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Volume 16 1962 Number 1
JOURNAL,
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
In This Issue
ANALYSIS OF PIERIS NAPI & BRYONIZ: IN EUROPE ECOLOGY OF DONNER PASS RHOPALOCERA BIOLOGY OF SANDIA AND DYOPS NOTES ON ERORA QUADERNA
(Complete contents on back cover)
30 August 1962
THE LEPIDOPTERISTS’ SOCIETY 1962 OFFICERS
President: Joun A. Comstock (Del Mar, Calif., U. S. A.) Ist Vice President: H. B. D. KettrLewe xu (Oxford, England) Vice President: Himosuti INovE (Fujisawa, Japan) Vice President: TARSICIO EscALANTE (Mexico City, Mexico) Treasurer: GrorcE EuLe (Lancaster, Penna., U. S. A.) Asst. Treasurer: SmpnEY A. HEssEu (Washington, Conn., U. S. A.) Secretary: Paut R. Exruicu (Stanford, Calif., U. S. A.) Asst. Secretary: Frioyp W. Preston (Lawrence, Kansas, U. S. A.) EXECUTIVE COUNCIL Terms expire Dec. 1962: Harry Krocerus ( Helsingfors, Finland) F. M. Brown (Colorado Springs, Colo., U. S. A.) Terms expire Dec. 1963: J. C. E. Riotre (Toronto, Canada) Da.ror Povotny (Brno, Czechoslovakia) Terms expire Dec. 1964: H. A. FREEMAN (Garland, Texas, U. S. A.)
S. L. pE La Torre y Cauieyas ( Havana, Cuba) and ex-officio: the above six elected Officers and the Editor-in-Chief
The object of The Lepidopterists’ Society, which was formed in May, 1947, and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches,...... to issue a periodical and other publications on Lepidoptera; to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed toward these aims (Constitution, Art. II). A special goal is to encourage free interchange among the lepidopterists of all countries.
Membership in the Society is open to all persons interested in any aspect of lepidopterology. All members in good standing receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer the full dues for the current year, together with their full name, address, and special lepidopterological interests. All other correspondence concerning membership and general Society business should be addressed to the Secretary. Remittance in dollars should be made payable to The Lepidopterists’ Society. There are three paying classes of membership:
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The Lepidopterists’ Society is a non-profit, scientific organization. The office of publication is New Haven, Connecticut (see address inside back cover). Application for Second-class mail privileges has been approved at New Haven, Connecticut.
JouRNAL OF
Tue Leprpoprerists’ SOCIETY
Volume 16 1962 Number 1
NOTES ON ERORA (LYCAENIDA‘)
by Kittan ROEVER
Prior to 1940 all the Erora occuring in the United States were con- sidered to be Erora laeta Edwards. The range at that time was known to be curiously disjunct. An eastern population occuring from Ontario and Nova Scotia south to Virginia and a southwestern population occuring in Arizona were known. Dos Passos (1940) described the population in Arizona and New Mexico as Erora leta sanfordi. He made no mention of Erora quaderna (Hewitson) so I presume no Mexican Erora were examined.
Fietp (1941) affiliated the Arizona population with Erora quaderna which was described from Mexico in 1868 and listed Erora lzta sanfordi dos Passos as a synonym. FreLp’s decision was based apparently on the original description and on the description found in the Biologia Centrali- Americana. He also made no mention of examining any Erora from Mexico. FieLp treated Thecla attalion Godman & Salvin as a dimorphic form of the male of quaderna, the name being based on a form lacking the green color on the ventral wing surfaces. He cautioned that such specimens lacking the green beneath may not occur in nature except in a worn condition. It has been my experience with quaderna in Arizona that the green overscaling is easily removed leaving a brownish-buff color on the undersurfaces. In view of this it seems that attalion can remain in synonomy.
CiencH (1943) revived sanfordi as the subspecies of quaderna oc- curing in the southwestern United States. His opinion was primarily based on the comparison of two quaderna specimens from Tancitaro, Michoacan, Mexico with Arizona material. CLENcH selected Tancitaro as the type locality because the type of quaderna, a female in the British
NS)
Rorver: Erora Vol.16: no.1
Museum, carried no accurate data. Unfortunately CLENcH failed to give any characters by which quaderna quaderna and quaderna sanfordi could be separated.
In an attempt to learn the diagnostic characters separating the subspecies I borrowed from the Museum of Comparative Zoology the two females (Tancitaro, Michoacan, Mexico, 7800’, VI-30-41 and VII-8-41, resp.; H. Hoocsrraat and R. Haac, collectors) which CLeNcH had examined. In addition Dr. Tarsicio EscaLantE kindly loaned me two females (Desierto de los Leones, approx. 7500’, Distrito Federal, Mexico, I-45 and IJ-23-61). These four specimens from central Mexico were compared with a series of Erora from various localities in southern Arizona. The variation of the Arizona material clearly encompasses the Mexican material. On the basis of my comparison I consider sanfordi to be a synonym of quaderna, as did Fie.p. I shall add that in a series of forty females from Madera Canyon, Santa Rita Mts., Santa Cruz Co., Arizona, there is considerable variation in the amount and shade of blue on the dorsal surface and in the size of the orange spots on the ventral surface.
In giving the distribution of quaderna within the United States Fretp (1941) lists Arizona, Utah, and New Mexico. I have been able to find no further record concerning the occurence of this species in Utah. Dos Passos (1940) records two specimens received from R. T. KELLOGG of Silver City, New Mexico. Ltoyp M. Martin (in litt.) states that there is one specimen in the Los Angeles County Museum collection bearing the label data: III-22-40, Providence Mts., San Bernardino Co., California, T. B. Buevins, JR., collector. This is an entirely different environment from that in which it occurs in Arizona.
In 1960 I located a specimen of quaderna in the collection at park headquarters of Big Bend National Park. The data on this specimen, a female, were Big Bend National Park, Brewster Co., Texas, VI-12-37, Roun H. Baker, collector. To my knowledge this represents the first record of this species from Texas and is a considerable eastward range extension.
The data of my Erora quaderna collections in Arizona are as follows: APACHE COUNTY: Trout Creek Road, 4-6 mi. S of Smith Park, approx. 7500’, VII-4-58, 3 males; VII-4-59, 4 males; VI-25 & 26-60, 4 males, 1 female; VII-22-61, 1 male, 3 females. COCHISE COUNTY: E. Turkey Creek Canyon, approx. 3 mi. E of Onion Saddle, Chiricahua Mts., 6400’ — IV-10-59, 2 males, 10 females, IV-15-60, 12 females. Pinery Canyon, Chiricahua Mts., IV-15-60, 2 males, 4 females. 1 mi. NW of the South- western Research Station, Chiricahua Mts., [V-15-60, 4 females. 1 mi.
1962 Journal of the Lepidopterists’ Society 3
N of Rustler Park, Chiricahua Mts., VI-19-60, 2 males. COCONINO COUNTY: Oak Creek Canyon, 7 mi. N of Sedona, III-28-59, 1 male. Todd’s Lodge, Oak Creek Canyon, VII-16-61, 1 female. GILA COUNTY: Peterson Ranch, 7000’, Sierra Ancha, VII-2-60, 2 females. Tonto Creek Fish Hatchery, VI-28-61, 1 male. GRAHAM COUNTY: Wet Canyon, 6000’, Pinaleno Mts., [V-23-61, 2 females. GREENLEE COUNTY: Rt. 666, 8 mi. S of Hannigan Meadows, VII-5-59, 6 males. Grays Peak Road Camp, Rt. 666, VII-5-59, 5 males, 1 female. PIMA COUNTY: Madera Canyon, Santa Rita Mts., 4400-4600’, III-8-59, 1 male; III-19-60, 3 males. Summerhaven, 7800’, Santa Catalina Mts., V-24-59, 1 male; VII-9-61, 1 female. PINAL COUNTY: Peppersauce Wash, 5000’, Santa Catalina Mts., IV-18-61, 1 female. SANTA CRUZ COUNTY: Madera Canyon, Santa Rita Mts., 5600-6400’, III-29-59, 3 males; [V-9-59, 4 males, 28 females; [V-10-59, 1 male, 32 females; [V-3-60, 8 females; IV-9-60, 3 females; VII-6-60, 2 males, 7 females; VII-9-60, 4 females; VII-13-60, 2 females; IV-6-61, 12 females.
All of the known records of guaderna in Arizona are from the isolated mountain ranges in the southeastern part or from the Mogollon Rim area which extends in an irregular line from near Sedona eastward to McNary, then southeast to the vicinity of Glenwood, New Mexico. In addition to the counties previously mentioned it has been recorded by others from several localities in Yavapai County.
The flight peaks seem to fall during the first two weeks of April and again during late June and the first week of July. Captures range in elevation from 4400’ to 7800’ although quaderna was encountered most frequently from 5500’ to 7500’ in the oak and pine zones. The collections I have made indicate that females are more abundant than males by a 3:1 ratio. In the case of lta, females are also reported more frequently than the males. This unequal sex ratio may be only a result of incomplete sampling methods.
Erora quaderna is readily attracted to the flowers of Ceanothus fendleri Gray. I have also taken it on the flowers of Prunus virens (Woot. & Standl.), Nolina microcarpa Wats., and Monarda sp. It is not unusual during early April in Madera Canyon to see several hundred specimens during the course of a day. At that time of year they are easily taken at damp spots along the stream or in flight along the road and trails. The flight has been noted to move down the canyon unless a strong wind was blowing. :
Several present day workers consider quaderna may be conspecific with Jzta. I am not prepared to comment on this relationship since I have examined only two speciments of the latter. A key character that
4 RoEveErR: Erora Vol.16: no.1
has been cited for distinguishing males of these two species is the absence of a blue tornal patch on the dorsal surface of the hindwing in quaderna and its presence in leta. This is not a consistant character as I have five males of quaderna which exhibit a conspicious tornal patch.
I have not been successful in locating the foodplant of quaderna. The foodplants of lata have been listed as Fagus (beech) and Corylus (hazlenut). According to KEARNEY & PEEBLES (1960) no members of these genera occur in Arizona.
The southernmost record of lata that has been published was of a female taken along the Little River, 3000’, Great Smoky Mountains National Park, Sevier Co., Tennessee, [V-15-38 by ArrHur SrupKa, the park naturalist (Field, 1941). While checking collecting records at the park headquarters of the Great Smoky Mountains National Park I found reference to a specimen of leta taken VII-17-36, on Andrew's Bald, 5860’, G.S.M.N.P., Swain Co., North Carolina, SrmBperr & Evans, collectors. This is the first record of this species in North Carolina. It also represents an altitudinal high and the southernmost record for the United States.
ACKNOWLEDGEMENTS
I am grateful to Mr. Harotp Bropericx, park naturalist at the Big Bend National Park, Dr. P. J. Darxincton, Jr. of the Museum of Comparative Zoology, and Dr. Tarsicio EscaLANTE of Mexico City for the loan of specimens, to Mr. Lioyp M. Martin of the Los Angeles County Museum for information concerning specimens under his care, to Mr. ARTHUR STUpPKA, park naturalist of the Great Smoky Mountains National Park for permission to examine the park collection, and to Dr. FLroyp G. WerNeER of the University of Arizona for comments con- cerning the manuscript.
References
Clench, H. K., 1943. A note on the Arizona Erora. Journ. N. Y. ent. soc. 51: 221-223.
dos Passos, C. F., 1940. A new subspecies of Erora leta Edw. from Arizona and New Mexico. Amer. mus. novitates 1052: 1-2.
Field, W. D., 1941. Notes on Erora leta (Edwards) and Erora quaderna (Hewit- son). Annals ent. soc. America 34: 303-316, 3 pls.
Kearney, T. H., & R. H. Peebles, with supplement by J. T. Howell & Elizabeth McClintock, 1960. Arizona flora, 2nd ed. 1085 pp., 45 pls.
Department of Entomology, University of Arizona, Tucson, Ariz., U. S. A.
962 Journal of the Lepidopterists’ Society 5
THE GENETICS AND REPRODUCTIVE ISOLATING MECHANISMS OF THE PIERIS NAPI-BRYONIAZ, GROUP
by Z. Lorkovic
I. INTRODUCTION
The case of Pieris napi L. and bryoniz Ochsenheimer has wasted much ink in arguments among taxonomists as to whether bryoniz is a distinct species or only a subspecies of napi, without having brought about a satisfactory explanation. Even after the extensive work of MULLER and Kautz (1938), a turning-point in the opinions on this problem, the situation remained rather unchanged for a more critical or biologically trained judge, familiar to some extent with the circumstances in this difficult group. This failure is mainly due to the fact that most early investigators as well as MuLLER and Kautz themselves have not used the genetic approach. The research of B. PETERSEN (1948-1955) offered a first refreshing exception, and his important conclusions anticipate some of those reached in the present paper. Nevertheless a more exhaustive genetic analysis of the napi - bryoniz# complex remained necessary. In fact, at the same time as I was about to publish my genetic research obtained some years ago, several articles of BOwDEN appeared (1955, 1956, 1957, 1958) containing also some genetical and experimental data about the questions involved, though a conclusive genetic survey has not been given. Besides this, his results differ in some respects so much from my own that it will be by no means unnecessary to show here the results of my investigations in the usual genetic way.
THE ProsLteM. The main attractive point for the taxonomist lies in the great color differences between the females of these two forms as well as in the geographic restriction of bryoniz to the Alps, Carpathians, Scandinavia, and to some less known parts of Asia and North America. On the other hand, napi is widely distributed in both the plains and the mountains of the Palzarctic and Nearctic Regions but usually does not appear in the areas of bryonizx. Because of its restricted boreal - alpine distribution and the occurrence of more or less common transitions towards napi, bryoniz was in the past considered to be an ecological race of P. napi. However since MuLLER and Kautz have in their extensive work offered certain arguments in-favor of the specific distinctness of the two forms, their view has gained more and more adherents among levidopterists, so that finaly Forster and Woutrart in their work Die Schmetterlinge Mitteleuropas (1955) quoted bryonix as a good species,
6 Lorxovic: Pieris napi — bryonizx Vol.16: no.1
although the problem has not reached its conclusive stage. My use here of the term “adherents” suggests that we are not concerned with a systematically simple case, but rather with one that is in some respect ambigous. Such cases of controversial opinions have much interest from the evolutionary point of view, because this fact itself suggests that we are dealing with an incomplete stage of speciation, which deserves a closer analysis, especially from the genetical and populational aspect.
The present paper refers to my research concerning the genetics of the morphological characters of napi and bryonizx, to the occurrence of recombinations of these characters in nature, to the possible and ex- perimental hybridisation and to the degrees of reproductive isolating mechanisms between the two forms.
II]. Generics or CoLor DIFFERENCES BETWEEN P. bryoniz# anv P. napi
MATERIALS AND BREEDING METHODS. The crossings were performed with the ssp. neobryonixz Sheljuzhko that is abundant in the Alpine parts of western Yugoslavia and with napi from the lowland districts of the environments of Zagreb, where no bryoniz occur. Some crossings were also obtained with Alpine napi captured in districts where they fly together with bryonizw (Krnica Valley, Ratecée, Planica). During 1947 - 1951 about 200 broods were reared to imagos and more than twice as many matings have been carried out. Previously (1926, 1931, 1954; broods from wild females occasionally captured at the lower situated localities of bryoniz (Fala, Savinja valley, Krnica below Prisojnik ) served rather as an orientation for further decisive research.
The matings were obtained always under the experimentator’s control either inside the laboratory, mostly at the windows, or outdoors in the net, sometimes also free in the field. Many pairings could be realised only by the authors’ artificial copulatory method using clamped females and immobilised males (Lorkovié 1948, 1952). The mating usually lasted 1 to 1% hours. The egg-laying females were kept in cages provided with a cotton net and the eggs were deposited on the potted food plant Roripa silvestris, where the larvee remained until the last molt. Sub- sequently the larve were transported to plants kept in water. The plants must be changed every one or two days whether they are eaten by the larvee or not. By this way the losses caused by diseases were reduced practically to zero. The pupation occurred on the walls of the cage or’ in cardboard boxes, where the larve were placed after they had stopped feeding. Emerged male imagos usually were kept in cardboard boxes about 15 X 15 < 30 cm for two or three days at a lower temperature (15°
1962 Journal of the Lepidopterists’ Society i
- 18°C) wherefrom they were liberated once daily for some minutes for feeding. In such a manner the males can be kept in good condition before they become ripe to mate; this usually takes two or three days before one may with great probability count upon the mating ability of a male. On the contrary, the females are able to mate as soon as their wings grow strong enough for flying. Older females are more convenient only in the case when the artificial pairing methods must be used.
GENES AFFECTING THE WING COLOUR OF bryonix# AND napi. The first clear and doubtless assertion about the inheritance of the main bryoniz character, i. e. the dark melanic markings along the wing veins in bryoniz females (these are missing in napi) has been expressed in PETERSEN'S paper (1955), where on the strength of my unpublished research he reported that the bryonizw character is dominant over that of napi. A short note appeared previously to the paper mentioned in the Proceedings of the IX. International Entomological Congress (1952), where I de- clared in the discussion of HEssELBARTH' report that the bryoniz pattern is dominant, whereas the inheritance of the brownish yellow color is intermediary. All other previous opinions about this matter, so far as is known to me, were uncertain, which is no wonder, for without reared F» generation or back-crosses such a decision could not be reached. In fact, nearly all previous breeders, as well as some recent ones, suffered great losses caused by diseases, owing to the faulty breeding methods, so that the poor F2 broods were not sufficient for an accurate genetical analysis. One exception was Main’s crossing between 1907 and 1909, but in spite of a satisfactory breeding method he reached a quite erroneous conviction “that the inheritance of bryonizw characters passed almost entirely through the female” (quoted after BowpEn, 1956). After this failure it was not earlier than in 1956 that BowbeNn published the first successful research on this matter, and his results agree in general with my own. In respect to the lacks in the genetic interpretation of the cross- ings in his paper I shall give here a genetical analysis of my crossings in connection with the question of the reproductive isolation of these two butterflies.
The following crosses have been carried out by the author:
napi & X bryoniz BRroop 1. 1932: & Podsused (Zagreb); @ Rogovilec, Savinja valley, Karawanken Alps RCE 2. 1947: § Maksimir (Zagreb); 2 VrSic, Julian Alps, 1400 m. OOF 3. 1948: g Maksimir (Zagreb); @ VrSié, ex larva 1947 1 bn
8 Beye, Lorxovic: Pieris napi — bryoniz Vol.16: no.1
bryoniex 3 X napi g 4. 1935: g Mangart, Julian Alps; 2 Podsused (Zagreb) M x P 0. 1948: ¢ Vrsié, ex larva 1947; 9 Periénik, Vrata valley, Julian Alps, 900 m. 3 nb 6. 1948: ¢ Krnica, Julian Alps, 1100 m.; 2 Mt. Sljeme (Zagreb), 600 m. (27)3
Besides the listed crossings, several bryoniz broods from various localities of the Julian Alps and the Karawanken Alps in the Yugoslay part of the southeastern Alps have been carried out, i. e. from: Zirovnica, 700 m.; Fala near Maribor, 300 m.; Rogovilec in Savinja valley, 650 m.; Vrata valley in the Julian Alps, 800 m.; VrSié and Mojstrovka in the Julian Alps, 1400-1800 m.; and Bovec in the Trenta valley, 600 m. P. napi were mostly used directly from nature, since napi is the species of the environment of Zagreb, ranging eastwards to the line Maribor - Zidani most approximately, where the last small populations of a light bryoniz can be found.
