluish-black color and it and the cocoon case is covered with a white powder. A similar powder occurs in certain Lasio- campidae but in no other Saturniid genus known to me. The full-grown larva is cylindrical, about 4-4.5 cms. long. Head black. Dorsally velvety-black, densely covered with reddish- brown and golden-yellow slightly raised minute dots many of which bear a short white hair. The colors of these dots are so arranged as to give the effect of a yellowish lateral line, above which is orange, then yellow again and finally an orange dorsal line. The chalazae are short, black, bearing dense spines which are black at the base with white tips. Spiracles inconspicuous. Ventrally, the ground color black, covered in grey slightly raised spots each bearing a whitish hair. Legs similarly marked. The duration of the final instar was 34 days and of the pupal stage (based on 4 individuals only — the rest of the pupae having been used for other purposes) — 60 days. The moths were noted to be diurnal flyers. 6(1): 53-58, 1967 REARING 55 EULEUCOPHAEUS RUBRIDORSA On receipt the larvae were offered F. sijlvatica which was at once accepted. R. pseudo-acacia was also found to be eaten. The final instar larvae are rather more active than those of lex, show the same tendency to drop when disturbed and also spin a flimsy cocoon in the corners of their cage or among the leaves of their foodplant. Similar in color and appearance; also covered with whitish powder. Only a few of the larvae died and although none produced any Apanteles parasites, three of them produced a dipterous parasite each. These were a single female Spoggosia (Spoggosia) floridensis (Tns.) and two male Leschenaultia sp. near leuco- phrys (Wied. ) and fusca Tns. These are the first records of Tachinidae from this host. The full-grown larva is cylindrical, about 4-4.5 cms. long. Head black with fine, sparse, whitish hair. Ground color dark grey, covered in lighter grey or silver round and oblong slightly raised, very small and numerous spots which, laterally and ventrally bear whitish hairs. These are so clustered as to form a distinct lateral lightish silver-grey line. The chalazae very short and with very dense spines, these being whitish-grey at base with black tips. The legs black with whitish-grey spots and whitish hairs. As with lex, most of the pupae were used for other purposes and only a few moths emerged. The first female to do so did not expand her wings until 18 hours after eclosion. This is the only instance known to me of such a long delay. It being my experience that unless wing expansion occurs within minutes of eclosion then it does not take place at all. This case was ob- viously exceptional and no other instance of it occurred either with this species, with lex, or amongst the many specimens of the closely related Hemileuca spp. that I have bred. The moths are diurnal, and shortly after this female had ex- panded her wings she commenced to “call” and was paired in the afternoon in bright winter sunlight. Copulation lasted be- tween 30 and 45 minutes. She commenced to oviposit just after dusk and laid 158 ova in two batches. She was very active the following day but died that night without laying any more. She was found to be empty of ova. Another virgin that was dissected was found to contain 155 ova. The ova are large for the size of the moth. When first laid they are jet black, but within 1-2 minutes fade to the brown color of milk chocolate, with a black j micropyle. They were laid in a regular mass on the side of the I i 56 BRIAN O. C. GARDINER J. Res. Lepid, cage containing the female. In nature probably in a ring round i twigs like H. maia does. The eggs were kept at 15°C. ±: 5°C. i and hatched in 59 days. The newly hatched larvae were transferred to 25 °C. and offered a choice of Salix sp. (which had been forced in a heated i' greenhouse) and various Graminae. After 48 hours they com- ^ menced to feed on the Salix, but two days later they at once | transferred to Mimosa dealhata when this was offered them. At [ this stage, and for the next few instars, they are strongly gre- 'i garious and move in single file columns. When feeding they form I, a tight cluster and feed gradually outwards from the base of the leaves towards the tip. When they reached the fifth instar the M. dealhata gave out and Quercus ilex was offered and ac- cepted. Unfortunately after the 7th moult the majority of the larvae died of what appears to have been a granulosis virus disease, although absolute confirmation of this has not yet been : possible. I; The fact that this disease occurred in a subsequent generation, ! and at such a great distance from any natural specific source, is ' strong evidence that this was a trans-ovarially transmitted disease. The duration of the various stages can be summarized as [ follows : — ' Egg stage at 15 °C. it 5°C. = 59 days 1st instar larva at 25 °C. = 10 ” 2nd ” ” ” 20°--25°C. = 11 ” 3rd ” ” ” = 7 ” 4th ” = 12 ” 5th " ” ” = 18 ” 6th ’’ ” ” = 11 ” 7th ” ” ” =: 15 ” 8th ” ’’ = 30 ” Pupal stage = 42 ” which gives a total of 215 days or seven months. DISCUSSION It has been possible to rear one species, ruhridorsa, right I through from egg to adult for the first time. The complete life- cycle period of seven months is an odd one and unlikely to be found in nature where the species is either double-brooded with a six-month cycle, or, perhaps rather more likely, at some stage or I; other diapauses, probably in the egg stage like H. maia. My eggs ( were kept comparatively warm at 15°C. ±; 5° and I have known 6(1): 53-58, 1967 REARING 57 maia eggs, when also kept warm hatch within two months in- stead of overwintering. The number of larval instars is eight, which is high for a Saturniid, many species of which have but 5 or 6 instars. Com- bined with this is the very long larval period of 114 days, which is nearly double that of Hylesia nigricans Berg, the only other Saturniid I am acquainted with which also has eight instars, and which when reared under comparable conditions, has a larval period of 64 days. The very slow and lethargic movements, combined with several days spent in moulting, supports the belief that this figure is a true one, probably corresponds to that found in Mexico, and is not attributable to the unfamiliar food that was being eaten. With the other species, lex, it was not possible to determine the complete life-cycle. Nevertheless, some useful information was obtained, the duration of the final instar and pupal stages being determined and also that this is a diurnal moth. In Mexico there seems little doubt that the two species are nearly con- temporaneous. Like all Hemileucine larvae the spines can give a painful sting when brushed against. The effect of both these species is similar, and not very painful, at any rate to the author! Very little appears to be known about the early stages or life- histories of the genus Euleucophaeus and several inaccurate and vague statements have been made about it. According to Michener (1952) most of the species are Mexican with a few ranging north to Arizona and Kansas. A study of the distribution of the various species given by Draudt (1930) confirms this. Crotch (1956) states, incorrectly, that they are all South American. Draudt ( 1930 ) also states that “all of them ( the larvae ) prob- ably live on grass”. Crotch ( 1956 ) that “the larvae are thought all to be grass-feeders”. However, the only species that have actually been found feeding on grass are mania Druce and oliviae Cockerell (Draudt, 1930). Both lex and ruhridorsa have to spin their oval white cocoons amongs the spines of the larva, now been found in the wild feeding on a tree or shrub which, although exact determination was not possible, was clearly of the family Leguminosae and the larvae were found subsequently to feed on other members of this family, Rohinia and Mimosa as well as various Fagaceae. (It is my experience of Saturniids that all species perhaps normally found on Leguminosae in the tropics will accept temperate zone Fagaceae). 58 BRIAN O. C. GARDINER J. Res. Lepid. The dipterous parasites were too few to give any indication of specificity, but is perhaps significant that only lex had been parasitised by an Apanteles species, since the two species were occurring together at the same stage. Nevertheless, as already surmised, it seems that the Apanteles must have an alternative host. ACKNOWLEDGEMENTS I would like to thank Dr. R. W. Crosskey of the Common- wealth Institute of Entomology for naming the parasitic Diptera. This work, including Dr. Blest’s and Dr. Collett’s expenses in the field, was supported by the United States Institute of Health (Project CM. 07109). LITERATURE CITED CROTCH, W. J. B., 1956. A silkmoth rear’s handbook. London. 165 pp. 26 plates. DRAUDT, M. IN SEITZ, A., 1930. The Macrolepidoptera of the World. Vol. 6, Stuttgart. 1304 pp. 197 plates. MICHENER, C. D., 1952. The Saturniidae ( Lepidoptera ) of the Western hemisphere. Bull. Amer. Mus. Nat. Hist. 98, pp. 339-501. Journal of Research on the Lepidoptera 6(1) : 59^64, 1967 1160 W, Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 NEW RECORDS, AND NOTES ON THE STATUS OF SOME HESPERIIDAE FROM MEXICO H. A. FREEMAN^ 1605 Lewis Drive, Garland, Texas While at the United States National Museum in Washing- ton, D. C. during July 1966 I had the opportunity of examining all of their American Hesperiidae and especially their specimens from Mexico. One particular reason for this trip was to check carefully their types and especially those of Dyar that W. H. Evans had placed in synomymy. The results were very interest- ing in that several of Dyar's "synomyms” were found to be valid species and I will make some remarks in this article pertaining to two of these. Others will be discussed in a later publication. In the American Museum of Natural History, New York, are located most of E. L. Bell’s types of tropical American Hesperi- idae, however there is one in particular in Washington that , has long interested me. This species is Mellana mulleri (Bell) from Mexico which Evans placed as a synomym of eulogius (Ploetz) in 1955. While Bell was active at the American Museum of Natural History we corresponded concerning the various species of Mellana (then Atrytone) that he had described and it sur- prised me to see that Evans had placed mulleri as a synomym of eulogius due to several reasons that I will discuss under that species. Among specimens sent to me by Dr. Tarsicio Escalante, Mex- ico, D. F., for determination were found two species previously unrecorded for Mexico. Bolfa Cyclops (Mabille) 1876 During 1953 Evans described sonda as a new subspecies of T would like to express my thanks to the National Science Foundation for Research Grant GB4122 which is making this study of the Hesperiidae of Mexico possible. 59 60 H. A. FREEMAN /. Res. Lepid. Cyclops from Orizaba, Veracruz, Mexico, based on differences i nthe coloration of the lower surface of the primaries. Appar- ently he failed to check the genitalia as there are specific differ- ences present between sonda and cyclops. For quick determin- i ation the coloration is sufficient as sonda is dusky ochreous on the lower surface of the primaries, whereas cyclops is a bright yellow. Previous records indicate that cyclops had been collected from Guatemala to Colombia, while sonda was more or less confined to the section of Veracruz in Mexico near Orizaba. Among specimens received from Dr. Escalante were five ex- amples of cyclops, one male, and four females, all from Sta. Rosa, Comitan, Chiapas, Mexico (May 1965). This is the first record of cyclops from Mexico. Piruna cyclosticta (Dyar) 1920 This species was placed as a synomym of hrunnea (Scudder) 1872 by Evans in 1955. Since that time 1 have collected rather widely over Mexico and found that actually there are two spe- ! cies involved that somewhat resemble each other superficially. While in Washington 1 examined the type of cyclosticta and found that it agreed with some specimens that I had from Tehua- can, Puebla (August 14, 1964, H. A. Freeman) and Aguas Cali- entes, Aguascalientes (August 1956, Stallings & Turner), and l also that the figure in Seitz of cyclosticta was very accurate. The information contained in Evans key to brunnea does not agree with cyclosticta. 1 have specimens of brunnea from Oaxaca, , Oaxaca (June 23, 1966, H. A. Freeman), and Tierra Colorada and Acahuizotla, Guerrero (August and September). There are ; specific differences in the genitalia, however the easiest way to separate the two species is by the following differences: (1) : brunnea is slightly larger, average total expanse, 26mm., whereas cyclosticta averages 22 mm.; (2) brunnea never has a second spot in space 2 located between the distal spot in that space and the base, while cyclosticta has such a spot present; (3) brunnea is darker than cyclosticta being nearly black, while cyclosticta is more grayish-brown; and (4) all spots are better defined in cyclosticta than they are in brunnea. Apparently the only material examined by Evans was the four specimens from Oaxaca, Mexico and the one from Guate- mala contained in the British Museum. These are all brunnea as all that 1 found in the Oaxaca area was that species. In the state of Puebla northwestward to Aguascalientes only cyclosticta was found. 6(1): 59-64, 1967 HESPERIIDAE 61 From the above mentioned information I am removing cyclo- sticta from the synomymy of brunnea and giving it full specific status. Dalla dividuum (Dyar) 1913 This species was placed as a synomym of ligilla (Hewitson) 1877 by Evans in 1955. After carefully examining the type of dividuum (Dyar) in Washington I found that it matched per- fectly the figure of that species in Seitz, and did not agree with his figure of ligilla. Recently I obtained a number of specimens of ligilla from Dr. Escalante that were collected at Catemaco, Veracruz, and Comitan and Ocozingo, Chiapas. These agreed perfectly with the figure in Seitz of that speces as well as Evans’ sketch of the male genitalia, so there is no doubt as to their status. During July 1967 I received from Robert Wind four males and a female Dalla that he had collected at Salada, Co- lima, Mexico during June 1967. These I mounted and noticed at once that they agreed with the figure in Seitz of dividuum, and also with the notes that I had taken of the type in Washing- ton. I dissected a male and noticed that the genitalia were not like those of ligilla so proving beyond a doubt that actually there are two separate species involved. The figures of the genitalia of these two species will be presented in a later publica- tion. There are several differences that are readily discernable between these two species: (1) ligilla is slightly larger averaging 30 mm. total expanse, while dividuum averages 28 mm.; (2) the elongated spot in space 3 of ligilla overlaps the spot in space 2 and reaches the outer edge of the cell spot, while in dividuum this spot is a tiny dot located beneath the apical spots and well separated from the spot in space 2 and the cell spot; (3) in ligilla the large yellowish central spot on the upper surface of the secondaries is not broken by dark veins, while in dividuum it appears macular due to the presence of dark veins; (4) on the lower surface of the secondaries ligilla has a dark brownish area between the discal spots and the margin which is not present in dividuum; and (5) on this same surface in ligilla there is one large yellowish spot locatel below the costa and situated over the large upper discal spot and the elongated cell spot, while in dividuum there are two smaller spots located in this same area. From the information presented there is no doubt as to the specific validity of the name dividuum and with this I remove the name from the synomymy. 62 H. A. FREEMAN /. Res. Lepid. Mellana mulleri (Bell) 1942 E. L. Bell described mulleri in 1942 from four specimens that i' were collected in the state Guerrero in Mexico. The type is in , the United States National Museum and I examined it while I I was there. Evans recorded this species under the synomymy i of eulogius (Ploetz) as “? mulleri Bell 1942: male Mexico: i genitalia figured”. The question mark indicating that he was not ; certain about this as apparently he had not examined the type. , Eulogius is a very variable species and I have collected it in a f number of areas in Mexico as well as at Brownsville, Texas. It is very common in parts of Guerrero, Puebla, and especially in the Valles area of San Luis Potosi. I have in my collection over fifty specimens of that species and even though it is variable it does not ever show the distinguishing characteristics of mulleri. During August 1962 I collected two males and five females of ; mulleri in the mountains just west of Ciudad Victoria, Tama- ulipsas, Mexico. Both males were dissected and compared with I BelFs drawing of the genitalia of mulleri and a number of the i genitalia of eulogius so as to make a careful comparison between ! the two. It was a simple matter to readily separate the two species since mulleri has one of the most distinctive genitalia of the Mellana. Godman and Savin present a fine drawing of the I genitalia of eulogius under the name of mellona Godman in I their Bioloia Centrali-Americana, plate 94, figure 19. Actually, , in appearance, mulleri more closely fits in with the nayana Bell j group than with the eulogius group but can be separated by the genitalia very easily. By superficial characteristics mulleri males i differ from eulogius in the following ways : ( 1 ) in mulleri, on the upper surface of the primaries, the fulvous coloration is more ; extensive than in the most extreme examples of eulogius as ; represented by the figures of mellona plate 94, figures 17 and -8 in the Biologia; (2) in mulleri, on the upper surface of the i secondaries, the discal fulvous spots are much broader than in eulogius thus producing a more narrow brown outer margin; ; (3) on this same surface, the easiest way to separate the two species is in the shape of the spots in spaces 5 and 6. In eulogius ! these spots are smaller than the rest of the discal band and the | spot in space 5 does not touch the cell spot, while in mulleri the spot in space 5 is the largest in the discal band and touches the cell spot, and the spot in space 6 is elongated and situated directly over the center of the spot in space 5; and (4) mulleri f has a somewhat blurred appearance due to the suffusion of 6(1): 59-64, 1967 HESPERIIDAE 63 fulvous scales over the dark areas of the upper surface of the primaries. Eulogim does not have this appearance even in the lighter specimens where the fulvous coloration is more ex- tensive than in the typical examples. In the females the differ- ences can be determined by the following characteristics: (1) muUeri is a much browner species, with rather indistinct fulvous maculation on the primaries, while eulogim is dark with a slight olive cast, and the maculation is distinct with the spots in spaces 2 and 3 white; (2) mulleri has a double fulvous cell spot over the inner edge of the spot in space 2, while eulogim rarely has any spots in the cell area, if present they are situated inward from the spot in space 2; and (3) in mulleri^ on the upper surface of the secondaries, the discal band is broad and rather indistinct, while in eulogim the discal band varies from none at all to a clearly defined row of yellow spots. Mulleri differs from nayarm (Bell) by being more orange-yellow on the lower surface instead of the bright yellow of nayana, nor does nayana have the blurred appearance of mulleri. From the present information I would like to remove mulleri from the synomymy and give it the full specific rank that it deserves. Meliana field! (Bell) 1942 This species was described from Guatemala. In the British Museum there are but three males and a female present and these came from Guatemala. Among the many specimens that I have received from Dr. Escalante for determination were present three males of fieldi from Catemaco, Veracruz, collected during December 1963, and one male and two females from Sta. Rosa, Gomitan, Chiapas, Mexico, collected during Sep- tember 1963. These are the first records for the occurrence of fieldi in Mexico. This species bears a slight resemblance to eulogim but can readily be separated by the orange-red maculation and orange- red fringes. It is a much darker species on the lower surface of both wings. The genitalia readily separate the two species as well. 64 H. A. FREEMAN /. Res. Lepid. REFERENCES BELL, E. L., 1941. New Species of Neotropical Hesperiidae. Rmer, Mus. Novitates. No. 1125, pp. 1-10, figs. 1-20. 1942. New Records and New Species of Hesperiidae from Mexico. Sohret. de los Anales de La Escuela Nacional de Ciencias Biologicas. Vol. IL No. 4. pp. 455-468. DRAUDT, M. 1924. Hesperiidae, in: Seitz. Macrolepidoptera of the World. Vol. 5. The American Rhopalocera. Stuttgart, vii. 1139 pp., 203 pi. EVANS, W. H. 1953. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part III, Pyrginae. Sec. 2. London: British Museum, 246 pp., pis. 26-53. 