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X = on THE LEFIJDOPTER^A Established in 1962 Edited by WILLIAM HOVANITZ Volume 3 1964 Published at 1160 W. Orange Grove., Arcadia, California, U.S.A. CONTENTS Volume 3 Number 1 March, 1964 The ettects of pH on the distribution ot the Megathymidae H. A. Freeman 1 Decapitation-initiated oviposition in Crambid moths C. S. Crawford 5 Book review—Fox and Fox: Introduction to Comparative Entomology 8 Life History Studies on Mexican Butterflies. 1. Gary N. Ross 9 Book review—Dos Passos: Synonymic list of Nearctic Rhopalocera 18 Melanie ’tendencies in Phalaenid and Geometrid moths in eastern Pennsylvania A. M. Shapiro 19 The genus Lycaeides in the Pacific Northwest J. H. Shepard 25 The origin of a sympatric species in Colias through the aid of natural hybridization W. Hovanitz 37 The southern limits of the range of Pieris napi and P. virginiensis B. Mather 45 Life histories of Papilio indra and P. oregonius E. J. Newcomer 49 Hybrids between Papilio memnon and Papilio protenor S. A. Ae 55 New gynandromorph of Colias philodice from Colorado T. C. Emmel 63 Volume 3 Number 2 June, 1964 The hidden wing-pattern of some Palearctic species of Gonepteryx and its taxonomic value Y. P. Nekrutenko 65 An evaporative cooling mechanism in Pholus achemon (Sphingidae) P. A. Adams and J. E. Heath 69 A new species of Riodinidae from Mexico H. K. Clench 73 Life history studies on Mexican butterflies 11. Anatole rossi Gary N. Ross 81 Review of the depicta group of the genus Annaphila J. S. Buckett and W. R. Bauer 95 102 Notice Bionomics of Agathymus (Megathymidae) K. Roever 103 W."!!. Edwards’ life histories of North American Coenonympha F. Martin Brown 121 Volume 3 Number 3 September, 1964 Revision of the North American Genus Behrensia J. S. Buckett 129 Larval habits of Agathymus marine H. A. Freeman 145 Instar determination of Agathymus larvae Kilian Roever 148 Early stages of Euphyes vestris John Richard Heitzman 151 The habits and life history of Amblyscirtes nysa ( Hesperiidae ) in Missouri John Richard Heitzman 154 Genetic relationships of Papilio indra and Papilio polyxenes John F. Emmel and Thomas G. Emmel 157 Adult oviposition responses in Pieris rapae William Hovanitz and Vincent C. S. Chang 159 Studies in life histories of North American Lepidoptera John Adams Comstock and Christopher Henne 173 Petaluma, a new genus J. S. Buckett and W. R. Bauer 193 Volume 3 Number 4 December, 1964 Oxygen consumption and metabolic rate of PapUio zelicaon pupae in a state of delayed eclosion L. LaDue 197 Life history studies on Mexican butterflies. Ill Nine Rhopalocera from Ocotal Chico, Vera Cruz G. N. Ross 207 The generic, specific and lower category names of the nearctic butterflies. Part 3. Argynnids P. McHenry 231 Volume 3 Number 1 March, 1964 THE JOURNJAL OF RISIAKCH ©N.' THE LEPlJDOPTEKA a quarterly published at 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. edited by: WILLIAM HO VANITZ 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.’' 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 editors 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 mailed air mail or registered at cost to the subscriber, if so desired. Journal of Research on the Lepidoptera 3(1) 1-4, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Coptjright 1964 THE EFFECTS OF pH ON THE DISTRIBUTION OF THE MEGATHYMIDAE H. A. FREEMAN 1605 Lewis Drive, Garland, Texas From my first capture of a member of the Megathymidae (a specimen of Megathymus yuccae stallingsi Freeman) at Vickery, Texas, April 12, 1938, a keen interest was developed in this group of butterflies. The close association of the Mega- thymidae with their host plants, Agave, Yucca and Manfreda, has led to the study of some very interesting habitats. At first, it was a matter of locating the adults on the wing and of trying to catch them with a net. That, however, was difficult and often resulted in damage to the specimen and sometimes to the collector. As knowledge about the life histories of the various species developed, techniques were devised for collecting the larvae and pupae. The net was then put aside and in its place such equipment as a fox hole pick, bayonet, and drain spade took its place. Records of the distribution of the various species of Agave, Yucca, and Manfreda were studied and from this in- formation, various collecting trips were made which resulted in the discovery of many interesting new species of Megathymidae. It was thought provoking that a perfectly fine stand of Yucca did not always result in the finding of a colony of Megathymus, whereas within a short distance a much less favorable-looking area would have plants attacked by the larvae. Now, it is known that there are several factors in the environment that govern the survival of a colony of these insects in a given habitat. One particular factor, the pH of the soil, attracted attention about six years ago and since that time soil samples have been carefully checked from all of the study habitats under con- sideration. The knowledge so obtained explains in part why success has not always been had in locating larvae in apparently fine stands of both Yucca and Agave. I would like to express my appreciation to the National Science Foundation for Research Grant GB-398 which is making my studies of the Megathymidae possible. 1 2 FREEMAN /. Res. Lepid. Along with Stallings and Turner, a rather extensive study of Agat hymns marme (B. & B.) and its related species has been made. In carrying on this present study, 33 locations were selected by the author; these extend from El Paso and Carlsbad National Park to Boquillas Canyon, and to the north of Bracket- ville, Texas, where Agave lecheguilla Torr. occurs. At each location, the following factors were checked, pH of the soil, type of soil, elevation, plant associates, average annual rainfall, and presence or absence of radiation with Geiger counter. The most significant factor seemed to be the pH of the soil. It was found that Agathymus mariae occurred where ever the pH ranged from 7.3-8. 9, with the average being slightly below 8, indicating that it is best suited to live where the soil is alkaline. In the southeastern section of the study habitats, the pH was around 7, indicating neutral soil; mariae was not found in these areas, but instead Agathymus estelleae ( Stallings & Turner ) and .three new species ( that are now in the process of being named ) were found. The following data has been found on the other species of Agathymus that occur in Texas. Agathymus florenceae ( Stallings & Turner) occurs where the pH of the soil ranges from 5.9-6.I, A. carlsbadensis (Stallings & Turner) around 7.9, A. judithae (Stallings & Turner) around 7.3, A. diabloensis Freeman around 7.4, A. mcalpinei (Freeman) around 7.4-7.6, A. chisosensis (Freeman) around 5.2. The last indicates the strongest acid soil relationship. In Arizona it was found that Agathymus aryxna (Dyar) is associated with soil where the pH ranges from 5.3-6. 1, and A. polingi (Skinner) from 5. 8-6.1. In Texas, there are several subspecies of what are now called Megathymus yuccae ( Bdv. & LeC. ) . It has been very interesting to see how the pH of the soil is related to the distribution of these subspecies. The first subspecies studied, stallingsi Freeman, occurs from north central Texas, Oklahoma to Caldwell, Kansas. In Texas, it is associated with limestone outcroppings where the pH is usually just under 8. East of the Dallas area, there is a distance of about seventy miles where the soil changes from alkaline, through neutral to distinctly acid. In the neutral area, no colonies of Megathymus have so far been located even though there are fine stands of Yucca present. At Canton, Ben Wheeler and Tyler State Park, the soil ranges from pH 4.9-5.1. In this area a different subspecies is found which was recently named 3 (l):l-4, 1964 EFFECTS OF pH 3 reinthali Freeman. This is a sandy soil subspecies associated with pines and oaks. In the Rio Grande Valley, the subspecies wilsonorum S. & T. is associated with sandy soil and scrub bushes but the soil ranges from pH 7.4-7. 8. In the Del Rio area, the subspecies louiseae Freeman is found. Here, the pH is very near pH 7, indicating a neutral soil relationship. In the extreme western part of Texas, the subspecies reubeni S., T. & S. is found in the Hueco mountains where the soil is alkaline, ranging from pH 7.3-7.6. Another subspecies (not yet named) was found in western Texas where the soil was very sandy and the pH was acid, averaging pH 5. Other Megathymus records for Texas indicate the following results. In areas where Megathymus t exanus B. and McD. have been collected, the pH ranges from 7. -7. 6. At and near the type locality of Megathymus violae S. and T. the pH was 7.5. In the Big Bend National Park, where M. violae is found the pH is 7.4. In the Tucson, Arizona vicinity, where the pupae of Mega- thymus ursus Poling were collected by myself the pH ran from 5.8-6. During August 1962, in association with the Stallings, a study was made of 22 selected habitats in north central Mexico, extend- ing from General Bravo down through Victoria to Antique Morelos, up through Saltillo, over to Torreon and then through Monclova to Allende. It was found that wilsonorum from near China occurred in areas where the pH was identical to its habitat in the Rio Grande Valley. The specimens collected of a violae- Hke species from El Tepeyac were associated with very dusty soil that had a pH of 6.1. Agathymus estelleae was found at several locations and the pH of those areas varied from 6.9-7.3. Agathymus remingtoni ( S. and T. ) was collected in areas where the pH was 5.