The most important genetic results were obtained from the cross “IY” which could be brought up to the 14th generation without introduc- tion of any new bryoniz blood, but with 7 refreshments of outdoor napi.
The recent analysis of these crossings confirmed the author’s previously mentioned remarks about the inheritance of the main bryoniz# and napi characters. The three taxonomic distinctions between these two forms are controlled by three independent pairs or groups of genes: 1) An autosomal gene pair controls the extension of the dark MELANIC COLOR along the wing-veins; its dominant allele, B, causes the bryoniz# females to be heavily infuscated along the veins, whereas with the recessive allele, b, characteristic for napi, the darkening along the veins is either absent or reduced to the outer (distal) part of the veins (Fig.l 2 2 ). In the male sex the manifestation of these alleles is only slight, it is similar in both forms and therefore markedly sex-controlled. 2) The BRownIsH- YELLOW GROUND COLOR of bryonix# females, and the WHITE one of napi controlled by another pair or group of genes, Y, y, and its expression is strongly sex-controlled, since the males are completely white. 3) A third pair of alleles, W, w, affects the WHITE or GREENISH-YELLOW GROUND COLOR of the underside of the hindwings and the apex of the forewings, especially in males, and is thus partially sex-controlled.
We shall now turn to the more detailed analysis of each of the men- tioned gene-pairs.
1962 Journal of the Lepidopterists’ Society 9
Fig. 1. Melanic wing color pattern in females and males of bryoniw (left) and napi (right) which are controlled by the allele pair B and b.
Tue B, b attetes. The main controversy concerning the allelic pair Bb which controls the principal feature of bryonizw against napi is the question of its dominance. Because of the fact that the bryonixz x napi hybrids show a less pronounced bryoniz pattern than the pure bryoniz it was believed earlier that this character behaves intermediately. Such opinions rest on a misinterpretation of the meaning of dominance. This term does not denote full identity with the parental phenotype, respective- ly the homozygous one, since there are all transitions between the com- plete dominance to none at all. In our case, by pairing homozygous bryoniz and napi, the F; generation of both reciprocal crossings is always of the bryoniz pattern, although a somewhat slighter one. In the F»2 or any later generation, when individuals heterozygous for the genes Bb are mated, bryoniz and pure napi segregate in a 3 : I| ratio. In back- crosses the segregation is likewise clear, and the ratio is 1 : 1. In Fig.2 the succession of 14 generations obtained in the stock “III” is represented diagrammatically. The genotypes of each brood or of the parents as
10 Lorkovic: Pieris napi — bryoniz Vol.16: no.1
D ee MAKSIMIR cg @ vRsi¢ 4. 28, Vil. ge Q zi 2, 28.Vil MAKSIMIR ° *@ oh 7 40:14 BR DEIN o 2 OrKsAvER WAL ate Z/ill OKRNICA 17 e zal sa 1948 ane on oo ea) 2 Toe 2/iu 33:30 5. 4 Sane oslo hohe) oe C oo9 oo FOG 29 ; oe os ee 628M as Os ° e 2 e o tus ° oe cial Samet % 31.Vil. c of & Se of oe © eG o” 9 6879 8. 27 Vill co SP Ce LY el cm LO al ©) 2/illg Si O/in 4/\\l pees TESOL CSL CLASP ; 46 53 44:0 0:0
40. FV. °° A se o Sous
18:0 mesic 10:0 aon 41. QOVIl STENJEVEC @ & @ CP eocld @
A534 A4 450 is ! 35:0 4:0 9:0 3:2
42. 10VIL MAKSIMIRG! @ oo og @
185 Lae. MAKSIMIR[ 45:14 1 of
13 6vil oO ©
| 205 206 MAKSIMIR 17:20 12:12
44 5. 1X 5 AGED oH RCiS Gf 2 ore
Fig. 2. Diagrammatic representation of 14 successive generations in the stock “III” of the napi 8 x bryoniz @ crossing. During this time only napi were introduced from outside. The actual ratio of phenotypes is indicated for each brood; the expected genotypes are indicated by black (BB), black and white (Bb) and white (bb). Male and female symbols show approximately the given ratio. The bold black lines indicate broods in which all individuals were of bryoniz phenotype, consequently at least half of them were homozygous.
indicated in the diagram result from actual ratios obtained in the progeny of each brood as well as by the symbols of male and female phenotypes. The four possible ratios which occurred in 54 broods are summarized in Tables 1 — 3, each brood containing not less than 15 individuals; broods with a lower number of descendants are not included, except a few which are of special interest.
1962
Journal of the Lepidopterists’ Society
11
Table 1. Bb x Bb. Expected 3 : 1 ratio. Ratio
Bé bé Be bgQ BS Q Mx P — 1935 14 2 8 3 23 Ill — 1947 2 0 4 3 6 6/IIL, — 1948 i 0 6 3 13 2/4b — 1948 13 2% 6 4 19 9 — 1948 12 3 11 5 93 128 — 1949 10 2 y) Ht 12 201 — 1949 11 3 i) 5 33 214 — 1949 1p 5 (2 if 25 247 — 1950 12 4 11 3 23 265 — 1950 WS) 6 14 6 29 289 — 1950 21 8 4 i 25 301 — 1951 NS) 3 10 5 25 307 — 1951 aut 4 4 2, 15 5bn — 1948 3 2 3 3 6 ional: 1L30/ 45 Lu 3 ais} Q74 Expected ratio: 151.5 BOD | WO 2 3205 | 278
A. BB x BB, BB x Bb, BB x bb. Expected 1
Brood
46 - 1949 53 - 1949 73 - 1949 45 - 1949
127 - 1949
_ 134 - 1949
167 - 1949 210 - 1949 231 - 1950 237 - 1950 241 - 1950 271 - 1950
- 1950 315 - 1951
O ratio.
SOoQQgogqaoqoqoeoqcqe®
bse
i)
Babb se bp:
1 ratio.
Expected 0
Brood
Ratio
3/IIL.
14/2/4b
213 - 1949 235 - 1950 255 - 1950
SQOQoqoee©S
p2 Lorxovic: Pieris napi — bryoniz Vol.16: no.1
Table 3. @ Bb x. 2 bb, and 2 Bb x @ bb. Expected 1 : 1 ratio.
Rat 10 Sex of
Brood bb parent
je) Os
a
— =
RxP,-1932 RxF,-1932 1/III-1947 3/TII,-1948 6/II1,-1948 1/II1,-1948 4/TII,6-1948 bbn-1948 165-1949 185-1949 905-1949 206-1949 236-1950 239-1950 264-1950 278-1950 984-1950 312-1951 313-1951
bo
=" SCWeOAGDWWUNARHRROWNE =I
t
_—
oe)
eown & © ow
NS)
0.000 0.076 0.091 0.052 0.032 0.666 1,125 0,036
— jd Nw
=) Os OS OS +O +O OS OS OS O38 OS +O OS OS OS +40 40 OS O3 OS
— — LhOwWwMAWOWATHWWAR
3 9 8 6 3 1 6 i 3 5 6
AUDOADPRE OD
le
— Ol —
Expected ratio: : : : Oneal Total: 140: : : 0.50 >P<0.70
1The greater part of young larve was abandoned. 2Several napi females were accidentaly lost. 3bryonizw 8 ex larva from Vrsié, 1947.
The Chi-squared test and the probabilities (P) of the ratios 3 : 1 and 1 : 1 convincingly prove that the difference between the bryoniz and napi pattern depends on only one single pair of alleles and that the bryoniz pattern is dominant over that of napi. Accordingly, the napi individuals are always homozygous, bb, and consequently pure-breeding; bryoniz can be either homozygous or heterozygous. It can be seen from the diagram of Fig.2 that the mother of the stock “III” was heterozygous, since her progeny segregated in 6 bryoniz and 3 napi, a quite unexpected fact for a high-alpine bryonizx, usually considered to be true-breeding. It must be pointed out that in all these broods a clear-cut discontinuity between the bryonizw and napi patterns was absolute, although bryoniz vary considerably. The best but by no means the only decisive diagnostic character which allows the clear distinction between bearers of B and b- has been found in the so called “margin-streak” (“Saumstrich” of MuLLER
1962 Journal of the Lepidopterists’ Society Lis)
and Kautz or Bowpen’s “bryo-streak” ), i. e. the dark suffusion along the rudimentary vein A, extending from the posterior discal spot to the outer wing margin (Fig.l, ms). This streak is so rarely to be found in napi, that when it is present it is more than probable that it originates from a previous mixture with bryoniz. It is present nearly always not only in each similarly patterned subspecies or species as adalwinda Fruhst., bicolorata Petersen, pseudobryonixz Vrty., camtschadalis Rob., and ochsenheimeri Stgr., but in Pieris melete Mén. too. Spring napi females which show a more marked gray suffusion along the veins than is usual in this species are also distinguishable by the lack of this streak. The distinction between a slightly marked heterozygous bryoniz# female and a more strongly marked napi would be sometimes difficult without the bryo-streak present in the former and absent in the latter. In general, there is a good correlation between the vein markings and the bryo-streak in bryoniz, the streak being a more reliable diagnostic character inasmuch as it is a qualitative one. One exception to this rule will be considered below. It may be also emphasized that the occurrence of the bryo-streak does not depend upon the presence of the posterior discoidal spot, since the streak can be well developed even when the spot is completely miss- ing.
Despite the fact that napi were three times introduced from outside into the stock “III” during the first six generations (not counting the parental crossing), no particular diminution of the bryoniz pattern could be noticed. On the contrary the homozygous BB individuals which appeared first in the 10th generation were obviously darker and had broader vein markings than the mother. This is also proof that the mother was, in spite of her high-alpine origin, heterozygous.
In spite of the well-proven monohybrid inheritance of the bryoniz - napi pattern one could hardly explain the extreme range of variability of bryoniz by only a twofold nature of its genotype. Moreover, the same phenotype can be either homozygous or heterozygous; crosses of napi with heavily dark bryonixz females give rise to hybrids which do not differ from specimens usually considered as normal bryonix (brood 3/III-3), while on the other hand several stocks of homozygous BB individuals bred from heterozygous parents show a less pronounced bryoniz pattern than the just mentioned primary hybrids. In respect to the clear monohybrid segregation there can be little doubt that this very variable shading of the bryoniz pattern depends on polygenic factors independent of the B gene. This is most clearly shown by the observation that among the progeny of the two similarly heterozygous bryonix sisters but with different napi fathers (back-crosses), both phenotypes
14 Lorxovic: Pieris napi — bryoniz Vol.16: no.1
bryoniz and napi may be considerably different in the dark shading, depending on the intensity of the dark pattern of the napi father mated to the bryoniz mother (brood 278). If the napi father has a dark pattern, the entire progeny which segregates into bryoniz# and napi, will have a dark one, too. Similarly a paler napi father gives rise to brighter in- dividuals of both forms, bryoniz and napi (Fig. 3). Although the shading varies in both crosses, the bryoniz pattern is preserved, since even in the palest females the bryo-streak is fully developed. Such pale females can always be reliably distinguished from every napi individual. One can compare the differences described to more or less exposed or developed copies made from the same photograph negative.
Thus, the intensity of this bryoniz character depends not only on the B gene itself but also on the genes which control the appearence of melanin in general. Apparently we have here an analogy to the expression of the dark markings in the Nun Moth, Lymantria monacha, as it was postulated by GotpscHMipt (1921, 1928).
Apart from such multiple chromogene genes there are also other genes which control the extent of the dark suffusion along the veins. It is well known that this suffusion varies from very narrow lines, hardly broader than the veins themselves, to such an extent that the dusky suffusion of two neighboring veins joins together and the whole surface of the wings becomes dusted with dark scales. A good support for the opinion that this suffusion depends on separate genes has been obtained in a cross between a heterozygous ¢& bryoniz X napi anda ¢ Pieris ergane female, reared in 1935. The dark pattern of P. ergane is identical to that of P. napi or P. rape; but similarly to P. manni the spots have less-defined contours and the outer part of the forewings is often largely dusted with dark scales. In the female hybrids (bryonizw xX napi) & X ergane @ the dark markings along the veins are strikingly broader on both the fore- wings and the hindwings than was the case in the father’s (bryoniz xX napi) sisters, that show the width of the suffusion usual for bryoniz x napi hybrids. |
Consequently, the expression of the dark melanic pattern in bryoniz would be dependent upon three independent groups of alleles: 1) the dominant allede B controls the extension of melanin along the veins, and its recessive allele b causes the absence of melanin; 2) an unknown number of additively acting genes (M, m) control the intensity of the darkening; whereas 3) a probably low number of genes would be responsible for the width of the vein-marking.
Now, after having got acquainted with all the possible genes influenc- ing the dark pattern of bryonizw females, we again must turn to the question of the dominance of the gene B. Bowpen (1956) was right
1962 Journal of the Lepidopterists’ Society 15
in his doubt about the dominance of this gene so far as some mixed low- land bryoniz - napi populations are concerned. Instead of a clear-cut distinction between the presence and absence of the bryo-streak, as was found to be the rule in our experiments, here its variability produces all transitions ranging from a common dark streak to a few scarcely visible dark scales or perhaps none. The same seems to apply also to the dark vein markings, which can be as narrow as in some more strongly marked napi specimens of pure napi populations. In such cases one cannot de- cide whether such a transitional specimen carries only two b genes or also a B gene. In my last crossing experiments no such specimens were available, and the crossings carried out many years ago were not complete. However, by a more thorough genetical investigation of this character perhaps very interesting detections could be done. Apparently, two possible explanations may be taken into consideration: either there exist two or even more multiple B genes with different manifestation of the bryo-streak, or the recessive gene b of these populations is not identical with the gene b of the pure napi populations, bringing about a stronger vein suffusion than commonly found in napi, so that sometimes a slight bryo-streak appears too. Such a supposition could lead to the conception that the gene b in bryonizx populations is not derived from hybridization with napi populations, but that it takes part in the specific gene pool of Pieris bryoniz as a distinct entity. The confirmation of such a supposition might turn our opinion in favour of the conviction of those authors who consider bryoniz as a separate species. However, in the absence of experimental evidence for any of these hypotheses we must for the present emphasize the genetic proof that the napi pattern segregates from heterozygous bryoniz.
Finally, it should be remembered that the intensity of the melanic color depends also upon environmental factors, especially on temperature and humidity. It is known that low temperature and high humidity lead in Pieridz to spread of melanin, so that it is more than probable that the form “concolor” R6b. is at least partially due to these environ- mental influences. Pupz kept in the refrigerator at the time of develop- ment gave imagos with extremly wide vein suffusion, while conversely, heat and dryness gave rise to pale and bright individuals.
Everything said so far applies only to the female sex. We must, however, also take into consideration the male sex, although it is not so important for our purposes, since here the phenotypic differences of the genes in question are only slight. Careless observers or novices mostly think that there are no differences between the males of bryoniz and napi, which may be true only exceptionally for the spring brood. In the
16 Lorxovic: Pieris napi — bryoniz Vol.16: no.1
summer broods bryoniz males are distinguished by gray streaks along the veins on the upperside, particularly on the hindwings, where these markings gradually diminish from the margin of the wing to half way toward the discocellular vein. Rarely the darkening is of such a width that the neighboring markings join near to the wing margin. Usually the markings are developed only as narrow “vein-streaks” as shown in fig. 1, left. In napi these gray vein-streaks are either absent cr are present as minute dark triangles at the ends of the veins (fig.1, right). In the spring brood the triangles can be prolonged about one millimeter or two at most; this is the only case when males of bryoniz cannot be distinguished with certainty from the napi ones.
The vein-streaks are present in every bryoniz male which carries at least one B gene, i. e. not only in homozygous but in heterozygous in- dividuals too. This circumstance is of great value in the establishment of more precise ratios of phenotypes in segregation, because it is possible to determine the presence or absence of the gene B in almost every brood in males as well as females, so that the numbers usable in deter- mining ratios are twice as large. The phenotypic differences of this character correspond in almost every case very well to the expected ratios of 3: 1 or 1 : 1 (see Tables 1 and 3).
Of course, the stronger vein darking in bryoniz males is present in the forewings as well as the hindwings, although in these the difference from napi is not so apparent, because the apical spot always sends shorter or longer vein-streaks towards the inside of the wing.
The demonstration that gene B, which controls the dark pattern of bryoniz, is dominant over the recessive napi gene, so that pure napi traits segregate from heterozygous bryoniz, is of decisive significance. It reveals that napi must be always homozygous recessive, bb, while phenotypic bryoniz can be either homozygous, BB, or heterozygous, Bb (Table 2). Mutter and Kaurz (1938) considered specimens with bryoniz pattern as pure bryoniz or as its ssp. flavescens because they did not know this. They also did not know or underestimated the fact that from heterozygous bryonix, although very dark, specimens of the napi phenotype segregate which do not differ from individuals of a pure napi population. MULLER even quoted from the literature some cases of the appearence of napi specimens within bryonizx broods, but he explained them as individuals accidentaly introduced with the food (l.c.: pp.36, 37). Corresponding examples in their monograph are: Tab.6, fig.15; tab.5, fig.11; but also 6, 7, and 8, except for their yellow color. It is true that Mutter and Kautz count such napi individuals as bryonix ssp. flavesc2ns Wagner, but there is no proof whether they belong to this subspecies or
1962 Journal of the Lepidopterists’ Society 17
rather are descendants of a cross with napi. It is only certain that such napi individuals belong to the population of Médling, which is composed of both flavescens and napi. Whether they belong to napi or flavescens cannot be resolved without breedings or crossings, as will be shown below. We shall learn also that bryonizw Xx napi heterozygotes are some- times present but concealed by the dominance of the gene B even in the so called “pure” mountain bryonizx populations.
THe Y y ALLELES. It was mentioned above that the factor for the brownish-yellow ground color of the bryonizw females is dominant, al- though it is not sure that it is really so, since this color shows graduated intensity in both the crossings and in nature. Owing to its strong sex- controlled manifestation it is not easy to decide whether this character depends upon one or more pairs of alleles. Since the males are entirely white, without any trace of a yellowish color, the choice of males in crossings is entirely by chance when pure stocks are not at one’s disposal.
The back-cross (R X P - 1932) between an intense yellow bryonix Bb female from Rogovilec (see below) and a pure napi male from Zagreb, where no yellow specimens occurred, seem to be a rather decisive one: 28 yellow and 30 white females appeared, the yellow specimens being re- presented by all transitions from nearly the same yellow color as the mother had to the very pale one; all white females had the same white color. This cross indicates that the gene for the yellow color is likely to be incompletely dominant, Y, its expression being variable, and that there is no more than one allelic pair for this color, a rather unexpected result, since such a gradual color variability usually is attributed to polygeny. However, the characters which are dependent on the cumulative action of polygeny can by no means result in a ratio of 1 : 1, providing, of course, that the estimation of color was correct.