1955. A catalogue of the American Hesperiidae indicating the classification and nomenclature adopted in the British Museum. Part IV, Hesperiinae and Megathyminae. London: British Museum. 449 pp., pis. 54-88. GODMAN, FREDERICK DUCANE, and OSBERT SALVIN. 1887-1909. Biologia Centrali-Americana. Insecta. Lepidoptera-Rhopalocera, II: 244-637; LII: pis. 112. 6(1) : 65-68, 1967 I Journal of Research on the Lepidoptera 1160 W, Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 THE GENERIC, SPECIFIC AND LOWER ^ CATEGORY NAMES OF THE NE ARCTIC BUTTERFLIES * PART 5 — The Genus Heliconius PADDY McHENRY 1032 E. Santa Anita, Burbank, California In addition to the generic names given here, the literature I includes certain names subjectively synonymous with Heliconius. J These names have not been included as they are not founded on type species occuring in the Nearctic area nor have they been applied to any of the Nearctic species and, furthermore, Heli- I conius has dating priority over them. Although I can find no author who has applied it to the Nearc- j tic species, the generic name Laparus Billberg has been included in order to provide a replacement for the preoccupied name i Migonitis Hiibner which has been used for certain Nearctic species. ® Linnaeus’ two spellings for charitonius (emendatio) have caused some confusion in the literature. This situation is dis- cussed in the data for that name. list of generic names used or available for heliconius HELICONIUS Klxik. ! Type, charitonius (Linnaeus). APOSTAPHIA Hhbner. ! Type, charitonius (Linnaeus). HELICONIA Godart.' ’ 1 Type, charitonius (Linnaeus). I heliconius" Latreille. Type, charitonius (Linnaeus). LAPARUS Billberg. i Type, erato (L innaeus). MIGONITIS Hhbner, [ I Type, erato (Linnaeus). 66 P. McHENRY J. Res. Lepid. MIGONITIS HUBNER. 1816. Verz. Bekann. Schmett. (l)j 12, no. 1. He included among others: "44. M. erato Linn. " | This name is preoccupied by Migonitis Rafinesque, 1815, j Analyse: p. 147. i Type, P[apilio]. eliconius]. erato Linnaeus. 1758. Syst. Nat. (10th. ed. ). 1; 467, no. S-f. Type Selection. Scudder. [ 8 Apr,] 1875., Proc. Amer. Acad. j Arts Sci, 10: 218-219, no, 698. He said: "Erato may be taken as the type, " LIST OF SPECIES AND LOWER CATEGORY NAMES USED OR AVAILABLE FOR HELICONIUS 1. HELICONIUS CHARITONIUS (LINNAEUS), charitonius (Linnaeus), tuckeri Comstock and Brown, vazquezae Comstock and Brown. 2. HELICONIUS PETIVERANUS (DOUBLEDAY), petiveranus (Doubleday), 1. HELICONIUS CHARITONIUS (LINNAEUS). charitonius, P[apilio]. eliconius]. (emendatio) Linnaeus. 1767. SystT Nat7~(12rh. ed. ). T(2); ' 757, no. "65; p. [ 1339], no. 65. "Habitat in America". No sex, series nor date data given. The name was given as charithonia on page 757 and as charitonia on page [ 1339], As first revisors, Comstock and Brown (16 Oct, 1942. Amer. Mus. Nov. (1467): 1-2) selected charitonia as the proper spelling. Correct Latin grammer would seem to dictate the spellings charitonius and charithonius for use with either Papilio or Heliconius as they are both masculine, Maynard. 1891, Butt. N. Engl. (2nd. ed. ): pi. 10 explanation page, for fig. 8, gave the name as Heliconia charionia, in error. tuckeri, Heliconius charitonius W. P. Comstock and F, M. Brown. 16 Oct. 1950. Amer. Mus, Nov. (1467): fig. 3, no. F; p. 13, no. F; pp. 15-16; p. 19 (in pt. ). "Holotype, Male: Winter Park, Florida, October 4". "Allotype, Female: Key Largo, Florida, July 15", "Paratypes: One hundred and twenty-five males and 99 females, all from Florida and Georgia". "The holotypes and allotypes of 6(1): 65^68, 1967 NEARCTIC BUTTERFLIES 67 HELICONIUS KLUK. 1802, Zwierz. Hist. Nat. Pocz. Gospod. 4: 82-84j no. 143. He included among others: "17. - charitonia". Heliconius Kluk was included in the Official List Gen. Names Zool. (1): 126, no. 923, Hemming. 1958. Type. P[apilio]. Hi eliconius]. charitonius (emendatio) Linnaeus. 1767. Syst, Nat. (12th. ed. ). 1{2):757, no. 65. Type Selection. Hemming. Oct. 1933. Entomologist. 66(845): 223. He said: "Type = Heliconius charitonia Linn. , Kluk. " APOSTRAPHIA HUBNER. 1816. Verz. Bekann. Schmett. (1): 13, no. 3. He included among others: "58. A. Charitonia Linn. . . " Fj^pilioL H[ eliconius]. charitonius (emendatio) Linnaeus. 1767. Syst. Nat. (12th. ed. ).~T(2): 757 no 65. , ' * Type Selection. Scudder. [8 Apr.] 1875 . Proc. Amer. Acad. Arts Sci. 10: 116, no. 106. He said: "Charithonia may be taken as type, " HELICONIA GODART. 1819. In Godart and Latreille in Latreille. Ency, Meth. 9( 1);203- 226. He included among others: "22. . . Heliconia Charithonia. " Type. Plapilio]. Hi eliconius]. charitonius (emendatio) LiEnaeus. 17"67. Syst. Nat. (12th. ed. ). 1(2): 757, no. 65. ■ . Type. Selection, Hemming. Oct. 1933. Entomologist. 66(845): 223. He said: "Type = Heliconia charitonia Linn. , God. " HELICONIUS LATREILLE. 1804. Nouv. Diet, Hist. Nat. 24(Tab. Meth. Ins.): 185, 199. He included only: "Papilio ricini, charitonia Fab. " which were originally of Linnaeus. This name is both a homonym and synonym of Heliconius Kluk. Type. P[apilio]. Hi eliconius], charitonius (emendatio) Linnaeus. 1767, Syst. Nat. (12th. ed.), 1(2): 757, no, 65. Type Selection. Hemming. Sept. 1933. Entomologist. 66(844): 198. He said: "Type = Heliconius charitonia Fab. ( = charithonia Linn. ), " LAPARUS BILLBERG, 1820. Enum. Ins. Museo Billberg: p. 77. He included among others: "Doris. . . Linn. " which is considered by Hemming (1934, p. 56, no. 102) as a synonym of P. H. erato Linnaeus, 17 58. Type. Plapilio]. Hi eliconius]. erato Linnaeus. 1758. Sy7t.”Nad7 ( U)thred7)7”l : '^677~no. 54. Type Selection. Hemming, 1934. Entomologist. 67: 37. He said: "Type - Papilio doris Linn. , 1771 (=Papilio erato Linn. , 1758). " 68 P. McHENRY /. Res. Lepid. all new subspecies [described in Comstock's and Brown's paper] are in the collection of the American Museum of Natural History", Mather and Mather, 6 June 1958. Tulane Studies Zool, 6(2): 69, no. 10, spelled as tucgeri, in error. vazquezae, Heliconius charitonius W. P. Comstock and F. M. Brown. 16 Oct. 1950, Amer. Mus. Nov. (1467): fig. 3, no. 6; p, 13, no, 6; p. 16; p. 19 (in pt. ). "Holotype, Male: Campeche, Mexico, November". "Allotype, Female: Allende, Vera Cruz, Mexico, July". "Paratypes: Eighty-four males and 53 females, all from Mexico and Texas". The three Texas specimens are: one male, Sarita, Willacy County; one female. Corpus Christi, Nueces County; one female. New Braxinfels, Comal County. " 2. HELICONIUS PETIVERANUS (DOUBLEDAY). z petiverana, Hel[iconia]. Doubleday. [4 Aug, 1847] . In Doubleday and Westwood. Genera Diurn, Lepid. 1(10): 103, no. 18. "Mexico, Honduras." No sex, series nor date data given. The authoi* cited fig. 2, pi. 4, in Petiver's Gazophylacium (1702-11). The name is modified to petiveranus when transferred to the genus Heliconius for purposes of Latin grammer. REFERENCES CITED Hemming, F. 1934. Generic Names Holarct, Butt. 1: i-vii, 1-184. FOOTNOTES 1. A copy of the work among the separates at the Allan Hancock Library (Univ. Sou. Galif. ) (ex library, Boston Soc. Nat. Hist. ) has the following printed label on the front wrapper: "Library of the Cambridge Entomological Club. Received April 8, 187 5, by gift from the author, " 2. Hemming. 1936-1943. Jour. Soc. Bibliog. Nat. Hist. 1: 335-464. Gave publication dates for the Genera Diurn. Lepid. NOTICES CHANGE OF ADDRESS: When sending a change of address, please include both old and new addresses. This will greatly facilitate your changeover. ZIP CODE: If your zip code is not present, or is incorrect, on your address plate, please inform us as the Post Office requires it on all domestic mailings. BUTTERFLIES OF THE DELAWARE VALLEY, by Arthur M. Shapiro. Special Publication of the American Entomological Society, Philadel- phia, Pa. i-vi, 1-79, 13 b. and w. plates, keys and check list. ^1.50 from publisher at 1900 Race St. , Philadelphia, Pa. AUTOMERIS AURANTIACA, living ova or cocoons urgently required to complete research project, contact Brian O. C. Gardiner, 18, Chesterton Hall Crescent, Cambridge, England. LEPIDOPTERA OF FLORIDA, by C. P. Kimball. Volume 1 of Arthropods of Florida and Neighboring Land Areas, 363 pp. , 26 plates. ^5. 00 from Division of Plant Industry, Florida Department of Agriculture, Gainesville, Florida 32601. IMPORTANT: Books and items listed are to be ordered from the company or person indicated under each listing, not from the Lepidoptera Foundation. Those available from the FOUNDATION are so indicated. The FOUNDATION will undertake to aid in the distribution of literature on the Lepidoptera for institutions or individuals at cost. 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WANT living Cymothoe coccenata. M. H. Ross, Div. Cancer Embryology, Biochem. Research Foundation, Newark, Delaware. WANT Papilio zelicaon, living stock females or pupae. Will pay for field work or stock. C. G. Oliver, Educational Services, Inc. , 108 Water Street, Watertown, Mass. 02172. Volume 6 Number 1 March, 1967 IN THIS ISSUE Spring Moths of a Natural Area in Northeastern Kansas Noel McFarland 1 A New Species of Nepticula on Bur Oak in Ontario T. N, Freeman 19 Fixation of the Type Locality of Lycaena phlaeas hypophlaeas and a Foodplant Correction Oakley Shields 22 Description of a New Species of Xylomiges from California John S. Buckett 23 The Life-Histories of South African Colotis erone, C. ione, C. vesta and Leptosia alcesta (Pieridae) Gowan C. Clark and Co G. C. Dickson 31 A New Species of Feralia John S. Buckett 43 Homonymy of the “New Genus” Petaluma and Proposal of the Name Petalumaria J. S. Buckett and W. R. Bauer 52 Rearing Euleucophaeus ruhridorsa and E. lex Brian O. C. Gardiner 53 The Status of Some Hesperiidae from Mexico H. A. Freeman 59 The Generic, Specific and Lower Category Names of the Nearctic Butterflies Paddy McHenry 65 THE J©URIMJAL ©F RESEARCH ©H THE LEF1D©PTERA a quarterly published at 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HOVANITZ THE PURPOSE OF THE JOURNAL is to combine in one source the work in this field for the aid of students of this group of insects in a way not at present available. THE JQURNAL will attempt to publish primarily only critical and complete papers of an analytical nature, though there will be a limited section devoted to shorter papers and notes. QUALITY WORK on any aspects of research on the Lepidoptera is invited. Analytical and well illustrated works are pre- ferred, with a minimum of long description. AUTHORS ARE REQUESTED to refer to the journal as an example of the form to be used in preparing their manuscripts. 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Second Class Postage Paid at Arcadia, Calif. Journal of .Research. on the Lepidoptera 6(2) : 69-178, 1967 1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006 © Copyright 1966 HILLTOPPING AN ECOLOGICAL STUDY OF SUMMIT CONGREGATION BEHAVIOR OF BUTTERFLIES ON A SOUTHERN CALIFORNIA HILL OAKLEY SHIELDS Department of Biology San Diego State College San Diego, Calif. THE LEPIDOPTERA RESEARCH FOUNDATION, INC. FOREWORD “Territoriality” in various vertebrate animals has been studied for many years, involving not only the fact that territoriality exists as a behavior pattern but also involving a functional need for this behavior in the life history of the animal. “Hilltopping” of butterflies as a form of territoriality has been observed by all persons who have collected, or studied, these invertebrates in the field, but no one has made such a concerted effort at ob- taining analytical data as has Mr. Oakley Shields. All experi- mental and/or analytical studies represent an effort on the part of a worker at obtaining significant data upon which to draw conclusions; there are apt to be as many different conclusions derived from the same set of data as there are independent analy- ses made of them but we can be sure that Mr. Shields has made every effort at drawing conclusions in keeping with the data he has presented. Others may be able to draw other conclusions from the same data. This is in the fulfillment of scientific spirit, and is as it should be. Mr. Shields has also given a very com- plete Bibliography of work related to this field. The literature of entomology is extremely large and he is to be commended for his efforts; such omissions as have occurred should be consider- ed in the context that it could happen to any of us. The Lepidoptera Research Foundation, Inc. is proud to be able to present this work of Mr. Oakley Shields in complete form in the thought that in this way it will prove to be more usable and satisfactory than if it were divided into sections for publication. William Hovanitz Editor HILLTOPPING 6(2):69~178, 1967 TABLE OF CONTENTS 71 ABSTRACT 71 INTRODUCTION 73 MATERIALS AND METHODS 89 Location and description of study area 91 RESULTS 93 Species present 93 Territorial and aggressive behavior 96 Daily residence 105 Diurnal periodicity 109 Migration 112 Mark-recapture study 112 Papilio zeUcaon release experiments 117 Percent virginity 122 Female rarity 128 Female behavior and mating 133 Behavior of sexes after uncoupling 141 Larval foodplant proximity 141 Species approach to the summit 144 Effects of wind at the summit 145 Feeding 146 Roosting 147 Predation 148 DISCUSSION 149 CONCLUSIONS 164 ACKNOWLEDGEMENTS 166 LITERATURE CITED 168 ABSTRACT / It is widely known that butterflies often congregate in num- bers on the summits of hills, ridges, peaks, and mountain-tops. However, few investigations have been made to determine why this behavior has developed. This study was conducted to try to answer this question. Initially, purely observational work was necessary to ade- quately describe the phenomenon. Dictionary Hill near Spring Valley, San Diego County, California, was chosen as the primary study area because it was readily accessible and the surrounding area was relatively undisturbed by man. To follow the behavior of individuals, a mark-release program was established. It was 72 OAKLEY SHIELDS J. Res. Lepid. found that the summit population consisted primarily of males that exhibited territorial or pathway-patrol behavior, with aggres- sion displayed toward other butterflies. These males sometimes spent most of the day in such behavior and repeatedly returned to the summit on subsequent days. During the study 21 species were found to congregate on the summit, while another 25 species flew “up-and-over” the summit without congregating or were generally distributed over the entire hill. Certain insects are known to congregate at topographic sum- mits to facilitate mating. To test this hypothesis for butterflies, the behavior of females that approached the summit was ob- served and females were collected. An abnormally high per- centage of females captured on the summit was virgin when compared to those captured elsewhere. This fact was determined by the absence of a spermatophore in the female's bursa copula- trix. Also, a substantial number of summit matings was observed for two species on the summit. The apparent scarcity of females was due to the inconspicuousness of in copula pairs and due to the fact that virgins stayed only long enough to mate and non- virgins only rarely approached the summit. Virgin females lingered about the summit until mated and readily mated with courting males. Females of two species were seen to depart from the summit after mating. Further evidence that this phenomenon serves to bring males and females together to insure fertilization comes from an ex- periment in which mated and unmated females of one species, Papilio zelicaon^ were released away from the hill. Only the unmated females were later captured on the summit, indicating that virgin females seek the summit. Evidence that males act- ively seek the summit comes from the fact that a substantial number of male P. zelicaon released at various directions and distances away from the hill later returned to the summit. The evidence strongly indicates that “hilltopping” in butter- flies is a phenomenon in which males and virgin (or multiple- mating) females instinctively seek a topographic summit to mate. The selective advantage presumably would be to central- ize isolated populations for mating and thus stabilize the gene pool. Other possible alternative explanations have been ad- vanced in the literature but all are here shown to be highly unlikely. The strongest alternative theory, that butterflies are concentrated on summits due to winds and updrafts, is shown to be unsatisfactory. Observations on feeding, roosting, and predation of hilltop- ping butterflies are also included, and the phenomenon is dis- cussed in relation to hilltopping insects. 6(2):69-178, 1967 HILLTOPPING 73 INTRODUCTION Students of insect behavior are aware that insects are largely at the mercy of their instincts. Insects do not behave a certain way because it is fun or because they want to but do so because their instincts compel them to. Thus when one sees butterfles flying about the summit of a hill and “playing tag” with each other, one may wonder what instinct brought them there and why the habit developed. This study was conducted to deter- mine the reasons for this summit congregation behavior in butterflies. Swarms of insects on the summits of ridges, hills, and moun- tains are frequently mentioned in the literature (Slosson, 1893, 1894, 1895; Bowditch, 1896; Guppy,. 1897; Newcomb, 1901; Currie, 1904; Poulton, 1904; Hudson, 1905; Meinecke, 1917; Howard, 1918; Van Dyke, 1919; Bryan, 1923, 1926; Scott, 1926; Swezey and Williams, 1932; Alexander, 1940; Chapman, 1954a; Edwards, 1956, 1957b, 1960, 1961; Mani, 1962). Sometimes large numbers of different insects are found dead on snow at summits (Bowditch, 1896; Caudell, 1903; Currie, 1904; Bryan, 1917, 1923; Meinecke, 1917; Fletcher, 1964:141). Such swarms are often composed almost entirely or exclusively of males that are con- gregating on a prominence from the surrounding territory (see, for example, Chapman, 1954a; Dodge & Seago, 1954; Catts, 1964; Shepard, 1966). Groups frequenting summits include the orders Coleoptera, Diptera, Hymenoptera, and Lepidoptera (Table 1). A number of Lepidoptera families contain species that show 74 OAKLEY SHIELDS /. Res. Lepid. TABLE 1. Families of insects with species that are known to hilltop Family Sources COLEOPTERA Byr rhidae Cerambycidae Chysomelidae Coccinellidae Edwards, 1956 Poulton, 1904 Slosson, 1893; Van Dyke, 1919 Slosson, 1693; Poulton, 1904; Hudson, Elateridae Scarabaeidae Rhynchophora DIPTERA Agromyzidae Anthomyidae Bibionidae Bombyliidae 1905; Van Dyke, 1919; Bryan, 1926; Balduf, 1935; Chapman, 1954a; Chapman, Romer, and Stark, 1955; Edwards, 1956, 1957a; Hagen, 1962 Edwards, 1956 Hudson, 1905 Van Dyke, 1919 Bryan, 1926 Bryan, 1926 Hudson, 1905 Currie, 1904; Van Dyke, 1919: Chap- man, 1954a; Dodge and Seago, 1954; Edwards, 1956, 1957b Borboridae Calliphoridae Culicidae Cuter ebridae Gastrophilidae Hypodermatidae Muscidae Bryan, 1926 Dodge and Seago, 1954; Edwards, 1956 Knab, 1907 Chapman, 1954a; Catts, 1963 Walton, 1930; Grunin, 1959 Grunin, 1959 Bryan, 1926; Scott, 1926; Dodge and Seago, 1954; Edwards, 1957b Oestridae Aldrich, 1915; Grunin, 1959; Catts 1964 Phoridae Rhagionidae Sarcophagidae Dodge and Seago, 1954 Shemanchuk and Weintraub, 1961 Chapman, 1954a; Dodge and Seago, Simuliidae Syrphidae 1954; Edwards, 1956, 1957b; Grunin, 1959 Edwards, 1956 Currie, 1904; Van Dyke, 1919; Bryan, 1926; Chapman, 1954a; Dodge and Seago, 1954; Edwards, 1956, i960 Tabanidae Chapman, 1954a; Dodge and Seago, 1954; Edwards, 1956, 1957b Tachinidae Currie, 1904; Van Dyke, 1919; Bryan, 1926; Chapman, 1954a; Edwards, 1956, 1960 HYMENOPTERA F ormicidae (winged ants only) Poulton, 1904; Hudson, 1905; Wheeler, 1905, 1917a; Van Dyke, 1919; Scott, 1926; Gregg, 1947; Michener, 1948; Chapman, 1954b; Fosberg, 1955; Chapman, 1957 Ichneumonidae Slosson, 1894; Bryan, 1926; Chapman, 1954a Psammocharidae Siricidae Bryan, 1926 Scott, 1923, 1924, 1926; Walsh, 1924; Sphecidae Chapman, 1954a; Edwards, 1957b Poulton, 1904 6(2);69-178, 1967 HILLTOPPING 75 hilltopping activity: Hesperiidae, Lycaenidae, Nymphalidae, Papilionidae, and Pieridae (Table 2). Welling (1958) reported the phenomenon in some nocturnal moths on a hill in Ohio. Certain hibernating Coccinellidae, Chrysomelidae, and Rhyncho-' phora beetle aggregations (Van Dyke, 1919; Chapman, 1954a) and aestivating Noctuidae moth aggregations (Common, 1952) have been reported from summits; such aggregations were not composed predominantly of males. The explanation of this phenomenon of insects congregating on summits, known as “hilltopping” in butterfly literature, is known for certain insects but remains unsettled for butterflies. Butterfly collectors often find that hilltops are profitable places to collect. Certain species that are rare or apparently absent in the surrounding countryside may congregate at such places in numbers (Van Someren, 1955). Barrett and Burns (1951:4-5), in referring to males of many species of hesperiids and lycaenids in Australia and New Guinea, said that certain hilltops will always have particular species at the proper time of year, Knudsen (1954) found three species of butterflies abundant on Kennesaw Mountain, Georgia. Here he found on the summit several hundred butterflies in an area of a few hundred square feet. On two hilltops in Australia, Waterhouse (1932:267) found butterflies clustered by the hundreds just before dusk or on the lee side of a shrub during wind. Minimum distances traveled by hilltopping butterflies from the nearest areas of their foodplants are reported as several thousand vertical feet for Papilio indra kaibabemis in Arizona (Emmel & Emmel, 1967) and 1,700 vertical meters for Pieris callidice in France (Muspratt, 1954). Various theories have been proposed for hilltopping among insects. Poulton (1904) discussed insects in general that fre- quent summits and concluded that they seek “conspicuous iso- lated features in the landscape” to mate, thus greatly reducing the area in which sexual encounters could occur. He offered as supporting evidence the ceramycid Dorcadion “pairing freely” on a summit. This theory has been particularly substantiated by work with winged ants and bot flies: observations by Forel (1874), Michener (1948), Fosberg (1955), and Chapman (1957) refer to matings among summit swarms of winged ants; bot flies on summits were reported to mate by Walton (1930), Grunin (1959), and Catts (1963, 1964). Dodge and Seago (1954) reported twenty-one mating pairs of Diptera from mountain summits of Georgia, and Chapman ( 1954a ) mentioned 76 OAKLEY SHIELDS /. Res. Lepid. TABLE 2.. A list of known butterfly '*Mlltopping*' species Species PAPILIONIDAE Realm Sources Papilioninae Battus philenor Nea, Weiss, 1927; Merritt, 1952 B. polydamus Neo, new (Brown) Chilasa clytia Ori. Tinkham, 1937; Best, 1954; Wynter-Blyth, 1957 C. c. dissimilis Ori. Best, 1954 Graphium^ most spec. Neo. new (Brown) G. marcellus Nea. Merritt, 1952; Knudsen 1954 Papilio^ many spec. Neo. new (Brown) P. anactus Aus, new (Wyatt) P. anchisiades Neo. new (Brown) P. androgeus Neo. new (Brown) P, angolanus Eth. Van Someren, 1955 P. arcturus Ori, Wynter-Blyth, 1957 P. bairdii Nea. Grinnell and Grinnell, 1907 new (many) P. b. brucei Nea, Rodeck, 1950; Newcomer, 1967 P, brevicauda Nea. Edwards, 1884; Tilden, 1964 P. crino Ori. Best, 1954 P. demodocus Eth. Longstaff, 1912r.225; Van Someren, 1955 P. demoleus Eth. Trimen, 1889 P. eurymedon Nea. Edwards, 1887; Merritt, 1952; Garth and Tilden, 1963 P. feisthamelii Pal. new (Wyatt) P. 