-5.5. Aegiale hesperiaris (Walker) was found where the pH was 6.9-7. Agathymus micheneri S., T. and S. was found where it was 7. There are several species involved in the mariae complex from various locations studied. All were found where the pH was on the alkaline side of the range or nearly neutral and never where the soil was acid. There were other species involved in this study; however, until their exact status is determined I will omit them from this discussion. No correlation was established between the presence or absence of certain species in a given environment due to radiation effects, because none of the areas studied showed to any pro- nounced degree the presence of radioactive particles. 4 FREEMAN /. Res. Lepid. The elevation of colonies was important to some extent in most species studied because the presence or absence of the host plants was correlated with elevation. Together with this was an indication of the presence of other plant associates. Correlation with type of soil appeared to be in direct relationship to the pH in most instances. The average rainfall seemed to have less to do with the presence or absence of certain species of Mega- thymidae than was previously thought. It was observed that west of Saltillo towards Torreon, the Agave lecheguilla were yellow or brown due to aridity yet the larvae of various Aga- thymus were present in the plants in more or less large numbers. From all the data collected over the past six years, the opinion has been dmwn that the pH of the soil is one of the most important factors governing the distribution of various species and subspecies of the Megathymidae, and in the selection of host plants. A listing of food plants of this group of insects together with the pH of the soil where found is given in Freeman ( 1963). REFERENCES FREEMAN, H. A. 1962. A new species of Agathymus from Texas. Amer. Mus. Novitates, no. 2097, 1~7, figs. 1-6. _______ 1963. Megathymus yuccae in Texas, with the description of two new subspecies. Journ. Lepid. Soc. 17 : 89-99. 1963. Type localities of the Megathymidae. /. Res. Lep. 2(2): 137-141. STALLINGS, DON B., J. R. TURNER and VIOLA N. STALLINGS. 1961. A new subspecies of Agathymus mariae from Mexico. Journ. Lepid. Soc. 15: 19-22. _____ 1963. Two new species and one new subspecies of Megathymidae from Mexico and Texas. Journ. Lepid. Soc. 17: 81-88. Journal of Research on the Lepidoptera 3(1) :5~8, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1964 DECAPITATION-INITIATED OVIPOSITION IN CRAMBID MOTHS CLIFFORD S. CRAWFORD Portland State College, Portland, Oregon The meager litfjiature on decapitation-initiated oviposition in insects has been partially reviewed by Chiang and Kim ( 1962 ) who observed the phenomenon in crane flies. M'Cracken (1907) had previously shown that mated female silkworm moths exhibit an early oviposition response to abdominal stroking when de- capitated; however spontaneous egg production was rare fol- lowing head removal. It has been observed in a number of instances that certain insects tend to oviposit in response to being trapped or wounded (Harwood, personal communication, 1963), yet there is little doubt that an inhibitory center in the subesophageal ganglion does exert a controlling effect on oviposition (Boeder, 1963). An opportunity to study the behavior of certain microlepidoptera with regard to the inhibitory role presumbably played by this ganghon over endogenous oviposition activity recently presented itself; the following being a report of some preliminary work suggesting areas where the nature of the control could be more fully explored. In a study of the reproductive biology of lawn moths in the family Crambidae it was found that decapitation of gravid females brought about oviposition generally within ten minutes for fall-generation Crambus bonifatellus Hist, and Euchromius californicalis (Pack.), the two species then available. Tactile stimulation following decapitation was not necessary to induce egg deposition. Decapitation was performed with a pair of fine scissors after moths had been cooled to sluggishness. Immediate- ly following the operation the headless moths stood quietly, and shortly thereafter the terminal part of the abdomen began to undergo rhythmic contractions which continued during ovi- position. In some cases the entire abdomen vibrated in a vertical plane as well. Decapitated male moths also exhibited 5 6 CRAWFORD /. Res. Lepid. abdominal contractions and periodically expanded their genitalia. Moths of both sexes showed no tendency to mate when held together in different positions after decapitation. The effects on oviposition activity of inflicting gross injury without damaging the subesophageal ganglion or nerve cord have not yet been in- vestigated. The time interval between head separation and oviposition proved variable for C. bonifatellus, the most abundant crambid and the only one used experimentally. Under room conditions, individuals of a group of ten females caught at light took from 50 seconds to nine and a half minutes to begin laying eggs after the operation. Directly after beginning to lay, these moths were dissected and subsequently divided into three classes according to the relative amount of eggs and fat body in the abdominal cavity. Since there was an obvious lack of correlation between abdominal class and time intervals concerned, it would appear that within about ten minutes after decapitation any mechanical pressure responsible for moving eggs out of the reproductive tract is not a direct function of the degree of gravidness. In order to determine whether the circadian oviposition rhythms known to occur in these species (unpubHshed data) are maintined by headless moths, a number of decapitated C. bonifatellm females were exposed to darkness at 28.0 ± 1.4° C and their eggs collected in a manner previously described ( Craw- ford, 1962). The resulting oviposition was sporadic, and though eggs were generally laid in batches, there was no semblance of the usual diurnal rhythm. This result is consistent with HarkeTs ( 1955 ) finding that subesophageal gland neurosecretory cells are responsible for activity rhythms in the American cockroach, Periplaneta americana L. It is therefore to be expected that successful subesophageal gland implanation into headless cram- bid females may re-establish the rhythm. The decapitated crane flies used by Chiang and Kim (1962) laid a greater percentage of fertile eggs than did controls, though it was not indicated whether this difference was statistically significant. In the present study crambid eggs were also tested to see if egg fertility was affected by decapitation. Twenty moths obtained from light ( and therefore of unknown age ) were cooled for at least 24 hours at about 5°C, then half of these were decapitated while cold. The other ten were used as controls. All moths were incubated at 28.6 ± 2.5 °C in petri dishes at the 3 {l):5-7, 1964 DECAPITATION OF CRAMBIDS 7 bottom of which were moist circular pieces of blotting paper. Decapitated females were not as physically active as controls, and remained alive ( in the sense that they were capable of body movement) an average of 3.9 ± 0.5 days, which was not sig- nificantly different from the 5.1 ± 0.5 days lived by controls. No records were kept of daily oviposition by individuals, but it was obvious that some decapitated moths laid for at least 48 hours. In calculating final percentages of fertility, eggs which remained yellow or slightly pink a week after being laid were considered infertile, while eggs obviously about to hatch but in danger of being eaten by cannibalistic larvae were considered fertile. Out of a total of 506 eggs laid by headless moths 11.1% were infertile, while 22.6% of the 974 control eggs were infertile. A chi square analysis of the pooled experimental data using con- trols as standard indicated the difference in fertility was highly significant (X^ = 52.28, 1 d.f. ) However a chi square analysis of heterogeneity (Snedecor, 1956) which was prompted by obvious differences in egg fertility among individuals was also significantly high (X^ = 93.91, 8 d.f.) The data therefore suggest that while decapitation may have had a positive effect on fertility, it would take a decidely larger number of moths to show clearly that this is true. ACKNOWLEDGEMENTS The author wishes to thank Dr. Q. D. Clarkson, Portland State College, and Dr. R. F. Harwood, Washington State Uni- versity, for critical advice given during preparation of the manuscript. For his assistance in obtaining specimens Mr. Peter F. Murray is also cordially thanked. REFERENCES CITED CHIANG, H. C. and Y. H. KIM. 1962. Decaptationdnitiated oviposition in crane flies. Ent. Exp. and Appl. 5: 289-290. CRAWFORD, C. S. 1962. An apparatus for measuring oviposition in some insects. Northwest Set. 36: 123-125. MARKER, J. E. 1955. Control of diurnal rhythms of activity in Periplaneta americana L. Nature, London. 175: 733. HARWOOD, R. F. 1963. Personal communication. MURACKEN, M. 1907. The egglaying apparatus in the silkwonn {Bombyx mori) as a reflex apparatus. Jour. Comp. Neurol, and Psychol. 17: 262-285. ROEDER, K. D. 1963. Nerve Cells and Insect Behavior. Harvard Uni- versity Press, Cambridge, Mass. 188 pp. SNEDECOR, G. W. 1956. Statistical Methods. 