The genetics of this character thus seems to remain somewhat obscure, too.
THE W, w parr. Especially interesting is the allele W which controls the white color of the underside of the hindwings and the apex of the forewings in the males of bryoniz populations. It is completely dominant over the greenish-yellow color of the recessive gene w. The same applies to the female sex, although the gene w is manifested only exceptional by the white color, mostly by a pale buff one. The phenotypic manifesta- tion of the recessive allele, w, is always the same in males and in females, in bryoniz and in napi. W is a remarkable gene in that male sex-controlled polymorphism is very rare in Lepidoptera, as pointed out by REMINGTON
18 Lorxovic: Pieris napi — bryoniz Vol.16: no.1
Table 4. Ww xX ww. Expected 1 : 1 ratio.
Brood |W ¢4 WWI WE Oe AO OW aes pane x? | P in % 1 bn 13 12 ili 17 30 : 29 aa S00 5 bn 2 3 p) 4 Ab toi 0.8182 | >30, <50 Ab a 1 11 6 18 13 8.8064 | >30, <50 4/4b 4 0 3 5 ui 5 0:3333 | S50) <70 2b 6 6 3 3 9 9 0.0000 | 100
24b 13 PD) 5 5 18 i 4.8400; >2, <5 14-2/4b 2 10 py) 1 4 11 3.2666| >5, <10 1 bbn 4 0 1 1 5 1 2.6666 | >10, <20 3 bbn 4 5 2 3 6 8 0.2857 | >50, <70 6/4b 3 0 PD) 3 5 3 0.5000 | >30, <50 5(27) 5 5 a = 5 5 0.0000| 100
55 9 9 9 10 18 19 0.0270| >80, <90 85 0 3 4 3 4 6 0.4000} >50, <70 60 2 iL ) 0 4 1 0.9000} >10, <20 Total Awe GS 63 61 sy BY os 0.6475: So, 50)
(1954). Although we do not deal here with a quite true sex-controlled dimorphism, since the difference appears in both sexes, the dimorphism is much more pronounced in the males than in the females, the latter being sometimes difficult to distinguish in respect to it.
It is worth nothing that the genetical identity of the white males and pale buff females has not been realized before the present analysis. Each of these two forms has been proviously considered to be a separate variant: ab. & “subtalba” Schima and ab. @2 “subtochracea” Kautz. Mutter and Kautz reported that “subtalba” can also rarely be found in males and females of napi, and they quote the find of a “subtalba” female in Pommerania in northern Germany. It is obvious that the white color of this female has nothing to do with the gene W, as was already suspected by Mutter (1938).
Another interesting point of this gene pair is that the author has not as yet been able to obtain a brood homozygous for the gene W. As is shown in Table 4, the segregation is the rule and the ratio 1 : 1 prevails. Of the two 3 : 1 ratios, broods 2/4b and 1/bbn, the first is only apparent- ly of this kind since the brood was raised from a back-cross, the mother of the second brood unfortunately escaped before her phenotype was noted. It is striking that 8 of 11 matings between Ww individuals were entirely sterile, whereas 3 others gave a total of only 5 individuals, 3 of them in one brood. Only the back-crosses were successful, and even
1962 Journal of the Lepidopterists’ Society 19
among these a great part produced no progeny. The writer is inclined to attribute this failure to inbreeding rather than to any other essential cause, except the F, hybrid sterility, which will be discussed below. No population of bryoniz is known in which the gene W is the exclusive allele. One of the most significant concentrations of this gene so far known to me occurs in the Julian Alps, where about 45% of all pheno- types are W bearers. According to Perer’s (1950) count the proportion of the “subtalba” males in the Allgauer Alps is 19%. No white males had been recorded in the western Alps, as some authors have claimed.
RECOMBINATION IN REARED POPULATIONS. The three gene pairs con- trolling the morphological features of bryoniz and napi, already dis- cussed above, have been found to segregate and completely recombine in the F2 generation and back-crosses. The dihybrid back-cross “R x P - 1932” already mentioned is especially illustrative. It yielded four female phenotypes: bryoniz pattern and yellow color (Bb Yy), bryoniz pattern and white color (Bb yy), napi pattern and yellow color (bb Yy), and napi pattern and white color (bbyy) in the ratio 14 : 14 : 16 : 14, which agrees very well with the expected dihybrid ratio 1 : 1 : 1 : 1. Besides these two gene groups free combinations with the Ww alleles have also been obtained, among them the formerly scantily-known combinations bYW and byW. Thus the three pairs of alleles are located in different chromosomes, all autosomes. This is only to be expected where the number of chromosomes is so large (n = 25). The recombinations are the same as certain “varieties” or “aberrations” from natural collections, already de- scribed, and it seems likely that the genetical explanation of these natural variants has therefore been found. I shall mention some examples of such natural “aberrations”: bryoniz# “obscura albida” (BB yy), bryonix “albida” Miill. (Bbyy), bryoniz “flavida reducta” Miill. (bDbYY or bbYy), and bryoniz “albida reducta” Miill. (bb yy). This shows at best how unsound it may be to give names to forms which are no more than re- combinations of a few gene pairs.
However, before starting with the comparison of the experimental genetic results and the circumstances in nature we must briefly get acquainted with the other morphological as well as physiological and ecological differences between bryonixz and napi.
(To be continued )
20 Vol.16: no.1
A BITING MIDGE ECTOPARASITIC ON ARIZONA LYCAENIDS
by Pau R. ExRiIcH
On 21 August 1961, I was using a Questar telescope to observe in- dividuals of Celastrina argiolus (=Lycznopsis pseudargiolus) sucking moisture from damp spots in the road running through the South Fork of Cave Creek (Chiricahua Mts., Cochise Co., Arizona, 5200 feet). Two small flies were seen to alight on the ventral surface of the hindwing of an individual under observation, and when the butterfly was carefully collected with forceps one stayed in place, while the other flew off. The fly remained attached while the butterfly was killed in a cyanide jar, and it was therefore possible to obtain the accompanying photographs of the parasite in situ.
Dr. W. W. Wirtn, of the U. S. Department of Agriculture, has identified the fly as Forcipomyia (Neoforcipomyia) baueri Wirth, a member of the dipterous family Ceratopogonide (—Heleidz) which is well known to most of us through the activities of the members which attack man (known variously as punkies, no-see-ums, sand flies, all-jaws, etc.). Forcipomyia baueri was described from a series of specimens taken by D. L. Bauer from the underside of the wings of Callophrys ( Mitoura) siva and Philotes enoptes on Mingus Mountain, Yavapai Co., Arizona. The only other record which has come to my attention of a Forcipomyia attacking a North American butterfly is that of F. mexicana Wirth attacking Pyrrhogyra otolais Bates (Nymphalide: Nymphaline) in Mexico (collected by BAvER). WirtuH's (1956) excellent paper on biting midges ectoparasitic on insects shows exotic ceratopogonids to be quite catholic in their tastes, with attacks recorded on Pieris (Pieride, Pierine), Eurema (Pieride, Coliadine), Danaus (Nymphalide, Danaine), Morpho (Nymphalide, Morphine) and _ Helicopis (Lycaenide, Riodinine ).
Ceratopogonids also attack larval butterflies, adult and larval moths, and a wide variety of other insects. In all cases they are presumably “bloodsuckers” (ingest the hemolymph). Many aspects of the relation- ship between the ectoparasitic midges and lepidopterans deserve study; it would be especially interesting to know the degree of host specificity displayed by the midges. Lepidopterists are urged to make detailed observations on any observed attacks, and to submit the flies to Dr. WietH for identification.
1962 Journal of the Lepidopterists’ Society PAL
Top: Forcipomyia baueri on ventral surface of right hindwing of Celastrina argiolus. Middle and bottom: close-up views of the midge. [Photos by M. A. Mor- tenson. ]
22 Enruicu: Biting midge Vol.16: no.1
I would like to thank Dr. W. W. Wirtu for identifying the midge, and Mr. Martin A. Mortenson for taking the photographs. This work was done while the author was at the Southwestern Research Station of The American Museum of Natural History, and was supported by Grant No. G-14740 from the National Science Foundation.
Reference
Wirth, Willis W., 1956. New species and records of biting midges ectoparasitic on insects (Diptera, Heleide). Ann. ent. soc. Amer. 49: 356-364.
Div. of Systematic Biology, Stanford University, Stanford, Calif., U. S. A.
GUIA PARA EL ENTOMOLOGO PRINCIPIANTE. [Guide for the beginning entomologist; in Spanish.] By Kenneth J. Hayward. 1961. 168 pp., 7 pls. Instituto Miguel Lillo, Universidad Nacional de Tucu- man. Argentina, Miscelanea, No. 22. Available from the Instituto for $3.00 USA.
This is a manual of entomological procedure aimed at beginners in the study of insects. These are discussions of the morphology and classification, of collecting techniques, and of the orders of insects. Professor Haywarp recognizes 34 orders. This is a larger number than seems ideal in the light of recent work, and especially good grounds exist for reducing to infra-ordinal status his Phasmida, Diploglossata, Megaloptera, and Raphidiodea. Probably the Isoptera should stand next to the Blattaria, and in the Apterygota the Collembola should be listed first (or better eliminated from the Insecta), Protura second, Entotrophi third, and Thysanura last; Thysanura are by far the closest to the Pterygota, and Collembola and Protura by far the most remote. But these old classificatory points are common to many current general texts, and the work under review cannot be singled out for them. Spanish-speaking insect collectors needing a beginner's guide will find this work valuable.
CHARLES L. REMINGTON
Dept. of Biology, Yale University, New Haven, Conn., U. S. A.
1962 Journal of the Lepidopterists’ Society 23
ECOLOGICAL STUDIES OF RHOPALOCERA IN A HIGH SIERRAN COMMUNITY — DONNER PASS, CALIFORNIA. I. BUTTERFLY ASSOCIATIONS AND DISTRIBUTIONAL FACTORS
by Tuomas C. Emmet and Joun F. EMMEL
The State of California, extending some 1,264 miles along the Pacitic Coast of North America, contains 158,693 square miles of dry deserts, moist coastlands, chaparral-covered hills, and great mountain ranges, with unparalleled opportunities for the ecologist and lepidopterist alike. Practically every combination of climatic factors may be found within the state’s boundaries. Six life zones, with approximately 4000 species of fern-allies and flowering plants, lie at elevations ranging from 282 feet below sea level to 14,495 feet at the peak of Mount Whitney. Tempera- tures and rainfall correspond generally with these life zones; average annual rainfall ranges from about 0 inches to 70 inches, and mean temperatures of about 40°F. in boreal regions to high summer tempera- tures of 130° in the deserts are recorded. Thus California provides many unique locations to study the ecological aspects of a species’ adaptation and preference for a particular environment.
One such location—the Donner Pass area in the Sierra Nevada, Placer County, California — was selected for intensive study during the summers of 1956 and 1960. This location encompasses the flora of three life zones and ranges in elevation from 6800 to 8300 feet. This paper reports the general ecological observations made on each of the 76 re- corded species, forms and races of Rhopalocera and the four habitats selected for study in the area. Factors involved in butterfly distribution are discussed using examples from this Donner Pass study and the literature.
INTRODUCTION
In this Donner Pass study and past investigations by other authors, two general factors in the ecology of Rhopalocera have been emphasized:
(1) Preference of many species for a particular type of habitat. (2) Effect of weather (extrinsic environmental conditions) on butterfly flight periods. .
A second paper will treat the latter subject. The present paper reports our data on the general ecology of the Donner Pass butterfly fauna.
24 EMMEL & EmMMEL: Donner Pass Vol.16: no<4
E. B. Forp (1957), in considering the geographical distributions and ecological preferences of British butterflies, notes that it is often difficult to separate these two distinctions. Many species will normally occur within a particular region and this region, though usually composed of varying habitats, is known as part of the general geographic distribu- tion of each resident or migratory species. However, within that general region, the ecological preferences of each species dictate the distribution of that butterfly.
As yet, few published papers have been devoted entirely to the ecology of butterflies in a particular locality. TitpEN (1959) has done compre- hensive studies on the butterfly fauna of Tioga Pass, California. He arranges the 43 species in groups based on their apparent abundance in various sub-alpine and alpine plant associations. Certain habits of alpine butterflies are noted. TILDEN concludes that range is usually de- termined by the presence of food plants, although food plants of some of these alpine species are presently unknown. In this author’s estimation, the occurrence of the adult butterflies is less effected by particular en- vironmental conditions than is that of the plants.
Many authors have noted habitat preferences for single species or groups of Rhopalocera. MuNROE (1951) in reporting collecting conditions in northern Quebec observed a seasonal succession and an altitudinal zonation of the various resident species. The flight season was so short that the time of appearance of species differed in days instead of weeks “as in more temperate climates.” Different Boloria and Plebeius species each had specific habitats, such as “rocky hilltops” and “only on grassy beds of dried-up lakes at from 2200 to 2400 feet.” Euriicn (1954; 1956) treats the Erebia in northwestern America from an ecological standpoint. Brown, Err and Rotcer discuss the ecological preferences of over 200 species in their book Colorado Butterflies (1957). Brown (1952) also discusses the restricted ecological preferences of Oeneis oslari Skinner in the South Park area of Colorado; the type of soil and terrain preferred is apparently directly related to the presence of the food-plant grass.
As in Brown’s paper (1952), many authors have attempted to give possible explanations for the basis of these “ecological preferences” of a species. Probably the most acceptable explanation for limiting factors in the distribution of resident species is the presence of food plants. The “hilltop” controversy of a few years ago brought out possible reasons for congregations of butterflies. SHoumatorr (1953) summarizes past authors’ hypotheses into two groups: ecological explanations, such as food-plant search, wind, and tropism, and intrinsic factors, such as “liking hilltops,” “social ambition,” and “gregariousness.” ARNHOLD (1952)
1962 Journal of the Lepidopterists’ Society DE:
notes that Euchloe olympia prefers to fly along the leeward side of ridges (near Dresbach, Minnesota); a narrow band of calm is created on the top or side of the banks according to wind direction. Thus wind, as well as food plant, appears to definitely be-a factor in the local habitat preference of a species. aastii
Since a species generally is very restricted to one group or species of food plant, it follows that the geographic range of the butterfly will correspond to the range of the required plants. However, climatic factors within the range may prevent the butterfly from existing with its food plant. Since butterflies are poikilothermic (“cold-blooded”) and _ there- fore dependent on solar heat for their activity, species cannot live where temperature and solar radiation do not meet their tolerances (Hovanitz, 1958).
EHRuicH (1956) lists climatic factors that could limit Erebia distri- bution in habitats of northwestern America, such as temperature fluctua- tions, amount of cold, snow cover, rapidity of runoff, and amount of spring flooding of habitat. The influence of any of these factors on butterfly distribution over a large territory would be the subject of a very interesting paper, but that would require an intensive investigation over many years.
The contribution of this paper towards studies of the latter type is to report information on the local habitat preferences of butterflies in a single locality. Perhaps some day we will have enough information to provide a generalized survey of climatic and vegetational factors, and their influence on butterfly distribution, for the entire Sierra Nevada range. However, towards the end of this paper, we have discussed the apparent reasons for habitat preference in the Donner Pass Rhopalocera, and these conclusions may prove to be a step on the way to full under- standing of the factors involved in butterfly distribution.
HABITATS OF THE DONNER Pass AREA
The Donner Pass area is located on the crest of the Sierra Nevada range at an elevation of 6800 to 8383 feet, 15 miles northwest of Lake Tahoe, in Placer County, California. Flora and fauna of both the east and west slopes of the Sierra meet along this crest. Adding to the desirability of the study area is the union of three life zones here — Transition, Canadian, and Hudsonian — with Canadian zone vegetation comprising most of the territory. Four habitats within the Donner Pass - area were studied: wet meadow, dry meadow, forest, and montane.
96 EMMEL & EMMEL: Donner Pass Vol.16: no.1
Fig. 1. The Donner Pass area, looking N.N.E. from Mt. Disney (7,953’). In center of photograph is Sugar Bowl Lodge, with the dry and wet portions of the Lodge meadow at lower center and at left. Lake Mary is at upper center; long railroad snowsheds and Highway 40 are at upper left. The actual Pass is slightly to the left of upper center.
DONNER PASS AREA PLACER COUNTY | CALIFORNIA
“V4 2 SCALE OF INSET MILE WVan Norden
Sketch map of the State of California showing the locality studied, and topo- graphical map (inset) of the Donner Pass area showing the habitats (numbered 1A through 4D) described in the text. Primarily from Geological Survey map, Norden Quadrangle, California, 7.5 Minute Series, U. S. Dept. of the Interior (1955).
1962 Journal of the Lepidopterists’ Society
bo ~l
1. WET MEAapow
A. SuGAR BOWL AND SoutH YuBA RIVER
The Sugar Bowl is a small bowl-shaped valley (elevation 7260 feet) formed by the confrontation of Mt. Disney, Mt. Lincoln, and the high ridge connecting the two peaks. A heavy snow pack on these slopes, coupled with scattered springs, provides a considerable flow of water into this area. Here also is the source of the South Yuba River, which flows into Lake Van Norden. Thus there is an abundance of plants due to the moisture and rich soil. Willows (Salix) are found in scattered thickets in the Sugar Bowl and along most of the length of the river. Grasses (Poa species) and sedges (Carex) are common throughout the area. Other common plants are Cow Parsnip (Heracleum lanatum), Sierra Sweet Cicely (Osmorrhiza occidentalis), Scarlet Gilia (Gilia aggregata), Common Monkey Flower (Mimulus guttatus), and Elephant Heads ( Pedicularis groenlandica).
B. Wet PortTIon oF LopcGE MrEApow
This area (elevation 7000 feet) is located at the base of Mt. Disney, where numerous springs provide a very wet environment. About one-half of it is covered with willows. In the other portions one finds a large number of grasses and flowers. Some of the more common plants are Indian Paint Brush (Castilleia miniata), Common Monkey Flower (Mimulus guttatus), Columbine (Aquilegia truncata), Larkspur (Delphinium pauciflorum), Tiger Lily (Lilium pardalinum), and Meadow Rue (Thalictrum fendleri).
2. Dry MEADOw
A. LopcE MEapOw AND ADJACENT PORTIONS
This flat area near the Sugar Bowl Lodge supports many flowering plants despite its dryness in July and August. Fireweed (Epilobium angustifolium), Common Yarrow (Achillea millefolium), and Pussy Paws (Calyptridium umbellatum) are common. Elevation: 6960 to 7000 feet.
B. EmicRANT MEADOW
Emigrant Meadow (elevation 7500 to 7600 feet) is a dry flat expanse with few plants; two species are Yellow-bud Penstemon (Penstemon letus) and Saxifraga zxstivalis.
28 EMMEL & EMMEL: Donner Pass Vol.16: no.l
C. SUMMIT VALLEY
Summit Valley (elevation 6800 feet) is a large flat meadow which is marshy in the spring, but dry throughout July and August. Grasses dominate most of the area but in some places Pussy Paws, Yarrow, and Pepper Grass (Lepidium perfoliatum) can be found.