'glaucus"””™” Nea. Scudder, 1887, 1889; Weed, 1901; Longstaff, 1912: 149; Weiss, 1927; Merritt, 1952 P. g. arcticus Nea. Wyatt, 1957a P. hector Ori. Longstaff, 1912; 380 P, indra Nea, Mead, 18 78; Edwards, 1884; Wright, 1906; Newcomer, 1910; Martin and Ingham, 1930; Garth, 1934; Emmel and Emmel, 1962; Garth and Tilden, 1963 P, indra fordi Nea. new (many) P, i. ka'ibabensis Nea. Emmel and Emmel, 1967 P, i. martini Nea, Emmel and Emmel, 1968 P. i. minori Nea. Eff, 1962 P, i, pergamus Nea. new (many) 6(2):69-178, 1967 HILLTOPPINC: 77 TABLE 2 (continued) Species Realm Sources Papilio leonidas Eth. Van Someren, 1955 P, liomedon Ori. Wynter- Blyth, 1957 P. machaon Pal, Seitz, 1909; Longstaff, 1912;46; deRhe- Philipe, 1932; Muspratt, 1954; Wynter- Blyth, 1957 P„ m. aliaska Nea. Cary, 1907; Leussler, 1935; Rawson, 1955; Wyatt, 1957a P, m, asiatica Pal. Hayward, 1937 P. m. centralis Pal, Peile, 1923 P. m. hippocrates Pal, Longstaff, 1912:46 P, macleayanus Aus. Waterhouse, 1932; new (Wyatt) P. paris Ori. Wynter- Blyth, 1957 P, pelaus Neo. New (Turner) P, podalirius Pal, Seitz, 1909; Muspratt, 1954 P. polymetus Neo, new (Kesselring) P. polyxenes asterius Nea. Weed, 1901; Weiss, 1927; Clark, 1932; Merritt, 1952; Knudsen, 1954; Tilden, 1964 P, rex Eth, Van Someren, 1955 P, rudkini Nea, new (many) P. rutulus Nea, new (Guppy, Scott, Tilden) P. thersites Neo, Avinoff and Shoumatoff, 1946; Shoumatoff, 1953; new (Turner) P. thoas Neo, new (Brown) P. torquatus Neo, new (Brown) P, troilus Nea, Weiss, 1927, 1928 P, zelicaon Nea, Mead, 1878; Osten-Sacken, 1882; Edwards, 1887; Snyder, 1 894; Wright, 1906; Newcomer, 1910; Grinnell, 1915; White- house, 1918; Van Dyke, 1919; Comstock, 1927; Garth, 1935; Merritt, 1952; Guppy, 1953; Emmel and Emmel, 1962; Garth and Tilden, 1963; Tilden, 1964 Papilio zelicaon- polyxenes hybrid (nitra) Nea, new (Scott) 78 OAKLEY SHIELDS /. Res. Lepid. TABLE 2 (continued) Species Realm Sources Teinopalpus imperialis OrL Seitz, 1927; Parsons, 1948; Bailey, 1951; Saunders, 1955; Wynter- Blyth, 1957 Zetides cloanthus OrL Bailey, 1951; Wynter- Blyth, 1957 PIERIDAE Coliadinae Eurema hecabe Pal. Ori. Longstaff, 1912:46, 388 Phoebis spp. Neo. new (Kesselring) Pierinae Anthocaris belia Pal. Gurney, 1907 A. cethura Nea. Morrison, 1883; Comstock, 1927; Garth, 1934 A, falloui Pal. Fountaine, 1906 A. midea Nea. Rawson, 1951; Arnhold, 1952; Merritt, 1952; Shoumatoff, 1953; Knud- sen, 1954 A. pima Nea. Beutenmuller, 1898; Wright, 1906; Comstock, 1927 A, tagis Pal. Gurney, 1907 Colotis bowkeri Eth, Trimen, 1889 C. eris Eth. Trimen, 1889 Daptonoura spp. Neo. new (Kesselring) Delias harpalyce Aus. new (Wyatt) D. nysa Aus, new (Wyatt) Euchloe ausonides coloradensis Nea. Garth, 1934, 1935; Garth and Tilden, 1963 E. belemia Pal. Longstaff, 1912:162; Peile, 1923 E. belledice Pal. Longstaff, 1912:162 E, charlonia Pal. Longstaff, 1912:162; Dover, 1922; Wynter- Blyth, 1957 E. c. transcaspica Pal. Peile, 1923 E„ hyantis Nea. Garth, 1934, 1935; Garth arid Tilden, 1963 E. h. lotta Nea. Garth, 1934, 1935 E. Olympia Nea. Clark and Clark, 1951; Arnhold, 1952; Merritt, 1952 E, o. rosa Nea. new (Justice) Piercolias huanaco Neo. Seitz, 1924 Pieris callidice Pal. Muspratt, 1954 P. daplidice Pal, Longstaff, 1912:33 P. occidentalis Nea. Snyder, 1894; Dod, 1908a; Shepard, 1966 P. o. calyce Nea. new (many) 6(2):69-178, 1967 HILLTOPPINC 79 TABLE 2 (continued) Species Realm Sources Pieris protodice Nea. new (many) P. rapae Nea. , Aus, Scudder, 1887, 1889; Meinecke, 1917; Bryan, 19^6 P. sisymbrii Nea. Tilden, 1964; new (many) NYMPHALIDAE Danainae Ceratinia vallonia Neo, new (Kesselring) Chittira fumata Ori. Longstaff, 1912:388 Euploea diocletiana Ori. Rawlins, 1949 Ideopsis gaura parakana Ori. Rawlins, 1949 Mechanitis nessaea Neo. new (Kesselring) Thyridia singularis Neo. new (Kesselring) Satyrinae Neominois ridingsii Nea. Comstock, 1927 Oeneis chryxus Nea, Snyder, 1894, 1897; McDunnough, 1927 O. c. ivallda Nea. Wright, 1906; Grinnell, 1915; Comstock, 1927; Martin and Ingham, 1930; Tilden, 1959; Garth and Tilden, 1963; Brown, 1965 O. c. Stanislaus Nea. new (many) O. marcounii Nea. Elwes and Edwards, 1893; Mast ers et al. , 1967 O. melissa assimilis Nea. Freemen, 1948; Mtinroe 1951 O. m. beanii Nea. Elwes and Edwards, 1893; Dod, 1901, 1908a, 1908b; Fletcher, 1905; Nicholl, 1906; Fletcher and Gibson, I9O8; Whitehouse, 1918; McDunnough, 1927 O. m. lucilla Nea. Mead, 1875; new (many) O, nevadensis Nea. Edwards, 1887; Danby, 1894; Fletcher, 1906; Wright, 1906; Guppy, 1962 O. n. iduna Nea, Edwards, 1884 O, polixenes brucei Nea. Klots, 1937 Pararge hindukushica Pal. new (Wyatt) P. megaera Pal. Fountaine, 1906; Peile, 1923; Scott, 1926; Temple, 1953; Muspratt, 1954 P. menava Pal. new (Wyatt) 80 OAKLEY SHIELDS /. Res. Lepid. TABLE 2 (continued) Species Realm Sources Pararg.e moera Pal. new (Wyatt) P harneuptvchia spp. Neo. new (Brown) Morphinae Brassolis spp. Neo. new (Brown) Caligo spp. Neo. new (Brown) Catoblepia spp. Neo. new (Brown) Dasyophthalma spp. Neo. new (Brown) Dynastor darius Neo. ne w (Brown) Eryphanis spp. Neo. new (Brown) Narope spp. Neo. new (Brown) Opsiphanes spp. Neo. new (Brown) Charaxinae Agrias spp. Neo. new (Brown) Anaea, most species Neo, new (Brown) A. ryphae Neo. new (Kesselring) Charaxes achaemenes Eth. Van Someren, 1955 C. castor Eth. Van Someren, 1955 C. epijasius Eth. Van Someren, 1955 C. fabius Ori. Best, 1953, 1954 C. polyena imna Ori. Best, 1953, 1954 C. viola Eth. Van Someren, 1955 Eriboea athamas Ori. Best, 1953, 1954 E. pyrrhus sempronius Aus. new (Wyatt) Euxanthe spp. Eth. Van Someren, 1955 Hamamumida daedulus Eth. Trimen, 1887; Van S 1955 Prepona, most species Neo. new (Brown) P. demophon Neo, new (Kesselring) Siderone nemesis Neo. new (Kesselring) Zaretes isidora Neo. new (Kesselring) Nymphalinae Adelpha, many spec. Neo. ne w (Brown) Ageronia februa Neo. new (Kesselring) A. feronia Neo. new (Kesselring) Argynnis hyperbius Pal. , Ori. Longstaff, 1912:100; Fraser, 1916; Wynter- Blyth, 1957 A, lathonia Pal. Wynter- Blyth, 1957 A. 1. issaea Pal. Longstaff, 1912:46, 64; Seitz, 1927 Boloria alberta Nea. Dod, 1901, 1908b; Nicholl, 1906; Wyatt, 1957b B. astarte Nea. Dod, 1901, 1908a; Fletcher, 1905; Nicholl, 1906; Skinner, 1908; Whitehouse, 1918; Wyatt, 1957b 6(2):69-17H, 1967 HILLTOPPING TABLE 2 (continued) Species Realm Boloria polaris Nea. gronlandica B, selene myrina Nea. B. toddi Nea. Chlosyne californica Nea. C. ismeria Nea. C. i. carlota Nea. C. leanira wrightii Nea. C. theona Nea. Doxocopa agathina Neo. D, kallima Neo, D. yacuna Neo. D, zunilda Neo. Dynamine mylitta Neo. D. argyrippa Neo. Eueides isabella Neo. Euphydryas anicia Nea. E. a. alena Nea. E. a. brucei Nea. E. a. capella Nea, Ek a. eurytion Nea, E. chalcedona Nea. E. c, sierra Nea. E, editha Nea. E. e. nubigena Nea. Euripus consimilis Ori. Hestina nama Ori. Historis spp. Neo, H. odius Neo, H. orion Neo. Limenitis archippus Nea. L. arthemis Nea. L. astyanax Nea. L. lorquini Nea. Melitaea didyma Pal. M. trivia Pal. M. t. persea Pal. Myscelia sophronia Neo. M. orsis Neo. Nymphalis antiopa Nea. N. californica Nea. N. cashmiriensis Pal. N, milberti Nea. N. urticae Pal. N. vau- album j- album Nea, Sources Munroe^ 1951 Scudder, 1887, 1889 Scudder, 1887, 1889 new (many) Knuds en, 1954 new (Justice, Scott) new (Breedlove) new (Shields) new (Brown) new (Brown) new (Brown) new (Brown) new (Brown) new (Brown) new (Brown) Whitehouse, 1918 new (Scott) new (Justice) new (Scott) new (Scott) new ( many) Brown, 1965 new (Scott, Thorne) new (many) Wynter-Blyth, 1957 Bailey, 1951 new (Brown) Avinoff and Shoumatoff, 1946 new (Kesselring) Scudder, 1887, 1889 Scudder, 1887, 1889 Weiss, 1927 Martin and Ingham, 1930 Peile, 1923 Peile, 1923 Peile, 1923 new (Brown) new (Brown) Scudder, 1887, 1889 Weiss, 1927 new (Breedlove, Dvorak) Longstaff, 1912:46 Scudder, 1863, 1887, 1889; Weed, 1901; Weiss, 1927; Garth, 1935; Garth and Tilden, 1963 Muspratt, 1954 Scudder, 1887, 1889; Weiss, 1927 82 OAKLEY SHIELDS /. Res. Lepid. TABLE 2 (continued) Species Realm Phyciodes tharos Nea. Poladryas pola Nea. P. p. arachne Nea. Polygonia egea Pal. P. faunus Nea. P, gracilis Nea. P, interrogationis Nea. Precis, 3 species Eth. P, orithyia Pal. Speyeria atlantis Nea. S. callippe comstocki Nea. S. c. meadii Nea. S, c. nevadensis Nea. S. coronis halcyone Nea. S, edwardsii Nea. S. egleis linda Nea. S. e. tehachapina Nea. S, hydaspe Nea. S. mortnonia Nea. S. zerene bremnerii Nea. S. z. platina Nea. Vanessa atalanta Nea., Neo., Pal. V. cardui Nea. Pal., Eth. , Ori., Aus. JY. c. kershawi Aus. _V, caryae Nea. V. dejeani Ori. Y. gonerilla Aus. V. indica Ori. V. itea Aus. V. myrinna Neo. V. tameamea Aus. Sources Scudder, 1887, 1889 new (Scott, Shields) new (Scott) Wynter-Blyth, 1957 Scudder, 1863, 1874, 1887, 1889 Scudder, 1887, 1889 Morrison, 1874; Scudder, 1887, 1889 Van Someren, 1955 Longstaff, 1912:46 Scudder, 1887, 1889 new (many) new (Scott) new (Shields) new (Scott) new (Scott) new (Ellis) Comstock, 1927; Emmel and Emmel. 1963a Shepard, 1966 Shepard, 1966 Edwards, 1887 new (Ellis) Mead, 1892; Dison, 1922; Moffat, 1922; Martin and Ingham, 1930; Shoumatoff, 1953; Muspratt, 1954; Emmel and Emmel, 1962 Edwards, 1884; Longstaff, 191Zt98, 162, 201, 385, 433; Dixon, 1922; Moff at. 1922; Seitz, 1924; Bryan, 1926; Weiss, 1927; Martin and Ingham, 1930; Tinkham, 1944; Tilden, 1961; Emmel and Emmel, 1962; Moucha, 1963 Longstaff, 1912’: 45 3 Edwards, 1884; new (many) Seitz, 1927 Hudson, 1898; Longstaff, 1912:482 Longstaff, 1912:385 Hudson, 1898; Longstaff, 1912: 448 new (Brown) Zimmerman, 1958 6(2):69-J78, 1967 HILLTOPPING 83 TABLE Z (continued) Species Vanessa virginiensis Realm Nea. Sources Mead, 189Z; Wright, 1908, 1930; Weiss, 1928; Martin and Ingham, 1930; Richards, 1931; Clark, 1932; Shoumatoff, 1953; Emmel and Emmel, 1962 V. V. brasiliensis Neo. new (Brown) Acraeinae Acraea ssp. Actinote, most spec. Eth. Van Someren, 1955 Neo, new (Brown) LYCAENIDAE Riodininae many species^ nearly 40 on one hilltop in Rio de Janeiro Neo. new (Brown) Dodona ouida Pal. Wynter-Blyth, 1957 Nymula calyce Neo. new (Brown) Lycaeninae Aphnaeus spp. Eth. Van Someren, 1955 Areas, many species Neo. new (Brown) Argiolaus spp. Eth. Van Someren, 1955 Atlides, many species Neo. new (Brown) A. halesus Nea. new (many) Callophrys dumetorum Nea. new (Shields, Thorne) C. viridis Nea, new (Gorelick) Celastrina argiolus echo Nea. new (Shields) C. a, ladonidas Pal. Longstaff, 1912:142 C. a. pseudargiolus Nea. Scudder, 1887, 1889 C. lanka Ori Longstaff, 1912:388 C. puspa Ori. Wynter-Blyth, 1957 C. singalensis Pal. Longstaff, 1912:46 Egumbia spp. Eth. Van Someren, 1955 Epamera spp. Eth. Van Someren, 1955 Evenus, many species Neo. new (Brown) Everes argiades Pal. Muspratt, 1954 E. comyntas Nea. Weiss, 1928 E. diporides Pal. Wynter-Blyth, 1957 Incisalia eryphon Nea. new (Dvorak) I. fotis Nea. new (Henne, Shields) I. iroides Nea. new (Thorne) I. iroides-fotis (hybrid) Nea. new (Henne) I. niphon Nea. Scudder, 1889 Lampides boeticus Aus. Bryan, 1926 Lepidochrysops spp. Eth. Van Someren, 1955 Leptotes marina Nea. new (Shields) Lycaena sallustius Aus, Longstaff, 1912:482 84 OAKLEY SHIELDS /. Res. Lepid. TABLE Z (continued) Species Realm Sources Miletis delic ia Aus. new (Wyatt) Mitoura spinetorum Nea. Shields, 1965 Myrina spp. Eth. Van Someren, 1955 Ogyris genoveva Aus. new (Wyatt) Polyommatus baeticus Pal. , Longstaff, 1912:64, Pratapa blanka Ori. Ori. 388 Wynter-Blyth, 1957 Pseudodipsas brisbanesis Aus. new (Wyatt) Satyrium behrii Nea. new (Emmel, Scott, Shields) S. calanus Nea. Weiss, 1927 S. saepium Nea. new (Emmel, Shields) S. titus Nea. new (Shields) Spindasis spp. Eth. Van Someren, 1955 Strymon columella Nea. new (Shields) S. melinus Nea, Tilden, 1964; new (many) Tajuria spp. Ori. Wynter-Blyth, 1957 Talicada nyseus Ori. Longstaff, 1912:98 Theda spp. Ori. Wynter-Blyth, 1957 Theclini, a great number of species Neo. new (Brown) Virachola spp. Eth. Van Someren, 1955 V. isocrates Ori. Wynter-Blyth, 1957 HESPERIIDAE Coeliadinae Bibasis sena Ori. Longstaff, 1912:357; Coeliades forestans Eth. Wynter-Blyth, 1957 Van Someren, 1955 Pyrrhopyginae many species Neo. new (Brown) Trapezitinae "The majority " of 51 species in Australia Aus. Waterhouse, 1932 Hesperilla idothea Aus, new (Wyatt) Toxidia peroni Aus. new (Wyatt) Trapezites iacchoides Aus. new tWyatt) T. iacchus Aus. new (Wyatt) T. phigaiia Aus. new (Wyatt) Pyrginae many crepuscular species Neo. new (Brown) Astraptes spp. Neo. new (Brown) Baracus vittatus Ori. Longstaff, 1912:388 Celaenorrhinus ambareesa Ori. Longstaff, 1912:391 Epargyreus spp. Neo. new (Brown) Erynnis afranius Nea. Snyder, 1894 (as lucilius); E, brizo Nea. Emmel and Emmel, 1962 new (Scott) 6(2):69-178, 1967 HILLTOPPING 85 TABLE Species Erynnie brizoj'bia.rgesBi E. b. lacustra E, horatiuB E, icelus E, martialis E. meridianus E. pacuvius E. £. callidus E, £. pernigra E, persius fredericki. E, propertius E. telemachus E. tristis Phocides spp, Sarangesa dasahara S. purendra Thorybes bathyllus T, confusis T, mexicana nevada T. py lades Urbanus spp, “Z^itusT’ dorus Hesperiinae Abantis paradisea A. tettensis Amblyscirtes vialis Hesperia Columbia ■H. comma H. harpalus ochracea H. harpalus yosemite H. juba H. leonardus _H. metea ^ FI, miriamae H. nevada H. pahaska H. p. martini H. p. williamsi H. pawnee H, sassacus H. uncas H. u. macswaini H. viridis 2 (continued) Realm Sources Nea. Freeman, 1951 Nea. new (many) Nea. new (Scott) Nea. Scudder, 1887, 1889; Tilden, 1964; new (Scott) Nea. new (Scott) Nea. preeman, 1951 Nea. Grinnell, 1904; new (Shields) Nea. Grinnell, 1904; new (Shields) Nea. new (La Due, MacNeill) Nea, new (Scott, Shields) Nea. new (Shields) Nea, new (Scott) Nea. Freeman, 1951; new (Shields) Neo, new (Brown) Ori. Longstaff, 1912*391 Ori. Longstaff, 1912:391 Nea. Weiss, 1927; Clark, 1936a -Nea. Clark, 1936a Nea. new (many) Nea. Weiss, 1927; Clark, 1936a Neo. new (Brown) Nea. new (Scott, Shields) Eth, Van Someren, 1955 Eth. Van Someren, 1955 Nea, Weiss^ 1927 Nea, new (Shields, Tilden) Nea, new (Grey) Nea, new (Scott) Nea, new (Shields) Nea. new (Roever, Scott) Nea, new (Grey) Nea. Clench, 1966 Nea. Garth and Tilden, 1963 new (Shields) Nea. new (many) Nea, new (Scott) Nea. new (MacNeill, Scott) Nea. new (Shields) Nea. new (Scott) Nea. new (Grey) Nea. new (Scott, Shields) Nea. new (MacNeill) Nea, new (Scott, Shields) 86 OAKLEY SHIELDS /. Res. Lepid. TABLE 2 (continued) Species Polites Sonora P. themistocles Stinga morrisoni Realm Sources Nea. new (MacNeill) Nea. Scudder, 1887, 1889 Nea. Tilden, 1961 new (Scott, Toliver) Nea. = Nearctic Neo. = Neotropical Pal. = Palearctic Eth. = Ethopian Ori. = Oriental Aus. = Australian Note: many of these species are listed from various sources which mentioned in passing that a species frequented topo- graphic prominences. Some of these may not represent hilltopping species and therfore further work to confirm or deny their validity is needed, in light of the mating- rendezvous definition of hilltopping. 6(2):69-178, 1967 HILLTOPPING 87 eight mating pairs of Diptera on Squaw Peak, Montana. Edwards (1960) noted mating activities in summit tachinid and syrphid flies. A more detailed analysis of this mating theory will appear in the discussion section. “Swarms” of male insects are associated with mating and are sometimes present on summits. Although Craig (1944) and Nielsen and Greve (1950) claim that certain Diptera swarms were not formed for mating, other workers have found that mating occurs when females enter male swarms in a variety of families (Knab, 1907; Gibson, 1942; Bailey, 1948; Downes, 1958; Blickle, 1959; Shemanchuk and Weintraub, 1961; Powell, 1964b; Thompson, 1967). Swarming of winged ants is also associated with mating (Wheeler, 1917b; Michener, 1948, 1960; Chapman, 1954b, 1957, 1963). In the biting Nematocera flies, the swarm develops visually to a “swarm-marker” such as a clearing or summit where both sexes respond to the marker; such swarms function to concentrate the population for the purpose of mating (Downes, 1958). Another theory often given to explain the presence of insects on summits is that wind bears them up, particularly warm air updrafts. This idea has been mentioned for insects in general by Bowditch (1896), Guppy (1897), Rowland-Brown (in Poul- ton, 1904 ) , Meinecke ( 1917 ) , Bryan ( 1926 ) , and Alexander (1940), and for winged ants by Wheeler (1905, 1917a) and Gregg (1947). As Chapman (1954a, 1954b, 1957) pointed out, however, updrafts may account for the presence of some but not all insects on summits (see Discussion). Current theories to explain “hilltopping” in butterflies are reviewed by Shoumatoff ( 1953 ) and Shepard ( 1966 ) . Appar- ently, Shepard (1966) is the only person to have made observa- tions of more than a casual nature on hilltopping butterflies by using a mark-recapture technique. It appears that little attempt has been made in the past to apply to butterflies the findings for other summit-frequenting insects. The different theories for the “hilltopping” of butterflies are listed in Table 3. Wind theories can be dispelled for the same reasons mentioned by Chapman (1954a) for insects (see Dis- cussion). The presence of larval foodplants on summits can be ruled out as a theory since many hilltopping species ascend far away from the foodplant area (see, for example, Clark, 1936a; Muspratt, 1954; Emmel & Emmel, 1967) as pointed out by Knudsen (1954) and Van Someren (1955). Males “liking” hill- tops as a playground, battleground, or assembly area or as a 88 OAKLEY SHIELDS /. Res. Lepid. TABLE 3. Butterfly hilltopping theories Theory Sources 1, Wind and/or updraft transported Newcomer, 1910 Guppy, 1925 Merritt, 1952 Beall, 1953 Knuds en, 1954 Rawson, 1955 Shepard, 1966 2. Wind on summit as enticement Beutenmuller, 1898 Muspratt, 1954 Emmel and Emmel, 1962 3. Unidirectional flight plus wind Beall, 1953 Shepard, 1966 4. "Liking" hilltops (urge to ascend) Scudder, 1863:620 Wright, 1906:86 Merritt, 1952 Shoumatoff, 1953 5. Ascend for "a s s embly ing " Beutenmuller, 1898 6. Male surplus Merritt, 1952 7, Male "playground" or "sporting ground" Comstock, 1927:20, 42, 265 Clark 1932:192 Clark, 1936a:24-25 Clark, 1936b:384, 405 Barrett and Burns, 1951:4-5, 144 8. Male "battleground" Wright, 1906:86. 