5th edition. Iowa State College Press, Ames, Iowa. 534 pp. 8 /. Res. Lepid. Book Review INTRODUCTION TO COMPARATIVE ENTOMOLOGY Richard M. Fox and Jean Walker Fox Reinhold Publishing Corporation, New York i - xiv, 1-450, one color plate, numerous illustrations. $9.50. This book has been designed primarily as a textbook for an elementary course in entomology. However, the reviewer finds it an excellent reference source, especially for the study of the comparative anatomy and morphology of insects. Unlike many standard entomology texts in which the classification of in- sects is the central theme, the Foxes have brought to this book a new approach. The general insect structural plan, and its deviations comprise the first two thirds of the book while classification is the last one third. The first eight chapters cover in order the skeleton, the appendages, the wings, maintenance systems (gut, circulation, respiration and reproduction), control systems (ner- vous, sensory and endocrine systems), early embryogenesis and postovarian embryogenesis. These subjects are covered in a masterful way, not duplicated by any other text, and long needed. The remaining five chapters cover compar- ative classification of the Arthropoda in such a way that a student can grasp the origin of insects from more primitive groups. There are chapters on Arachnids and Myriapods. The final chapter presents a comparison of the orders of in- sects, and some of the major families. The reader should not obtain this book if he wishes an identification man- ual. It has not been designed for this purpose. He will find here however a well organized, truly comparative insight into the insect structure. There will be found little on insect ecology, habits, physiology, genetics, geographical distribution or fossil record. But one book cannot be everything to everybody. In physical appearance, the book is put up very neatly and has excellent illustrations. The Editor Journal of Research on the Lepidoptera 3(1) :9=-18, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1964 LIFE HISTORY STUDIES ON MEXICAN BUTTERFLIES I. NOTES ON THE EARLY STAGES OF FOUR PAPILIONIDS FROM CATEMACO, VERACRUZ GARY N. ROSS Department of Entomology, Louisiana State University, Baton Rouge, Louisiana During the summer of 1962, while engaged in collecting butterflies near Catemaco in the Tuxtla Mountains of southern Veracruz, Mexico, I was able to rear the immature stages of four species of the family Papilionidae: Graphium helesis (Bates), G. epidaus epidaus (Doubleday, Westwood & Hewitson), Papilio androgeus epidaurus Godman & Salvin, and P. anchisiades idaeus Fabricius. References to the immature stages of these are scanty and not readily available. Recent illustrative works, therefore, are of interest and benefit to students of the Lepidoptera. Scale lines in all photographs represent .5 inch or 127 mil- limeters. All measurements are based on living material. GRAPHIUM BELESIS Bates The only reference made to the immature stages of this butterfly, to my knowledge, is a brief description of a mature larva (Dyar, 1912). The present photographs represent the first published illustrative material. EGG. (Description based on 2 eggs.) Duration of stage, 4 days (74°-90° F. ambient temperature). The egg is a sphere having a diameter of 1.0 mm. and being light yellow in color. Superficially, it appears per- fectly smooth but upon close examination, tiny facets or reticulations could be discerned. Eggs are attached to the undersurfaces of the leaves of Annona muricata L. (Annonaceae) called “Guanabana” by the local resi- dents. A related species, A. reticulata, grew in the same vicinity, but females never seemed to be attracted to it. FIRST INSTAR LARVA: Fig. 2A. (Measurements based upon one larva.) Beginning of stadium-length, 1.7 mm.; greatest width, 1.0 mm.; head diameter, .9 mm. Head black with tiny, fine setae. Body expanded anteriorly into a 'Talse head”; color velvty black with a white transverse band (saddle) on sections of fourth and Hth abdominal segments. Legs and prolegs concolorous with body. Osmateria yellow-orange in color. Second instar larvae similar. 9 ROSS /. Res. Lepid. Fig. 1. Adults. A. Graphium belesis (Bates). B. Graphium epidaus epidaus (Doubl., Westw. & Hew.). C. Papilio androgens epidaurus God. & Sal. D. Papilio anchisiades idaeus Fabr. Photographs by J. H. Roberts. LAST INSTAR LARVA: Figs. 2B & G. (Measurements based on two specimens.) Beginning of stadium-length, 21.1-21.4 mm.; width, 9.0-9.2 mm.; head diameter, 4.1 mm. Termination of stadium-length, 32.1-34.2 mm.; width, 10.9-12.0 mm. Head as before. Body with anterior portion slightly less expanded than before. Color black with an irregular transverse white band (saddle) occupying major portions of fourth and fifth abdominal segments dorsally but only fifth segment laterally. Segments with longitudinal rows of smooth, dull red, paired tubercles: first and second thoracic segments with 1 lateral and 1 suprastigmatal pair; third thoracic and all abdominal segments with 1 subdorsal, 1 lateral, and 1 suprastigmatal pair. Legs and prolegs as before. Osmateria more orange. 3 (1):9-17, 1964 LIFE HISTORIES I 11 PUPA: Figs. 4A & B. (Measurements based on one specimen.) Length, 20.0 mm.; greatest width, 11.1 mm. Duration of stage, 13 days. The pupa of this species represents quite an interesting form (figs. 4A & B). Color a light, uniform green (slightly lighter on the “belly” portion). Four longitudinal rows of tubercles, also light green but slightly darker than the surrounding are^s. Head portion with a ventral projection or “snout” 5.0 mm. in length. Girdle around constricted thoracic portion. Adults (fig. lA) were numerous around the edges of the secondary forests bordering Laguna Catemaco. The range of this species according to Hoffman (1940) is the warm regions of the eastern Sierra, Chiapas, and the southern and western Sierra as far as Colima and Jalisco. GRAPHIUM EPIDAUS EPIDAUS (Doubleday, Westwood & Hewitson) Immature stages (second and sixth instars and pupae) of the western subspecies tepicus Rothschild & Jordan, have been pictured by Comstock & Vazquez (1960). I observed no sig- nificant deviation from their illustrations in the eastern sub- species epidaus. However, I include it here because of the taxonomic difference and because of a difference in duration of the pupal stage. EGG. (Comparisons based on 5 eggs.) Eggs are attached to the undersurfaces of die leaves of Annona reticulata L. ( Annonaceae ) . As with G. belesis, females never were seen to oviposit on the related A. muricata. LARVA. (Comparisons based on 11 larvae.) All instar larvae showed no deviation from the descriptions of Comstock & Vazquez ( 1960 ) for tepicus. A fifth instar larva is pictured in figures 3 A & B. PUPA. Figs. 4C & D. (Measurements based on 4 specimens.) Length, 26.1-26.5 mm.; greatest width, 10.3-10.6 mm. Duration of stage, 201 days. No deviation from tepicus was apparent. However, duration of pupal stage is quite different. Comstock & Vazquez state that the adults of tepicus emerged nine days after pupation whereas four epidaus imagos reared did not emerge from the chrysalids for 201 days. It should be mentioned here that my epidaus larvae pupated during early August, almost the exact time as the tepicus larvae of Comstock & Vazquez. However, these chrysalids were under natural (field) conditions for only SYz months of the total time since I departed Mexico in early December of the same year and carried the pupae with me back to Louisiana. Then on a morn- ing late in March, after having been exposed to laboratory conditions (about 28°C. ) for nearly 3 months, 3 adults emerged. The following day, the fourth adult emerged. It seems reasonable to conclude that the larvae which I reared were representatives of the final brood of the year (after late August, no adults were seen in the field). It is likely that the eastern subspecies epidaus found in Veracruz undergoes a pupal diapause through the period of heaviest rains ( September-January ) and emerges as an adult after drier weather begins (February or March). To be sure, half of the time my pupae were under artificial conditions. However, I believe that the 3’/4 month diapause under natural, field conditions is justification for the above suggestion. Since Comstock & Vazquez did not report any diapause in tepicus, I assume that it either undergoes no diapause or else diapause begins at a later date than mid or late August. 12 ROSS /. Res. Lepid. Fig. 2. Larvae. A. G. helesis, first instar. B. G. belesis last instar (lateral view), C, G. belesis, last instar (dorsal view). D. P. anchisiades idaeus, last instar. Photographs by R. F. Andrle. 3 (1):9-17, 1964 LIFE HISTORIES I 13 Adults (fig. IB) were fairly abundant in the general vicinity of the town of Catemaco during the months of April- August. After August, no individuals were observed. The range of the nominate species ( Hoffman, 1940 ) is Veracruz, Tabasco, nothern Chiapas, Campeche, and Yucatan. PAPILIO ANDROGEUS EPIDAURUS Godman & Salvin Four authors, Merian (1705), Moss (1919), Sepp (1855), and Burmeister (1879) have mentioned the immature stages of this species. SECOND INSTAR LARVA (probably). Fig. 3C. (Measurements based on 3 specimens.) Beginning of stadium-length, 7. 2-7.6 mm.; great- est width, 1.5-1. 6 mm.; head diameter, 1.3 mm. Termination of stadium- length, 16.1-16.7 mm.; width, 3. 