3. FOREST
Dense forest covers about one-half of the total study area, particularly between 6800- and 7600-foot elevations. This is almost all Canadian zone forest. Some of the forest trees and plants near Lake Mary and Lake Van Norden are typical Transition zone inhabitants, and the higher montane forest is typical of the lower Hudsonian zone. The majority of the forest trees are Red Fir (Abies magnifica), Lodgepole Pine (Pinus contorta), Silver Pine (Pinus monticola), and Mountain Hemlock (Tsuga mer- tensiana ).
On the forest floor a great variety of plants are found. Collecting was usually good in small clearings or near the forest edge; many butter- fly species were found beneath an aerial transportation system (used in winter), which cuts a swath through the forest from Highway 40 to Sugar Bowl Lodge. Some of the forest plants are the Spotted Coral Root Orchid (Corallorrhiza maculata), Fireweed, Corn Lily (Veratrum californicum), Yarrow, Pine Drops (Pterospora andromedea), and Alpine Shooting Star (Dodecatheon alpinum).
4, MONTANE
A. Mount JupaH
Mount Judah (8,243’) appears barren due to the large granite boulders scattered over its talus slopes. The dry, almost soilless environment dis- courages most trees. The flora includes Alum Root (Heuchera micrantha), Rabbit Brush (Chrysothamnus nauseosus), Sage Brush (Artemisia tridentata), and Mule-Ears (Wyethia mollis).
B. Rocky SLOPE LEADING TO MOUNT JupaH
Seemingly a jumble of small granite rocks, this area (elevation 7160 to 7400 feet) supports many plants; included in this flora are scattered Lodgepole Pines (P. contorta), Wyethia mollis, Lupinus, and Sedum species.
1962 Journal of the Lepidopterists’ Society 29
C. Mount LINCOLN
Mount Lincoln (8,383') is quite similar to Mount Judah. One minor difference is the presence of Mountain Hemlock (Tsuga mertensiana).
D. Mount DIsNEY AND Crow’s NEST
This twin-peaked mountain is part of a long ridge and supports a flora similar to the other peaks, although Mt. Disney (7,953’) has more species. Found here are Red Fir, Lobb’s Buckwheat (Eriogonum lobbii), Mountain Alder (Alnus tenuifolia), Chrysopsis brewerii, and Western Pennyroyal (Monardella odoratissima).
Loca DistTRIBUTION AND ECOLOGICAL PREFERENCES OF SPECIES RECORDED
The phrase “flight period” refers herein to the first and last records for the species. Observations were made from June 17 to August 26 (1960, unless otherwise noted); obviously, some species were already flying before the first day of observation. Nomenclature of species follows EuruicH (196la). Subspecific names are used for clarity with certain polymorphic western species, such as in Speyeria.
A. PAPILIONIDA&
1. Papilio zelicaon Luc. Flight period: June 17-August 6. Uncommon; males found typically on peaks of area (bare rocks) and never observed visiting flowers. Females on lower slopes of Mt. Judah, flying around food plant (Cymopterus terebinthinus) and ovipositing single eggs (ob- served fresh female depositing eggs, around 10 a.m. on June 27); one female seen visiting Western Pennyroyal (Monardella odoratissima).
2. Papilio indra indra Reak. Flight period: June 17-July 10. Males occasionally seen around 3 peaks of area and lower slopes of Mt. Judah (landed on rocks and opened wings to “sun” themselves ). Males frequent- ly seen at muddy places. Females occasionally (June 27, June 30) seen ovipositing single eggs on Cymopterus terebinthinus. L. M. Martin has found several hundred indra larve on this Cymopterus at Kaiser Peak (10,000'), Fresno Co., Calif.
3. Papilio rutulus Luc. Flight period: June 17- July 15. Males occasionally seen around willows and in forest; one female observed laying eggs on willow in Summit Valley.
4. Papilio eurymedon Luc. Flight period: June 17- July 24. Males common in wooded areas; females on lower slopes of Mt. Judah.
30 EMMEL & EMMEL: Donner Pass Vol.16: no.1
Reported food plants available: Ceanothus velutinus and prostratus, and Prunus emarginata.
5. Parnassius clodius baldur Edw. Flight period: June 27-July 25. Found almost everywhere, but particularly abundant on the lower slopes of Mt. Judah (females laying eggs on or near Sedum in this location).
B. PIERIDA:
1. Neophasia menapia F. & F. Flight period: August 8-August 15. Only on lower slopes of Mt. Judah and Lake Van Norden area; males flying around upper tips of Pinus contorta; no females seen.
2. Pieris sisymbrii Bdv. Flight period: June 20 - June 28. Lower slopes of Mt. Judah, but only seven adults observed during summer (1960). Species of Cruciferze (recorded food plants) present in area.
3. Pieris protodice L. Flight period: June 23-August 25. Most common in dry meadows; known food plants (Cruciferz), such as mustard species, were present; double-brooded.
4. Pieris rape L. Flight period: June 18- August 14. Dry and wet meadows; reported food plants (Crucifere ) abundant.
5. Euchloe creusa hyantis Edw. Flight period: June 21- July 13. Only on lower slopes of Mt. Judah; food plants (Cruciferz ) present.
6. Anthocaris sara form julia Edw. Flight period: June 18 - July 16. Lower slopes of Mt. Judah and open wooded slopes; rarely seen in meadows. Food plants (mustard species) present; several mature larve found in mid-June (1956). Males identical to typical white reakirtii were occasionally taken in fresh condition. A. julia is a yellow- tinted, high-altitude form of sara, while the lowland, spring form of sara is reakirtii.
7. Colias eurytheme Bdv. Flight period: June 17 - August 26. Found in all habitats, but most abundant in Summit Valley meadows. Females seen ovipositing on clover (Trifolium).
8. Colias eurytheme form amphidusa Bdv. Flight period: July 1- August 26. Appeared in early July and by August this strongly-marked form was dominant over typical eurytheme.
9. Colias philodice eriphyle Edw. Flight period: (July 1). Only 3 found — in dry meadows of Summit Valley.
C. DANAIDZ&
1. Danaus plexippus L. Flight period: June 23 - August 25. No milk- weed species in area and individuals seen were likely strays from lower elevations.
1962 Journal of the Lepidopterists’ Society 31
D. SATYRIDA:
1. Caenonympha tullia californica Westwood. Flight period: (late June). Seen only in 1956 (wet winter) in late June on Mt. Judah (grassy areas). Flight period in 1960 (dry winter) may have finished by fumes i.
2. Cercyonis sthenele oetus Bdvy. Flight period: July 11 - August 20. Found only on high western slope (above 8000 feet) and around peak of Mt. Judah; landing on sagebrush stems (Artemisia tridentata). Food plant likely to be grasses, which-were abundant on mountain. Adults (both sexes) visited Western Pennyroyal blooms.
E. NYMPHALIDA®
1. Speyeria cybele leto Behr. Flight period: July 15- August 3. Five fresh specimens seen on lower slopes (6,980') of Mt. Disney, visiting Western Pennyroyal flowers. Possible food plants (Viola species) for leto and the following Speyeria species were ‘present in wet meadows.
2. Speyeria zerene zerene Bdv. Flight period: August 2 - August 25. Uncommon; both sexes visited Western Pennyroyal on lower slopes (6,980) of Mt. Disney. |
3. Speyeria coronis snyderi Skin. Flight period: July 3- July 15. Females proportionately more common than males; flying around Western Pennyroyal on Mt. Disney (6,980').
4. Speyeria atlantis irene Bdv. Flight period: July 12- August 26. Both sexes almost as abundant as S. mormonia arge; adults visited Western Pennyroyal. Females occasionally seen in wooded areas.
5. Speyeria mormonia arge Stkr. Flight period: June 29 - August 26. The most abundant Speyeria and found throughout the Pass area; adults avidly visited Western Pennyroyal.
6. Boloria epithore Edw. Flight period: June 17- July 22. Very abundant in forest and around Pine-mat Manzanita (Arctostaphylos nevadensis ). Food plant is reported as Viola (violets ).
7. Chlosyne palla Bdv. Flight period: (July 7). Only 1 worn male seen. Food plants (Castilleia and Aster species are reported) abundant in area. As palla is recorded from higher elevations than Donner Pass, some unknown climatic factor may prevent its establishment, or possibly the species has not extended its range to this particular area yet.
8. Chlosyne hoffmanni hoffmanni Behr. Flight period: June 25-August 6. Found throughout the area; very abundant where its food plant (Chrysopsis breweri) grows, especially on open slopes of Mt. Disney and Mt. Judah. Mature larve fairly common at end of June; females
32 EmMMEL & EMMEL: Donner Pass Vol.16: no.1
laid egg masses, each containing 30 to 110 eggs. These eggs hatched in approximately 3 weeks, and larve grew quite slowly, staying together in communal webs. By last week in August, every larva observed had assumed a lethargic state. Larve were still in webs, though plants were dying.
9. Phyciodes campestris montana Behr. Flight period: June 18- August 25. Practically as abundant as C. hoffmanni; found most com- monly where Aster species (recorded food) were growing in meadows. Females were observed hovering around Aster integrifolius.
10. Phyciodes mylitta Edw. Flight period: June 20-August 5. Scarce in area, although its reported food plant, thistle (Cirsium), was fairly abundant along Lake Mary road (dry location).
ll. Polygonia zephyrus Edw. Flight period: June 20-August 18. Over-wintering individuals collected in late June and early July; fresh males collected in early August on Monardella odoratissima. Food plant (Ribes viscosissimum — Sticky Currant) common in Lodge meadow. Mature larve found by second week of July.
12. Nymphalis californica Bdv. Flight period: June 17-August 20. Over-wintering individuals numerous during late June. Swarms of larve found on Ceanothus in first two weeks of July on peak of Mt. Disney (lone location). About July 15, individuals began to hatch and on July 21, the species reached its peak of abundance with an estimated 5,000 seen in one hour in a 100’ x 100’ area on Mt. Judah. On this date, the butterflies began flying in a westerly direction at about 10 a.m. On July 22, population counts were about “normal” again.
13. Nymphalis milberti Godt. Flight period: June 17-August 6. Apparently a stray species in this area. Adults visited Western Pennyroyal. Urtica, the usual food for milberti and also atalanta, was not growing in the Pass area; it is abundant along streams at nearby, lower elevations.
14. Nymphalis antiopa L. Flight pericd: June 19-August 15. Over- wintering individuals seen during late June and early July. Fresh speci- mens seen during late July and early August. Food plant: willows.
15. Vanessa cardui L. Flight pericd: June 22-August 16. Seen around rocky peaks of the 3 mountains. Food plant: thistle.
16. Vanessa atalanta L. Flight period: June 22-July 21. Only several males seen, flying around peaks with cardui. No food plant in area.
17. Vanessa virginiensis Dru. Flight period: June 18-August 25. Also found around peaks with cardui. Larvee were found during July on Gnaphalium palustre (Cudweed ); this food plant grew only in one clump at the base of Mt. Disney.
18. Vanessa carye Hbn. Flight period: June 18-August 19. Found frequently throughout the area. Food plant probably Lupinus here.
1962 Journal of the Lepidopterists’ Society oo
19. Precis lavinia Cramer. Flight period: June 18-August 18. Males common in dry meadows; females common in wet meadows, and oc- casionally seen hovering over Plantago (Mimulus also in the area).
20. Limenitis lorquini Bdv. & Lec. Flight period: June 23-August 10. Frequently seen in willows (food plant) along the South Yuba River.
F. LYCAENID/:
1. Satyrium californica Edw. Flight period: July 20-August 10. A distinct inhabitant of the area along the Lake Mary road. Adults flew around and landed on Eriogonum nudum blooms. Reported food plant is Quercus; Huckleberry Oak (Quercus vaccinifolia) was found in this area.
2. Satyrium sylvinus Bdv. Flight period: July 18-August 26. Found most commonly around willows (recorded food plant) on the lower slopes of Mt. Judah; adults landed on Eriogonum nudum flowers.
3. Satyrium sepium Bdv. Flight period: July 20-August 16. Found only along Lake Mary road; females observed ovipositing on Ceanothus velutinus. Adults landed on Eriogonum nudum and_ Elderberry (Sambucus velutina).
4. Satyrium behrii Edw. Flight period: (August 4). One found in 1960; fairly abundant in 1956 on rocky slopes of Mt. Judah. Adults landed on Eriogonum blooms. Probable food plant: Lupinus species.
5. Satyrium fuliginosa Edw. Flight period: July 10-August 9. Fairly common on lower slopes of Mt. Judah. Females seen ovipositing on lupines (food plant).
6. Strymon melinus Hbn. Flight period: July 7-August 15. Found flying around Eriogonum blooms; fairly abundant. Reported food plant: Polygonum alpinum (Knotweed ), present in several areas.
7. Callophrys johnsoni Skin. Flight period: (June 28). Probably a resident of the area, but extremely restricted colony. Female observed flying around Arceuthobium (a mistletoe and recorded food plant) on Mountain Hemlock at lower end of Sugar Bowl.
8. Callophrys nelsoni Bdv. Found rarely in 1956; foodplant (believed to be Incense Cedar — Libocedrus) not found in Pass area.
9. Callophrys augustinus iroides Bdy. Flight period: June 28 - July. Found rarely in late June and July on rocky slopes; adults fed on Eriogonum. Food plant: Ceanothus and Sedum reported by authors.
10. Callophrys eryphon Bdv. Flight period: June 28- July 10. Only oecasionally captured (on semi-forested, talus slope) but all fresh specimens. Food plant: probably Pinus contorta.
34 EmMMEL & EMMEL: Donner Pass Vol.16: no.1
11. Callophrys dumetorum perplexa B. & Benj. Flight period: June 22-July 9. Occasionally found in the high montane areas (above 7,500’). Food plant probably an Eriogonum species.
12. Lycena arota virginiensis Edw. Flight period: July 20-August 11. Found only along the Lake Mary road; fairly abundant. Males hatched in mid-July, females appeared in early August. Females were ovipositing on Ribes montigenum (Gooseberry) during first two weeks of August (eggs hatch the following spring).
13. Lyczna editha Mead. Flight period: July 15-August 19. Fairly common throughout the entire area, especially in dry meadows. A possible food plant (Potentilla drummondii) occurs in the Lodge meadow.
14. Lycena nivalis Bdv. Flight period: July 3-August 17. Rather scarce; males more abundant than females. Found in dry meadows. Food plant is unknown.
15. Lyczena cupreus Edw. Flight period: June 17-July 15. Found in dry meadows. Adults frequently visited flowers of Calyptridium um- bellatum (Pussy Paws). In early July, mature larve were found feeding on the blossoms of this plant; being pink in color, they blended well with the flowers.
16. Lyczna heteronea Bdv. Flight period: August 6-August 16. One small colony found near a tiny stream (elevation 7160’) on Mt. Judah. Observed food plant was Eriogonum nudum, and the adults frequently visited the flowers of this buckwheat.
17. Everes comyntas amyntula Bdv. Flight period: June 17-July 1 One colony found on the top of Mt. Judah (8,234'). Here the species’ observed food plant, Astragalus whitneyii (Milkvetch or “locoweed’ ) grew on top of the narrow ridge connecting the two peaks of Mt. Judah. A constant wind blew; beth sexes landed on the plants and flew up only occasionally.
18. Plebejus anna Edw. Flight period: June 26-August 19. Found throughout the area but common only on the moist lower slope of Mt. Disney. Food plant unknown in spite of the species abundance.
19. Plebejus seepiolus Bdy. Flight period: June 19-August 17. Found only in wet portions of Lodge meadow and Summit Valley (very abundant). Females (all rufescens form) oviposited on Trifolium hybridum (Alsike Clover) throughout late June and July. 7
20. Plebejus icarioides Bdv. Flight period: June 22-August 3. Found throughout the area; especially common in areas with wide-spread patches of Lupinus (food plant)—usually on rccky talus slopes.
1962 Journal of the Lepidopterists’ Society 35
21. Plebejus shasta Edw. Flight period: June 30-August 18. Found throughout the area, and very common in dry meadows on mountain slopes. Some adults seen landing on Monardella odoratissima blooms. Food plant is unknown.
22. Plebejus acmon West. & Hew. Flight period: June 18-August 15. Found only in Lodge meadow in late June and along Lake Mary road in early August. Females oviposited on Eriogonum nudum (food plant).
23. Plebejus acmon lupini Bdy. Flight period: June 25-July 25. Found only in Sugar Bowl and on Mt. Judah. Food plant probably Eriogonum. TILDEN (1959) suggests this insect may be a species distinct from acmon.
24. Agriades glandon podarce F. & F. Flight period: June 27-August 3. A small colony found in wet portion of Lodge meadow; fairly abundant here. Food plant perhaps Vaccinium nivictum (Sierra Bilberry), found in same area. :
25. Glaucopsyche lygdamus behrii Edw. Flight period: June 17-July 16. Found throughout the area in late June, but most common on the moist lower slope of Mt. Disney. Food plant here probably Lupinus.
26. Philotes enoptes Bdv. Flight period: June 26-July 13. Males found landing on moist ground and flying on lower slopes of Mt. Judah. One female seen ovipositing on Eriogonum nudum.
27. Philotes battoides intermedia B. & McD. Flight period: June 25- July 18. Found on moist stream banks on lower slope of Mt. Judah. Several females seen around Eriogonum nudum.
28. Celastrina argiolus echo Edw. Flight period: June 17-July 15. Fairly common in all habitats. Food plant probably among Ceanothus, Vaccinium, Spirza, and Actinomeris species (all recorded food plants ) found in the pass area.
G. HESPERIIDZ:
1. Thorybes nevada Scud. Flight period: June 17-July 13. Abundant in dry and wet meadows. Food plant unknown.
2. Pyrgus ruralis Bdy. Flight period: June 17-July 11. Found in every habitat. Food plant probably Potentilla drummondii (larvee known to feed on P. tenuiloba elsewhere).
3. Pyrgus communis Grt. Flight period: June 17-August 19. Abundant in every habitat. Particularly found around muddy places. Females seen ovipositing on Sidalcea glaucescens (Mallow Family) and larve found all summer.
36 EMMEL & EMMEL: Donner Pass Vol.16: no.1
4. Erynnis jucenalis Fabr. Flight period: June 19-July 25. Males collected on the 3 peaks and occasionally elsewhere; no females seen. Food plant unknown for this area.
0. Erynnis afranius Lint. Flight period: June 17-August 5. Males collected on the three peaks; 1 female collected in Lodge meadow. Food plant possibly columbine (food plant of the related E. lucilius)— Aquilegia grew in the meadows.
6. Hesperia juba Scud. Flight period: June 17-July 1. Occasionally seen in both dry and wet meadows.
7. Hesperia nevada Scud. Flight period: June 22-July 10. A large thriving colony was found at the very peak of Mt. Judah. Males appeared in late June and females appeared in early July. Females were observed Ovipositing on a species of grass.