112 Rawson, 1951 9. Tropism Merritt, 1952 10. "Phototropic urge" (attraction to warmth and light) Van Someren, 1955 11. Foodplant on summit Wright, 1906:112 Arnhold, 1952 Merritt, 1952 Emmel and Emmel, 1962 12. Congregation point for mating Seitz, 1909:13, 14 Moffat, 1922 Peile, 1923:62 Clark, 1932:192 Guppy, 1962 Emmel and Emmel, 1967 6(2):69-178, 1967 HILLTOPPING 89 response to a tropism urge does not go far in explaining the phenomenon because natural selection would eradicate this tendency for males to congregate away from females (Guppy, 1962). Surplus males congregating on summits due to the “extrovertive” male flight behavior is probably a misconception (Shepard, 1966). The idea that updrafts congregate butterflies due to the hilltop interrupting the butterfly’s undirectional flight (Shepard, 1966) is inadequate because the event occurs also on still days (Knudsen, 1954) and because of the "up-and-over” tendency observed for nonhilltopping species reported here. The ideas that hilltops are meeting areas for the sexes to mate, shown to be the explanation for certain other insects, will be presented and discussed in detail in this paper as the best ex- planation of the phenomenon of hilltopping in butterflies. MATERIALS AND METHODS Observations were made continuously from March 1966 to April 1968 (except for the months of July and August), with emphasis on the winter and spring months. A total of 245 hours on 170 days was spent observing and collecting on the summit of Dictionary Hill, San Diego County, California. Some observa- tions were made for an entire day, but most were for one to four hours, generally between midmorning and midafternoon. Visual sightings of species in the field were recorded. Activ- ities such as perching, feeding, chasing, courtship, mating, roost- ing, and predation were noted as they occurred. The occurrence and behavior of females on the summit were particularly check- ed to determine the validity of the summit-mating theory. Careful note was made of all species present on the summit and whether they exhibited hilltopping behavior or ''up-and-over” flight. The proximity of larval foodplants of these species to the summit was checked. Flight behavior in relation to wind and updrafts was noted. Butterflies were collected with an insect net in the air or when they alighted. All identifications were made to species. Exten- sive marking of male hilltopping butterflies was done to deter- mine the habits of individuals of each species. A total of 1,011 specimens was marked. The “1-2-4-7” system (Ehrlich & David- son, 1960) was soon found to be the best method for marking and satisfied most of the criteria for an ideal marking technique described by Gangwere et al. (1964). Eight different colored “Marks-A-Lot” felt-tip pens were used. The dye stain left on the wing was quick-drying, waterproof, and permanent without 90 OAKLEY SHIELDS J. Res. Lepid. affecting flight. Specimens could be marked quickly and recog- nized individually by this method. The maximum number that could be marked at one time with one color was 154. Papilioni- dae and Nymphalidae were held gently with wings shut between thumb and forefinger for application of the marks to the wing underside. The more delicate Pieridae, Lycaenidae, and Hesperi- idae were marked through the netting after being gently con- fined. Marked specimens were not placed in glassine envelopes except in release experiments. In most cases butterflies had to be recaptured for their numbers to be read. Marking had no apparent injurious effect on the individual. Released specimens either alighted, flew rapidly downhill, or resumed their previous activity. Many specimens marked that initially flew downhill soon returned. “Normal” activities were seen to resume in many marked specimens. For example, often two males that were chasing each other when netted continued doing so when released. Also, males frequently returned to i perch in a previous territory or “fly” area after marking. Speci- mens were not always released at the point of capture but were : always released on the summit immediately after marking. Number, sex, wing condition, and time of capture were recorded for each marked specimen. The Papilio zelicaon males used in release experiments away ‘ from the hill were netted, marked, placed in glassine envelopes, and stored in a cigar box which was kept in the shade. These experiments were carried out during periods when their popu- lation density was low so that their recapture was easier. P. zelicaon was used because its presence was conspicuous and it was relatively easy to recapture. Females were collected on tbe summit whenever possible to check if they were mated or virgin. Male Lepidoptera deposit sperm in membranous saclike spermatophores. The bursa copu- latrix of each female was later dissected under a binocular dis- secting scope to check for the presence of a spermatophore (see Ouye et ah, 1964; Burns, 1966; Taylor, 1967). Also, bursae from females of certain species from nonhilltop areas were dissected for comparison. A total of 548 bursae was examined during the survey: 281 from hilltop areas and 267 from nonhilltop areas. The two assumptions that Burns (1966) made are also assumed here: (1) the male transfers one spermatophore per mating, and (2) the spermatophore walls can be recognized even when collapsed. Females lacking a spermatophore were assumed to be virgin; such females were usually in fresh wing condition. 6(2):69-178, 1967 HILLTOPPING 91 Collected females were either stored under refrigeration or their abdomens were transferred to 80 percent ethyl alcohol for later inspection. Dried females were checked from previous collections. Their abdominal tissues were reclaimed for dis- section by the technique described by Van Cleve and Ross (1947), which consists of soaking the abdomens for 24 to 72 hours in a solution of trisodium phosphate diluted to 0.5 percent with distilled water. Weather readings were taken when in copula pairs were ob- served and at the time that hilltoppers first arrived for the day. Air temperature, relative humidity, and wind velocity were recorded on the summit by taking head-high readings with a Dwyer anemometer and a Bacharach sling psychrometer. Pacific Standard Time was used throughout the study. For comparative purposes, certain other summits in San Diego County were occasionally visited to mark and observe hilltop- ping species. These included a hill 997 feet in elevation, 1.7 miles south-southeast of the El Cajon Post Office; a hill 842 feet high 0.9 miles east of Dictionary Hill; Cowles Mountain; Tecate Mountain; Monument Peak; Mount Kentwood; and “Two Mile Hill,” one mile west of Scissors Crossing, 6.5 air miles east of Julian. Location and Description of Study Area Dictionary Hill is 1,064 feet in elevation and is located one mile south of Spring Valley and 1.75 mile north of Sweetwater Reservoir (T. 17 S., R. 1 W., Sects. 5, 32, 33), San Diego County, California (Fig. 1). The nearest higher hills are Mount Helix (1,373 feet), 2.9 air miles to the north-northeast; Mother Miguel Mountain (1,527 feet), 3.3 air miles to the south-southeast; and San Miguel Mountain (2,565 feet), 3.9 air miles to the southeast. Three lesser knobs on the eastern slopes of Dictionary Hill are 924 feet, 842 feet, and 677 feet high. The surrounding low- lands range from 200 to 800 feet below the summit of the hill. The hill consists of Jurassic-Triassic meta-volcanic rocks, with a rough stony land soil-type. The climate of the area is dry summer Mediterranean. La Mesa, California, at 560 feet, located three miles to the north of Dictionary Hill, has the fol- lowing climatic features: the average maximum temperature is 75.0° F, the average minimum temperature is 50.0° F, the aver- age mean temperature is 62.5° F, the average number of days with 0.01 inches or more precipitation is 44, and the average growing season is 327 days (Felton, 1965). 92 OAKLEY SHIELDS /. Res. Lepid. Fig. 1. Dictionary Hill (A) and Hill 842 (B), as seen from the southeast. 6(2):69-178, 1967 HILLTOPPING 93 The plant community on the slopes of Dictionary Hill is Coastal Sage Scrub, according to the classification of Munz and Keck (1965). Indicator plants present include California Sage- brush ( Artemisia calif ornica ) , California Buckwheat ( Eriogonum fasciculatum) , Golden Yarrow (Eriophyllum confertiflorum) , White Sage (Salvia apiana), and Black Sage (S. mellifera). Four conspicuous shrubs present are Broom Baccharis (Bac- charis sarothroides) , Monkey-flower (Mimulus puniceus) , Red- berry (Rhamnus crocea), and Laurel Sumac (Rhus laurina). Other plants on the slopes include Locoweed (Astragalus sp.), Slender Wild Oat (Arena harbata). Black Mustard (Brassica nigra). Blue Dicks (Brodiaea pulchella). White Forget-me-not (Cryptantha intermedia), Dodder (Cuscuta sp. ), Tansy-Mustard (Descurainia sp. ), Sweet Fennel (Foeniculum vulgar e), certain grasses, Deerweed (Lotus scoparius) , Plantain (Plantago Hook- eriana var. calif ornica) , Figwort (Scrophtdaria calif ornica) , Wild Pansey (Viola pedunculata) , and others. The summit of Dictionary Hill was graded at one time and presents an elliptically shaped flat area measuring about 230 by 275 feet (Fig. 2). This area is covered by Broom Baccharis. Most of the observations were made near sumac clumps at the northeast and southwest edges, where hilltopping activity was most intense. In the fall of 1958 a fire destroyed the vegetation on the western slope of Dictionary Hill. The area has since grown back con- siderably, but the fire probably destroyed a colony of. Chlosyne leanira wrightii, a species that was common on the summit from 1955-1958. Three males were collected there on April 3, 1960, but none have been taken since that time. Its foodplant, CaHil- leja sp., grew on the west slope. C. 1. ivrightii specimens used to fly in areas where Papilio zelicaon and Euphydryas chalcedona hilltopped on the summit ( Breedlove, personal communication ) . Thus C. 1. wrightii may have been another hilltopping species. RESULTS Species Present Forty-five species of butterflies were collected on the summit of Dictionary Hill during 1966-1968. A forty-sixth species, Chlosyne leanira wrightii^ has not been collected there since 1960 (Table 4 and Fig. 3 and 4). Twenty-one of these were considered as hilltoppers because of their territorial or patrol behavior of males confined to the summit, as opposed to the twenty-five non-hilltopping species that flew “up-ond-over,” fed without remaining, or did not congregate at the summit (Table Dictionary Hill summit with Broom Baccharis, 6(2). -69-1 78, 1967 HILLTOFFING 95 TABLE 4. Svimmit of Dictionary Hill, Spring Valley San Diego Coxinty, California. Species present collected by Oakley Shields Hilltoppers PAPILIONIDAE~ Battus philenor ■Papilio eurymedon Papilio zelicaon PIERIDAE Anthocaris cethura Pieris protodice NYMPHALIDAE Chlosyne leanira wrightiT””” Euphydryas editha "Euphydryas” chalcedona Speyeria callippe comstocki Vanessa atalanta Vanessa cardui Vanessa caryae Vanessa virginiensis LYCAENIDAE Atlides halesus Callophrys dnmetorum Celastrina argiolus echo Incisalia iroides Leptotes marina Satyrium saepium Strymon melinus HESPERIIDAE Erynnis hristis Total number of species = 46 0/0 of total that are hilltopper Non-hilltopper s PIERIDAE Anthocaris sara Colias eurytheme Colias harfordii Zerene cesonia Eurema nicippe Phoebis senna e Pieris rapae NYMPHALIDAE Agraulis vanillae Chlosyne gabbi Coenonympha tullia californica Danaus plexippus Danaus gilippus berenice Junonia coenia Nymphalis antiopa LYCAENIDAE w Apodemia mormo virgulti Brephidium exilis d-Everes comyntas Glaucopsyche lygdamus australis Lycaena helloides « Philotes battoides bernardino HESPERIIDAE Erynnis funeralis Heliopetes ericetorum Hylephila phyleus » Ochlodes sylvanoides Pyrgus communis = 45. 7 4^ = reported in the literature as hilltopping » = species that rarely showed a territorial or aggressive activity but did not concentrate on the summit 96 OAKLEY SHIELDS /. Res. Lepid. 5). Of these non-hilltopping species persent, both Apodemia mormo virgulti and Everes comyntas remained on and near the summit and one in copula pair was observed at the summit for both. However, these species were not particularly concentrated at the summit and occurred in about equal numbers on the slopes of Dictionary Hill as well. During 1960 K. Roever found 28 butterfly species on “A” Mountain summit, Tucson, Arizona (Table 6). The proportion of hilltopping species to non-hilltop- ping species at “A” Mountain and Dictionary Hill was remark- ably similar (46% at both locations were hilltopping species). Certain species were present in the canyon bottoms to the east, southeast, and south of Dictionary Hill (2,000-5,000 feet away from the summit) that were not collected on the summit: Pieris heckerii, Philotes sonorensis, Poanes melane, and Urbanus proteus (one). All but the last named were regular residents of the canyon bottoms, along with the non-hilltopping species Anthocaris sara, Nymphalis antiopa, Junonia coenia, and Ohio- syne gahhii that were occasionally present on the summit. Territorial and Aggressive Behavior Most of the specimens that showed hilltopping behavior were males. This fact was determined by visual observation and capture of the 21 hilltopping species. In those species where the sexes were difficult to determine visually (e.g., Vanessa and Erynnis)^ captured specimens were nearly always males. All of the hilltopping males except two pierid species estab- lished perch sites (F. protodice and A. cethura milled about without landing except occasionally to feed). The preferred perch site of V. atalanta, V. cardui, V. carye, and C. dumetorum was the ground or vegetation close to the ground. V. atalanta seemed especially partial to perching on rocks in clearings (Fig. 5). V. virginiensis was the only Vanessa species to consistently perch off the ground on vegetation (one to three feet up), al- though the others did so sometimes. Other species that perched on vegetation well above ground were B. philenor^ P. eurymedon, P. zelicaon (Fig. 6) (on the ground occasionally), A. halesus, S. melinus (Fig. 7), S. saepium, C. a. echo, and E. tristis (Fig. 8). E. chalcedona perched on the ground, ground vegetation, and rocks in clearings and sometimes on vegetation one to two feet up. B. philenor and F. eurymedon rarely alighted and flew almost continuously in milling over the summit. Once landed, perched species usually faced perpendicular to the sun’s rays. The pattern of establishing and returning to perch sites was noted. F. zelicaon flew back and forth over an area and alighted 6(2):69-178, 1967 HILLTOPPING 97 -fe §10 03 i7il 3 to^l o ^ .eo « K ^ 5^1 'g)2 S a Q ?3 ■Si i| l-s®!’-^ «• Q Qj - “ «~~sl °'C SrtOfe ■y S e„ 111 §> ^ ' ^ S 03 =l|ii'~' ! I-I^ i s /. Res. Lepid. ^ O S C73 CO Q •— --g fc --I oq =o S C « •IS' ^ Co'' S lll^ll -H o' 3 ^ .. O a ® a. c o S ^ ^-S S Co jj. .§■ s as s: 5,; - ^ ^oq s aS ffi I O ^ ^ - I o— •§ S oj S ■5N a -2 a. S a. o 'Sb» s Co O a bj , ^ i-H Co 3 Sj S r>q , 5 § in § CO S '^ , • g f-t 3 V. w § C/2 :S' ^ ■ 0^§) ^ S't' 3 §' ^ ;S o -§::.l|s.s S S^ . § o-^ •S a aO a ‘^ ' ' « '^ ' 0-- S oq^S^ C ^31-. -a. 1 3 Q CiT SQ o oq oq § Mh' 1 "S 00 s Q CO Co'' Co' 3 o "Si <50 C/3 S 'a 3 <:a ,Co "S "Ss CU Co ^•2 CO o " 5ca o, S-^ <50 ■ 2 i-i oq -c fc( -^Bs ■ £§- i^c OAKLEY SHIELDS 6(2):69-178, 1967 HILLTOPPING 99 TABLE Non-hilltopper s present on the summit of Dictionary Hill during 1966-1968 Species 1. Anthocaris sara 2. Colias eurytheme 3. Colias harfordii 4. Zerene cesonia 5. Eurema nicippe 6. Pieris rapae 7. Phoebis sennae 8. Danaus gilippus berenice 9. Danaus plexippus 10. Coenonympha tullia californica 11. Agraulis vanillae 12. Chlosyne gabbi 13. Junonia coenia 14. Nymphalis antiopa 15. Apodemia mormo virgulti 16. Brephidium exilis 17. Everes comyntas 18. Glaucopsyche lygdamus australis 19. Lycaena helliodes 20. Philotes battoides bernardino 21. Erynnis funeralis 22. Heliopetes ericetorum 23. Hylephila phyleus 24. -j-Ochlodes sylvanoides 25. Pyrgus communis Comments Both sexes flying over, mostly females, occasional ovipost- tion behavior. Both sexes flying over. One male flying over. One male flying over. Some seen flying over, one female collected. Both sexes flying and feeding. Two males flying over. One male flying over. Several seen flying over. A few males flying over. Two males collected, other specimens seen flying over. Male and female collected. A few seen flying over. A number seen flying over. Two males territorial. Both sexes present in numbers on summit but no concentra- tions; two in copula pairs collected near summit. Male and female feeding. Males and females flying near summit, one in copula pair collected near summitT Males flying over, females occasionally present. A female feeding. Two males territorial. A number seen, several males and a female collected flying over and feeding. Several males flying over, one male collected. One male feeding. Two males territorial. Occasional specimens flying over. T Not enough present to say if males preferred the summit or not. 100 OAKLEY SHIELDS /. Res. Lepid. TABLE 6. Sxommit of "A" Mountain, West of Tucson_ Pima Coixnty, Arizona. Species present during I960, collected by Kilian Roever Hilltopper s Non-hilltoppers PAPILIONIDAE PIERIDAE Battus philenor Anthocaris sara Papilio polyxenes Colias eurytheme asterius Eurema nicippe Papilio rudkini Nathalis iole Phoebis senna e PIERIDAE Zerene cesonia Anthocaris pima NYMPHALIDAE Euchloe hyantis lotta Pieris protodice Agraulis vanillae Asterocampa leilia NYMPHALIDAE Chlosyne lacinia Danaus gilippus strigosus Vanessa atalanta Danaus plexippus Vanessa cardui Vanessa caryae LIBYTHEIDAE Vanessa virginiens is Libytheana bachmanii LYCAENIDAE LYCAENIDAE Atlides halesus Strymon melinus 4* Leptotes marina HESPERIIDAE HESPERIIDAE Hesperia pahaska Erynnis funeralis williamsi Pyrgus communis Total number of species = 28 0/0 of total that are hilltoppers= 46. 4 4* = reported in this pape: * as hilltopping. 6(2):69~178, 1967 HILLTOPPING 101 Fig. 5. Male Vanessa atalanta in perched position. 102 OAKLEY SHIELDS J. Res. Lepid. Fig. 6. Male Papilio zelicaon in perched position. 6(2):69~178, 1967 HILLTOPPING 103 Fig. 8. Male Erynnis tristis in perched position. 104 OAKLEY SHIELDS J. Res. Lepid. on plants within the same area after leaving the first perch in chases. Three P. eurymedon observed on different days flew lazily about the northeast side of the summit for about ten minutes before landing on the same sumac bush. E. chalcedona usually perched at various places in a particular clearing after each chase from a perch. All the Vanessa species seemed in- different about where they would perch. They were not drawn to a particular spot or clearing and readily established a new perch area after a chase. All of the three A. halesus males perch- ed on a particular sumac on the northeast top. From these perch sites, investigative flights directed at the the same species, different species, and sometimes different in- sects were taken. When an individual of the same species pass- ed a perched male, the perched male flew up to investigate, gave pursuit, and frequently engaged in a “battle” with the other male. Perched P. zelicaon males rapidly pursued passing P. zelicaon for eonsiderable distances before breaking away and returning to the vicinity of the perch. Sometimes three or fuor P. zelicaon pursued in a chain. Frequently two P. zelicaon males would engage in a “battle,” especially when males first establish- ed their territories during the morning. They flew in tight circles around each other and climbed high into the air, sometimes locking legs and audibly beating wings. (Eff [1962] has observ- ed that Papilio indra minori males may physically damage each other in such encounters by beating their wings and falling to the ground while their legs are locked together. ) E. chalcedona, S. melinus, and C. dumetorwn were often seen to spin about each other in these high, climbing flights before breaking away and returning to the perch area. When two males of a Vanessa species met, they climbed at a slight angle to the ground, usually into the wind, with the pursuer behind the pursued. In all cases where identity could be determined by identifiable wing defects or marked specimens, it was the original male that returned after leaving the perch in the territorial species. This is what has been observed with territorial males in Hesperia (MacNeill, 1964), Hesperia metea (Shapiro, 1965), Agathymus evansi (Roever, 1964), and in hundreds of instances in the bot fly Cuterebra latifrons on hilltops (Catts, 1963). Investigative flights by milling pierids were noted. Typically when two A. cethura met, one chased the other, with the pur- suer often becoming the pursued. Frequently other A. cethura and sometimes a P. protodice would join. Up to eight A. cethura were seen in such a “cloud.” The two pierid species usually 6(2):69~178, 1967 HILLTOPPING 105 followed a stereotyped, predictable pathway in approaching and flying on the summit. Marked P. protodice and A, cethura did not remain on the summit for long periods but were recap- tured at intervals during the morning and early afternoon on the summit along their patrol pathways. Investigative and chase flights against different species by perched and patrolling males frequently occurred, but inter- specific battles were rarely seen. Often the other species was considerably smaller or larger or had a different color