2-3.4 mm. Duration, 5 days. Head tan with numerous tiny, fine setae. Body expanded anteriorly into a false head. Segments with rows of paired bristly tubercles: first thoracic segment with 1 reduced subdorsal, 1 prominent supralaternal, and 2 less prominent pairs ( 1 lateral and 1 sublateral); second third thoracic segments similar but with supralateral pair reduced; abdominal segments as above but with lateral and sublateral pairs nearly indistinguishable. Color glossy tan-brown with a slight cream mottling; ^so, a cream colored saddle on the third and fourth abdominal segments, and an extensive creamy area on the last three segments. Legs and prolegs tan. Osmateria light orange. LAST INSTAR LARVA. Fig. 3D. (Measurements based on 3 specimens.) Beginning of stadium-length, 41.1-43.5 mm.; width, 6.9-7.9 mm.; head diameter, 3.9 mm. Termination of stadium-length, 74.0-75.9 mm.; width, 19.1-19.7 mm. Head as before. Tubercles on all segments reduced to slight knobs with basal crescent shaped blue markings. Ground color dark grey to black; numerous white streaks and blotches (the latter being particularly extensive on the lateral sections of all thoracic segments and second-fourth and seventh-ninth abdominal segments). Legs and prolegs dark brown. Osmateria orange. PUPA. Fig. 5B. (Measurements based on 3 specimens.) Length, 42.1-42.6 mm.; greatest width, 7.2-7.3 mm. Duration of stage, 51 days ( only 1 adult emerged, the 2 other pupae having died ) . Color dark brown with creamy, longitudinal lines and bands; also, a slight green shading on the wing cases. Head portion with three horn- like projections directed anteriorly. Girdle around non-constricted thoracic section. Larvae were found resting on the upper surfaces of the leaves of Zanthoxylum elephantiasis Macf. (Rutaceae). To my knowl- edge, this is the first mention of this tree being the larval food plant of P. androgens. Adults (fig. 1C) were not common in the Tuxtlas. Only two individuals were seen and captured around the lake where the food plant was abundant along the margins of fields; two individuals were seen ‘liill-t opping” above the peak of Volcan San Martin (5400 ft.) and one individual was seen engaged in the same activity above the peak of Cerro Tuxtla (2700 ft.). 14 ROSS /. Res. Lepid. Fig. 3. Larvae. A. G. e, epidaus^ fifth instar (dorsal view). B. G. e. epidaus, fifth instar (lateral view). C. F. androgens epidaurm, second instar. D. P. androgeus epidaurm, last instar. Photographs by R. F. Andrle. 3 (1):9~17, 1964 LIFE HISTORIES I 15 The range of the species (Hoffman, 1940) is southern Tamaulipas southward and the western Sierra as far as Sinaloa and Durango, PAPILIO ANCHISIADES IDAEUS Fabricius The stages of P. anchisiades are probably the best known of any of the four species under consideration here. This is probably due, in part, to its wide distribution (see below). Bates (1861), Caracciolo (1891), Dewitz (1886), Jones (1883), Moss (1919) and Stoll (1781) have mentioned the various stages in the life history of this insect. LAST INSTAR LARVA. Fig. 2D. (Measurements based on 5 larvae.) Beginning of stadium-lengffi, 27.0-27.8 mm.; width, 6.1-6.8 mm.; head diameter, 3.9 mm. Termination of stadium-length, 42.2-43.9 mm.; width, 7.0~7.2 mm. Duration, 8 days. Head medium brown with tiny tubercles and setae. Body more or less cylindrical, only slightly expanded anteriorly. Segments with pairs of subdorsal, supralateral and lateral tubercles (as described under P. androgens) bare and concolorous with ground color. Color greenish brown with numerous streaks, lines and flecks of white or cream. Legs and prolegs brown. Osmateria yellow. Larvae are nocturnal feeders on lime {Citrus sp.). During the day- light hours they rest together in groups on the trunk or stems of the food plant. PUPA. Fig. 5A. (Measurements based on 3 specimens.) Length, 34.1-35.0 mm.; greatest width, 10.2-10.4 mm. Duration of stage, 13 days. Color light brown with light green mottling beginning on wing cases and terminating at cremaster, giving the chrysalid the appearance of having a lichen encrustation. Slight dorsal protrusion anteriorly. Tubercles still evident. Girdle around non-constricted thoracic portion. Adults (fig. ID) were common around the citrus groves bordering Laguna Catemaco. According to Hoffman ( 1940 ) the range of the species is the entire eastern area of Mexico. Ehrlich & Ehrlich (1961) list the range as ‘'southern Brazil to Mexico and southern Texas.'' ACKNOWLEDGEMENTS I wish to express my appreciation to the following people: Mr. R. F. Andrle (Buffalo Museum of Natural Science, Buffalo, New York), Dr. J. H. Roberts (Louisiana State University, Baton Rouge, Louisiana), and Dr. S. M. Russell (Louisiana State Uni- versity in New Orleans, New Orleans, Louisiana) for photographs and film processing; Dr. V. E. Rudd (U. S. National Museum, Smithsonian Institution, Washington, D. C. ) for food plant identi- fications; Mr. K. H. Wilson (University of Kansas, Lawrence, Kansas ) for specimen identifications and bibliographical ci- tations; Mr. S. L. Warter, Dr. M. S. Blum and Dr. L. D. Newsom for suggestions regarding the manuscript and lastly, Christopher Andrle (Buffalo, New York) for help in assisting me with the larval rearings. 16 ROSS /. Res. Lepid. Fig. 4. Chrysalides. A. G. belesis, lateral view. B. G. belesis, ventral view. C. G. e. epidaus, ventral view. D. G. e. epidaus, lateral view. Photographs by R. F. Andrle. Fig. 5. Chrysalides. A. P. anchisiades idaeus. B. P. androgens epidaurus. Photographs by R. F. Andrle. LITERATURE CITED BATES, H. 1861. Contributions to an Insect Fauna of the Amazon Valley. Papilionidae. Jour. Ent. 1 : 218-245. BURMEISTER, H. 1879. Lepidopteres. Atlas de la Description Physique de la Republique Argentine Contenant des cues Pittoresques et des Figures d Histoire Naturelle. 5 ( 2 ) :■ 4. CARACCIOLO, H. 1891. Notes and News. Ent. News 2: 51-55. COMSTOCK, J. and L. VAZQUEZ GARCIA. 1960. Estudios de los Ciclos Biologicos en Lepidopteros Mexicanos. An. Inst. Biol. Mex. XXXI (1&2): 339-448. DYAR, H. 1912. Descriptions of the larvae of some Lepidoptera from Mexico. Proc. Ent. Soc. Wash. 14: 54-58. DEWITZ, H. 1878. Entwickelung einiger Venezuelanischer Schmetterlinge nach Beobachtungen von Gollmer. Archw. fur Naturgesch. 44: 1-36. EHRLICH, P. and A, H. 1961. How to know the Butterflies. Dubuque, Iowa: Brown, 262 pp. HOFFMAN, C. 1940 Catalogo Sistematico y Zoogeografico de los Lep- idopteros Mexicanos. An. Inst. Biol. Mex. 11 (2) :639-739. JONES, E. 1882. Metamorphoses of Lepidoptera from San Paulo, Brazil. The Nomenclature and Description of New Forms. Proc. Lit. Philos. Soc. Liverp. 36: 327-377. MERIAN, M. 1705. Histoire Generate des Insectes de Surinam et de Toute LTurope. VoL 1. Des Plantes de Surinam. Amsterdam, 72 pp. MOSS, A. M. 1919. The Papilios of Para. Novitat. Zoolog. 26: 259-319. SEPP, C. 1855. Papillons de Surinam. Natuurlijke Historic van Suri- naamsche Vlinders. VoL 3. Amsterdam, 328 pp. Stoll, C. ITSI. in Pieter Cramer De Uitlandsche Kapellen Voorkomende in de Drie Waereld-Deelen Asia, Africa, en America. Supplement 3. Amsterdam, 384 pp. 18 /. Res. Lepid. Book Review A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA Cyril F. dos Passos The Lepidopter ists ' Society, New Haven, Connecticut i - V, 1 - 145, including index, ^ 4. 50 This is the first list of Nearctic butterflies printed since 1938. As such, it is an excell- ent compendium of names because the meticulous care of the author, no doubt, insures that all names are included. An innovation for a checklist is the placing of the year of publication after a name. An excellent retention is the list of infrasubspecific names. The reviewer does not be- lieve that such names need or ought to be in Latin, but in view of the fact that most such names in butterflies were proposed as species, or 'varieties', they may be nomenc lator ially valid: in addition, there is yet too much uncertainty as to the nomenclator ial position to which many names belong. For example, the name hagenii, is listed as a seasonal form un- der 286b as Colias eurytheme keewaydin whereas the reviewer uses this name as a sub- species of Colias philodice.' The problem is that lists like this do not always represent the latest biological knowledge of the group in question. As used in the present list, the name hagenii is not valid, but as used by the reviewer, it is. Dos Passos has done an excellent nomenclatorial job, but there are serious defects cre- ated by limited study of the biology of many groups of Nearctic butterflies. It has been shown that C. harfordii (289), C. occidentalis (290), C. alexandra (293), are subspecies of one spec- ies and that C. barbara (288) is most probably a seasonal form of one of these. Also, it has been shown that C. interior (291) and C. pelidne (294) are geographical or altitudinal counter- parts of one species (thus, are subspecies). Other obvious problems arise with Zerene eury- dice (298) and Z. cesonia (299), which are subspecies. Pieris occidentalis is a species dis- tinct from P. protodice (277); Oeneis ivallda (662) is a subspecies of O. chryxus (665). Lim- initis arthemis (516), L. weidemeyerii (519), L. lorquini (520) and probably L. astyanax (517) are geographical races of one species; many of the Papilios (248-254) need detailed bio- logical work for many of these are subspecies; Plebejus icariodes (458), P, pardalis (459), and P. pheres (460) are subspecies; Melitaea gabbii (574), M. malcolmi (575), M. acastus (576), M. palla (582) and possibly M. neumoegeni (57 3) are subspecies; Melitaea leanira and M. alma are subspecies; the races of Euphydryas chalcedona (593), E. anicia (594) and E. editha (595) are badly mixed up as there are probably only two species here; G. lygdamas (479) and G. xerces (480) are one species. This list could grow and grow depending on the extent of one's knowledge. The information necessary to have put the names mentioned in a natural order has been published for all the above named, and no doubt for many others. It is unfortunate that the great efforts and the meticulous work of the author did not extend to the biological aspects of taxonomy, for a list of names such as this is intended to indicate phylogenetic relationships and it will be used as such. If it were not to be used in this way, no great catastrophy would be assumed. Another deficiency in the present work is the failure to include a complete list of Nearc- tic generic names. This comes