8. Hesperia harpalus Edw. Flight period: (August 8-August 11) A small colony was found on the lower slope (7,240') of Mount Judah. Two females were seen ovipositing on a grass species, and other females were observed hovering over the grasses in this small area.
9. Polites sonora Scud. Flight period: June 23-August 25. Found in every habitat but most common in dry meadows. Food plant: likely grasses, as in other Hesperiine.
10. Polites sabuleti tecumseh Grin. Flight period: June 19-August 19. Distributed in every habitat but particularly common in the dry meadows (especially in Summit Valley). Most abundant in late June and early July, though the species was collected throughout the summez.
H. PossisLe— FururE ADDITIONS TO THE BUTTERFLY FAUNA OF DONNER PASS
Papilio brucei Edw. could be established in the area and overlooked in 1956 and 1960. Its food plant on Kaiser Peak (10,000 to 10,300 feet; Fresno County, California) reported by L. M. Martin to be Cymopterus terebinthinus, which was present on Mt. Judah (8,243’).
Colias eurytheme form autumnalis Cockerell might have been flying right after snowmelt in early June.
Euphydryas chalcedona macglashanii Rivers has its type locality as “Truckee” (10 miles east of the Pass) and Comstock (1927) notes it as occurring in the mountains north of Lake Tahoe in late June and early July.
Euphydryas chalcedona sierra Wright is common in the Lake Tahoe region, 15 miles from Donner Pass, and has been collected at Gold Lake, 30 miles north of the Pass.
1962 Journal of the Lepidopterists’ Society 37
Phezdrotes piasus Bdv. is recorded from similar elevations in northern California and its food plant (Lupinus) is common in Donner Pass.
Ochlodes sylvanoides Bdv. was common in Squaw Valley (August 11, 1960), 10 miles southeast of the Pass. Apparently-similar habitats are available in the Lake Van Norden meadows.
Speyeria callippe juba Bdv. was found at Emigrant Gap, 15 miles west of the Pass area. Other callippe forms have been reported from the Lake Tahoe and Gold Lake regions.
Table I. DISTRIBUTION OF RHOPALOCERA SPECIES IN THE DONNER PASS AREA.
Species Habitat
A. PAPILIONIDZZ oe 2AG 2B Er 2D en s3 4A Papilio zelicaon Be see ee a ear IRD eet ge TOK Papilio indra cae Papilio rutulus Xe es Papilio eurymedon Xe ae Parnassius clodius TEMP ONG ng yep y ee Se os | OK
B. PIERIDZ Neophasia menapia es Pieris sisymbrii at Pieris protodice x Pieris rape xX Euchloe creusa = Anthocaris sara x Colias eurytheme xX Colias philodice a,
C. DANAID Danaus plexippus IX, We Oe OE ONS FING te XW ON) OM
D. SATYRID#= Canonympha tullia Ua ae Ste ee ees Ppa a ee) aks Pe ee Cercyonis sthenele RN ele eat len ALLL NG | NLL || se )Xereh hu seal gE
E. NYMPHALID Speyeria cybele Speyeria zerene Speyeria coronis Speyeria atlantis Speyeria mormonia Boloria epithore Chlosyne palla Chlosyne hoffmanni Phyciodes campestris Phyciodes mylitta
PA rs |
rs
mA | | PS PS PS PS
PA A OK |
A |
LAX Mel mm | bd bd |
bd bd bd | bd pd pa | dd | | D4pd] dopardl pal | ey pedo
| dod] pede od lb | bd bd bd | bd bd Dt PS Dd |
\
38 EMMEL & EmMMEL: Donner Pass Vol.16: no.1
| | 1 Ly | pepe
Polygonia zephyrus xX Nymphalis californica x Nymphalis milberti X Nymphalis antiopa x Vanessa cardui = Vanessa atalanta = Vanessa virginiensis _ Vanessa carye
Precis lavinia
Limenitis lorquini
F, LYCANIDE Satyrium californica Satyrium sylvinus Satyrium sepium Satyrium behrii Satyrium fuliginosa Strymon melinus Callophrys johnsoni Callophrys nelsoni Callophrys augustinus Callophrys eryphon Callophrys dumetorum Lycena arota Lycezna_ editha Lycena nivalis Lyczna cupreus Lycezna heteronea Everes comyntas Plebejus anna Plebejus szpiolus Plebejus icarioides Plebejus shasta Plebejus acmon Plebejus acmon lupini Agriades glandon Glaucopsyche lygdamus Philotes enoptes Philotes battoides Celastrina argiolus
G. HESPERIID/EZ Thorybes nevada Pyrgus_ ruralis Pyrgus communis Erynnis juvenalis ss Erynnis afranius — Hesperia juba = Hesperia nevada —_ Hesperia harpalus = Polites sonora xX Polites sabuleti c
re mr
bd > PS PS ~ xX
x Xx xX
rs PS OM | PA PS PX PS PS |
XS ex
| MK | | MrMrs] | | | pe | | Krone] |
xX |
| | | PAM bd bd bd be | |
| Parts) pd] Pt napa] P11 1 1 I beep be oe | bd bd | | rarspal | Pal P| pd] | pepe dd oe | a] Los] Pal | pl | | P| bday) | Od Pane saaie eal
| 54 | I | |
PPI Ps od PPS | | | pa bd] Pd nad od | Pd bd og] pa Pd! | be pede Parr dl PII Dede] Dd] Dd LT Lode] dd] Deb] Dede] be be Dd |
am
| PA PA bd bd be |
MAM MM
PM PM 4
PSP PS PS Od
Pad | bd bd bd bd bet bet be
wx | ~ A | ~~ | MM 4 | ~~ ~
1962 Journal of the Lepidopterists’ Society 39
DISCUSSION
From this study several factors important in determining the distribu- tion of a species within a local habitat have emerged. The explanations for the “ecological preferences” of the butterflies in the Donner Pass area (and presumably elsewhere ) fall ideally into the following categories.
I. Larvat Foop PLANT
As pointed out in the introduction, the specific larval food plant required by most species limits the distribution of the butterfly to areas where the food plant grows. But within the geographical range of the food plant, two quite different distributions of the plant are possible: (1) a “closed” range, where plant and butterfly are confined to very restricted areas within a habitat; (2) an “open” range, where the plant and butterfly are widespread in a given habitat; or, alternately, an “open” range where the food plant is widespread but the butterfly is confined in distribution.
A. CLosED RANGE
In some butterfly species, such as Euphydryas editha in the San Francisco Bay region, the adults seem to “choose” to remain within a very specific area despite the fact that its food plant is available in a somewhat greater area. This apparent “choice” factor EHriicH (1961b) terms “intrinsic barriers” to dispersal.
In the Donner Pass area also, some species were found with apparent intrinsic barriers to dispersal. Everes comyntas amyntula and Hesperia nevada were restricted to the very top of Mt. Judah, and each species was always found within a few feet of its respective food plant, which was also restricted to the ridge (8,240') here. With a constant, strong wind and the weak flight of Everes, any dispersal away from the food plant would seem to be dangerous for the survival of the species there. Hesperia nevada seemed just to “choose” to remain in this territory.
Examples from lower elevations also come out of the Lycenidz and Hesperiidae. Certain lycenids (Lycena heteronea and Satyrium cali- fornica, for example ) and skippers, such as Hesperia harpalus, were found only in the immediate vicinity of food-plant colonies. Such restricted plants are found only in areas with particular soil and moisture re- quirements. Willows, the larval food plant of Limenitis lorquini, are found only in wet areas and Limenitis stays among the willows, despite its strong flight. One might hypothesize that a specifically de- veloped physiology (weak flight and a single food plant with specific
40 EMMEL & EMMEL: Donner Pass Vol.16: no.1
soil and moisture requirements) or an intrinsic “choice” factor (plus a single food plant) has caused these butterfly species to be restricted even within a habitat the size of those studied (refer to map).
B. Oren RANGE
In this category we include food plants that enjoy a wide distribution throughout one or more habitats, and that have their corresponding butterfly species occurring with them. Examples are numerous: Pieris protodice, Anthocaris, Colias, Chlosyne hoffmanni, Precis lavinia, Boloria epithore, Plebejus szpiolus, and many others.
C. OrpEN Foop PLANT RANGE BUT RESTRICTED BUTTERFLY DISTRIBUTION
This is a very common circumstance in temperate North America. The food plant will occur over a wide region, yet the butterfly species will be distributed in only part of the habitats available to it. (This, of course, is assuming the same elevation and general climate prevails throughout these habitats.) The following explanations are offered:
(1) The species is a weak flier and is only increasing its range very slowly. EnriicH (1954) postulates that species of this nature (e.g. Erebia)may have wind as an important dispersal agent.
(2) The species may be a fairly strong flier but it never strays far from its its colony in a particular habitat. EHriicu (1961b) has offered an intrinsic barrier explanation for this phenomena.
(3) The species (a weak or only fair flier) exists in a habitat surrounded by an almost insurmountable barrier, such as mountains, a forested area, or even a wide river. BAUER (1959) notes that the south- ward spread of Erebia vidleri Elwes and Boloria chariclea Schneider from Washington into Oregon has been barred by the Columbia River gorge.
(4) To the casual cbserver, habitats may appear to be suitable be- cause of similar climate and elevation, but microclimates in the area may preclude any settling of the butterfly species. Besides severe cold in the winter, there may be flooding from spring runoff, lack of sufficient solar radiation due to frequent cloudiness or northerly exposure, an abundance of parasites and predators, feeding heavily on butterflies of the area (single strays from elsewhere would be destroyed. too frequently for permanent settlement), and extreme temperature fluctua- tions within the immediate area of the food plants (bare rocks with sun reflection, for instance).
1yez Journal of the Lepidopterists’ Society Al
Three examples of this “open but restricted” phenomena in the Donner Pass area are offered from the hairstreak group. Satyrium sylvinus was found around willows at the base of Mt. Judah, but in Summit Valley (same elevation—about 6800 feet) the species was absent despite the abundance of willows. Food plants for Callophrys eryphon and C. augustinus iroides were quite numerous and widely distributed, but these hairstreaks were found only rarely in both 1956 and 1960.
II. Aputt FEEDING PLANT
This distributional factor has not been noted by most authors, but in the Donner Pass area, the adult feeding habits of some butterfly species dictated the apparent distribution of the imagos. The most obvious examples of this intrinsic ecological factor were the members of the Speyeria group. The fritillary species were very rarely found in the moist areas where the larval food plant (Viola species) grows (and then only females). This is the complete opposite of the first case considered (I-A) where certain species were extremely restricted to the food-plant area. Instead, Speyeria occasionally would be seen flying very rapidly through dry meadows or on rocky slopes where no larval food plants were known to occur. We are indebted to L. M. Martin for pointing out that single-egg-laying species, such as Speyeria, often must fly a considerable distance between each oviposition. Usually, the adult Speyeria congregated around flowering Western Pennyroyal to feed during most of the day.
Thus preferences of the adult butterfly for a certain feeding plant can be very important in determining the distribution of some species within a habitat that may not even contain the larval food plant!
Il]. Stronc Furers or INTER-HABITAT MIGRANTS
Examples in this group are familiar to every North American lepidopterist: Vanessa cardui, V. atalanta, V. virginiensis, Nymphalis californica, Danaus plexippus, and Polygonia zephyrus. Members of this category often fly many miles and one or more thousand feet in elevation away from the nearest food plants.
Vanessa cardui and V. atalanta were found only at elevations over 8,200 feet, while thistles (food plant for cardui) grew to around 6,800 feet and no food plants were found for atalanta. No larval food plant for Danaus plexippus was available for many miles around, but in- dividuals of this renowned migrant were seen rather frequently above
7,000 feet.
42 EMMEL & EmMMEL: Donner Pass Vol.16: no.1
For two species (Nymphalis californica and Polygonia zephyrus), the larval food plant was found only in one locality, while the adults flew in every habitat. The strong flight of these species has granted considerable freedom of movement to the adults, and thus the butterflies often appear far from their larval food plants. Yet this “migrating” tendency does not imperil the survival of the species, for the adults hatch close to their food plants and can mate and lay eggs before flying off, or else can find new food-plant sources in their movements.
IV. Mountrain-Tor ATTRACTION
No paper of this type would be complete without entering the “mountain-top controversy. We have already mentioned (III) the Vanessa species flying about the peaks. There were two other strong fliers in the study area that were restricted mainly to the highest peaks: Papilio zelicaon and P. indra. Elsewhere in California, P. zelicaon is found wherever a suitable food plant (Umbellifere) grows. On the Donner Pass peaks, the available food plant for both zelicaon and indra was Cymopterus and this plant’s range included the tops of the peaks down to 7,100 feet. For these strong fliers, the winds along the high ridges and peaks may prove enticing, if one concedes that the species can indeed “choose” to remain there. On the other hand, the closeness of their food plant would make the adults’ appearance around the peaks seem but natural.
CONCLUSIONS
Using the ecological data obtained from our research in the Donner Pass area, we have found first that the distribution of butterflies in a Sierran community is apparently governed by the inherent physiology of each species; in other words, by intrinsic factors. This physiology may be dependent upon one or more of the following factors: (1) the larval food plant; (2) the adult feeding plant; (3) the development of wing muscles for flight, i.e., how strongly or weakly the butterfly flies; (4) other unknown intrinsic factors, such as the preference of some species for mountain-tops and the lack of any dispersal tendency, as in Satyrium sylvinus.
Secondly, we have found that the distribution of butterflies depends upon various extrinsic factors which work on the intrinsic factors of each species. Among extrinsic factors we have noted climate, food-plant distribution, elevation above sea level, natural barriers to dispersal, wind, and presence of parasites and predators.
1962 Journal of the Lepidopterists’ Society 43
Thus in the search for explanations of butterfly distribution we must consider all the above factors, for each factor has its examples and many butterflies are not easily categorized. The real basis for the distribution and “ecological preferences” of a species must be sought in the evolution and adaptation of physiological mechanisms for exploiting a particular habitat in nature.
SUMMARY
The local distributions, ecological preferences, and flight periods of 76 species, forms, and races of Rhopalocera in the Donner Pass area, Placer County, California, U. S. A., are reported. Factors involved in determining butterfly distribution are discussed. It is concluded that the distribution of a butterfly species is determined by (1) intrinsic factors, including larval food plant, adult feeding plant, strength of flight, and unknown factors, such as intrinsic barriers to dispersal; and (2) extrinsic factors, including climate, food-plant distribution, elevation above sea level, natural barriers to dispersal, and presence of parasites and predators. These extrinsic conditions work on intrinsic physiological factors, which have evolved for exploitation of a particular environment, and so determine the geographical range and local dis-
tribution of a species.
ACKNOWLEDGMENTS
The authors are deeply indebted to Mr. Witx1am N. Goopatt of the National Audubon Society, Director of the Audubon Camp of California at Donner Pass (Norden, California), where full cooperation for collecting data was extended to us during our service as assistants on the staff. Drs. THomas Harvey and KENNETH TANKSLEY, members of the Camp staff, kindly furnished plant identifications. Thanks are due to Mr. Lroyp M. Martin, Los Angeles County Museum, who helped in determination of some Rhopalocera species and read the manuscript.
References
Arnhold, F. R., 1952. Notes on collecting Anthocharis midea and Euchloe olympia. Lepid. news 6: 99-100.
Bauer, David L., 1959. A new geographical subspecies of Chlosyne hoffmanni (Nymphalide) from Washington State. Journ. lepid. soc. 13: 207-211. Brown, F. Martin, 1952. Oeneis oslari Skinner, rediscovered (Lepid.: Nymphalidz ).
Ent. news 53: 119-123. - hee , D. Eff, & B. Rotger, 1957. Colorado butterflies. Denver: Denver
Museum of Natural History. 368 pp. Comstock, John A., 1927. Butterflies of California. Los Angeles, California. 334
pp.
44 EMMEL & EMMEL: Donner Pass Vol.16: no.1
Ehrlich Paul R., 1954. Notes on Erebia rossii Curtis (Lepidoptera: Satyride). Ent. news 5d: 225-297.
ee ee , 1956. Ecological observations on Erebia (Lepidoptera: Satyrida) in northwestern America. Ent. news 67: 29-36.
Btn. een , & A. H., 196la. How to know the butterflies. Dubuque, Iowa; Brown. 262 pp.
Pee rere , 1961b. Intrinsic barriers to dispersal in checkerspot butterfly. Science 134: 108-109.
Ford, E. B., 1957. Butterflies. London: Collins. 368 pp.
Hovanitz, William, 1958. Distribution of butterflies in the New World, in Hubbs, Carl L. (ed.), Zoogeography: 321-368. Publication No.51 of the American Association for the Advancement of Science. 509 pp.
Jepson, Willis L., 1957. A manual of the flowering plants of California. Berkeley: Univ. of Calif. Press. 1238 pp.
Munroe, Eugene, 1951. Field notes on the butterflies of Knob Lake, northern Quebec. Lepid. news 5: 7.
Munz, Phillip A., 1959. A California flora. Berkeley: Univ. of Calif. Press. 1681 pp.
Shoumatoff, Nicholas, 1953. The excelsior complex. Lepid. news 7: 38-40.
Tilden, J. W.. 1959. The butterfly associations of Tioga Pass. Wasmann journ. biol. 17: 249-271.
Wiltshire, E. P., 1957. The Lepidoptera of Iraq. London: Nicholas Kaye Limited. 162 pp:
(TCE) Reed College, Portland 2, Oreg. U. S. A. (JFE) 5341 West Blvd., Los Angeles 43, Calif., U. S. A.
SATYRIUM BEHRII (LYCAANIDA) IN OREGON
Prompted by the recent record of Satyrium behrii Edwards from Nevada (Philip, Journ. lepid. soc. 15: 56; 1961) the Reverend A. I. Goon, of Wooster, Ohio, wrote to inform me of his capture of the species in Oregon, another of the states whence behrii was previously unreported (cf. Clench, in Ehrlich & Ehrlich, How to know the butterflies: 192; 1961).
Rev. Goon took a single fresh male on 5 July 1961 at John Day (Grant Co.), the only individual he saw in nearly a month of collecting in the area. He has generously presented the specimen to Carnegie Museum.
E. J. Newcomer, of Yakima, Washington, has also reported in corres- pondence the capture of S. behrii in Oregon. He took it at two localities: Camp Sherman (Jefferson Co.), 22 July; and Pringle Falls (Deschutes Co.), 23 July. There is also a record of the species from Ft. Klamath (Klamath Co.) in the Museum of Comparative Zoology at Harvard.
Harry K. CLencu, Carnegie Museum, Pittsburgh 13, Penna., U. S. A.
1962 Journal of the Lepidopterists’ Society 45
THE AUTHORSHIP OF THREE SCIENTIFIC NAMES OF NEARCTIC RHOPALOCERA VARIOUSLY CREDITED TO BOISDUVAL OR LUCAS
by Cyr F. pos Passos
In a recent paper by Dr. LincoLn P. Brower (“1958” [1959]: p.101) it is stated in a footnote “I have been unable to ascertain whether BorspuvaL or Lucas first described P. eurymedon and P. rutulus in ¥852.”