or different flight pattern. Table 7 lists some of the species that hilltopping species pursued. Investigative and chase flights of other orders of insects were noted for P. zeiicaon (black dragonfly), V. atal- anta (red dragonfly), E. chalcedona (black reduviid and black syrphid), S. melinus (flies, red dragonfly and yellow wasp), and E. tristis (flies). The behavior of a particular S. melinus male with chipped wings was observed in detail for 55 minutes on October 23, 1966. The entire time he perched on a sumac bush with wings half open and did not fly except to investigate flying insects. One continuous sitting period lasted 14 minutes. He was twice seen to pass the forelegs over the eyes and antennae while twist- ing the head, much as flies do when “cleaning.” This same in- dividual was noted to return after one elapsed day to the same sumac bush to perch. On following days his perch area was occupied by other S. melinus males. Daily Residence Various males of hilltopping species remained on the summit for long periods of time during the day (* = undisturbed ar- rival and departure observed ) : Species Battus philenor Papilio eurymedon"^ P. zeiicaon Atlides halesus^ Callophrys dumetorum Strymon melinus Celastrina argiolus echo No, I 1 1 1 1 2 1 Time in continuous residence on summit ( min. ) 70 37 110 82 91 30, 55 116 Most males recaptured more than once on the same day were not observed continuously. Maximum times between when first and last collected on the summit during the day for these were: 106 OAKLEY SHIELDS J. Res. Lepid. TABLE 7. Investigative and/ or pursuit flights observed among different butterfly- species on Dictionary Hill (Pursuer species listed first) P. zelicaon P. eurymedon A. cethura E, chalcedona V. caryae E. tristis A. cethura P. zelicaon P. protodice A. sara C. tullia P. protodice P. rapae A. cethura E. chalcedona P. zelicaon V. atalanta V. atalanta V, cardui V. caryae V. virginiensis V. virginiensis P. zelicaon V. atalanta V, caryae S. melinus V. caryae P. zelicaon P. protodice C. tullia V. virginiensis E. tristis S. melinus N, antiopa V. cardui V, virginiensis P. b. bernardino E. tristis S. saepium V. virginiensis C. dumetorum E. chalcedona Vanessa sp. E. tristis V. caryae 6(2):69-178, 1967 HILLTOPPING 107 Species Anthocaris cethura Pieris protodice Speyeria callippe comstocki Vanessa atalanta V. cardui V. caryae V. virginiemis Leptotes marina Erynnis tristis Elapsed time (min.) 213 140 150 123 12 87 82 25 45 An attempt was made on May 16, 1966, to recapture Euphy- dryas chalcedona males in the afternoon that were marked in the morning. E. chalcedona were marked and recaptured during two 80-minute periods, from 0800-0920 and from 1223-1343 hours. Thirty-one individuals were marked, 19 in the morning and 12 in the afternoon. Only about 10 individuals were seen that were not marked. Five specimens were recaptured: after 15, 44, 50, 246, and 318 elapsed minutes. Only two from the morning period were recaptured in the afternoon. The data suggest that there was a high rate of turnover during the course of the day. One marked P. zelicaon male was seen in territorial behavior on the summit on eight different days over a 20-day period ( De- cember 19, 1966, to January 8, 1967). This male frequented a particular area on the south side of the summit from which it “attacked” other male zelicaon in battles and pursuit flights. Fig. 9 is a summary of the times during the day it was observed. Numbers of specimens recaptured at different times on a given day (after at least ten minutes of elapsed time) on summits during this study are as follows : Battus philenor 1 Papilio eurymedon 13 P. indra pergamus 3 P. rtidkini 5 P. zelicaon 66 Anthocaris cethura 29 Pieris protodice 13 P. sisymhrii 4 Euphydryas chalcedona 13 Speyeria callippe comstocki 3 Vanessa atalanta 7 V. cardui 2 V. caryae 1 19 Dec 108 OAKLEY SHIELDS /. Res. Lepid. te* ^ ^ eo esi CM «M **» $ 3 1 V a 3 a n 1 d V33 H Fig. 9. Summary of times when a marked male Papilio zelicaon was observed over a 20-day period on Dictionary Hill summit. Dots represent individual recaptures and bars represent periods when continuously seen. 6(2):69-178, 1967 HILLTOPPING 109 V. virginiemis 5 Atlides halesus 3 Callophrys dumetorum 9 Celastrina argiolus echo 8 Leptotes marina 1 Strymon melinus 3 Erynnis tristis 2 These numbers indicate that at least a substantial number of hilltopping males remain for periods of time on a summit on a given day. Daily arrival as reflected by continuous collecting on a summit on a given day was determined for Papilio zelicaon, P. eury- medon, and Anthocaris pima (Table 8). In each case a majority arrived within the first hour after the first individual for the day had arrived, with the remainder arriving two to two-and-one- half hours after the first hour. Diurnal Periodicity Initial sightings of males in hilltopping activity were observed for the four Vanessa species. These first flew to the summit to hilltop only in the afternoons between 1220 and 1515 hours. Generally all four species were present on a given day. On six days, two to three of the species initially hilltopped together. The first species to be sighted in hilltopping behavior varied from day to day: First species No. days V. atalanta 19 V. virginiensis 15 V. cardui 12 V. caryae 11 Only the first Vanessa of the day to remain onthe summit in circling, aggression, and alighting behavior was recorded; those individuals feeding or flying over the summit were not con- sidered. For these Vanessa males, time of initial sighting, ambient air temperature, and relative humidity were recorded on 51 days during various times of the year. The time of initial sighting is correlated with air tempertaure and relative humidity ( Fig. 10 ) . The warmer the air temperature, the later they will arrive; the cooler it is, the earlier they will arrive. If the temperature remains constant, they would arrive earlier with decreasing relative humidity and later with increasing relative humidity. No correlations were apparent when relative humidity was plot- TABLE 8. Daily arrival reflected by continuous collecting of butterfly species on a summit 110 OAKLEY SHIELDS J. Res. Lepid. 0£bI-00bT 00H"0££I 0££I"00£T 00£T“0£ZT 0£ZT“00ZT 00ZT"0£n 0£n“00n 00IT“0£0T 0£0I-000T o 2 H g a r- xO ox i“H S rt fx. xO ox 0 s' 1 .s r«- xO fd 0 >x ’S 0 >x fl F—l 0) o £ ox H ?-l g 1—) o 0) N O rt o 0 u u fsS u O 0 ^ d ftf o ■d U d © o * B © 'd o 0 ?-i 03 ®?=l +j o © h h rt Pj o "i-i u ■■■ pH o • H u P^ ■*± © Cfl rt .H ^ CM o ^ r-, ^ CO nO r- CO m C« ^ CO ^ r- « CO ^ r- fNi' CO %J o o +J m 9 o s ca § © I O • H ?^ ffl o CO d o +j d ^ ' f3. © © ro 0 p. © • H d '.(H d ’3 1 s c d .m pH d i~i cti 4J tJ u rd >\ s! I’m u • H Cfl “rt rH © d u rtf W d u O “1 > >. P *f=4 u Cti to rd CO nl m (4 m N f% d o «H +j s ffl ce m m IM d ro « d ® d © d © a O pH pH d >. (d f«J fd rt Cti N © m > > > > O m o bfl • H d 'd N rd O. © X ® CQ ® ®H ^ o ■y ® ? m rH pH "d © « f=H ® « d £ ^ 'S p4 ^ 'd g : o O o u u ^ o « « » ^ O «H ♦ 114 ^3 c« 0) , rd 0) Cfl a (d o (d . ^ ^ cu I - I - A o 0) CQ Q) u 0^ Cb w OAKLEY SHIELDS sO fNJ in rg vO sD O ^ 00 (NJ m rg /. Res. Lepid. (d 0 +J . u Pu o pH cd CO o "h +J a 41 m > W m 0 Q o m xO fU ^ ^ xO S S o ® u rS 4J O m BO d ■d ‘ d * o >x . ^ 1 O -d © s © fa td u o r- • • o ^ rO S rH ' « d ' xo d 'g‘’2 -s' " «3 O r-t h ® ^ rd ft m W >' S ^ U-'OA*^ S ^ ® 3 -d « P O ^ o ^ C S -s 3 ^ m O m oj (M m I r- Ox d >0 h m a xp nx xO 0) 'P +J P-i © Q) y* X Li m © ^ fH “ s Q) 2 I Q o ^ h o m cti d bO O ^ U £ < m s 0x « r- . r- 6 ^ r*- o u d © TO ca d CO m d m ra g © u © ca cd 'd © o P4 rt f4 nJ CO 'd «H fi o .2 ® 'd o TO 4J • rt ■d © +j ca d bo d ‘3 ■d u o S4 d o ®H S3 « 2 TO TO cn ®6=l u O C0 d rt © o 14 •r-S Cti & • »4' A fd m ffl >x h 'd m a >x •d cu rt CD rt u d TO o TO d g g ca o s=H O (ti TO TO © 0 d «r=l 43 © © u +4 o « p. c d Si © ai d d cd fH d a (4 r™l o fa < W w W X > > fa fa xO b ® g .s ^ o © a TO 116 OAKLEY SHIELDS /. Res. Lepid. TABLE 12, Elapsed days from marking to recapture for certain hilltopping species on Dictionary HilL including separate entries for specimens taken on two or more different days. Number days P. zelicaon A. cethura P. protodice E. chalcedona S. callippe C. dumetorum S. melinus | E. tristis 1 1 38 1 6 6 1 3 1 1 2 18 5 6 1 2 3 13 5 3 5 1 4 16 2 1 3 1 1 1 1 5 15 2 2 2 6 4 1 7 22 1 3 2 2 8 16 2 4 1 9 8 1 2 2 10 8 1 11 1 12 3 13 7 1 14 5 1 15 4 16 1 1 17 1 18 1 1 19 3 20 2 21-26 (none) 27 1 28 2 29 1 total specimens 126 17 10 22 4 9 3 3 recaptured no. taken > once 35 3 1 6 2 4 2 1 O/O of recaptures 00 17. 6 10. 0 27. 3 1 ♦ 44. ‘ 4 ♦ « taken > once ^ = entry not very significant. 6(2):69-178, 1967 HILLTOPPING 117 first marked and had extremely worn wing condition and weak flight when finally recaptured. Papilio zelicaon Release Experiments One hundred and forty-three male P. zelicaon collected on the summit of Dictionary Hill were marked and released at various distances and degrees of the compass to see if they would ''home in” on the summit (see Table 13 and Fig. 11). One-third did return, about the same percentage that were marked on the summit and were recaptured there. In one in- stance two males that returned from the north also returned to the summit when released from the south. One male that was released nearly three miles away flew into a wind to return. P. zelicaon released far away from the summit homed in on a nearby hilltop. Ten male P. zelicaon and one male P. eurymedon collected on the summit of Dictionary Hill were transported four miles to the northeast and released. Two of the P. zelicaon and the P. eurymedon homed in on the summit of a hill 997 feet high 3,400 feet to the southeast of the release point. Three of 19 P. zelicaon males transported to the desert about 40 air miles away hilltopped on “Two Mile Hill” about one mile to the north- west of the release point. There is some evidence that P. zelicaon males will return to their own hill when given a choice of hills. Two groups of males, one group from the summit of Dictionary Hill and the other from the summit of a hill 924 feet high and 2,200 feet southeast of the summit of Dictionary Hill, were released together at loca- tion number 12 (Fig. 11): Date No. Elapsed days released No. released recapt. to recapture 111-19-67 6 (from Dictionary) 4 1, 1, 1, 6 5 (from 924^) 1 1 IX-27-67 4 (from Dictionary) 3 1, 1, 1 1 (from 92T) 1 1 On both dates all the recaptures were from the hilltop where the specimen was originally collected. The specimen collected on Hill 924 on September 28, 1967, was released on the summit of Dictionary Hill and returned the next day to the summit of Hill 924. Minimum speed of return to the summit of Dictionary Hill was estimated for four P. zelicaon males that were recaptured on the same day: 118 6 sb! vO r- Lfl fsT S o 9 m o tn in lo vO ^ O' -H ^ o in r- n- m m m m in 00 vO 00 o vO vO tn fsj (SJ (M m m vO nO cn so m vo O' o /. Res. Lepid. Specimens released at locations 7, 8, and 9 were recaptured on the summit of the 842 foot hill (distance = feet from summit of 842 foot hill): 7 560 3, 200 1-27-67 8 1 8 300 3, 200 I- 8-67 5 1 9 140 3, 800 II- 2-67 4 1 6(2):69-178, 1967 HILLTOPPING 119 Fig. 11. Release points for reared female ( $ ) and male ( $ ) Papilio zelicaon and the summits they “homed in” on: A == Dictionary Hill, B r= Hill 924, C = Hill 842, D =: crest near Hill 677. 120 OAKLEY SHIELDS J. Res. Lepid. Distance travelled Time elapsed Speed (ft.) ( min. ) (ft. /min. (1) 7,200 155 47 (2) 2,600 34 76 (3) 2,600 31 84 (4) 2,600 19 137 Specimen ( 1 ) travelled the distance when no wind was blowing, and specimens (2)~(4) travelled at right angles to a strong westerly wind. Eighty males from the stock used in the Papilio zelicaon female release experiment shortly to be discussed, were released on February 21, 1968, at location 6 (Fig. 12). Twenty-one (26.3%) of these were recaptured on nearby summits: Location Vertical feet Horizontal feet No. recapt. Elapsed days to recapture A 685 4,000 11 2, 2, 2, 2, 2, 3, B 545 3,200 5 3, 3, 3, 4, 4 4, 7, 7, 11, 18 C 460 5,500 1 9 D 220 1,300 4 7, 7, 8, 8 Each recaptured male 21 was collected and was not released back into the population. The results show that newly emerged males do not move to the same nearby summit when released from the same location. The individual at location C had to cross a ridge that blocks off location C from the release point. One wonders why the percentage of recapture (26.3%, N = 80) should be so similar to recapture for P. zelicaon resident males released away from the summit (33.6%, N = 143), recaptured resident males (32.4%, N = 389), and for recaptured resident males of Papilio rudkini (26.7%, N =: 15). An experiment was devised to test whether or not virgin fe- males as opposed to recently mated females seek the summit. Papilio zelicaon adults reared from eggs were used; the stock came from five females collected in copula on the summit of Dictionary Hill, January 11-14, 1968. Their progeny were reared indoors on Sweet Fennel, Foeniculum vtdgare. Forty-five fe- males were marked and released. Females emerged over a seven-day period prior to the release date of February 21, 1968. After emergence the adults were kept in boxes under refriger- ation. One-half (22) of the females were hand-paired to wild- caught and reared males by the technique described by Clarke and Sheppard (1956) and half were left virgin. These two 6(2):69~178, 1967 HILLTOPPING 121 Fig. 12. Fourteen release points from which Papilio zelicaon males collected on Dictionary Hill summit returned to the summit of Dictionary Hill (1064 feet) and Hill 842 (842 feet). 122 OAKLEY SHIELDS /. Res. Lepid. groups were released together late in the afternoon 2,000 feet (385 vetrical feet) to the northeast of the summit (Figure 11). During the subsequent four days the summit was intensively collected; the results were as follows: Date No. females unmated No. females mated Feb. 22 10 1 Feb. 23 3 1 Feb. 24 0 0 Feb. 25 0 1 13 3 In other words, 59.1 percent of the virgin females and only 13.6 percent of the mated females were recaptured, yet all of these 16 recaptured females proved to be virgin when dissected. Thus the three “mated” females recaptured apparently did not ade- quately mate when hand-paired, but the high number of un- mated recaptures suggests that the mated females not recaptured very likely received a spermatophore when hand-paired. Percent Virginity The percent virginity of species from non-hilltop areas varied from 0 to 30 percent, mostly under 10 percent (Table 14). In addition, T. E. Pliske kindly supplied the following unpublished data on percent virginity for species from non-hilltop areas: Papilio palamedes, 0.0 percent (N = 32); P. troilus, 0.6 percent (N = 358); P. glaucus, 0.0 percent (N = 171); Ascia monuste, 0.0 percent (N = 100); and Danaus gilippns berenice, 1.5 percent (N =: 194). These females were randomly collected at flowers at various places in Florida during the summer of 1961. O. R. Taylor (unpublished) dissected females of many United States and Costa Rican species from non-hilltop situations and found 5 to 20 percent virginity. Colias virgin females constituted 10 to 20 percent of the populations (N > 100). He was able to obtain virgins of most species where the sample was greater than six. The high percentages of virgins collected on summits (Tables 15 and 16) are in sharp contrast to samples taken in non-hilltop areas. In 21 hilltopping species where N was less than six, 15 (71.4%) were 67 percent or more virgin. The percentages of virgins in seven species, where N was nine or greater, varied from 0 to 97 percent. (There is some question whether or not Celastrina argiolus cinerea, whose percent virginity was zero, is a hilltopping subspecies.) Burns (letter dated June 28, 1967) says that the only virgin female Papilio glaucus he has examined (N > 100) came from “atop a pronounced escarpment” in Dallas County, Texas. TABLE 14, Spermatophore counts for species collected in non-hilltop areas 6(2). -69-178, 1967 HILLTOPPING 123 O h o > ^ (0 tfl CO o 0 ^ ^ f| ^ ■§ Sa fl o' O P( •u fn ^ ^ X X X •TJ T3 PI P! 4) 4) X m cn rt* cn fM CO oa m o m (NJ CO ra vO CO CO 00 l> CO 00 LfJ h- sO fM o ^ M fi- ^ 2 l-H pt 0. o < X •H 'd Pi o TO 43 ■d cd TO 43 d N-l 43 d d 43 d o cd u fd h X •H Pi rtf • fH 1—1 pH 'O rH s (ti 0 d 43 • H P. Pu Pi N cuja(|CL*|PL,jC^|a.| e ^lO QO ^ CO -4 04 ^ ^ 00 <-H csj m o 1=0 j4|Wi£ TABLE 19 (continued) 132 OAKLEY SHIELDS /. Res. Lepid. ^1 (C 0) 4) II I— I gj 4) SI I I 4) IP^1 Lf) uT) m •H* "i-r vo sD so a £3 i I 1 s ’ C 4) 4) O CO cn CO CO GO DO bO &0 GO GO GO GO 4) 4) 4; 4) *1^ ^ nJ S ^ g I 2 o3 3 CO p. GO O GO a. 43 ^ sO —I F-l P-H fSj P-) CO 0- f- ^ 04 vO CO If) > a. . o <\U CO 4) p (ti CO CO 0 CD U 43 > P ’’0 ’•4-> d >- ca ^ . rt +-> CO 0 d pj pd CO S CO CO 43 GO 4) E ra ■M w GO s fH 0 43 43 pd w 0 S H 43 ^ n 43 P 4J S 43 - .g '• II r' L 6(2):69-178, 1967 HILLTOPPING 133 sighted for a total of 32 females, or 56.1 percent of the total male population. During February 11, 1967 to March 12, 1967, there was a period of high male density; 123 were marked on 15 different days. During that time only five females were cap- tured, or only 4.1 per cent of the total male population. Thus, during times of low density populations there may be more than half as many females as males present. Due to the difficulty of finding females in copula, as many females as males may actu- ally have been present. The population of Papilio eurymedon on Dictionary Hill is low in density compared to certain other summits in San Diego County, where males are frequently plentiful. Dictionary Hill is probably on the periphery of its range. During January to May 1967 only 12 different males were marked, collected, or seen on the summit, and three females were collected and one seen for a total of four females, or 33.3 percent of the total male popu- lation. Female Behavior and Mating Papilio zelicaon. These observations are based primarily on 77 females (58 virgin, 19 non-virgin) collected on Dictionary Hill summit. They were collected while in flight or alighted; 17 in copula pairs were collected and other courting and mated pairs were observed. Wing condition, as expected, showed a high correlation with mated condition; virgins almost exclusively had a fresh wing condition while most non-virgins had a fair to worn wing condition. Only one of the non-virgins had a freshly deposited spermatophore; the remainder had spermatophores that were collapsed. Frequently virgin females’ genital regions were expanded when captured. All females collected appeared on the summit during the time when males were resident and well after the males had established territories for the day. Females on the summit usually flew quite slowly; virgin females with fresh wing con- dition characteristically had a slow, fluttering flight, generally in areas occupied by males. When all or nearly all of the resi- dent males were collected, females flew back and forth over the summit, occasionally alighting on the ground or vegetation; three females stayed on the summit for six, seven, and eleven minutes respectively in these lingering flights. Females that were frightened off the summit by a swing of the net flew downslope but generally returned within a few minutes. (Edwards [1884] reported the flight behavior of a female 134 OAKLEY SHIELDS /. Res. Lepid. 1' i;' Papilio brevicauda on a mountain top at Topsail, Newfound- land: “It made long flights, rarely alighting, but apparently reconnoitering the whole mountain.” The female was probably a virgin since she layed infertile eggs when confined. ) Virgin females, including two taken in copula, often had pollen adhering to their bodies and were twice noted to visit flowers before flying to the summit. Courtship stages were seen in twelve pairs. The females flies into an area where males are perched or patrolling; the male immediately investigates her and responds by fluttering close behind her in a bobbing flight. At times this flight is done im- medately in front of the female. Presumably during this flight the male is emitting a pheromone. Males as old as four weeks have a sweet pungent odor, while freshly emerged males do not. Copulation itself may take place downslope or on the summit. In copula pairs were found either slightly downslope (as far as 125 yards from the summit) or on the summit. The female alights at the edge of the summit or usually slightly downslope. The time the pair first meets to when their genitalia are in contact usually takes about one to three minutes. The male alights behind and parallel to the female or underneath the females and curves his abdomen until their genitalia are in con- tact. He then establishes a position to that each is facing in opposite directions and oriented with their backs to the sun. In most cases the female’s wings are shut or partially open and the male’s wings are opened flat. Pumping motions of the male’s abdomen last for a few minutes. In one pair the female used her back legs to stroke the male’s abdomen and genitalia inter- mittently. Pairs found in copula were resting on vegetation one to four feet off the ground (Fig. 13). The female carries the quiescent male when the pair is disturbed. The male initiates the uncoupling of the in copula pair. One attempted courtship by more than one male was rejected by a virgin female. From one to four males chased the female. The female flew rapidly rather than slowly fluttering when pur- sued. Once a female broke into a fluttering, falling flight when closely investigated by a male Vanessa atalanta. Anthocaris cethura. Fourteen females were collected on or near the summit of Dictionary Hill; of these, nine were virgins and five were non-virgins. Three females were taken in copula. Virgin females were mostly taken while in flight or when they had alighted in areas where males patrolled. One virgin female was collected feeding on a flower on the summit. 