about by the e.xpedient of using a recent name in North Am- erica (Speyeria) for a world wide genus (Argynnis) having an older established name. How many other times older names with priority have been omitted can only be conjectured; all our common generic names, Papilio, Pieris, Colias, Melitaea, Vanessa, Parnassius, etc. etc, , could be eliminated from North America by the same procedure and will be in the fut- ure by "splitters". Names propounded by splitters do not have to be used merely because they have been proposed. Eventually, every species will be placed in a different genus by somebody; this is inevitable. The balance of perspective must not be lost and it is up to biologists in general to demand that we look at the forest, from a distance, not only at the trees. It is hoped that this list will not be used as an inviolate text; the knowledge of phylogene- tic relationships in the butterflies is advancing into a new level of excellence and taxonomic relationships are being brought up to date daily. Unlike the methods used in developing lists of the past, modern scientific methods of experiment and analysis are being used; the rela- tionships shown by names should reflect such knowledge, or their purpose has been thwarted. To the end that dos Passos has produced an excellent and complete list of names, he is to be commended. Now, Lepidopterists owe it to him, and to science in general, to indicate by the study of relationships between populations, the natural arrangement in phylogeny of the in- sects designated by those names. The Editor Journal of Research on the Lepidoptera 3(1) : 19-24, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1964 MELANIC TENDENCIES IN PHALAENID AND GEOMETRID MOTHS IN EASTERN PENNSYLVANIA ARTHUR M. SHAPIRO 7636 Thouron Ave., Phttadelphia, Pa. Although Industrial Melanism has come to be a standard citation in the literature of genetics and evolution, relatively little has appeared in the United States on the subject. Few descriptions of melanics on this side of the Atlantic have found their way into print, nor have the few that have been remarked received much attention from the geneticists. This paper gives records of 15 species showing melanic tendencies in Pennsylvania, with frequency data for the commoner forms; several of these species have not been reported previously as having melanic forms. The evolutionary implications of the melanism are dis- cussed following the enumeration of species. GEOMETRIDAE Several of the classic Palaearctic mutant melanics occur in this family, especially in the tribe Cleorini of the Ennominae, and the best-known of the American ones, namely Amphidasys cognataria Gue. and Phigalia titea Cram., are congeneric with Old World species having melanic forms (A. betularia L. and P. pedaria Fabr. ). In A. cognataria the black form^ (which is solidly black, without markings ) is decidedly more frequent than the “normal” in eastern Pennsylvania. The species is very com- mon and data are available for a number of localities, but too little is known of the movements of individuals and the char- acteristics of the population of this, as of most nocturnal moths, for a really meaningful analysis to be made of the variation in frequency of the melanic form noted from different (urban vs. rural) sampling stations. The over-all frequency of the melanic phenotype appears to be 60-65% of both sexes. In the very abundant spring species, P. titea, the melanic form is less frequent than die “normal”, but a marked rise in frequency has been observed in recent years in Pennsylvania. The increase was especially notable in March- April, 1963, in which season the species was also unusually abundant in general. As pointed out by Ford and others, unusual abundance of a species is as a rule accompanied by extraordinarily wide latitude in variation, and this was the case here. While the number of 19 20 SHAPIRO /. Res. Lepid. total melanics exceeded 25% of the males for the first time at nearly all sampling stations, several partial melanics of previously unknown types also appeared. The usual melanic form as in cognataria, is solid black. The partial melanics were of two types: either black, with the submarginal band on all wings normal gray; or dark, uniform charcoal-gray, with the darker markings indistinctly visible above and beneath. Examples of all of these types are in the collection of the author. The fre- quency of total melanics in 1959-61 was 10-15%, and in 1962, ca. 18%. Melanism occurs in a large number of Cleorini in Pennsyl- vania. Aside from these two common, well-known species, several of the less common forms have shown melanic tendencies. The distinctive Epimecis virgirmria Cram, has developed both partial and complete melanism. The partial melanics are dark, brownish-gray, with the three principal lines of the wings un- suffused over their light edging (basad on the postbasal line, and marginad on the median and submarginal ones). The total melanie is solid black with the exception of these same three light lines, which are even more conspicuous on the black ground.^ E. virginiaria is uncommon at the latitude of Philadelphia and northward, but the black form seems to be occurring with con- siderable frequency, the combined values of the two dark types probably reaching at least 50% ( of which at least 35% is '‘total” ) . One melanic specimen of Paraphia suhatomaria Wood has been taken in Chestnut Hill, Philadelphia (28 July 1962); it and the related P. unipuncta Haw, are both uncommon to rare. Another related species with melanic tendencies is the scarce Nacophora quernaria Ab. & Sm. This insect is too uncommon for any reliable frequency figure to be available for the melanic form in eastern Pennsylvania. Less than a dozen area speci- mens have been seen by the writer. The melanics^ have the ground-color full black, with the white edging of the postbasal and postmedian lines (basad and marginad, respectively) con- spicuously unsuffused. No melanics have yet turned up in the rare spring species, Lycia ursaria Walk., but they may be ex- pected. Two more possible instances are found in the Cleorini, namely Cleora pampinaria Cue. and Ectropis crepuscularia L. Both of these common insects have a wide range of variation from very light to very dark gray forms. One fully black pampinaria was taken at Flourtown, Montgomery Co., Pa., 8 September 1963. A single melanic specimen of crepuscularia has been described.^ The writer has not seen any black Pennsylvania individuals of this species. The important pest, Palaeaciita vernata Peck, is quite variable in color, and a good case for melanism can be made when the a (l):19-24, 1964 MELANIC MOTHS 21 darkest forms are compared with the lightest. No fully black individuals have been taken, and the character is apparently not so clear-cut as in other Cleorines. The darkest individuals seem to be more frequent in recent years than previously, however. One more indisputable case of melanism in the Geometridae concerns a member of the Sterrhinae, Cosymbia lumenaria Gue. Two black specimens were taken in 1963 (Flourtown, 7 July; Conshohocken, 11 Aug., both Montgomery Co.), both with the discal spots and the pastmedian line on all wings the usual pale gray, contrasting with the sooty ground-color. The species is rather uncommon in the area, and no frequency figure can really be given. PHALAENIDAE The best-known species with melanic tendencies in the Phalaenidae is Charadra deridem Gue. It is common and the frequency of the black forms in Pennsylvania is at least 50% and probably somewhat higher. Some of the melanics are solid black; most have the postmedian line white, and a few have the postbasal line white also. The second of these was described in 1923.® All have the normal black markings faintly discernible. A few females seem to be intermediate between the ^normal” and melanic phases, being moderately suffused with black scales.® Two scarce species contribute additional records. Panthea fur- cilia Pack, has a nearly melanic form which is considerably darker than the typical form, but the dark markings remain visible. Folia latex Gue. has a black form, previously unrecorded, also with the dark markings still identifiable. Both are too un- common for frequency figures to be available. Dark furcilla have been taken for several years in Philadelphia, and the first melamc latex was taken at Conshohocken, Montgomery Co., 11 May 1963. One more possible case of melanism is Marasmalm inficita Walk., but the citation rests on but a single melanic specimen (Norristown, Montgomery Co., 9 August 1961). The species is quite uncommon. NOTODONTIDAE Two Notodonts appear to have melanic forms, namely Cerura ckwrea Walk, and Lophodonta ferruginea Pack. Both appear to have a relatively hi^ frequency, although the second species is generally much commoner than the first, and hence more readily analyzable. In both, the melanism is quite noticeable, although in cinerea it is a dark charcoal gray, not a black, form. The frequency in ferruginea has been about 60% for several years, indicating that, as in A. cognataria, the darker has become the statistically "normal” form. In 1927 Chermock described melanics of Heterocampa um~ brata Walk.