This problem actually involves an additional name to those mentioned above i.e., Papilio zelicaon. The authorship of these three names proposed in 1852 has been shifted back and forth over the years between BortspuvaL and Lucas and the time has come when this uncertainty should be put finally to rest.
All three names were published by BortspuvaL (1852a) in a paper entitled Lépidoptéres de la Californie and by Lucas (1852) in a paper entitled Descriptions de nouvelles espéces de Lépidoptéres appartenant aux collections entomologiques du Musée de Paris. BotspuvaL appears to have presented his paper on the Lepidoptera of California at a meeting of the Entomological Society of France on 25 February, but it does not appear to have been published until August. These facts appear from a separate (1852b) of the paper and a reprint (1852c) bearing new pagination (pp.1-52), both of which are in the author's library. The latter paper has a title page dated 1852 and states that it is an “Extrait des Annales de la Société entomologique de France (aoft 1852)”. On the other hand, Lucas’ paper has at the top of every even numbered page “(Mars 1852)”. Hence, on the intrinsic evidence all three names must be ascribed to Lucas.
_ The foregoing evidence appears to settle the matter insofar as intrinsic evidence is concerned and it would prove difficult to upset that con- clusion by any extrinsic evidence, if it exists.
- It should be observed that Brower, in his references, ascribes the Lucas paper to “M. H. Lucas”. The full name of Lucas is PIERRE Hierotyte Lucas, but he usually signed himself H. Lucas. The “M.” stands of course for Monsieur. It was customary at that time in France to use the abbreviation M. before authors’ names. Also the pagination of Lucas paper is 128-141, not 138-141 as Brower cites it. ©
46 pos Passos: Papilio names Vol.16: no.1l
The foregoing study was made some time ago at the suggestion of Mr. Pappy B. McHenry of Burbank, California, who inquired about the matter while the present author was preparing his Check List of Nearctic Rhopalocera and I am indebted to him for calling the problem to my attention.
References
Brower, Lincoln P., “1958” [1959]. Peale’s Lepidoptera Americana and the correct name for Papilio multicaudatus. Lepid. news 12: 101-102.
Boisduval, Jean Baptiste Alphonse Déchaufour de, 1852a. Lépidoptéres de la Californie. Ann. soc. ent. France (2e série) 10: 275-324.
Boia , 1852b. Lépidoptéres de la Californie. ibid. 10: 275-324. [Separate.]
Pema teas , 1852c. Lépidoptéres de la Californie. 52 pp. Typographie Felix Malteste et Ce, Paris. [ Reprint. ]
Lucas, Pierre Hippolyte, 1852. Descriptions de nouvelles espéces de Lépidopteres appartenant aux collections entomologiques du Musée de Paris. Rev. mag. zool. pure appl. (2e serie) 4: 128-141.
Washington Corners, Mendham, N. J., U. S. A.
THE FIRST RECORD OF COLIAS NASTES IN THE UNITED STATES (PIERIDA*)
A single female of Colias nastes Boisduval was collected in northern Washington while the author was working on a forest lookout tower. This specimen was collected on July 13. A single male of this species was sighted five days later but eluded capture. The female agrees with HoLuanp’s figure of the species in every way except that the yellow coloring has a more greenish undercasting. The previously recorded range of this species is Labrador northward and westward (Klots, Field guide to the butterflies: p.33; 1951) and British Columbia. The exact locality of capture follows: Bunker Hill Lookout, near edge of a northern facing cliff, 7000 feet elevation, (T 40 N, R 19 E, S 16), Okanogan County, Washington, July 18, 1961, Jon SHEparp collector. Both the male and female were discovered in association with Red Heather (Phyllodoce empetriformis (Smith) D. Don.). The specimen is in the authors collection.
Jon H. Sueparp, 2315 Jackson, Corvallis, Ore., U. S. A.
1962 Journal of the Lepidopterists’ Society AT
NOTES ON NEOTROPICAL LEPIDOPTERA. 1, THE EARLY STAGES AND COMPARATIVE MORPHOLOGY OF TWO SPECIES OF DYOPS (NOCTUIDA‘) HITHERTO CONFUSED
by E. P. WILtsHIrE
Perhaps the most peculiar trees of the Neotropical forests, though not the most beautiful or majestic, are those of the genus Cecropia, known in Brazil as Imbauba; in English it is sometimes referred to as the Chandelier-tree, from its aspect, or the Ant-tree, from its myrmecophily.
During an all-too-brief stay of 18 months at Rio de Janeiro (see my general paper — Journ. lepid. soc. 13: 79-88; 1959), I decided to pay especial attention to the Lepidoptera and other insects attached to this genus of tree, and in a later article I hope to deal with the relations between various orders of insects on it. In this, my first special article on the Lepidoptera of the New World I shall deal only with two species of noctuid moth which I obtained in the forests fringing Rio only by breeding larve found on Cecropia. I suppose them to be monophagous on this genus, but of course cannot be sure.
The appearance of both these species of larva is of some taxonomic interest, as it differs from that hitherto considered to characterise the Quadrifid subfamilies of the Noctuidae. These subfamilies are primarily classified by their hindwing neuration, but it is considered that they are also characterised by the less complete development of the larval abdominal legs. The Trifid group, of course, have five completely de- veloped pairs, of roughly equal size. Many Catocalinz (a subfamily of the Quadrifids) have five pairs, but the foremost pair, on abdominal segment 3, is smaller than that on segment 4. It now appears that the Dyopsine are an exception amiong the Quadrifids, as the pair on segment 3 is equal to that on segment 4, to judge from these two species. Both kinds of larva, moreover, have long single hairs of a sort never seen in the Old World on larvze of Quadrifids.
Both I and Dr. H. B. D. Kerriewe.i, who was visiting Brazil when I found the first of the two (species no.2615), fully expected the larva to hatch into an Acronyctine or perhaps Arctiid moth; we were fooled by the long conspicuous single hairs, and the fully developed prolegs. We did not at first perceive that it lacked the rather dense tufts of hair of the genus Apatele, etc.; it reminded us of the Palearctic Apatele alni. A detailed description is given later of this larva, which is solitary.
48 WILTSHIRE: Dyops Vol.16: no.1
When the moth emerged and proved to be a Dyops (there was little difficulty about determining the genus) it was a great surprise. Further examples were bred later in my stay, but the assistance of the British Museum was required to determine the species correctly.
As for the second species (no.2742), I only found the larve on one occasion, but as it was gregarious I was able to obtain more adults of it than of the first. This larva too, in its last instar, recalled an Apatele larva, but its earlier instars were so completely different from those of the first species, that it was again a great surprise when another Dyops hatched out.
On studying the material of this genus in the British Museum I found both forms confused in a series over the name Dyops ocellata Cramer; but the type of Dyops cuprescens Hampson, placed later, was in fact indistinguishable from the first, larger, species (no.2615). I found only slight differences in the male genitalia, and none in the female; it requires an experienced eye to distinguish the imagines by their pattern and superficial appearance, and without the clue given by the strikingly different larvae the confusion in the Museum series might not have been rectified for many years. I must acknowledge the assistance of Mr. D. S. FLETCHER, without whom I would not now be able to give my conclusions on the correct names of these two species, with comparative morphology.
The two species, by priority of description, will come in the opposite order to that given above.
1. Dyops ocellata Cramer (no.2742) —
(1) Dyops ocellata Cramer [Phalzena ocellatad Cramer (Papillons exotiques 3: pl.276, D, E; 1780)]. Dr. A. Draxonorr kindly informed me that Cramer’s type could not be found at Leiden. I select CRAMER'S figure E as the LECTOTYPE, as I consider the two moths figured as not conspecific. CRAMER thought them the female (D) and the male (E) of the same species, but in fact the sexes do not differ in aspect or size in this genus, in my experience.
I also suggest that the British Museum’s series from French Guiana should be neotypes (Plate 1: figs. 1 and 2).
Synonym: Bacula chromatophila Walker, List lepid. ins. British mus. 15: 1669; 1858) for which I select as LECTOTYPE a ? in Hope Dept. of Entomology, University Museum, Oxford, “BRAZ. 276” (the type series of chromatophila is a mixture, but this selection disposes of the name).
PLATE 1 DY OPS
1) — Dyops ocellata (French Guiana); NEOTYPE in British Museum (x 1); 2) — same (x2); 3) — same, full-grown larve (enlarged); 4) — Dyops cuprescens, Rio de Janeiro, Brazil, (<1); 5) — same, Asuncion, San Lorenzo, Paraguay, leg. H. Pearson (x1); 6) — same, penultimate instar larva (enlarged); 7) — same, full-grown larva (enlarged). 49
50 WILTSHIRE: Dyops Vol.16: no.1
GUENEES remarks on this species are an additional factor leading me
to select ocellata as above, and are worth reproducing (Hist. des insectes, Lep. 6: p.283): —
“1088 Dyops ocellata Cr. 27 DE.
“38 mm. Ailes supér. d'un gris verdatre, avec les lignes médianes géminées, ondées et denticulées. Une grande tache foncée, subcarrée, saupoudrée, et entourée de clair dans la cellule et, au bord terminal entre les 2me et 3me inférieures, un petit oeuil noire 4 pupille blanche, double. Partie de la ligne subterminale qui le précéde, d'un vert doré. Quelques points blancs prés de l’apex. Coudée marquée de blanc au milieu. Ailes infér., d’un gris noiratre luisant, avec un double trait blanchatre a langle anal et deux petites taches ocellées, dont l’extérieure bipupillée et surmontée de vert-doré, entre les 2me et 4me inférieures. Derriére toutes les ocellées on voit, a certains jours, une bandelette cuivrée. Dessous des quatres ailes d'un gris jaunatre, avec les nervures plus claires, une forte tache cellulaire, une ligne médiane et une ombre postérieure noiratres, tres marquées. Dessous de l’abdomen, avec une bandelette noire. Femelle semblable.
“Brésil. coll. Lefebvre, Saunders et Gn. On I’a envoyé en abondance des environs de Pernambuco.
“Nota, Cramer la dit de Cayenne et ses figures D et E paraissent présenter quelques différences, mais elles sont si grossiéres quil ne faut pas sy arreter. Il sera bon de vérifier toutefois par la suite si !espéce de la Guyane est complétement identique avec celle du Brésil, que je décris ici.”
The “green grey’ fore-wing colour, referred to by GuENEE for his Brazilian species, characterises the gregarious smaller species (2742) taken by me. The larger, solitary species (2615) has a purple-brown ground.
The British Museum has a good series of both from French Guiana and Brazil; and after making preparations of the genitalia of several of each, Mr. D. S. FLercuer found that the male’s valve-apex afforded a good character, and kindly sent me sketches of this. My own slides, made from my material, corresponded well to those sketches, and figures 8 and 9 were made from them; fig.10 was made from Mr. FLETCHER’S sketch of a species I do not possess, which can also be distinguished by the valve apex. It will be seen that the valve terminates in a short mem- branous flap with short hairs: in ocellata (fig.8) this flap continues in a straight line from the costa to its point, from which its ventral border curves away at an acute angle to merge in the valve’s chitinous ventral border; in cuprescens (fig. 9) the flap’s point is roughly midway be- tween costa and ventral border, forming a right angle and being obtusely angled to the costa; in dotata Walker (fig. 10) the costa and ventral borders taper to a very acute tip.
1962 Journal of the Lepidopterists’ Society ol
10
8) — Dyops ocellata, & genitalia, ventral open view, with edeagus detached; 9) — D. cuprescens, & right valve tip; 10) —D. dotata, @ left valve tip. All drawings to same scale.
EARLY STAGES (2742).
On November 2, 1958 I found on a single low leaf of a Cecropia sapling on the side of the Rua Redentor, Corcovado Mountain, Rio, at about 500 m. height, a gregarious mass of nearly fifty larve nearly 1” long, slender and glossy brown, with black heads and scanty black hairs which were inconspicuous. The head and tail were slightly raised; all feet were black; there were five equal pairs of abdominal feet. The larvee rested side by side touching each other, and usually fed in the same formation. They resembled a putrescent jelly-like mass and were repulsive, both in this and the next instar. On November 6 they moulted gregariously; after this they were orange-brown in colour, 22-25 mm. long.
On November 18 they again moulted gregariously; and immediately after, the larva-mass appeared darker, as though mouldy, this impression being given by the fresh white hairs on the new dark skins. A little later, they separated to feed and ceased to be gregarious.
52 WILTSHIRE: Dyops Vol.16: no.1
Mature larva (fig.3 of Plate): black, with three fine white transverse dorsal lines on each somite and an orange-brown lateral stripe running the length of the body; head glossy black; feet black; ventral surface, with a few transverse short lines.
Foodplant: Cecropia, and occasionally if nothing else was offered, the foliage of two other trees.
Cocoon: silken, firm, with debris adhering to the outside; I am uncertain where it is placed in the wild state, but presume on the ground, perhaps below the surface.
The moths started emerging on November 27, and about forty had hatched by the end of the month. Many were under-sized due to the difficulty of providing enough of the right foodplant daily; no more moths emerged after December 30. Probably the phenology of this moth is like that of the following species, i.e. multivoltine with “winter pause’, but as only one generation has been bred, this is a guess. In an equatorial climate the “winter pause” would, I suppose, not be made.
The adult moth, when disturbed, falls on its back and flutters jerkily vibrating its wings and body and revealing the lighter, more orange and banded under-side colouring; this is possibly a mimicry of Hymenoptera and may be in an early stage of evolution. It might, alternatively be an exhibition of deceptive or “flash” colouring before suddenly settling.
2. (no.2615) Dyops cuprescens Hampson 1926
This was named by Hampson in Descriptions of new genera and new species of Noctuinae 128. The ? of Cramer's ocellata (P1.276 D) appears identical with both sexes of this species. Hampson’s type, from Chaqui- mayo, S. Peru, is in the British Museum.
This species is on the average larger than the preceding and can usually be distinguished from ocellata by its purple-brown fore-wing ground-colour. See also figure 9 for the valve-tip of the male, and remarks under ocellata above.
EARLY STAGES (2615).
I have found the half-grown and full-grown larve on the underside of larger leaves of young Cecropia trees, feeding by day but hidden. They are usually found singly, but when small are sometimes found two together.
Fourth (?) instar: about 34” long, resembling bird-excrement, brown, slender, shiny, with dorsal white patch on somites 4-9, and paler brown
1962 Journal of the Lepidopterists’ Society 53
ventrally. Head yellow brown. Sete black, white-ringed, with rather long black hairs. A slightly larger larva, observed on another occasion was bluish white, suffused with yellow near the spiracles and on somite 11; the head was yellow with four black spots as big as the sete (see Plate 1, fig.6).
Next instar. the head is glossy black, some of the hairs have thick tips, and the length of the larva is 1”-114” (see fig.7). On the slightly enlarged 10th somite there are two dark patches above two lateral warts; these patches may be brown or black. The dorsal and ventral areas are white with heavy black seta, recalling European Cucullia larve; be- _ tween the two posterior setze of each somite is a short black transverse bar; there are also other transverse black bars of varying length on each somite and these tend to become broader laterally; in the darkest forms these black markings coalesce so that the setee no longer appear as distinct black spots, and the white colouring is reduced to the extent of forming two creamy transverse dorsal bands, and a pair of creamy lateral spots on each somite. One larva found full-grown was feeding on the underside of a large leaf with only its head shewing. When I picked the leaf it stopped, threw back its head in alarm and assumed a dead, squashed appearance, looking rather limp and mouldy. A small dipterous fly was noticed trapped in the hairs of this individual and appeared lifeless. A pre-pupal change of colouring affects the sublateral and ventral areas which turn deep crimson, leaving the dorsal area creamy
white dappled with black.
Cocoon: silky, firm, with debris adhering; pupal period, about three weeks.
Phenology: multivoltine with a probable diapause in the cool season; apparently there are four broods during the warmer half of the year at Rio. Larve of the first are full-grown in early November and produce moths in early December. A full-grown larva was also found in early January and would have produced a moth in mid- February had circumstances permitted completion of rearing. Half- grown larve found in April produce moths in late April and early May. Half-grown larvae found in mid-May produce moths in late June. Whether there is really a diapause in July-October, and in what stage, is uncertain.
Pupa: glossy dark brown, heavily chitined, of normal noctuid aspect, without bloom. The wings cover part of the first five abdominal somites, leaving the next five clear and entire cremaster, consisting of seven short, straight, sharp, well-separated spines, on a blunt, rounded rump.
54 WILTsHIRE: Dyops Vol.16: no.1
3. Other Dyopsinz
Although I have personally come across only the above two species, it may be of interest to add a few words here about other species in this sub-family which appears to be endemic Neo-tropical.
The whitish-banded form Dyops dotata Walker has been placed in the British Museum as an aberration in the D. ocellata series, but Mr. D. S. FLETCHER after investigating the male genitalia came to the con- clusion that it was also distinct (see fig.10 for its valve tip. ).
The remaining Dyops species in the British Museum are easily dis- tinguished from the above three; their names are: chlorargyra Hampson, cyanargyria Hampson, pupillata Felder, subdifferens Schaus; and, placed in a separate genus (Eudyops Hampson), xantholepis Dyar.
Immediately following GuENEE’s description of ocellata quoted above are the descriptions of two further Dyops species: 1089. Dyops oculigera Gd. (38 mm.) and 1090. Dyops hatuey Poey (58 mm.); of these two I have no knowledge and can express no opinion.
SUMMARY
Two species of Dyops, widespread in Central and South America and hitherto confused, have been studied and bred in Brazil.
Dyops ocellata is the smaller species; fore-wing greenish brown in ground-colour. The larva is gregarious at first and in its last instar is black with creamy white hoops and orange lateral stripes, and has long single hairs. A lectotype and a synonym of ocellata are given.
Dyops cuprescens is the larger species; fore-wing purplish brown in ground colour. The larva is solitary or found in couples at first; it is whitish with black spots and bands and long hairs thickened at the end.
Both larve have ten abdominal prolegs equally developed; this, together with the long hairs, distinguishes them from the larvz of other sub-families with which this genus has been placed. The foodplant is Cecropia, a myrmecophilous tree. |
Political Agency, P. O. B. 114, BAHRAIN
1$62 Journal of the Lepidopterists’ Society 55
PRELIMINARY NOTES ON THE LIFE HISTORY OF CALLOPHRYS (SANDIA) MAC FARLANDI
by Down B. Srauines, J. R. TurNer, and Paut R. Exnruicn
It is evident that the proper description of the life history of Callophrys (Sandia) macfarlandi Ehrlich & Clench will have to wait until detailed comparative studies of related Hairstreaks can be made. It therefore seems that a preliminary report of our knowledge is in order at this time.
The egg appears to be typical of the Lyczenide: a pitted oblate spheroid. In the single observed instance of egg laying, the female deposited the egg on a lateral stem near the main stem of the flower stalk. Other eggs were observed on the sheath at the base of the stems cf the flower stalks. Undoubtedly all instars of the larva have been observed and preserved, although no single individual has been raised from egg to adult. Therefore we do not know the exact number of instars.