6(2):69-178, 1967 HILLTOPPING 135 136 OAKLEY SHIELDS /. Res. Lepid. A virgin female was observed on the summit from 1009-1018 hours on April 27, 1967. During this time the female alighted four times (for periods of two minutes, three-and-one-half min- utes, one minute, and one minute respectively) on vegetation in areas along a 100-foot strip where males in the past had patrol- led. When perched, the female flew up to meet two passing male Pieris protodice and once flew up after a male Speyeria callippe comstocki. In both cases the protodice briefly courted the female but separated after five to ten seconds. At 1018 the female was collected. A pair in coptda was taken resting three feet up on vegetation on the summit at 1209, March 16, 1967. The female was posi- tioned downward and the male upward. The male carried the quiescent female when the pair was disturbed. Another pair was taken in copula on March 21, 1968, at 1050. The female contain- ed a collapsed speratophore indicating that she was mating for the second time. Mating was seen with one female on February 28, 1967. At 1023 and 1102 hours fresh virgin females appeared at a rock outcropping area which males frequent on the summit. One of these was immediately collected and the other was observed. The observed female fluttered in an area with many nectar sources for about one minute and alighted on these plants with- out feeding. The female then alighted on some grass, perched facing upwards, and remained there, with wings shut, one foot off the ground. A marked male with fair wing condition, which had been recaptured three times on the same day, flew by the area and immediately dropped down to the female, alighting parallel to her, and initiated copulation. The female then posi- tioned herself so that the male faced up and she faced down. The pair remained in this position with wings shut for six min- utes, during which time a spermatophore was passed as shown by later dissection. Four different males, two of which were collected, flew by the area. In each case, the male suddenly dropped down to the mated pair, attempted copulation for about ten seconds, then flew off. The female may have emitted a pheromone that attracted males who were in close proximity, or the male’s response may have been strictly visual. After six minutes the pair was collected. On March 25, 1967, two non-virgin females were collected very near the summit during oviposition behavior. One oviposited on tansy mustard, Descurainia pinnata, a known foodplant that grows up to the summit of Dictionary Hill. The other female 6(2):69-178, 1967 HILLTOPPING 137 hovered over black mustard, Brassica nigra plants, alighted on one small mustard plant, and curled her abdomen toward the plant; however, no egg was found. Pierk protodice. Observations are based on eleven females collected on Dictionary Hill summit: two virgins and nine non- virgins. One of the virgins was captured feeding on a flower, and the other was taken in copula. Two of the non-virgins were taken in oviposition behavior directed toward young Brassica nigra plants. Abbott (1959:288) described in detail the courtship and mat- ing of P. protodice in Texas. One non-virgin female on Diction- ary Hill summit displayed acceptance behavior toward a hover- ing male (0854, May 14, 1968): i.e., wings spread and abdomen elevated while resting on vegetation. A female with a freshly deposited spermatophore exhibited rejection behavior toward a courting male (1258, February 3, 1967, on Hill 842, east of Dic- tioary Hill). The female, pursued by a male, alighted on the ground. The male alighted alongside of the female and tried unsuccessfully to make genitalia contact. He then walked around in front of the female, faced her, and flew off. An in copula pair taken at 1058, February 20, 1967, on Dictionary Hill summit, was located because two males alighted by the pair. In two in copula pairs the male carried the quiescent female when the pair was disturbed. Vanessa species. Despite numerous observations of male en- counters in hilltopping Vanessa species, only two courtships of females were observed. This inability to observe mating pairs was probably due to the formation of in copida pairs in the late afternoon when few observations were made. The two court- ships occurred at 1513 and 1533 hours on January 6, 1967, only two hours before sunset. Vanessa mating times have been re- ported as 4:15 p.m. for V. atalanta, 6 p.m. for V. cardui (both Pronin, 1964), and 8:30 p.m., D.S.T. for V. cardui (Temple, 1953). The two pairs observed in courtship were V. cardui and V. caryae. In both cases the courting pair alighted on the ground with the male behind the female and were then col- lected. Strymon melinus and Incisalia iroides. No matings of these two species were observed. I. iroides males were only occasion- ally seen, but S. melinus males were frequently encountered in the afternoons. Again, probably the reason no matings were observed was that pairs may have formed in the late afternoons when few observations were made. Reinhard ( 1929 ) states that 138 OAKLEY SHIELDS J. Res. Lepid. S. melinus mated in Texas cotton-fields “late in the evening or shortly before sunset.” Powell (1964a) noted that four pairs of 1. iroides mated on a lemon tree only after 4:00 p.m., P.S.T. Callophrys dumetorum. Two in copula pairs were collected on the summit of Dictionary Hill. At 1018, March 25, 1967, an in copula pair, both sexes with fresh wing condition, was found on an unopened composite flowerhead two feet above the ground. Both individuals were horizontal with wings folded, broadside to the sun. Their resting site was at the edge of a clearing where a male was exhibiting territorial behavior. Courtship and mating were observed with a pair at 1010, March 16, 1967. A virgin female with fresh wing condition flew from the north into a male’s territory on the summit. The male, which was perched and had a worn wing condition, flew up to investigate the female and hovered behind her. The female then alighted almost immediately two-and-one-half feet up on vege- tation and quivered her partly opened wings. The male alighted parallel and slightly behind the female, curved his abdomen around, engaged genitalia, and positioned himself so that he faced downward and the female upward. All these actions hap- pened rapidly. The pair faced the sun and had their wings closed. After nine minutes the pair was collected; a spermato- phore had been passed during that time. Erynnis tristis. Females were difficult to distinguish from males in flight. They were recognized as females only when in copula and during courtship. Essentially all of the 35 females collected on the summit were virgin and had fresh wing con- dition. All but one of the 14 females with freshly deposited spermatophores were taken in copula on the summit. All females appeared when males were actively hilltopping. Courtship stages were seen partially or completely in six different pairs. The virgin female flies into a male’s territory on the summit. The male flies off his perch to investigate the female and flutters behind and below her, occasionally rising up and making apparent contact. The female responds by alighting on vegetation one to four feet above ground ( a rock in one instance served as the substrate). The male immediately alights parallel and slightly behind the female and curves his abdomen to make genitalia contact. Upon engagement the male positions himself so that he is facing down and the female faces up. Females which were closely examined at this stage had numerous scales along the shaft and nudum of their antennae. These were prob- ably androconial scales from the male’s coastal fold deposited 6(2);69-178, 1967 HILLTOPPING 139 during the brief “contacts” before alighting. Burns (1964:88) predicted that these scales “probably serve a communicative function during courtship.” The female next grasps the substrate rapidly with her legs and begins to brush oflE the scales along the shafts of her antennae with her front legs. Next, either the left or right leg of the female twitches rapidly, sometimes out and back and sometimes in apparent contact with the pair’s genitalia region. The tip of this leg has a tuft of hairs not present on the other hind leg. I did not notice where these hairs were taken from. This leg twitching is intermittent and lasts about three minutes. Finally, the female cleans all the scales off her antennae with her front legs, at the same time rubbing her eyes and coiling and uncoiling her proboscis. The male is quiescent during all of these motions except for rhythmic contractions of his abdomen. Soon the pair ceases all motions and rests in a position about 30 degrees from the horizontal (Fig. 14). These motionless pairs could be approached quite closely, although several times such a pair took flight after prolonged close obser- vation. Such flights extended 10 to 50 feet away, the female always carrying the quiescent male. All of the in copula pairs noted ( 18 ) that were not seen in courtship were found resting in or near a known male territorial area. Frequently, passing males would investigate such a pair and attempt copulation but would leave very shortly. All pairs were perpendicular to the sun’s rays. On sunny days a quiescent pair would have their wings closed, but on overcast days or dur- ing the passage of a cloud across the sun their wings were ex- panded. Excessive numbers of males apparently terminated one court- ship. At 1355, February 12, 1967, a female was seen pursued by two males. The female alighted several times on branches but each time flew off. A third male joined the chase when the group passed through his territory. At this point the female flew rapidly off with the three males in pursuit. Momentarily the group was lost from sight, but shortly the three males returned chasing each other. The whole encounter took about three min- utes. A rejection flight by a freshly mated female was seen at 1400, January 27, 1967. A male fluttered behind the female. The female rose high into the air, and the male rose behind her, touched her, and returned to ground level. Immediately the female returned to ground level, where the same process was repeated. The female then alighted on a bush and was collected; she contained a freshly deposited spermatorphore. OAKLEY SHIELDS Fig. 14, In copula pair of Erynnis tristis. 6(2):69~178, 1967 HILLTOPPING 141 Behavior of Sexes after Uncoupling The behavior of eight Papilio zelicaon pairs after uncoupling on the summit was observed. If conditions were warm or mild, the females flew rapidly and continuously downhill until lost from sight; one such female was followed for one-third mile. On cool days the females unsually sunned for awhile before flying downhill; their flight was much slower with frequent alightings on vegetation to sun themselves. In all cases the females depart- ed from the summit and did not fly around the summit like the virgin females. Departing females flew into and at random angles to the wind. Males resumed hilltopping behavior shortly after uncoupling. In one instance the male that had uncoupled from a pair 375 feet from the summit was found 20 minutes later in copula with another female at the summit. Two Erynnis tristis pairs were observed after uncoupling on the summit on a cool, overcast day (February 19, 1968). In both cases the female sunned on the ground for several minutes before rapidly flying off the summit. With one of these pairs, arrival, copulation, and departure of the female was seen during a 61-minute period; the male resumed territorial behavior in his original territory two minutes after uncoupling. Larval Foodplant Proximity Tables 20 and 21 give the known larval foodplants for hill- topping and non-hilltopping species and whether they are locat- ed near or far away from the summit of Dictionary Hill. There was no difference in the percentage of hilltopping and non-hill- topping species whose foodplants were near or far, as measured arbitrarily from the 900-foot contour line (distance from the summit to the line is 400 to 1,400 feet and 164 vertical feet): Larval foodplant present above 900-foot contour Present Absent % Absent Hilltoppers 14 7 33.3 Non-hilltoppers 17 8 32.0 Certain hilltopping species had to fly a moderate distance to hilltop. Papilio eurymedons nearest foodplant source is a colony of about twenty Rhamnus crocea bushes 1,600 to 2,100 feet to the east (300 vertical feet). Erynnis tristis feeds on Quercus agrifolia (Burns, 1964). The nearest Q. agrifolia tree is 2.6 miles away, the first area of concentration is 3.1 miles away, and the main center of concentration is 3.8 miles away to the east. Quercus dumosa also grows in the area and may be a possible foodplant. The first bushes (five) of Q. dumosa are 1,900 feet away and 142 OAKLEY SHIELDS /. Res. Lepid. TABLE 20. F oodplants for hilltopping species on Dictionary Hill Species Larval foodplant present on slopes above 900' contour Known possible foodplants avail- able in the area 1, Battus philenor absent Aristolochia sp. 2, Papilio eurymedon absent Ceanothus sp. , 3. Papilio zelicaon absent Rhamnus californica, R. crocea, Prunus sp. Foeniculum vulgar e. 4, Anthocaris cethura present Daucus sp, Descurainia sp. , 5. Pieris protodice present ma)fee Brassica nigra Brassica nigra. 6. Chlosyne leanira present Raphanus sp, Castillija sp. 'X wrightii 7. Euphydryas editha present Plantago Hookeriana 8. Euphydryas chalcedona present var. californica Scrophularia californica 9. Speyeria callippe present Viola pedxincxilata comstocki 10. Vanessa atalanta absent Urtica urens, thistles 11. Vanessa cardui present Boraginacea, 12 Vanessa caryae present Compositae, Malvaceae (Malva parviflora preferred) Malva parviflora 13 Vanessa virginiensis present preferred, other Malvaceae Gnaphalium sp. , other 14. Atlides halesus absent Compositae Phoradendron flavescens 15, Callophrys dumetorum present Eriogonum fasciculatum. 16. Celastrina argiolus present Lotus scoparius Ceanothus sp. , echo Lotus sp. 17. Incisalia iroides present Cuscuta sp. , 18. Leptotes marina present Ceanothus sp. , Sedum sp. Astragalus sp. , 19. Satyrium saepium absent Plumbago sp. Cercocarpus betuloides. 20. Strymon melinus present Ceanothus sp. many 21. Erynnis tristis absent Quercus agrifolia Foodplants compiled from Comstock (1927), Klots (1951), Emmel and Emmel (1963b), Garth and Tilden (1963), Burns (1964), and Thorne (pers. comm. ). Higgins (1949) was used as an aid to finding certain plant locations. "A Butterfly no longer present. 6(2):69A78, 1967 HILLTOPPING 143 TABLE Zl. Foodplants for non- hilltopping species on Dictionary Hill 1. Anthocaris sara present Brassica nigra 2. Colias eurytheme present Medicago sativa. 3. Colias harfordii present less frequently other legumes Astragalus sp. 4. Zerene cesonia present Amorpha sp. , 5. Eurema nicippe absent Trifolium sp. Cassia sp. 6. Pieris rapae present Brassica nigra. 7. Phoebis sennae absent cultivated Cruciferae Cassia sp. 8. Danaus gilippus absent Asclepias sp. 9, berenice Danaus plexippus absent Asclepias sp. 10. Coenonympha tiillia present Gramineae 11. californica Agraulis vanillae absent Passiflora sp. 12, Chlosyne gabbi present Corethrogyne sp. , 13. Junonia coenia present Hazardia sp. Plantago sp. , 14. Nymphalis antiopa absent Gnaphalium sp. Ulmus sp. , Salix sp, , 15. Apodemia mormo present Popxilus sp. Eriogonum fasciculattrm 16, virgulti Brephidium exilis present Atriplex sp, , 17. Everes comyntas present Chenopodium sp. Astragalus sp. , 18. Glaucopsyche lygdamus present Trifolium sp. Lotus scoparius. 19. Lycaena helloides absent Astragalus sp. Rumex sp, , 20. Philotes battoides Ternardino present Polygonum sp. Eriogonum fasciculatum 21. Erynnis funeralis present Lotus scorparius. 22. Heliopetes ericetorum present Medicago sativa, Nemophila membranac( Malvastrum sp. , 23. Hylephila phyleus absent Amaranthus sp. Cynodon Dactylon 24. Ochlodes sylvanoides present Gramineae 25. Pyrgus communis present Malvaceae 144 OAKLEY SHIELDS J. Res. Lepid, the first area of concentration is 4,600 feet away to the east. Erynnis tristis belongs to a superspecies of Quercus feeders; no other foodplants are known for the group (Burns, 1964). Freshly emerged Papilio zelicaon males hilltopped as far as 5,500 feet away from the release point. Three hilltopping species were seen to oviposit on the summit: Anthocaris ceihura, Pieris protodice, and Euphydryas chalcedona. Since they do oviposit as well as mate on the summit, the per- centage of virgin females for these species would be less than expected if only mating took place. These three species did have the lowest percentages of virginity of the hilltopping spe- cies present (see Table 15) yet were still above the highest percent virginity for non-hilltop areas. Species Approach to the Summit Instances of actual flight to the summit by hilltopping species were watched for although rarely observed. Particular attention was given to local wind conditions when species did approach the summit, in view of the emphasis placed by various authors on updrafts and winds as causative agents in hilltopping. Male Papilio zelicaon were twice noted at distances greater than 100 feet from the summit to approach rapidly in a straight line flight about four to five feet off the ground. In one case there was no wind noticeable; in the other case, a fair breeze was blowing at right angles to the butterfly’s flight. On Febru- ary 13, 1967, 27 marked P. zelicaon males were released 2,600 feet to the north of the summit. Three of these were recaptured on the summit about a half-hour later. During this time a steady, fresh breeze was blowing from the west, presumably at right angles to their flight paths. Pierids were easily watched in their summit approaches be- cause of their conspicuous white color. Pieris protodice males were twice noted to approach the summit from the east along a ridge and once from the south slope. These flights occurred when no breeze was noticeable and when a slight breeze was present; one flew at right angles to a breeze. Anthocaris cethura males mostly approached the summit of Dictionary Hill from the south or southeast slopes. One male approached the summit from the east side while a wind was blowing from the northwest. On February 18 and 21, 1967, on Two Mile Hill, many cethura were seen to approach the summit on the south slope while fresh breezes blew from the southeast. Like protodice^ cethura males approached summits over a particular, stereotyped flight pathway. 