^ and in 1929 of Fentonia marthesia Cram.,* both 22 SHAPIRO J. Res, Lepid. from Pittsburgh. Both species occur in the writer's sampling area, but no melanics have been found of either. One black specimen of Schizura apicalis G. & R. has been taken (Flour- town, Montgomery Co., 14 June 1963). The species is un- common. A possible second melanic of the same species, in very poor condition, was taken 8 July, 1959 (Norristown). DISCUSSION There seems to be adequate evidence to support the notion that melanism is fully as widespread in America as in Europe. Of particular interest is the regularity with which melanism occurs in species having a certain facies in the 'normal” type condition, i.e. silvery-gray moths with black markings or dusting, irrespective of family. It becomes possible to predict the occur- rence of melanics in various species; the writer actually did predict that melanic Cleora would be found, a year before the first specimen known to him turned up. In several cases where melanism has not appeared within a species where it might be expected on the basis of wild-type coloration, a black sibling species exists instead; for example, Tolype velleda Stoll and its dark sibling laricis Fitch ( Lasiocampidae ) . Many of these melanics seem to recur sporadically in widely separated localities. The evidence would suggest that the large number of apparently new melanics turned up by the author in recent years is not really unusual, but rather the result of more assiduous collecting. Many species which show melanism today have had melanic "aberrations” described in the past, mostly on the basis of single specimens, and the genetic continuity of some of these with present-day melanics in the same species is often dubious. The melanic form of Phigalia titea is perhaps the best documented, and seems to have arisen spontaneously in various parts of the range at various times, meeting with variable success in different localities. In at least some of the species concerned, the rate at which the mutation to "melanic” occurs must be fairly high. The more often a specific mutation occurs, the more likely it is that it will at one time or another occur in a situation or environment where it will be advantageous, or it least not deleterious. Under such circumstances it is likely to be pre- served and propagated. It is specifically this fact that the orthogenecists have seized upon to support their inference of a “direction” or "purpose” in evolution. All we can say at present is that certain genes seem to show an instability (probably chemical), and that in the cases here considered, that instability leans in the direction of melanism. 3 (1): 19-24, 1964 MELANIC MOTHS 23 Several of the lesser known of the examples cited have been found only quite recently, and in the absence of prior records it is impossible to state to what extent they have been present in the past, either genetically continuous with the current form or as a result of previous mutation of the same type. It is of some interest, however, to consider the future of any given melanic mutation which may be assumed to have just arisen de novo. Most “industrial” melanics whose genetics have been studied have proved to be unifactorial, autosomal dominants. This is true of at least A. cognataria in the U. S., as well. The selection against deleterious dominant mutations is most effective, so that for a dominant mutation to be successful it is not even enough for it to be adaptively neutral; it must have some adaptive ad- vantage attached to it. What are the factors determining whether or not a given moth melanic will succeed biologically (be selected “for” or at least not selected “against” ) ? The conventional explanation of the development of “in- dustrial melanism” in England was that the melanics had a concealment advantage in the industrial areas. This may be true under conditions of extreme contamination by soot, where the landscape is physically blackened, but it is of dubious value under other circumstances. It is true that in England at least, the highest percentages of melanics have been found to coincide with urban-industrial centers. This has led to an alternative explanation, namely, that the urban-industrial conditions have discouraged predators which would weed out the conspicuous melanics, and that in consequence the pressure of selection has been decreased, allowing the dominant melanics to spread. Further, several authors have reported that the melanics in several Palaearctic species possess a higher survival value under adverse conditions (e.g., semi-starvation) in the larval stage. This higher viability complements the second hypothesis nicely, in that it augments neutrality by giving an advantage which accounts for the rather steep frequency increases observed. TTiis differential viability would be especially significant if it were to be demonstrated that the melanics are more resistant to urban- industrial atmospheric pollutants, such as SO3. It is difficult to state to what degree the melanism in American species is associated with urban-industrial conditions. As prev- iously remarked, there are not suflficient data available for most species to support a meaningful and truly significant analysis. The two exceptions, A. cognataria and P. titea, so far as the 24 SHAPIRO /. Res. Lepid. presently available Pennsylvania data would indicate, seem to follow the British trend of correlation well. Since the urban- industrial complex in question— Philadelphia and vicinity— is not characterized by extensive soot contamination, we may reasonably inquire into the means of maintenance of melanism thereabouts. The only concealment advantage for melanics which is apparent as a result of civilization there would be on asphalt highways; and it is rare indeed to find a moth of any kind resting horizontally on the ground. Most of the species having melanic forms rest by day on tree trunks, and several sit down among dense foliage in the ground-cover story. These facts suggest that of the three possibilities suggested for the maintenance of the melanism, concealment value may be the least significant. We do not, of course, know whether the pleiotropic increased viability exhibited by Cleorine melanics ( at least in Europe ) is shared by melanics in other species groups; it would be expected that the melanism in various families and subfamilies would have arisen by mutation of various non- homologous genes in somewhat different genetic systems; in- creased viability would not, then, be likely consequential gen- erally. We do not know enough about moth biochemistry to state whether the viability increase is a direct result of the in- crease in melanin, or vice versa, or whether the two effects are interlocking but not in direct sequence ( as where an enzyme may play a key role in two sets of concurrent reactions). Similar melanizing effects in different species may be produced in quite different ways, by different enzymes in different reaction se- quences. The writer would like to see the progress along these lines coming, at last, from the laboratories of American re- searchers. ACKNOWLEDGEMENT The author s thanks to Mr. WILLIAM F. BOSCOE of Phila- delphia, Pa., for granting him access to his collection and for furnishing several instances of melanism with which the author was personally unacquainted. 1. A. cognataria “ab. swettaria” Bames & McD., 1917 Contr. Nat. Hist. N.A. Lep., Ill (4), p. 246; also figured, pL 27, no. 6. 2. E. virginaria “form carbonaria” Haimbach, Ent. News XXVI (1915), p. 321. 3. N. quernaria “form atrescens” Hulst, Can. Ent. XXX (1898), p. 162. 4. E. crepuscularia “ab. fumataria” Minot, Proc. Bost. Soc. Nat. Hist. XHI (1869), p. 84. 5. C, deridens “form fumosa” Draudt, in Seitz, Macrolep., N.A. Noct. VII (1923), p. 19. 6. C. deridens “var. nigrosuffusca” Strand, Archives Naturgeschicter, A(2) (1917), p. 46. 7. H. umbrata “ab. nigra” Chermock, Bull, Brook. Ent. Soc., XXH (1927), p. 118. 8. F. marthesia “ab. nigra” Chermock, Bull. Brook. Ent. Soc., XXIV (1929), p. 20. Journal of Research on the Lepidoptera 3 (1) : 25-36, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1964 THE GENUS LYCAEIDES IN THE PACIFIC NORTHWEST JON H. SHEPARD Department of Entomology, Oregon State University, Corvallis, Oregon A CRITICAL AND THOROUGH REVISION of the Noith American forms of the genus Lycaeides Hubner 'was published by Nabokov (1949). The published names of the species and subspecies in the genus were clarified and the basic distribution of the differ- ent forms was presented. In this paper, he mentioned the lack of information on species from the Pacific coast, especially north of California. It is the purpose here to provide additional in- formation on the distribution of the species of Lycaeides in this area and to present the possible means of the formation and origin of subspecies of Lycaeides in the Pacific Northwest. The territory included in this study covers Oregon, Washington, Idaho, southern British Columbia and western Montana. The author looked at and identified the specimens in every major Pacific Northwest collection of butterflies, with the exception of the collection at the University of Idaho. The genus Lycaeides is composed of three species. Two of these are found in the Pacific Northwest. The species L. ar- gyrognomon Bergstrasser is found in the mountainous and forested areas; being distributed from approximately 3000 feet to above timberline. L. melissa Edwards is a lowland form, occurring in the Great Basin and Columbia Plateau east of the Cascade mountains and is found from 300 to about 3000 feet. Melissa is most commonly associated with deciduous forests and the ponderosa pine plant associations (figure 2). The male genitalia was used to separate argyrognomon from melissa. Nabokov (1949) showed that this was the best method for distinguishing specimens of heavily marked argyrognomon from the normally heavily marked melissa. Measurements of the genitalia were not made because of a lack of long series from any one area. Nabokov (1949) attempted this in an effort to separate different subspecies. But as was pointed out by Brown (1950a,b), the measurements were not statistically valid and thus not pertinent for separation of subspecies. Submitted as part of requirement for Bachelor of Science, with Honors, Oregon State University, June, 1963. 