The larvae were characteristically slug-shaped and were extremely variable in color. The main color range was from a human flesh color to dark maroon. This range of color included over 75% of the specimens. Several larvze were of a distinct pink color, and a single specimen was light green. In pattern some were immaculate and others had several rows of chevron stripes and spots. More than 200 larvzee were examined.
The larve feed primarily on the blossoms of Nolina mircocarpa S. Watson (Agavacez), a Beargrass. However, we observed some of the larger larvae chewing pits into the stem of the bloom stalk. Most larvze were found at rest on the main and lateral stems of the bloom stalk and often concealed themselves in the sheath at the base of the stems. Even those larve feeding on the blossoms were camouflaged well enough, by their color and markings, that they were not seen at first glance. An unidentified species of ant was attending the larve.
No pupz were found in nature, so the pupation site is unknown. The form of the pupa is typical of the genus.
The flight of the adults is rather rapid and erratic, and our observations indicated that they remained close to the location of the larval food plant. The flight during the day is rather limited. Nearly all specimens found in flight were sighted between 9:00 A.M. and 10:30 A.M. Adults were observed to spend the night deep in the basal rosette of the food plant. The main flight in 1959 appeared to be during the first three weeks of May. Some of the mature larvae that we collected were permitted to
56 STALLINGS, et al.: Sandia biology Vol.16; no.1
Callophrys (Sandia) macfarlandi: top left — upperside; top right — underside; middle left — Nolina microcarpa, the foodplant; middle right — larva feeding on blossoms (dorsal); bottom left — two larve feeding (note attending ant above arrow); bottom right — larva feeding (lateral).
pupate. Some of these then emerged the last part of May and early in June; the remainder did not emerge until the following year. Some fresh specimens of the second generation were taken in flight with the more worn specimens of the first generation.
The type locality is on the dry west slope of the Sandia Mts., in Bernalillo County, New Mexico, just north and east of Albuquerque. The exact locality is the La Cueva Camp Ground in La Cueva Canyon. The holotype was caught some 10 yards east of the stone shelter house at the camp ground, and many in the type series were taken in the
1962 Journal of the Lepidopterists’ Society 57
Type locality of C. macfarlandi; Paut ExruicH and Jack Sraxurincs collecting in foreground.
immediate vicinity, About the only other butterflies common at the type locality at the time the type series was collected were Apodemia mormo Felder & Felder and Callophrys (Mitoura) siva Edwards. C. macfarlandi appears to be more abundant in Tijeras Canyon, a short distance from the type locality, and collectors are urged to seek it in this area rather than at the type locality.
References
Ehrlich, P. R., & H. K. Clench, 1960. A new subgenus and species of Callophrys (s. 1.) from the southwestern United States. Ent. news 71: 137-141.
Stallings, D. B., J. R. Turner, & P. R. Ehrlich, 1959. A “hit and run” trip to Texas and New Mexico. Journ. lepid. soc. 13: 99-100.
[DBS, JRT] Caldwell, Kansas, U. S. A. [PRE] Div. of Systematic Biology, Stanford University, Stanford, Calif., U. S. A.
58 BOOK REVIEW Vol.16: no.1
WILD SILK MOTHS OF THE UNITED STATES, SATURNIINA, EXPERIMENTAL STUDIES AND OBSERVATIONS OF NATURAL LIVING HABITS AND RELATIONSHIP. By Michael M. Collins & Robert D. Weast. Collins Radio Co., Cedar Rapids. 138 pp. Available from: Wild Silk Moths, 201 34th Street, Cedar Rapids, Iowa, U. S. A.; price $4.25.
This little book (may be read at a single sitting) contains a great deal of useful information on the species of giant silk moths occurring naturally in the United States of America. This includes moths of the saturniid subfamily Saturniinz mainly, but the genus Automeris (Hemileucine ) is included also because of its popularity with amateur breeders.
The book is copiously illustrated with excellent photographs (although somewhat light in tone, no doubt from the reproduction process) and is organized into two main parts: Part I, United States species, under which are discussed, for each species, (1) habitat, (2) breeding habits, (3) breeding in captivity, and (4) collecting; general remarks are given also under each genus or subgenus; Part II, Studies and experiments, under which are discussed, (1) population dynamics, (2) breeding flights, (3) parasites, (4) diseases, (5) breeding, and (6) hybrids.
There is a nearly complete coverage of species and subspecies and the discussions are detailed and mostly based on personal observations by the authors. Much new information is recorded also. In these respects lies the greatest value of the book both to the non-professional collector and breeder — he may find reliable information to aid him in obtaining and rearing not only the better known eastern species, but also the lesser known southwestern species as well —and to the research worker — he may utilize the first-hand biological data in comprehensive studies on phylogeny, physiology of sexual attraction, geographical distribution, etc.
Critically speaking, I find a few dubious or meaningless statements, such as, “They [the larve of two species] have gaudy, scarlet tubercles which make them appear poisonous or dangerous. (p.80), and “Once the scent is detected, the male flies against the wind. This action suggests the use of eyesight for determining the wind direction.” (p.91) [italics mine]. Also disturbing are the sometimes loose style, incompletely cited and scarce source references, non-italicized latin names, and capitalization of the initial letter of species-group names. Some of these points may be excused because the book is not intended as a scientific work.
All in all, I would say that the authors’ purpose is well served by their book, “ ... to present various aspects of the subject with the hope of creating enthusiasm and to be of help to the breeder or student who plans to do work with these fascinating insects.”
CuHarxLes L. HocugE, Dept. of Entomology, University of California, Los Angeles, Calif., U. S. A.
1962 Journal of the Lepidopterists’ Society 59
FIELD NOTES
AN ALBINO LYCAANA PHLAZAS IN CONNECTICUT
In August 1960 I had a rather interesting experience while hunting in a field in Portland, Connecticut. I was working in a large open field which had a depression at one end. In the depression grew a heavy mass of a pinkish red flower about 2 feet tall. In this depression and on the flowers were swarms of Lyczena phlzxas americana. No sorrel was apparent; consequently I stood wondering what the attraction was. While I was still pondering the question, a small white butterfly appeared on the scene which I did not immediately recognize. To my amazement it turned out to be a white L. p. americana. By “white,” I mean pure white; I have previously found very light pinkish forms. This specimen is identical with the normal form as to spots, etc.
To me the more interesting feature was the fact that it settled on the only white flower in the patch, which makes me ask: did it know it was white, and choose the white flower intentionally? Are butterflies that colour conscious? Further afield there were large patches of the same white flowers, but a search turned up no more white freaks.
The specimen in question is in my collection at Cassadaga.
STANLEY V. FuLLER, Box 81, Cassadaga, Fla., U. S. A.
A WHITISH LYCAENA PHLEAS IN OHIO. (LYCAENIDA:)
One of the more accessible collecting areas near the community of Wellington, Lorain Co., Ohio, is Findley State Park, located two miles south of the village on Ohio State Route 58. The park consists of nearly 900 acres, a large part of which was originally woodlands. In 1954 a shallow valley, through which a small stream flowed, was cleared and an earth fill dam was constructed at the north side of the park. This created an artificial lake of about 90 acres. With access roads, cleared picnic areas, and an artificial bathing beach, the park has become a center of summer outdoor recreation in this area. Plantings of native and introduced trees, made about 30 years ago, are in various parts of the park. These are mostly species of Quercus (oak), Betula (birch), Liquidambar (gum), and various conifers.
The afternoon of August 10, 1961 was sunny with setae cumulus clouds, temperature in the middle eighties, but with a gusty wind of
60 FIELD NOTES Vol.16: no.1
about 16 to 18 miles an hour from the south-southwest. I was collecting on this afternoon in the above described park, following a service road which led through a brush-filled area and into an open meadow-like field of about 20 acres in area.
Abcut 2:30 P.M. (EDT) and at the point where the drive entered the field, I saw a small butterfly darting around in a belligerent manner. Not reccgnizing it, I attempted to net it on the wing, but failed. It returned almost immediately, and this time alighted upon a small patch of bare ground in the drive. Thinking it to be some species of Blue not common to this area, I carefully netted it as soon as it arose from the ercund, papered it, and continued my collecting.
Returning home, I examined my catch. To my utmost surprise, I found this to be a specimen of Lyczena phleas Linnzus. A perfect male, it lacked completely any red or coppery red color, both on the upper and under surfaces of the primaries and secondaries. Instead, this color was replaced entirely by white or silvery white areas, although the dark gray and black portions were of normal color and design. A rather striking appearance was created by this coloration, or rather, lack of it.
Examination of the white areas under a microscope at 200 reveals that these scales are curled or rolled, giving them an almost quill-like appearance. Similar areas on a normally colored specimen reveal flat shingle shaped scales. This suggests that the loss of refractive power of the curled scales results in the loss of the coppery red color, leaving a colorless or white area.
At the present time, this specimen is in the writer's general collection. Subsequent intensive collecting in the area of capture failed to produce any similar individuals, although many normal specimens were taken.
The complex genetics regarding this type of aberration is not com- pletely understood by the writer; but it is felt that a report of such a specimen should be a matter of record, perhaps helping to contribute to information concerning the frequency and distribution of such a form.
I would like to thank Harry CLeNcH of the Carnegie Museum and Prof. A. B. Kuots of the American Museum of Natural History for their interest expressed in this specimen and the encouragement for this article. Special thanks to P. StvireR SmitH of Birmingham, England, for his suggestion of the microscopic examination of the structure of the colorless scales and the observed results. Also the information that this may be the rare form “alba” (Tutt), although this form name is apparently not well known to general collectors in this country.
LeLaNnp L. Martin, 409 Herrick Ave., West, Wellington, Ohio, U. S. A.
1962 Journal of the Lepidopterists’ Society 61
LARVAL NOTES ON CHLOSYNE LACINIA AND C. CALIFORNICA
On 8 Sept. 1961 a few larve of Chlosyne lacinia Geyer were found among a larval colony of Chlosyne californica Wright, apparently at home in this communal association. The few orange caterpillars of lacinia were conspicuous among the dark californica larve.
At the time, colonies of C. californica in various instars were plentiful on Desert Sunflower, Viguiera deltoidea var. parishii in Sentenac Canyon on Highway 78, ten miles northeast of Julian, San Diego County, Cali- fornia. However, these few lacinia caterpillars were found only in one gregarious assemblage of perhaps a hundred fourth instar californica larvee. The butterflies were reared out by WiLt1amM HeEnces; hence iden- tification is confirmed.
Chlosyne lacinia is not common in San Diego County although con- ditions appear quite favorable for it in Borrego Valley. Normally the caterpillars prefer Helianthus annuus, and can be found in large quanti- ties in the fall in Imperial Valley on roadside patches of sunflowers. This record on Viguiera deltoidea probably represents a new host plant record. |
Chlosyne californica on the other hand is one of the dominant species of the desert canyons of this area whenever rainfall is favorable, and the butterflies appear in large numbers at times. The caterpillars accom- modate readily to Helianthus annuus in captivity but I have never ob- served them on anything but Desert Sunflower in nature.
FreD THORNE, 1360 Merritt Dr., E] Cajon, Calif., U. 8. A.
TWO NEW FOODPLANTS OF SOUTHWESTERN SATURNIIDA&
The hosts of Agapema galbina anona Ottol. and Citheronia splendens Druce were discovered September 1 and 2, 1961, in Brown’s Canyon of the Baboquivari Mountains of Pima Co., Arizona. While making a detailed search for Rothschildia jorulla cinctus Tepper, I quite accidentally came upon a Cluster of cocoons of A. galbina anona. The host is a species of Lycium, which is a very thorny, sparsely-leaved shrub. Previously I had spent considerable time trying to locate the moth in its known areas of occurence in the Santa Rita and Santa Catalina mountains without success. Once the precise foodplant was determined I was able to find the cocoons in considerable numbers. —
62 FIELD NOTES Vol.16: no.1
The cocoons are spun in a cluster towards the center of the shrub, being completely surrounded by dense thorns. This affords excellent protection from birds and rodents. The clusters frequently contain more than a dozen cocoons. The gregarious larvae must sometimes completely defoliate the host. I collected cocoons along the canyon’s slopes, which are hotter and drier than the canyon’s floor, where the cocoons can also be found. The adults fly in November and early December and are usually taken at “black light”. Lycium, where found, is common, but it does not seem to be as widespread as are other common plants of the area. Lycium grows in association with various cacti, agave, acacia, mesquite and ocotillo, but it is frequently missing from habitats containing these plants.
A mature larva of Citheronia splendens was taken on wild cotton, Gossypium thurberi. This plant grows along canyon streams and in other moist situations. This striking larva had evidently completed its full growth on the plant. It pupated in mid-September. C. splendens is also reported to feed on walnut trees which grow throughout the mountain ranges. The species flies in July.
Rosert D. WeEast, 4818 Seneca St., Des Moines 10, Iowa, U. S. A.
ADOPCEA LINEOLA (HESPERIIDAE) NOW ALSO IN NORTHERN ONTARIO
On July 9, 1960, the author observed on a roadside near Sudbury, Ont. (Richard Lake), a skipper which looked to him like Adopeea lineola. It was, however, then not possible to secure the specimen. On July 16, 1961, the first specimen of Adopeea lineola was quite accidentally caught about 300 yards away from the spot of the 1960 occurrence by JOHN Dencuuck, a Sudbury High School student interested in Lepidoptera and accompanying the author on a field trip. This brought the indisput- able proof that this little skipper is extending its range now into northern Ontario. On July 17, 1961, the author took a second specimen and this at the spot where the species was observed a year ago. It certainly will be interesting to see at which rate Adopeea lineola, which is now still quite rare, will increase in coming years.
The specimens are in the collection of the American Museum of Natural History in New York.
J. C. E. Riorre, 278 Bathurst St., Toronto 2-B, Ont., CANADA
1962 Journal of the Lepidopterists’ Society 63
MIGRATION OF NYMPHALIS CALIFORNICA IN WASHINGTON (NYMPHALID®)
During the week of July 16 to 22, 1961, an immense migration of the butterfly Nymphalis californica was noted in the high mountain country of the State of Washington. Reports indicate that on all the major mountain passes in the state this butterfly was in such great numbers as to slow traffic and clog radiator grills. The greatest numbers were observed at an altitude between 3500 ft. and 4000 ft. although reports have been received of observations up to at least the 5000 ft. level. This area, at least in the eastern part of Washington, is at about the upper level of the Transition Zone or even the lower part of the Canadian Zone. Yellow Pine, fir, Western Larch, spruce, Jack Pine, poplar, alder, willow, and aspen abound in this area and in the open spaces wild lilac, spirea, and Buck Brush (Ceanothus thyrsiflorus) are in bloom then.
On a trip from Oroville to Spokane on July 16, we first saw this butter- fly swarming in a large sunlit rock cut on the summit of the pass be- tween Disautel and Nespelem at about 9:30 in the morning. This pass is on Washington state highway 10A. There were mud puddles at the edge of the road and many butterflies were drinking or resting on the ground along the side of the road. There were many dead and feeble ones on the ground but we were unable to determine the cause, whether from the passing cars, fatigue, disease, or some other cause. There did not seem to be any predators or parasites bothering them at the time. We took pictures and gathered specimens at this time and noted that taking them was much easier than is normal. Having no net we merely picked them up from the road side. Their flight was abnormally feeble.
On the return trip on July 20, over Sherman Creek Pass on State Highway 3P we took specimens, although at that time the flight was not nearly so great as it had been early in the week.
I have reports of this butterfly in great numbers on other passes and mountain country in the State of Washington. They were seen on Blewett Pass on July 17, in the Tiffany Lake area of the Okanogan National Forest west of Omak Wash on July 23, and in the Lost Lake and Bonaparte Lake area east of Chesaw, Wash., and north of Wauconda, Wash., at various times during the same week.
On July 24, we returned to the Republic and Swan Lake vicinity and could find no trace of the butterflies. From this fact I believe that this was a migration rather than a large hatch, I was unable to determine in which direction they were migrating.
Westey H. Rocers, Route 1, Oroville, Wash., U. Sik
64 Vol.16: no.1
ESPECIALLY FOR FIELD COLLECTORS (Under the supervision of FRED T. THORNE, 1360 Merritt Dr., El Cajon, Calif., U.S.A.)
COLLECTING CENEIS NEVADENSIS (SATYRINAZ) AND OTHER GENERA ON VANCOUVER ISLAND, WITH A THEORY TO ACCOUNT FOR HILLTOPPING
by RicHarp Guppy !
The discussion on “hilltopping” butterflies, which has somewhat died out in recent years, was in 1960 revived in my mind by a rather good season for CEneis nevadensis. This butterfly is by all accounts the most inveterate hilltopper in this part of the world.
The spot in which I collect most of my CE. nevadensis Felder is actually the top of a ridge, rising quite steeply up from the seashore. It is by a rough estimate, perhaps 500 ft. high. On the landward side the slope is much more gentle; from my home it takes me about ten minutes to walk to the top of the ridge, although the ascent from this point is probably no more than 200 to 300 ft.
The butterflies are found always along a short stretch of the ridge, about 50 yards altogether, which I suppose must be the highest point. This area is divided into three “compartments’, small clearings separated by clumps of Arbutus and stunted Douglas Firs. In 1960, for about two weeks in the latter part of June, I could almost depend on finding each of these compartments occupied by a male CE. nevadensis, which when collected, would be replaced by the following day.
If it were not for their stubborn insistence on staying in their chosen small area, these butterflies would not be easy game by any means. With most of the more active and wary butterflies, a missed swipe means that the insect will clear out in a hurry, and give the collector no second chance. But these hilltop CG£neis will circle repeatedly; often the collector is able to get in five or six successive tries, and even if he finally frightens the butterfly off, only a little patience is needed; in ten minutes or so it will be back. :
These hilltop CEneis are all males. Females I very seldom find, and always in localities at some distance from where the males like to disport themselves. Having considered all this evidence, I have arrived at the conclusion that this hilltopping business is simply a means by which the males and females are brought together. The problem of finding mates
1962 Journal of the Lepidopterists’ Society 65
that are other than siblings, must trouble most insects, and it must be admitted that not all solve the difficulty by hilltopping. Among butter- flies some, notably the Lyczenidze, would not meet with the problem, be- cause they tend to form colonies, which remain in one small area, where the necessary host plant is plentiful.
In spite of these objections, it seems certain that, in the case of species that wander a great deal, some instinct that would draw all newly emerged individuals towards a central point would serve them in good stead. Suppose that every (Eneis, on emerging, is possessed of an urge to fly uphill. The males, which are always on the wing before the females, will become concentrated at certain high spots. The females will then begin to arrive at these meeting places. It is a known fact that the be- haviour pattern of a female insect may change after mating. We can suppose then that since the female CFneis, on reaching a hilltop, will immediately meet a male, she will shortly lose her uphill instinct, which will be replaced by an urge to search for a suitable place for oviposition. The strongest support for this part of my theory lies in the fact that, if females did not fly up to the hilltops, the males which persisted in staying there would get no mates. That being so, natural selection would inevit- ably eradicate the habit. The short stay necessary for the females to com- plete business and move on would mean that a collector could easily miss seeing them, if he were in the habit of stopping only long enough to collect any males in residence at the time.