6(2):69~178, 1967 HILLTOPPING 145 The four Vanessa species approached the summit of Diction- ary Hill from the east and sometimes from the west. Their straight approach flight, observed from the summit, changed to circling and alighting behavior once the top was reached. Erynnis tristis males were frequently seen at the summit in apparent approach flights from the east. Only one was seen to approach from any distance from the summit. This male was seen about 200 feet down the north slope in a steady uphill flight and became territorial on the summit one minute later. In May of 1966 and 1967, four species were hilltopping abund- antly on Dictionary Hill between 0700-0800, two hours after sunrise, when updrafts, if present, would have been slight. In summary, approach flight by butterflies was noted both into and with winds of various strengths and at various random angles to the wind, and during times when updrafts were both maximal and minimal. Effects of Wind at the Summit Winds had some effect on the summit distribution of hilltop- ping species. Males of the four Vanessa species, Papilio zelicaon, and Euphydnjas chalcedona were noted at times to remain on or close to the summit during fresh breezes (17-21 knots on the Beaufort Scale for wind) by flying into the wind. During these winds, hilltopping species mostly conflned their activities to the leeward side. The Vanessa species often flew into fresh after- noon breezes blowing from the west to maintain position along the east edge of the summit. P. zelicaon males on Dictionary Hill often conflned their hilltopping activities to the leeward side of the summit but did not buck fresh breezes to remain on the summit when the breezes were generally over the whole hill. E. tristis males conflned their territorial activities along the east edge of the summit during west winds. Normally they spaced themselves over much of the summit. Females of Papilio eunjmedon, P. zelicaon^ Anthocaris cethura, Euphydryas chalcedona, and Speyeria calUppe comstocki flew usually quite rapidly into the winds in approaching or flying on the summit. One female zelicaon flrst arrived on the summit with the winds, but when blown from the summit it flew into the wind to return to the hilltop. The effect of winds above the hilltop was seen when two P. zelicaon females were swept up high from the summit between 1130-1200 on January 15, 1967 (calm at ground level), and when a small migration of Vanessa cardui headed to the northeast in 146 OAKLEY SHIELDS /. Res. Lepid. the afternoon of March 30, 1966. The cardui flew out about 50 to 100 yards from the summit and then were carried up and out of sight; the wind was from the southwest at a steady 7 to 9 mph. Feeding Actual feeding on flowers by hilltopping butterflies was not often seen. Frequently species showed hilltopping activity when no flowers were available on or near the summit, such as during the fall and winter months. Perching behavior on flowers in a territory often occurred and was distinguished from feeding behavior in which the proboscis entered the flower. Papilio zelicaon territorial males on the summit often had pollen adhering to the underside of their bodies. Eight marked male zelicaon captured between 1100-1400 hours had adhering pollen. These males varied from fresh to worn wing condition which suggests that feeding probably occurs throughout their life span. Occasionally males were seen feeding on flowers at at the summit before resuming hilltopping behavior. One male in the early morning was seen to leave a roosting site on the summit and to feed down the east slope without exhibiting hilltopping behavior (0725, May 9, 1967). Another male (0853, April 16, 1967), the first one to show hilltopping behavior that day, sipped water while perched on a sumac leaf. One male was seen feeding on the summit as late as 1522 (May 1, 1967); no other zelicaon males were seen on the summit after that time on that particular day. Presumably male zelicaon feed in the morning before hilltop- ping and probably later in the afternoon before roosting. They may also feed away from the summit during periods of the day when actively hilltopping since certain marked males were absent from the summit at times. Some feeding by hilltopping males on the summit was seen but generally no flowers were available there. Perhaps these zelicaon have feeding territories, as was shown to exist in a Papilio glaucus population in Marvland (Fales, 1959). Virgin females of Papilio zelicaon occurring on the summit, including several taken in copula, often had pollen adhering to their bodies. Several were seen to feed on or near the summit before performing their characteristic “circling” flight on the summit. One female fed intermittently on flowers for 30 minutes on the north slope about one-quarter mile from the summit; it had an uncollapsed spermatophore in the bursa. This particular female had no pollen adhering to its body; thus individuals 6(2);69-178, 1967 HILLTOPPING 147 taken on the summit with no pollen could still have fed. Virgin females apparently feed for a time after pupal emergence before seeking the summit to mate. The Vanessa species fed extensively on flowers in the morning in home gardens surrounding Dictionary Hill, especially on lan- tana. At 1150 on April 13, 1966, many Vanessa individuals fed on yellow composites on the north slope, but none were hilltop- ping up to the departure time of 1310. Occasionally Vanessa individuals fed on or near the summit but always before start- ing their afternoon hilltopping behavior. One V. caryae male fed on the summit as early as 0725, two hours and twelve min- utes after sunrise (May 2, 1967) when the temperature was 54 °F. During the height of Vanessa hilltopping activity, indi- viduals did not feed, although when they began to hilltop there was some overlap of feeding individuals and hilltopping individ- uals. Numerous Euphydryas chalcedona males on the summit were sometimes seen feeding on yellow composites and at the same time displaying aggression toward each other and passers-by. One mated female was collected while feeding on the summit. Other species noted feeding on the summit included a male and a virgin and mated female of Anthocaris cethiira, a male and a virgin female of Pieris protodice, a female Pieris rapae, a male Speyeria callippe comstocki, a female Chlosyne gabbi, a virgin female Leptotes marina, a male Callophrys dumetorum (feeding in its territorial area), two male Erynnis tristis, and a male Erynnis funeralis. Roosting Crane (1967) mentions that five species of Heliconius in Trinidad roost gregariously, the same individuals returning to the same bushes or vines on successive nights. McFarland (1965) says that the lycaenid Callophrys macfarlandi roosts in clumps of Nolina, its foodplant. Ministrymon leda also roosts on its foodplant (Mesquite) in southern Arizona. Such gregarious roosting and foodplant roosting were not noticed on the summit of Dictionary Hill. Rau & Rau (1916:251-257) describe sleep postures of some butterflies and include an annotated bibliog- raphy on the subject. Roever (personal communication) says that Vanessa species roost in palo verde trees on the summit of “A” Mountain near Tucson, Arizona. The vegetation on the summit of Dictionary Hill is extensive so that roost sites were difficult to locate. During April and May, 148 OAKLEY SHIELDS /. Res. Lepid. 1967, eleven attempts were made to locate species roosting by inspecting the summit early in the morning and late in the after- noon. Both hilltopping and non-hilltopping individuals were found roosting. Non-hilltopping individuals included a male Nymphalis antiopa, two female Euphydrijas chalcedona, and a female Papilio zelicaon (all three females contained spermato- phores). Roosting males of hilltopping species included two Vanessa atalanta, one V. cardui, one Speyeria callippe comstocki, one Euphydryas chalcedona, and 14 Papilio zelicaon; these roost- ed mostly in weeds along the edge of the summit. The one E. chalcedona male and four of the P. zelicaon males were re- captures from previous days. One P. zelicaon male was recaptur- ed roosting on three successive mornings along a 55-yard strip of weeds. On April 30 this specimen sunned itself at 0940 and was captured in hilltopping behavior at 1020. Roosting specimens were found most often in the early morning on the summit vege- tation with closed wings or with wings open to sun themselves. P. zelicaon males when disturbed from sunning positions flew quite rapidly and alighted again on vegetation almost immed- iately. Many hilltopping species of flies were noted roosting on the summit on Broom Baccharis. Catts (1963) found that at least some Cuterebra latifrons ( Cuterebridae ) males spent the night in chaparral on a hill summit and resumed territorial behavior on the summit at temperatures of about 19° C. Catts (1964), working with the oestrid flies Cephenemyia apicata and C. jellisoni, mentions that males roosted in and near hilltop aggre- gation sites. Predation Knudsen (1954) reported several hundred butterflies of three hilltopping species in an area of a few hundred square feet on the summit of Kennesaw Mountain, Georgia, and Waterhouse (1932) collected over 300 butterflies of three species on top of Kosciusko in Australia in one day. About 25 to 100 butterflies are usually encountered on a given day during late February, March, and April on the summit of Dictionary Hill. Concen- trated predation on such large numbers of butterflies was ex- pected but not seen. Vespula wasps have been known to prey on a hilltop swarm of winged ants ( Chapman, 1963 ) , and there is a report of sparrows attacking a “mud-puddle club” of Papilio glaucus canadensis (Rawson & Bellinger, 1953), but no such attacks were observed during this study. 6(2);69~178, 1967 HILLTOPPING 149 Predators reported attacking adult butterflies include the fol- lowing: robber flies (Fryer, 1913; Dover, 1920; Field, 1938; Klots, 1951; Price, 1961), dragonflies (Scudder, 1889; Fryer, 1913; Dover, 1920; Field, 1938; Klots, 1951; Price, 1961), Hymen- optera (Scudder, 1889; Ford, 1957), Hemiptera (Field, 1938; Macy & Shepard, 1941; Klots, 1951), mantids (Field, 1938), crab spiders (Field, 1938; Klots, 1951; Voss, 1953), web weaver spiders (Klots, 1951), frogs (Klots, 1951; Price, 1961), toads (Klots, 1951), lizards (Dover, 1920; Annandale & Dover, 1921; Field, 1938; Klots, 1951; Ford, 1957), tree-snakes (Dover, 1920; Annandale & Dover, 1921), many different birds (Scudder, 1889: 1612; Fryer, 1913; Dover, 1920; Annandale & Dover, 1921; Car- penter, 1937, 1941a, b; Field ,1938; Klots, 1951; Knowlton, 1953; Ford, 1957; Urquhart, 1960:209; Olson, 1962; Petersen, 1964), bats (Field, 1938), chipmunks (Morris, 1953), and monkeys (Field, 1938; Klots, 1951). During this study no butterfly predation was seen. However, certain potential predators such as lizards and birds were oc- casionally present on the summit of Dictionary Hill, Also, terri- torial male butterflies often had beak-marks or “beak injuries” from birds (see Carpenter, 1937), although such predation at- tempts may have occurred away from the summit. A flycatcher (Myiarchm cinerascens) was sighted on eight different days over a 25-day period (presumably the same indi- vidual). The bird perched for extended periods and frequently darted out after passing insects. A whippoorwill {Chordeiles sp. ) was sighted on the summit one late afternoon for one-half hour and one morning briefly. Swift lizards and horn-toad lizards were occasionally present. Crab spiders {Misumenoides forma- sipes, determined by B. Kaston) were present on the summit in curled sumac leaves in late September, 1967, and were not pres- ent in sumac bushes checked below the summit. Some of these spiders fed on hilltopping flies. Conceivably these spiders might also eat butterflies that perch on the sumac leaves, such as Vanessa virginiensis and Strymon melinus. DISCUSSION In this section a number of topics will be discussed in rela- tion to hilltopping behavior in butterflies. The probable function of hilltopping in butterflies as a means of congregation to facili- tate mating will be advanced, and other methods that butterflies may use to congregate for mating will be pointed out, along with some discussion of the courtship devices used. The func- 150 OAKLEY SHIELDS J. Res. Lepid. tion of territoriality in relation to hilltopping will be developed, and comparisons will be drawn between hilltopping insects and hilltopping butterflies. Special attention will be given to wind and updrafts in relation to hilltopping. In addition, possible methods of orientation in approaching summits and the extent to which hilltopping has developed in butterflies will be describ- ed. Shepard (1966) defined a “hilltopping” butterfly as a “species which is found on a hilltop in only one stage of development [adult]; the food source of the larvae is not present; the insects are not forced there by a macro-environmental factor such as strong winds.” In light of the results of the present study, this definition should be modified: ''Hilltopping' in butterflies is a phenomenon in which males and virgin or midtiple -mating fe- males imtinctively seek a topographic summit to mate. The gen- eralized behavior pattern of hilltopping species is given in Fig. 15. Hilltopping apparently does occur in some species whose larval foodplants are present on or near a summit as well in species that fly to a summit a considerable distance away from the nearest foodplant. Territorial or patrol behavior at the summit is the most conspicuous aspect of the hilltopping male butterfly. The idea that hilltopping butterflies congregate on summits for the purpose of mating is mentioned occasionally in the liter- ature. Seitz (1909) said that the sexes in Papilio machaon and F. podalirius meet on summits for mating but did not cite sup- porting evidence. Moffat (1922) suspected that Vanessa cardui and V. atalanta individuals on a hilltop at Wexford, Ireland, were males; each “attacked” all newcomers “to keep the ground to himself in the hope that ultimately a female would appear.” Moffat also thought that hilltopping was a peculiarity of the mating season since during a different time of the year Vanessa individuals were in lowlands but not on summits, while neither species was seen in the lowlands when hilltopping. (However, V. car did and V. atalanta hilltop only in the afternoons, so that this observation was probably a reflection of the time of day rather than the time of year.) Peile (1923) noted that Melitaea trivia persea males in Mesopotamia frequent ridge crests; he found one pair in copula on a crest and concluded that ridge crests are probably their mating sites, “the males waiting there to waylay the females as they come by.” Clark (1932:192) men- tioned that Papilio polyxenes asterius mates on summits but gave no details. Temple (1953) reported that Pararge megaera mated after “all moved uphill to a stony arid place.” Wyatt (1957a) 6(2):69~178, 1967 HILLTOPPING 151 found two in copula pairs of Papilio machaon aliaska on a ridge summit where hilltopping males were active. Guppy (1962) concluded from observations on Oeneis nevadensis that hilltop- ping serves to bring the sexes together. He did not report any actual matings. Guppy developed certain theoretical implica- cations of this mechanism. He pointed out that hilltopping is probably an instinct to fly to a central point in newly emerged individuals; females would lose this instinct once mated and would depart to oviposit. Natural selection would eradicate the habit if no mating occurred. An important point to stress that Guppy mentions is that females would be on summits only for the brief time to mate and thus a collector could easily miss seeing them. Emmel and Emmel (1967) reported that Papilio indra kaibabensis males “hilltop” on the rims of the Grand Gan- yon, Arizona, where presumably virgin females fly up to mate, several thousand feet higher than their nearest foodplants. Other methods for bringing the sexes together for mating besides hilltopping probably include males flying to treetops; over rockslides; through bogs; in canyons, stream courses, and gullies; and around the larval foodplant. Jackson (1961) sus- pected that members of the lycaenid subgroup Lipteninae in Uganda ascend to treetops to mate since most of the females collected there had fresh wing condition. Examples of each type are given in Table 22. It remains for future investiagtion to establish the importance of these methods for mating in butterflies. Catts, Garcia, and Poorbaugh (1965) found that males of the oestrid fly Hypoderma lineatum aggregate probably for mating along the open margins of streams in pastureland ravines. Cer- tain other members of this family mate at summit congregation sites. Mating of Sirex noctilio, a siricid wasp, occurs on treetops where males periodically swarm (Morgan, 1968). Certain other species in this family are known to “hilltop.” In moths certain members of the Lasciocampidae, Bombycidae, Saturniidae, and Arctiidae mate by the males homing in, sometimes over long distances, on a pheromone that the virgin female releases (But- ler, 1967; Ewing and Manning, 1967). No such long-range method has been noted for butterflies. Certain butterfly congre- gations probably serve no mating function, such as male congre- gations for feeding (Reinthal, 1966) and butterfly migrations (Williams, 1930). If hilltopping is a mating mechanism in insects, one would ex- pect this behavior pattern to be developed to a greater extent 152 OAKLEY SHIELDS J. Res. Lepid. TABLE 22. Probable methods of bringing sexes together for mating in butterflies besides hilltopping Method Examples 1. Tree-top seeking Delias spp. (Seitz, 1927:123), Eriboea spp. , Charaxes spp. , Euripus consimilis, Euthalia garuda (last four from Wynter- Blyth, 1957). 2. Rock- slide fliers Erebia magdalena, Chlosyne damoetas, Lycaena snowi, L. hypophlaeas. 3. Confined to bogs Colias scudderi, Oeneis jutta (Klots, 1951), Boloria eunomia (Klots, 1951), Speyeria nokomis, Incisalia lanoraieen- sis (Klots, 1951). 4. Canyon, gully, and stream fliers a. Territorial species Asterocampa celtis, Chlosyne gabbi, Euphydryas chalcedona ssp. , Junonia coenia, Limenitis spp, , Lycaena hermes, Apodemia mormo ssp. , Lephelisca wrightii. b. Patrollers Papilio eurymedon, P. multi _ caudata, P, rutulus, Anthocaris lanceolata, A. sara, Pieris beckerii, Philotes sonorensis. 5, Confined to larval foodplants (recognition of foodplant by males) Neophasia menapia, N, terlooti, Adelpha bredowii, Brephidium exilis, Philotes spp. , Plebejus icarioides, Habrodais. grunus, Hypaurotis crysalus, Mitoura gryneus, M. loki , M, siva, Satyrium dryope, S. sylvinus, Ochlodes yuma, Pseudocopaeodes eunus. Examples from personal experience except where otherwise noted. 6(2):69-178, 1967 HILLTOPPING 153 in groups that are rare, parasitic, predaceous on ephemeral prey, or whose larval foodplants are scattered or rare than in common species or species that mate at the larval foodplant. Such groups are more likely to need a common meeting ground to facilitate mating. The following evidence supports this hypothesis. 1. Chapman (1954a) concludes from an insect survey of the summit of Squaw Peak in Montana that “parasitic and not usually abundant groups of insects were well represented.” 2. Dodge and Seago (1954) report that flies collected by net on Georgia summits produced many undescribed or little-known species for which the females and life histories of most were un- known. Their entire net-collected Sarcophagidae represented more than 16 saprophagous species and more than 36 parasitic species. 3. Hilltop congregations are reported for all the major groups of parasitic bot flies (Catts, 1964); Cuterebridae, Gastrophilidae, Hypodermatidae, and Oestridae. Grunin (1959) points out that large concentrations of male bot flies on summits indicate the scarcity or absence of males in the surrounding territory and since bot flies generally have scattered, low density populations, these gathering points are probably mating places. A number of matings of bot flies on summits has been reported by Gatts (1963, 1964). 4. Hagen (1962) summarizes the characteristics that aggre- gating Goccinellidae beetle species have in common; most of these congregate on some prominent object that forms a silhou- ette on the horizon, including peaks and hills. The species that aggregate are associated with “ephemeral” prey, chiefly aphids, and not with sessile Homoptera. Four large tribes which do not have many species that aggregate feed mostly on scale insects, mealybugs, or fungi — all sessile prey. Species that do aggregate in these tribes have ephemeral prey. Aphid-feeding species of Goccinellini that do not form aggregations feed mostly on colon- ial, arboreal, non-migrating aphids. Hagen suggests that “ephem- eral” food led to the selection for long dormancy in the absence of food and for aggregation to bring the dispersed sexes together. Mating occurs mostly at these aggregation sites before the beetles disperse at the end of dormancy. 