25 26 SHEPARD /. Res. Lepid. llie initials after the names are used with the locality data to indicate in which collection the individual specimen can be found. In addition, the abbreviations (Nab) and (BVL) were used to indicate the records of distribution published by V. Nabokov and Ben V. Leighton. It should be noted here that B. V. Leighton's “Butterflies of Washington” (1946) has been much critized. The author does not feel that these criticisms are justified, as it is more realiable than any other published list of the butterflies of the Pacific Northwest. The above published records were included here when the author did not personally see material from the specific localities. Since this paper at- tempts to clarify the distribution of the Lycaeides in the Pacific Northwest, every available and reliable locality record was used. Lycaeides argyrognomon ricei (Cross) Plebeius scudderii ssp. ricei Cross, 1937, Pan-Pacific Ent., 13:88. Lycaeides argyrognomon ricei occurs in the Cascade moun- tains from Crater Lake, Oregon to the Hope Mts. of British Columbia (Table 1). At the southern end of its range it blends into the Sierra Nevada subspecies L. a. anna Edwards, The mountainous area in Oregon from Crater Lake to Mt. Shasta, Calif, is the area of intergradation of anna & ricei. In subspecies anna the underwings are chalk white with distinct black mark- ings, whereas ricei is characterized by pale chalk- white ground on the undersides of the wings and by very light black marking on the underwing (fig, 1, nos. 14 & 15). The populations south of Crater Lake have individuals with both characteristics. The Lakeview specimen is an example of this (fig. 1, no. 13). It would be easy to dismiss this problem by labeling the specimens L. a. anna near ricei, however specimens of ricei from Blue Slide, Washington, Paulina Lake and Gilchrist, Oregon (fig. 1, nos. 10, 11, 12) also show the heavier marked underside characteristic of anna. These specimens (fig. 1, nos. 10, 11, 12) come from isolated populations along the eastern edge of the Cascade mountain system. The climate there is different from that of the central Cascades. The summers are longer, hotter and drier. The sun shines almost constantly with little summer precipitation. This is in contrast to the weather of the central Cascades. Here rain, thunder and snow storms are not un- common, even in July. The Cascade mountains of southern Oregon are broken and the dry eastern weather reaches further west to Medford. Thus several questions are raised. One, are 3 (l):25-36, 1964 GENUS LYCAEIDES 27 specimens from the drier eastern edge of the Cascades reacting phenotypically to the environment or are these populations just isolated local races? If they are reacting to the environment then are the populations from southern Oregon also examples of this or do they represent an intergradation with the California anna? This is a problem not immediately answerable. The author suggests that for convenience all western Oregon ar- gyrognomon be referred to as ricei and all California argyrog- nomon from the northern part of the state be referred to as anna. At the northern extent of its range, ricei is much less com- plicated than near the southern boundary. In southern British Columbia there is a distinct break between ricei and scudderi Edwards. In northwestern Washington and southwestern British Columbia the underwings of ricei often have some heavy mark- ings. From Lake Chelan, Washington to the Hope Mts. of British Columbia and from the Olympic Mts. of Washington to Vancouver Island, the populations of ricei have a varying per- centage of specimens with scattered and irregular black mark- ings (fig. 1, nos. 19-22). However the pale chalky underside is retained in all of the males and most of the females. The popu- lations of scudderi have an underside ground color that is char- acteristically light brown (fig. 1, nos. 1, 2). The distinction between the two is definite. Nabokov (1949) was not sure of where the Vancouver Island, British Columbia populations should be placed. They seem to represent the most northern extension of ricei. These speci- mens all have the chalk white underwing of the typical ricei. Each specimen has a strong pattern of dots on the underside of the wing but those dots are irregular and diffused (fig. 1, nos. 20, 21). The dot pattern of scudderi is not irregularly dis- tributed and the black dots are definite spots (fig. 1, nos. 1, 2). Blackmore (1927) and Jones (1951) have confused Vancouver Island argyrognomon with melissa. This mistake should be noted and corrected in any future paper on British Columbia Lepidoptera. In addition, the southern British Columbia popu- lations of argyrognomon have been constantly referred to as anna (Blackmore, 1927, Jones, 1951). This is due to the heavy marking found on the underwings. However the pattern of these markings is very different from anna (fig. 1, nos. 14, 20, 21). Lycaeides argyrognomon scudderi ( Edwards ) Lycaena scud- derii Edwards, 1861, Proc. Acad. Nat. Phil., p. 164. As mentioned above, the separation of ricei and scudderi 3 (l);25-36, 1964 GENUS LYCAEIDES 29 is distinct. However, separation of scudderi and melissa in southern British Columbia is not as simple. Both forms have a definite pattern of black dots on the underside of the wing. Only an examination of the male genitalia will assure adequate separation. The genitalia from specimens of the Okanogan Lake region are always distinctly scudderi or melissa even though the populations of the two species may be found within a few miles of each other. This is in contrast to the Wyoming area where the genitalia of some specimens are intermittent (Nabo- kov, 1949). This would suggest that the two species are not interbreeding in the Okanogan area of British Columbia. The subspecies scudderi occupies all of southern continental British Columbia except for the Hope Mountains, the narrow sagebrush-filled valleys, and the extreme southeast corner of the province (Table 1). The Hope Mountains are populated by L. a. ricei, the sagebrush by L. melissa, and the southeast by L. a. ferniensis Chermock. Nabokov (1949) credited British Columbia with the sub- species ferniensis from Ferney and Cranbrook B. C. No speci- mens were examined from either of these specific localities. However a specimen from Kaslo, just north of Cranbrook, is definitely scudderi. Again, as with the anna-ricei problem, many more specimens need be collected from the area. Is ferniensis Fig. 1. Lycaeides argyronomon scudderi, 1, 2,.; melissa, 3, 6, 9; atraeprae- textus, 4, 5, 7, 8; ricei,_ 10-13, 1^-24; anna, 14. 1. Salmon arm B.C. 2. Lytton B.C. 3. Oak Creek, Wash. 4. Cornez Creek, Ore. 5, Lake Wallowa, Ore. 6. Pearrygin Lake, Wash. 7. & 8. Oregon Butte, Wash. 9. Kennewick, Wash. 10. Paulina Lake, Ore. 11. Gilchrist, Ore. 12. Blue Slide Lookout, Wash. 13. Lakeview, Ore. 14. Wolverton Mdws., Calif. 15. Tombstone Prairie, Ore. 16. Mt. Rainier, Wash. 17. Crater Lake, Ore. 18. Sheep Lake, Wash. 19. Bunker Hill Lookout, Wash. 20. & 21. Mount Malahat, B.C. 22. Bunker Hill Lookout, Wash. 23. Bradley Creek, Ore. 24. Mt. Cheam, B.C. 30 SHEPARD /. Res. Lepid. a local form or something of wide enough distribution and dis- tinct enough to deserve a subspecific name? Southeastern British Columbia is composed of many small mountain chains separated by deep valleys. There are many areas where local populations could be isolated and differentiated. Further collecting will be necessary to clarify the concepts of subspecies in this area. Lycaeides ar gyro gnomon atrapraetextus (Field) Plebejus atrapraetextus Field, 1939, Jour. Kansas Ent. Soc., 12:135-136. This subspeeies is found in the Rocky Mountains of Idaho and Montana and in the mountainous areas of eastern Oregon and southeastern Washington (Table 1). This subspecies bears an artificial resemblance to melissa because of the heavily marked underwing and the brown color of the underside of the wing. The two forms can be adequately separated only by an examination of the genitalia. The subspecies atrapraetextus closely resembles its northern relative scudderi. The two are separated by a character on the underside of the hind wing. There is a band of light, almost white, chevrons basal to the outer band of orange markings (fig. 1, nos. 7, 8) in atraprae- textus. Scudderi has this band reduced to a series of light chevrons, resembling check marks (fig. 1, nos. 1, 2). When Nabokov (1949) discussed the distribution of this subspecies, he limited it to the mountains of northern Idaho and western Montana. Nabokov had one isolated specimen of argyrognomon from near Boise, Idaho which he could not place as to subspecies because he lacked any other specimens from near this locality. The specimens from eastern Oregon and Washington are not sufficiently different from the Idaho and Montana forms to be separated as a subspecies. Thus a connect- ing link is supplied between Boise and northern Idaho. The author suggests that the name atrapraetextus be used to apply to argyrognomon from all of Idaho, western Montana, eastern Oregon and southeastern Washington. A question might arise. Why is there no intergradation be- tween populations of atrapraeteodus from the Ochoco Moun- tains, Oregon and populations of ricei from just west in the Cascades? The Ochoco's seem to receive all their faunal ele- ments from the Rockies. A narrow but permanent band of juniper separates the Cascades from the mountain ranges of eastern Oregon. This is also an effective barrier for such groups as Speyeria Scudder and Euphydryas Scudder. 