5. There is some evidence to show that rarer species of butter- flies are more likely to hilltop than abundant species. J. Scott (in manuscript) collected butterflies randomly in Gregory Gan- yon, Boulder Gounty, Golorado, from March to May, 1966, to see if species present that are known to hilltop were rarer than 154 OAKLEY SHIELDS J. Res. Lepid. non-hilltopping species. Using number of individuals collected per hour as an index, he found for 32 hilltopping species and 34 non-hilltopping species that hilltopping species averaged a third as abundant as non-hilltopping species. Although there was considerable overlap in the amount of abundance, the upper and lower ends of the scale differed significantly: Number of Number of non-hilltopping hilltopping species species One or more per hour 10 3 One-fourth or less per hour 11 21 Of the three most abundant hilltopping species, Polites themis- todes and Incisalia eryphon apparently are only weak hilltoppers, and Celastrina argiohis cinerea may not hilltop as evidenced by the high percentage of mated females (see Table 15) collected by Scott, although C. a. edw definitely does hilltop in southern California. Another advantage of hilltopping besides the congregation of adults that occur in low numbers or are scattered is the develop- ment of a stabilized gene pool from the centralization of isolated populations during mating ( Catts, 1963 ) . Wind and updrafts as causative agents for hilltopping in butterflies seem unlikeb for the following reasons. Some butter- flies approach summits into the wind or at random angles to the wind, as well as flying with the wind. Butterflies actively seek and stay on hilltops during windless days, sometimes before updrafts would have much effect. Butterflies maintain a position on or near the summit even during fresh breezes. Euchloe olijm- pia (Arnhold, 1952) and Boloria polaris gronlandica (Munroe, 1951 ) have been noted to return to summits after being blown off the tops by winds. Butterflies that are non-hilltoppers fly up-and-over the summit and show no such “staying” behavior. Vanessa species hilltop in the afternoons when winds would have much more effect than updrafts; these species often ap- proached the summit into the wind as well as with the wind. Shepard (1966) noted the behavior of hilltopping Pieris oc- cklentalis males in relation to winds on Slate Peak, Washington. No specimens were seen approaching the top from the west where winds blow directly up out of the valleys. All arrived from the east slopes where winds were limited until the summit 6(2):69-178, 1967 HILLTOPPING 155 was reached. They had to fly against the wind to reach the very summit. Click ( 1965 ) states that certain species of butterflies are wind- propelled by their manner of flight when flying directly into a strong wind. They keep their wings vertical and open and close them alternately to offer the least wind resistance. The butterfly is driven forward by the wind eddying against the wing under- surface. Winds and updrafts in relation to other hilltopping insects parallel the findings with butterflies. Chapman (1954a) found on Squaw Peak, Montana, that “early morning summit activity was frequently considerable by 8:00 a.m. when updraft currents would be expected to be slight if present at all.” Dodge and Seago ( 1954 ) noted summit activity of flies during moderate to strong winds. Aldrich (1915) reported that hovering hot flies actively maintain themselves on summits during strong breezes. Hurd (1920) mentioned that some beetles were carried by ascending winds and precipitated on a summit against their flight powers. However, Edwards (1956, 1957a) suggests that winds do not draw Coccinellidae beetles to summits. He men- tions that they sometimes fly to and stay on summits on windless days as well as flying against the wind in approaching summits. Chapman ( 1954b ) noted in detail the effects of winds on the summit behavior of winged ants on Squaw Peak, Montana. Maximum numbers of ants were present on warm, calm days when updrafts would have their greatest effect. Sometimes the ants had difficulty maintaining themselves on the summit against the wind but were not noted to swarm in calm eddies present just below the summit. He states that air currents may play a role in transporting ants to summits but are not entirely re- sponsible for summit swarms of ants. The evidence for this is that ( 1 ) ant swarms are localized to the actual summits and are not present along approaches where wind eddies exist, (2) look- out stations reporting “tens of thousands” of ants included grassy or brush and forest covered ridges and hills where thermal up- drafts are not well developed, and (3) updrafts strong enough to be a transportation factor would carry ants up and away from summits. In his study of Squaw Peak summit insects, Chapman (1954a) listed the following reasons why updrafts do not account for the presence of all insects on summits: (1) updrafts would not be symmetrical enough to concentrate insects over a peak as opposed to slightly lower points, (2) updrafts may carry insects 156 OAKLEY SHIELDS J. Res. Lepid. up and away from the peak, (3) updrafts are unlikely to be so selective for species and sex, and (4) updrafts would not ac- count for “the continued presence of the large strong and fast fliers” which could easily leave the peak at any time. Mani (1962) distinguished between insects actively seeking summits and insects passively transported by wind in the Hima- layas. Vertical convection currents lift many insects including heavy-bodied forms up from the plains, where they are trans- ported by upper-air winds, chilled, and dropped on snow and glaciers. Winged insects alive on summits are visitors from other biota lower down and have been observed ascending even when no updrafts are present and sometimes against prevail- ing winds, according to Mani. Thermal updrafts on mountains are discussed by Malone (1951:663), Chapman (1954b), and Geiger (1965:407-408). Updrafts are most noticeable on southern slopes and are negli- gible on northern slopes. The are clearly defined on warm calm days. They are strongest at 20 to 40 meters above the ground where they average two to four meters per second. Updrafts are strongest in canyons and gullies and are weak on projecting ridges. In the northern Rocky Mountain region, updrafts gen- erally start about 9:00 a.m. and continue to late afternoon. Updrafts reach their greatest intensity at one-and-one-half to two hours after maximum insolation. Orientation flight toward the summit by both sexes in butter- flies is almost certainly visually directed. Release experiments with Papilio zelicaon males on Dictionary Hill suggest that they can “home in” on the summit when released at various distances and directions and in various wind directions and velocities. Given a choice, male zelicaon could discriminate between which hilltop they were collected from and some other hilltop. When male zelicaon were released at considerable distances away from the hilltop, one could predict which new hilltop in the vicinity they would seek. Female zelicaon are probably responding like the males in seeking a summit and are not brought there by random flight because of the large numbers that are virgin and because only virgins sought the summit in a release experiment with mated and unmated females. Certain insects are known to orient to distant objects (silhou- ettes) on the horizon. lersel and Assem (1965) found that fe- males of the wasp Bembix rostrata return to their nests by orienting to tree tops as far as 80 meters away on the horizon. Schneider (1962) reports that the beetle Melolontha vulgaris 6(2):69-178, 1967 HILLTOPPING 157 after hibernation selects the area on the horizon with the great- est average height (hypotaxis) within a radius of about 3.2 kilometers during a spiral flight. Once the area is located, the beetles maintain a steady flight to the area, which is a forest or hill. Presumably a similar type of orientation and directional flight exists for males and virgin or multiple-mating females in hilltopping insects. Kennedy (1939), investigating the upwind flight of the mosquito Aedes aegypti, found that the mosquitos maintained upwind flight orientation by visually referring to the apparent movement of the ground. In hilltopping insects, a combination of visual orientation to the ground and to the horizon would seem necessary to approach summits. Once the summit is reached, the hilltopping butterfly begins territorial or patrolling behavior. In essence, the whole summit can be considered a macro-territory for the butterflies present, with micro-territories and patrol pathways established within this territory. Beusekom (1948) found that the female wasp Philanthus triangulum perceives micro-topographic landmarks in finding the nest, a behavior learned during one or more orien- tation flights. A similar learning process probably takes place in butterflies that establish territories on summits. Nobel’s (1939) definition of territory as “any defended area” has been applied to a wide variety of animals. Alexander ( 1961 ) points out that two phenomena are widely associated with territorial animals: (1) a tendency to stay in restricted areas, or to repeatedly return to specific areas, or both; and (2) aggressive behavior displayed toward intruding individuals. Carpenter (1958) reviewed territoriality in vertebrates — it is exhibited by fish, amphibians, reptiles, birds, and mammals. A general discussion of territoriality as it applies to insects is given by Alexander (1961:172-174) and Catts (1963:83-84). Catts (1963) states three criteria for proving the existence of territoriality in tagged male Diptera on summits: (1) the stationing of marked males at summits, (2) frequent intraspecific encounters with in- truding males, and (3) diurnal permanence of male stations within a specific area. Presumably conditioning by familiarity with the microenvironment allows the male to dominate in his territory (Nice, 1941; Hinde, 1956). Possible advantages due to the spacing of a congregated, territorial male population in- clude: (1) decreased chances of mass predation by a few predators, (2) less time spent in intraspecific aggression, (3) in- 158 OAKLEY SHIELDS J. Res. Lepid. creased frequency of male-female encounters, and (4) decreased interference to courting and mating pairs by other males (Catts, 1963; Lin, 1963). Catts (1963:83) emphasizes the need to ob- serve marked individuals before territoriality can be shown to exist in a species. Territorial behavior has been proven by mark-recapture methods for certain insect species. In Diptera territoriality is exhibited by the families Cuterebridae (Catts, 1963) and Oestridae (Catts, 1964). In Hymenoptera it is exhibited by Megachilidae (Jaycox, 1967) and Sphecidae (Lin, 1963). In Odonata territoriality is exhibited by certain dragonflies (Jacobs, 1955; Ito, 1960; Kormondy, 1961; Johnson, 1962b, c; Moore, 1964) and damselflies (Bick and Bick, 1963). In Orthoptera it is exhibited by Gryllidae (Alexander, 1961). Strong circumstantial evidence for territoriality ( a perch area combined with aggressive behavior in males) also exists for the following insect groups: Diptera in the families Gastro- philidae (Grunin, 1959) and Hypodermatidae (Grunin, 1959); Hymenoptera in Andrenidae (Linsley, 1958; Cazier & Linsley, 1963), Apidae (Janzen, 1964; Bennett, 1966; Gruden, 1966), and Scelionidae (Wilson, 1961); Coleoptera in the genus Necrophorus (Pukowski, 1933); and Odonata in dragonflies (Moore, 1952; Johnson, 1962a, 1964; Pajunen, 1962; Young, 1965) and damselflies (Bick & Bick, 1965; Bick & Sulzbach, 1966) . Strong circumstantial evidence exists for territoriality in various Lepidoptera species as well (Table 23). Ross (1963) marked males of two species of Hamadryas in an effort to prove or disprove territoriality in these species and found that they did not frequent definite perch areas but did display aggression. Pugnacity is known in a large number of butterfly species. Shapiro (1966) reported pugnacious males for one papilionid, two pierids, seven nymphalids, four lycaenids, and 18 hesperiid species in Delaware Valley, Pennsylvania. Characteristically, aggressiveness, or pugnacity, of territorial male insects can be divided into four parts: (1) investigation flight toward intruders into the male’s territory, followed by (2) chase of the intruder, or (3) intraspecific fight, or (4) a return to the perch area (Lin, 1963; MacNeill, 1964). Generally the male is perched, or resting, at a perch site in his territory until he investiagtes passing intruders (MacNeill, 1964). However, some species “patrol” a territory (e.g., the butterflies Papilio thersites, Battus philenor, Anthocaris cethura) and perch briefly 6(2):69-178, 1967 HILLTOPPING 159 TABLE 23. Butterfly species that have territorial behavior, presumed from circumstantial evidence (perch area or site combined with aggressive behavior) Species Sources PAPILIONIDAE Chilasa clytia Wynter- Blyth, 1957 Papilio thersites Shoumatoff, 1953 NYMPHALIDAE Apatura iris Richards, 1927:304 Charaxes species Van Someren, 1955; Wynter-Blyth, 1957 Eriboea species Wynter-Blyth, 1957 Hypolimnas misippus Stride, 1956 Lethe creola Klots, 1951 L, portlandia Klots, 1951 Limenitis astryanax Shapiro, 1966 Nymphalis antiopa Hargitt, 1915; Clark, 1937; Shapiro, 1966 Oeneis nevadensis Guppy, 1962 Phyciodes tharos Klots, 1951 Polygonia comma Tietz, 1952 Vanessa gonerilla Hudson, 1898 V. itea Hudson, 1898 LYCAENIDAE Lycaena phlaeas americana Klots, 1951 Satyrium titus Shapiro, 1966 HERPERIIDAE Abantis paradisea Van Someren, 1955 A. tettensis Van Someren, 1955 Agathymus evansi Roever, 1964 Atrytonopsis hianna Shapiro, 1965 Bibasis sena Wynter-Blyth, 1957 Coeliades forestans Van Someren, 1955 Epargyreus clarus Saunders, 1932 Hesperia species MacNeill, 1964 Hesperia metea Shapiro, 1965 Pyrgus communis Clark, 1937 Thorybes pylades Klots, 19 51 160 OAKLEY SHIELDS /. Res. Lepid. if at all. Presumably the perched individual is conserving energy between aggressive encounters. MacNeill (1964:24) divided intruder-insects investigated by Hesperia into three categories on the basis of male behavior: (1) insects not of the same or a closely related species, (2) females of the same or a closely related species, and (3) males of the same or a closely related species. He found that male Hesperia will often investigate other butterflies and insects but would chase and fight only with males of the same or closely related species. The primary function of territoriality in insects seems to be to increase the probability of mating (Alexander, 1961; Kor- mondy, 1961; Wilson, 1961; Johnson, 1962b; Catts, 1963, 1964; Cazier & Linsley, 1963; Lin, 1963; Johnson, 1964). Increased mating probability was the selective force necessary for '‘the origin and maintenance of territoriality” in dragonflies (John- son, 1962b). Alexander (1961) found that territoriality in field crickets appeared in low density populations and was absent in high density populations, with both conditions producing a maximum degree of insemination of females. Movement is necessary to elicit the courtship approach in all male butterflies that have been investigated ( Marler and Hamil- ton, 1967:258). Some work has been done to isolate the stimuli that males are responding to. Tinbergen (1965) tested the Gray- ling, Eumenis semele, and found that bigness, darkness, near- ness, and dancing activity of the moving object stimulated the male to respond. Swihart (1967) found that certain butterflies respond maximally to colors most like their wing pigmentation. In the nymphalid Hypolimnas misippus, the early stages of courtship are visually controlled and the later stages are prob- ably “behavioral” and chemosensory (Stride, 1956, 1957, 1958). Pheromones emitted by androconia hairs and scales are used by many male butterflies to stimulate the female during court- ship. Families containing species with pheromones include Papilionidae, Pieridae, Nymphalidae, Lycaenidae, and Hesperi- idae (Clark, 1927; Ford, 1957). The male odors are often per- ceptible to man, but the female “directive” odors are not and their presence is inferred from behavior (Ford, 1957). Once the female is mated, certain barriers to further insemin- ation may develop. Labine (1964) found that there may be a female “awareness” of the presence of a spermatophore that makes her resistant to courting males. In the butterfly genera 6(2):69-178, 1967 HILLTOPPING 161 Parnassius, Euphydryas, Speyeria, and Acraea, the male secretes a plug (sphragis) on the female during copulation that helps prevent further mating (Labine, 1964). In the moth Atteva punctella, the presence of a spermatophore does not initiate oviposition but seems responsible for an inhibition of receptivity to males (Taylor, 1967). Various authors have noted that an ascending flight by mated females of certain species is an avoid- ance behavior to male courtship (e.g., Stride, 1958). Table 2 lists species that are known to hilltop. Those species reported by Scudder (1887, 1889) and Weiss (1927, 1928) were merely listed as present on summits and thus need to be verified. The genus and species names listed were brought up to date with current nomenclature where known. Certain genera in North America are apparently devoid of hilltopping species (list not exhaustive): Colias, Cercyonis^ Erebia, Phyciodes (one possible exception), Apodemia, LepheP isca, Lycaena, Philotes, Plebejus, Ochlodes, Pholisora, Pyrgus, Agathymus^ and Megathymus. K. Brown (in letter) says that certain members of the genus Doxocopa strongly hilltop in the early afternoons in Brazil. Doxocopa is probably congeneric with Asterocampa, although no members of Asterocampa ap- parently hilltop in Brazil or North America. Table 24 lists eight North American genera that contain many hilltopping species. The genera Erynnis and Hesperia probably contain additional hilltopping species as yet unrecorded. Hill- topping genera containing certain species that weakly hilltop include Speyeria (except callippe and egleis), Boloria (except astarte), Limenitis, and Incisalia. Ten of the eleven species of Vanessa (—Pyrameis) in the world are reported to hilltop (see Table 2). Only the species samani has not as yet been reported to hilltop. V. cardui is reported to hilltop in all the realms of the world except as yet in the Neotropical. Hilltopping is widespread in butterflies both geographically and taxonomically. Representative species are present in all the geographic realms of the world (see Table 2). Tropical regions have large numbers of species which hilltop (see Van Someren, 1955; Brown, Table 2), although little work has been done to catalogue and study these. Sixteen of the 25 subfamilies of butterflies and five of the six families contain species which hilltop (Table 25). Of the nine subfamilies that are, at present, not known to hilltop, four ( Baroniinae, Pseudopontiinae, Galina- ginae, and Styginae) contain only one species each and the TABLE Z4* Genera from North America north of Mexico that contain many hilltopping species 162 OAKLEY SHIELDS /. Res, Lepid. nO O' o r- sO vO 4) P. m 0) Cfi o Cfl to m O 03 0) o 43 H ^ g P u o « » s i 03 erf fi X . 03 P,j g o o “ m 03 U 43 43 P. CO 43 o I ■ 6(2):69-178, 1967 HILLTOPPING 163 TABLE 25. Butterfly subfamilies in which species are known to ''hilltop'' * i PAPILIONIDAE Baroniinae Parnassiinae * Papilioninae PIERIDAE •*> Coliadinae * Pierinae Dismorphiinae Ps eudopontiinae NYMPHALIDAE Ithomiinae * Danainae * Satyrinae * Morphinae ■*> Charaxinae Calinaginae Nymphalinae * Acraeinae LIBYTHEIDAE Libytheinae LYCAENIDAE * Riodininae Styginae * Lycaeninae HESPERIIDAE Megathyminae * Coeliadinae * Pyrrhopyginae * Trapezitinae * Pyrginae * Hesperiinae 1 This subfamily classification for true butter- flies (Papilionidae through Lycaenidae) follows Ehrlich (1958) and Ehrlich and Ehrlich (1967) and for skippers (Hesperiidae) follows Brues, Melander, and Carpenter (1954). 164 OAKLEY SHIELDS /. Res. Lepid. subfamily Libytheinae contains only ten species. Of the four remaining subfamilies that are not known to hilltop, Megathymi- nae fairly certainly does not (Roever, pers. comm.) and Parnassi- inae, Dismorphiinae, and Ithomiinae have so far not been re- ported to do so. The fact that most subfamilies contain species which hilltop and certain genera contain species which do strongly while other genera have species which do not or only weakly suggests that the hilltopping instinct has very likely developed independ- ently in various groups at various times. CONCLUSIONS Hilltopping in butterflies is so widespread that it must serve an essential function and have survival value. This study offers the following evidence that this phenomenon serves to bring males and females together to insure fertilization and that other explanations for this behavior lack validity, 1. From many sources it is apparent that males of hilltopping species occur in far greater numbers on the summits of favorable hills than they do in the surrounding areas; therefore, some mechanism is operating to create relatively high densities of male populations, with some degree of stability, in very limited areas. 2. The summits offer little to attract the males that they could not find elsewhere in the way of nectar sources, moisture, larval foodplants around which the females might be emerging, shelter, and warmth. 3. Males on summits adopt territorial behavior including “patrolling” and aggression which are sexually oriented behavior patterns. 4. Females, however, do not remain on the summits unless they are unmated and males are absent or few in numbers. An abnormally high percentage of females captured on summits are virgin when compared to those captured elsewhere. 5. A substantial number of actual courtships and matings have been observed for two species on the summit. 6. Unmated Papilio zelicaon females, when released below the summit of Dictionary Hill, flew to the summit while mated females did not. 7. Male F. zelicaon released at various distances and directions from the hill were recaptured on the summit in such numbers that they must have “homed in.” A summary of the behavior patterns of hilltopping butterflies is given in Figure 15. 6(2):69~178, 1967 HILLTOPPING 165 (fl - 5 < Q p, - ^ CuO fd C ^ -r^ OJ S d c tti XJ 03 o S ..H OT ^ "o 03 o3 o X O p: 03 03 P P 03 • i-i 03 ^ 1 r-l* a Xi 03 U >. o 03 03 CuO m p— t •rH !h X £ r-H 0 P X Q U I" ‘■J 6(3) : 19-9-202, 1967 NATURAL HABITATS 201 m 0) o 0) (U (X P- -M cn f-s :0 0) t," ^ ^■O '-M if'rt ^ 2 ^ f,^ L nj ^ z ^ Li OJ ^ 4^; a !•'“ ^ L'•>^ U -03 P 5S S :'(U >. -P rfti ^ ■' Cfl ■■rH hH ■ ■a o 1, •i ■I .. ^ ^ o3