3 (l):25-36, 1964 GENUS LYCAEIDES 31 ^ Lycaeides ar gyro gnomon ricei Cross Q Lycaeides argyrognomon scudderi Edw. A Lycaeides argyrognomon ferniensis Chermock ^ Lycaeides argyrognomon atraepraetextus Field H Lycaeides melissa Edw. Figure 2. The Distribution of Lycaeides in the Pacific Northwest. I would like to thank the following persons for providing me with access to their private or public collections: Dan Carney, E. J. Dornfeld (EJD), Clarence J. Duffy (CJD), M. T. James Washington State University) (WSU), J. D. Lattin (Oregon State University (OSU), David Mays, David McCorkle (DM), Miss Olga Meixner (Vancouver City Museum) (VCM), E. J. Newcomer (EJN), and G. G. E. Scudder (University of British Columbia ( UBC ) . 32 SHEPARD J. Res. Lepid. TABLE 1. DISTRIBUTION RECORDS L. a, ricei: Oregon- Clacl^as Co.: Government Camp, Hwy, 26, July 2, 1934, S, Jewett Jr., 1 cf, (OSU) ; Mt. Hood, July 2, 1934. 1 d, (OSU) ; East of Toll Gate, Hwy. 26, Jvdy 15, 1906, 2 d'd', (OSU). -Deschutes Co.: Paulina Lake, July 28, 1934, S. Jewett Jr., 2 cfcf and 2 (OSU). -Douglas Co.: Diamond Lake, July 14, 1937, 1 cf and Aug. 16, 1935, 1 iontana: De Borgia (1), Gallatin Valley (1). L. a, scudderi: British Colvunbia- Cariboo Dist. : Cariboo, July 10, 1920, Id, (UBC); Quesnel, (Nab) ; Stanley, (Nab). -Kamloops Dist.: Armstrong, June 29 - July 10, 1914, W. Downes, 4 dddd and 3999, (UBC) ; Chase, Aug. 9, 1919, W. B. Anderson, Id and 1 9, (UBC) ; Enderby, June 18 - Aug. 15, 1920 - 1922, J. Wynne, 4 9999, (UBC) ; Hefferly Creek, (Nab) ; Nicolo Lake, June 17, 1922, W. R. Buckell, 1 9, (UBC) ; Salmon Arm, April 25 - Jmie 17, 1914 - 1921, W, R. Buckell, 8 males and 1 female, (UBC) ; Vaneby, elev. 4500', Aug. 10, 1921, 1 9 and June 21, 1921, 1 9, T. A. Moillet, (UBC) ; Vernon, June 22 - Aug. lo’ 1904 - 1919, 4 dddd and 4 9999, (UBC). -Kootenay Dist. ; Carbonate, (Nab); Kaslo, 1897, Id, (UBC) ; Revelstoke Mt. , Aug. 14, 1923, W. R. Buckell, 6 males and 2 females, (UBC). -Lil- looet Dist, : Lillooet, July 10, 1922, K, F, Anderson, 1 d, (UBC) ; Jesmond, (Nab) ; Pavilion, Jtily 18, 1933, W. Downes, 2 dd and 1 9, (UBC). - New Westminister Dist.: New Westminister, (Nab), -Osoyoos Dist. ; Kelowna, (Nab). -Peace River Dist. : RoUa, (Nab). Genitalia Examined: British Columbia; Armstrong (2), Chase (1), Kaslo (1), Lytton (1), Revelstoke Mt. (1), Salmon Arm (1), Vernon (1). 34 SHEPARD /. Res. Lepid. Lycaeides melissa ( Edwards ) Lycaena melissa Edwards, 1873, Trans. Amer. Ent. Soc., 4:346- 348. L. melissa is found in the Pacific Northwest only in the cen- tral Columbian Plateau, the Snake River drainage lowlands, and the Great Basin area of southern Oregon (Table 1). It can be confused with certain subspecies of argyrognomon. The dis- tinctions have been discussed under those respective subspecies; scudderi and atrapraetextus. The habitat is much more varied for melissa than for argy- rognomon. On the east side of the Cascades it can be found in ponderosa forests. From there it invades the lowland sagebrush country only along streams where deciduous growth is supported. It is not found directly in the dry sagebrush. Also it is common- ly found in the valleys of Oregon and Washington where irri- gation and farming occur. Along the Snake River, Columbia River and in southern Oregon it is found near streams and irri- gated regions. The species melissa can succeed in invading mountains only to the limit of continuous ponderosa pine dis- tribution. Then the species argyrognomon is dominant. Nabokov (1949) pointed out several places where argyr- ognomon and melissa were sympatric i.e. Brewster, Washington and Fort Klamath, Oregon. This sympatricity has never been found by the author or by any of the Pacific Northwest collectors with whom he has communicated. The Brewester records are not especially reliable for specimens that are normally mountain forms. Remington (1963) has recently brought attention to the labels of T. C. Hopfinger (of Brewster). The ricei specimens labeled Brewster could never have been taken exactly there. The same is probably tru for the specimens from Fort Klamath. That is, they were not taken together but at close localities separated by wide ecological differences. In Wyoming, where Nabokov (1949) has demonstrated the interbreeding of the two species, argyrognomon and melissa, it would appear that the two are able to occupy the same micro- habitat and thus interbreed. In the Pacific Northwest, other than perhaps extreme southeastern Idaho, the two species have definite and separate ecological preferences. The distribution of the genus Lycaeides in the Pacific North- west is very similar to that of the bird genus Zonotrichia Rand ( 1948 ) . Rand discussed the origin of this and several other bird genera in relation to the effect of the Wisconsin glacial period GENUS LYCAEIDES 35 on the isolation of species of birds to form subspecies. Rand proposed that at the glacial maximum there were four refugia where animal species were isolated. These areas were northwest Alaska, the Pacific coast, Rocky Mountains, and eastern United States. The various groupings of the subspecies of ar gyro gnomon correspond well to these refugia. L. ricei, anna, and lotis Linter, found on the Pacific coast, are one group characterized by an underwing with a chalky white groundcolor. Alaskensis Cher- mock and scudderi, distributed from southeastern Alaska to southern British Columbia are another natural grouping. These two subspecies blend into each other and specimens from the intermittent area are not easily assigned to either subspecies. Atrapraetextus from the central Rockies and aster Edwards from eastern United States are the two other groupings corresponding to Rand's bird distribution. In addition there is a fifth group including sublivens Nabokov and longinus Nabokov. These are found in he Rocky Mountains south of Montana. There was an extensive glacial sheet covering northwestern Wyoming during the Wisconsin period (Kamp, 1963). This was likely the factor that separated argyrognomon to form sublivens and longinus. Since argyrognomon formed subspecies as a result of glac- iation during the Wisconsin, it and the species melissa must have been separated as species prior to this time. SUMMARY There are two species of Lycaedis in the Pacific Northwest, argyrognomon and melissa. Argyrognomon has formed three subspecies ricei, scudderi, and atrapraetextus. These subspecies probably resulted from a break-up of the species during the Wisconsin glacial period. LITERATURE CITED BLACKMORE, E. H. 1927. A check list of the Macrolepidoptera of British Columbia. B. C. Provincial Museum Pubs. BROWN, MARTIN F. 1950a. Measurements and Lepidoptera. Lep. News, 4: 51-52. 1950b. In reply to Prof. Nabokov. Lep. News, 4:76. EHRLICH, PAUL R. and ANNE H. 1961. How to know the butterflies. Wm. C. Brown Co., Dubuque, Iowa. HEMMING, FRANCIS. 1954. Opinion 270, Opinions of the International Commission on Zoological Nomenclature, vol. 6 (part 2) :25-40. 36 SHEPARD /. Res. Lepid. KAMP, J. W. 1963. Description of two new species of Grylloblattidae and of the adult of Grylloblatta harheri with an interpretation of their geographical distribution. Ann. Ent. Soc. Amer., 56:53-68. LEIGHTON, BEN. V. 1946. The butterflies of Washington. Univ. of Wash. Press {Occ. Papers), 9:47-63. JONES, J. R. J. L. 1951. An annoted check list of the Macrolepodoptera of British Golumbia. Ent. Soc. B. C., Occ. papers no. 1. NABOKOV, V. 1949. The nearctic members of the genus Lycaeides Hubner (Lycaenidae, Lepidoptera ) . Bull. Mus. Comp. Zoo., 101: 477-540. 1950. Remarks on F. Martin Brown’s ‘measurements and Lep- idoptera’. Lep. News, 4:75-76. 1952. The female of Lycaeides argyrognomon sublivens. Lep. News, 6:35-36. RAND, A. L. 1948. Glaciation, an isolating factor in speciation. Evolution, 4:314-321. Journal of Research on the Lepidoptera 3 (1) : 37-44, 1964 1140 W. Orange Grove Ave., Arcadia, California, U.S.A. © Copyright 1964 THE ORIGIN OF A SYMPATRIC SPECIES IN COLIAS THROUGH THE AID OF NATURAL HYBRIDIZATION WILLIAM HOVANITZ ( continued from volume 2, page 223 ) V POPULATION COMPARISONS Section IV of this series showed by means of scatter diagrams, the relationship between the variations of two characteristics in arctic populations of Colias. This section (V) will show the general distribution within each population of each of the two characteristics so as to give a better idea of the nature of the introgression within the populations in a way that the conelation diagrams could not indicate. The characteristics are those previously illustrated on figure 8 (page 271, vol. 1) and fig. 9 (page 206, vol. 2), each being graded in a series of nine classes, 0, 1, 2, 3, 4, 5, 6, 7 and 8. Using these classes, histograms have been constructed based upon the nu- merical values for each class. The histograms so constructed are illustrated in figures 43 through 49. to be continued Figures 43 through 49. Histograms illustrating the variation in pigmentation (left) and border pattern grade (right) of various arctic populations. The grades have been illustrated in earlier sections of this paper. Note that some populations (such as Lake Harbour) show no in- trogression but only Colias hecla or CoUa nastes, others (such as Reindeer depot) show no introgression but only one species, either C. nastes or C. hecla, and others show a wide range of introgression with or without many parental species ( Spence Bay, Coral Harbour, Repulse Bay, Chester- field Inlet). 37 38 HOVANITZ /. Res. 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Lepid. 01 2345678 ORANGE 0.1, 2, 3, 4, 5, 6, 7, 8 REINDEER DEPOT MEADE RIVER L_J L—l I I I I I I 01 2345678 BORDER