An interesting aspect of the problem, for which I have no good ex- planation, is the way in which males can establish and hold a territory. On most peaks, only one is present at a given time, unless there is quite a plateau, in which case several will remain about 50 yards apart. At that distance they apparently cannot see one another. The tree clumps on my ridge, by screening the butterflies from sight of one another, allow three males to occupy quite a small area. When a male comes into a rival's territory, the two flutter around each other for a time, then one, which I cannot say of course is always the trespasser, will clear off out of sight. Why it should do so it is difficult to say, since the other male obviously cannot hurt it. But the utility of the system is evident. Each male is under the necessity of investigating every butterfly which comes into its range of vision, in case it should prove to be a female of his species. Two males remaining continuously in sight of one another, would create an intoler- able situation. 7
It seems probable that some species that are not hilltoppers use the rendezvous system, although with these the means by which the meeting place is selected remains a mystery. I know of spots near my home where,
66 Guppy: (Cneis hilltopping Vol.16: no.1
year after year during the correct season, I can go each day to collect the current male Limenitis lorquini Bdv., which will be replaced each time just as are the CEneis males on the ridge. But these Limenitis spots have, so far as I can make out, absolutely nothing to set them apart from the rest of the scenery. They are not at the tops of hills, but often on a slope.
While on the subject of hilltoppers, I would like to add some further information that has come to light, regarding our other inveterate hill- topper, Papilio zelicaon Luc. In this journal (Vol.7: 43-44; 1953) I stated that on Mt. Arrowsmith there were no suitable host plants for P. zelicaon, and that to reach the summit from the nearest stand of umbelliferous plants, they would have to fly about five miles over, or through, dense forest. In 1960, while on my annual collecting trip to Mt. Arrowsmith, I noticed a butterfly of this species, behaving in a suspicous manner. So certain was I that it was engaged in oviposition, that after an un- successful attempt to collect it, I returned to examine what had appeared to be a patch of rocks supporting a few lichens and mosses. Close ex- amination revealed a number of tiny plants, just coming into bud. So small were they, and so closely appressed to the rock, that any one of them could have been hidden by a silver dollar, yet in the aggregate there were enough to feed several P. zelicaon larve. I took home a sample, and in water the buds opened into tiny yellow umbels. They were undoubtedly a species of Lomatium, very probably L. martindalei C.& R. I could find no ova on any of the plants, but it is, after all, im- material whether the butterfly was ovipositing or not. By looking as if it were laying eggs, it had led me to discover on Mt. Arrowsmith an um- belliferous plant which I had not noticed before, and which certainly could account for the presence of P. zelicaon in the vicinity.
\
R. R. 1, Wellington, B. C.,. CANADA
The artist who prepared the figure for the front cover for Volume 16 is again WiLL1AM Vars, of Yale University.
1962 Journal of the Lepidopterists’ Society 67
COLLECTING IS STILL GOOD IN THE NORTHWEST
by E. J. NEwcoMER
The increasing use of insecticides and herbicides in the West, the construction of dams and super highways, the occurrence of forest fires, all are reducing the good collecting areas for the lepidopterist. But many very rewarding areas remain, and will continue to exist for many years.
Instinctively the butterfly collector looks for open places — natural meadows, gravel-storage areas, railroad rights-of-way, abandoned highways, skiing slopes. For the combination of sunlight and the numer- ous flowering plants that grow in such areas are very attractive to butterflies. It is my purpose here to describe three such places that I have found to be very productive.
1. Skookum Meadow. Looking at a road map of Oregon, I became curious about the Walker Rim, a mountainous area in the Fremont National Forest just east of U. S. Highway 97 and about 50 miles south of Bend. A “rim” is the same as an escarpment, a steep slope caused by an ancient upthrust of the land. So I went up there. On top, the land levelled off toward the east, the soil was mostly pumice, making driving easy, but covered with a thick growth of Lodge-pole Pine, under which practically nothing grows. A worse place for butterflies can hardly be imagined.
But the Forest Service map had shown some meadows. So I pushed on. Suddenly, the trees opened up and here was a gem of a meadow, a half mile long and a couple of hundred yards wide, covered with flowers, and literally swarming with butterflies. I could hardly believe it.
Collecting in this meadow for less than three hours, I found I had 100 specimens, after discarding the poor ones, and they added up to 33 species. Commonest was Phyciodes campestris and it was a_ pest, at times being mistaken for something else. Then came Euphydryas editha and then Cercyonis ctus. Next to these in abundance I would put the Speyeria, and Paut Grey has identified six species for me: coronis ssp., zerene near conchyliatus, callippe ssp., egleis unnamed ssp., hydaspe, and mormonia, the latter perhaps erinna. There were nine species of lycaenids, the most interesting one being Plebejus shasta. Scarcest was Danaus plexippus, a single specimen of which was being
68 Newcomer: Collecting in Northwest Vol.16: no.1
hustled and harried all over the meadow by a frantic and frazzled fritillary, which Paut tells me is S. callippe.
There were a few clouds in the sky and when the sun got behind one of them the Speyeria disappeared. I soon found where they were, spread out flat on a barren sandbank facing the sun, soaking up as much heat as they could. As soon as the sun came out, back they went to the blossoms.
There was a very noticeable number of dwarfed specimens in this meadow — Boloria, Euphydryas, Cercyonis, Phyciodes and some of the lyczenids. Could this be the result of not enough food or perhaps of inbreeding in this isolated place?
This meadow is not shown on the forest map, — it is too small. But I have named it Skookum Meadow, because it is just south of Skookum Butte, which is on the map, and because in the Chinook Indian language “skookum” means “mighty good.” The elevation is 5200 feet. I collected here late in July and am curious to see what it will yield a month earlier.
2. Satus Ski Area. This is near Highway 97 also, but in Washington between Toppenish and Goldendale. It is part of the Brooks Memorial State Park. The latter is a few miles south of Satus Pass, but to get to the ski slope you must turn off the highway right at the pass and go west for a couple of miles.
A considerable area has been cleared and is now growing up to flowers, grass and low shrubs. The top of the slope is perhaps 500 feet higher than the bottom and the average elevation is about 3700 feet. Because range cattle sometimes graze the slope, it is not always good. But an area of seepage at the bottom is worth looking at, and Speyeria may be found in season on a patch of dogbane (Apocynum) just above the lodge.
Because time is important as well as place, this area does not yield much in June. But here, on a July day in 1960, I collected 30 species. Altho not as numerous as at Skookum Meadow, many were plentiful. Notable were five species of Speyeria, the same basic ones as at Skookum Meadow, except for egleis, but the subspecies were different; also Euphydryas colon and editha, Chlosyne palla (whitneyi and sterope), and hoffmanni manchada, all past their prime; nine species of lyceenids; and CEneis nevadensis. DAN CaRNEY took CEn. chryxus here a few days earlier, but I did not find it.
The great popularity of skiing has opened up slopes like this one in many places and collectors would do well to examine them. No skiers will crash into you while you are doing it.
1962 Journal of the Lepidopterists’ Society 69
3. Bear Canyon. West of Yakima, in Washington, the Tieton River comes down from Rimrock Lake and combines with the Naches River, both carrying the precious stored-up melted snow that makes the Yakima Valley so rich agriculturally. Collecting is not especially good
along either river. But go 8 or 10 miles up the Tieton and if you look sharply, you will see a narrow forest road going into the hills to the right. This is an old logging road going up Bear Canyon, and maintained for access in case of fire. There is a small stream coming down the canyon, but the Forest Service refers to this as a canyon rather than a creek, probably because there are already two other Bear Creeks in Yakima County.
And this is a canyon, narrow, with steep sides,—in many places there is just room for the creek and the road. The sun shines down into it much of the day, there are a few small side ravines, and every- where there are flowers,—asters, spireea, pentstemon, Eriogonum, yarrow, Erigeron, mallow, Mimulus, Senecio, Erysimum, Arnica, lupines, and the little Woolly Sunflower ( Eriophyllum) which is especially attractive to butterflies. Here, in a stretch of about three miles, with an average elevation of 2500 feet, is a butterfly paradise. Collecting is good from May to August. And here on a July day, Dr. A. I. Goop, of Ohio, and I took what is so far a record for me — 36 species. They are worth listing:
Papilio eurymedon P. rutulus
P. multicaudatus Colias occidentalis Pieris rapzx
P. protodice
P. beckeri Caenonympha tullia Cercyonis pegala
C. sthenele
Speyeria cybele leto S. callippe semivirida S. coronis simztha S. hydaspe sakuntala Chlosyne_ palla Phyciodes muylitta P. campestris Polygonia satyrus
Nymphalis antiopa N. californica Limenitis lorquini Satyrium behrii
S. sepium
S. californica Strymon melinus Lyceena heteronea
L. nivalis
L. helloides
Plebejus acmon
P. icarioides Heliopetes ericetorum Pholisora catullus Erynnis persius
E. icelus
Hesperia juba Ochlodes_ sylvanoides
70 NewcomMer: Collecting in Northwest Vol.16: no.1
In addition to these, 26 other species have been taken at other times in Bear Canyon. They are:
Parnassius smintheus Callophrys eryphon Papilio indra C. fotis
P. zelicaon C. augustinus (iroides ) Colias eurytheme C. spinetorum Anthocaris sara Plebejus melissa Euchloe ausonides Glaucopsyche lygdamus Neophasia menapia Everes amyntula (Eneis nevadensis Celastrina argiolus Chlosyne hoffmanni Apodemia mormo Euphydryas colon Pyrgus communis
E. editha Erynnis propertius Nymphalis cardui Thorybes pylades Polygonia zephyrus Amblyscirtes vialis
Thus a total of at least 62 species occur in this small area, of the 100 so far found in Yakima County. Two species were especially abundant in June 1961. These were Chlosyne hoffmanni and Lyczna_ nivalis. Heliopetes ericetorum was numerous in 1960, the caterpillars feeding in the bush mallow, but scarce in 1961. Papilio indra and multicaudata are always scarce, and Neophasia menapia and Apodemia mormo were taken there for the first time in August 1961, altho I have been collecting in this canyon for four seasons.
There are many other good collecting places in the Northwest: — in Oregon, the Ochoco and Maury Mountains in Crook Country, Tomb- stone and Lost Prairies in Linn County, the resort area around Bachelor Butte in Deschutes County, Diamond Lake in Douglas County, and Summit Prairie east of Lakeview; in Washington, Mt. Spokane, O’Brien Creek near Republic, the Blue Mountains, some of the forest lookouts in Yakima County, many places in Okanogan County, and some western Washington places; in northern Idaho, the higher elevations in the Priest River Experiment Station in Bonner County, and of course many others. But the three described above have been the best. Forest Service people are always happy to help with suggestions, with advice about roads, and with maps of their forests.
1509 Summitview, Yakima, Wash., U. S. A.
1962 Journal of the Lepidopterists’ Society 71
EDWIN PAUL MEINERS (1893-1960)
We regret to report the death on October 28, 1960 of the veteran entomologist and physician Dr. Epwin Paut MEINERS of St. Louis, Missouri. Dr. Mrrners was a Charter Member and long a. Sustaining Member of the Lepidopterists’ Society and contributed several articles to the pages of the Lepidopterists’ News (see biblicgraphy at the end
12 Meiners Obituary Vol.16: no.1
of this obituary). From early boyhood he was an ardent lepidopterist and continued this interest until his death.
EpwIN MEINERS was born in Troy, Illinois, on February 11, 1893. His grandfather emigrated to America in 1849 and came first to St. Louis, later locating in Troy, a small town of about 2000, where he remained for the rest of his life. He owned and operated the White Horse Tavern which became famous as a stopping place in the stage coach days when the old National Road was the main artery of travel between the East and the West. Epwin’s father, FREDERICK MEINERS, Jr., was born and raised in Troy and continued the family business which expanded into the Central Hotel. His mother was Exiza Prarr and he was the oldest of five children born to WitL1AM and Exiza MEINers. He is survived by three brothers.
In a detailed and careful autobiography and family genealogy which EDWIN MEINERS wrote a few years before his death, he says of his father: “His life exhibited very markedly the characteristics of the German as we are wont to think of him in the days when he was loved by all; non- aggressive, modest Hanoverian Teuton, a sincere lover of home and children and all the comforts of life as best summed up and expressed in the German “Gemiitlichkeit”. His devotion to his children was very deep. The days of my childhood that have left their deepest impression cn my mind and character, that today remain the fondest of memories, were those spent in the company of my father. It was his guiding hand although untutored, that first pointed out the beauties of nature to me. It was he who aided me in the problems of gathering and preserving my natural history history specimens, who helped me in my work and play.” I would like to say here that many of the endearing qualities which Dr. MEINERs attributed to his father were also manifested in large measure by him. In addition there was a deep scholarliness and absolute integrity which caused me to recommend his services whenever a friend asked me for the name of an honest physician.
Dr. MEINERS received his medical training in St. Louis and interned at St. Luke’s Hospital. While still an intern he was the first to develop a method of injection of intravenous glucose to combat post-operative shock. Thereafter it became a regular procedure in the treatment of shock. In World War I he served in Nashville, Tennessee, with the United States Public Health Service. While at St. Luke’s Hospital he met and married Liww1an Mar McGrmsey, who survives him. Children of this marriage are Dr. Epwin Pau MeErners, Jr., Dr. THEODORE M. ME ners, and Mrs. Mary EvizaBeTH BoucHNov. There are twelve grand- children living.
1962 Journal of the Lepidopterists’ Society
~I
Loy)
Epwin Merners always felt keenly that St. Louis, whose metropolitan area has a population well over one million, could and should support a genuine natural history museum comparable to those in other large cities of the country, where the student could go to consult study collections and carry on biological research. It was his great ambition to see this established, but he died without seeing his dream realized. This was his great disappointment. In his efforts to arouse support for a museum he was a member of several embryo scientific clubs and of societies in St. Louis, some of which had brief histories: the Heink Entomological Club, the St. Louis Entomological Club, the Webster Groves Nature Study Society, the Greater St. Louis Museum of Natural History, the St. Louis Naturalists’ Club. and the Academy of Science of St. Louis. He was one of the organizers in 1934 of the Entomology Section of the Academy, which flourished for about ten years. Members of this section, many of whom have since published significent con- tributions on entomology were: O. ZreELinski, RICHARD FROESCHNER, L. Prnxus, E. D. BUEKER, WENDELL SHAY, Huco Press, Martin Gorpon, Harotp O'Byrne, Puit Rav, P. S. and C. L. Remincton. Writing of these meetings MEINERS says: “One night Suay electrified us by appear- ing rather late loaded down with a two foot long setting board and two quart jars filled with the most wonderful moths. He not only took our breath, but he broke up our meeting by offering to take those of us with him who were willing to go at this hour of the night and to share his discovery with him.” Dr. MEINERS was chairman of this section at different times. He also served the Academy as Curator, Librarian, Second Vice-president, Editor of the Bulletin, member of the Board of Directors. At one time a gift to the Academy gave it a very large brick residence, in a fine area of the city, which was turned into a museum. Dr. MEINERS organized and arranged the first insect exhibits, using much of his own collection. However, this effort eventually died out and the museum now maintained by the Academy is virtually a Childrens’ Museum. Dr. Merners felt that this was not the proper place for study collections and so he willed his fine collection of insects to the University of Missouri. It is now under the care of Dr. WitpurR Enns, who tells me that the collection consists of 330 boxes and drawers of Lepidoptera, 96 of Coleoptera, 13 of Hymenoptera, 15 of Orthoptera, and 38 of mis- cellaneous groups. A conservative estimate places the number of speci- mens at a total of 30,000 to 40,000. The collection is now in the Ento- mology Museum at the University of Missouri.
In recent years Dr. Merners developed an interest in other branches of entomology besides Lepidoptera and made a good general collection
74 Meiners Obituary Vol.16: no-1
of the insects of Missouri as shown by the above items. His favorite — collecting haunts around St. Louis were Cliff Cave, Creve Coeur Lake, and especially Ranken, about which he wrote a fine article (Lepid. news 10: 163). He was greatly interested in the history of science and prepared excellent papers on outstanding Missouri entomologists. He collected all the published papers of Cuartes V. RILEY, first State Entomologist of Missouri, and of Mary V. Murrrevpt, an early lepidopterist of Kirk- wood. He owned a collection of letters and writings of early and con- temporary entomologists and a notable collection of rare books on entomology. This was purchased by the General Library of the University of Missouri. Part of the purchase price was used by his widow to estab- lish the Dr. E. P. Meiners Memorial Scholarship fund at the University. Mrs. Merners tells me that the Doctor was working at the time of his death on a book to be entitled “The Effect of Insects on Civilization.” An effort will be made to ascertain if some of the material in it can be published.
Dr. MEINERS was regarded as an authority on medical entomology. He was certainly the dean of entomologists in St. Louis. His first love was always Lepidoptera. He was always willing to present interesting and scholarly papers on topics connected with butterflies and other in- sects. He wrote: “I spent about four hours one Sunday afternoon showing my collection and lecturing on entomology to a class of students from Harris Teachers College. I doubt whether any of the class had ever sat through a lecture that long!” He well recognized the importance of encouraging young people in the study of insects and several of the young men to whom he gave encouragement are now professional en- tomologists, including the Editor of this publication.
He was a member of the American and Missouri Medical Societies, the Entomological Society of America and the American Association for the Advancement of Science. Just before he died he was instrumental in the starting of a new entomological society in St. Louis. Plans for this were continued after his death and the society took in his honor the name of The E. P. Meiners Entomological Society. The society has held monthly meetings under the chairmanship of Mr. Pinxus and is thriving.
Dr. MEINERS will be remembered by his many friends for his unfailing kind efforts to help even the layman gain some understanding cf entomology and to show them his collection and encourage them to learn of the wonderful world of insects. He tried to give them an appreciation for the truly scientific viewpoint toward all matters biological. His name may be found in pos Passos forthcoming Check List
1962 Journal of the Lepidopterists’ Society 75
of the Nearctic Rhopalocera, for GuNpDER named an aberration of Strymon melinus and FieLtp named the spring form of Everes comyntas for him. Dr. MEINERS was not particularly proud of this, for he held that aberra- tions and forms should not be named. Indeed, he long felt that there were too many species names and believed that when the biology of Lepidoptera is better understood, many names will disappear into the synonomy.
Many pleasant evenings were spent by this writer in Dr. MEINERS’ study or in mine as we chatted about insects. On one occasion I recall saying to him a little facetiously: “Doctor, you know the world thinks you and J are “nuts” to devote so much time and effort to collecting and studying bugs!” He replied, without the familiar twinkle in his eye: “Mr. REMINGTON, what the world is more “nuts” like you and me! There would be a lot less tension and trouble in the world if this were so.”
BIBLIOGRAPHY OF PUBLICATIONS BY E. P. MEINERS
1936a. Notes on Papilio asterius Cram. and a record of var. americus Kollar. Ent. news 47: 269.
1936b. The dimorphic blue female of Everes comyntas Godt. Annals ent. soc. Amer. 29: 620-621.
1939. The life history of Euchloe olympia Edwards with some notes on its habits. Proc. Missouri acad. sci. 4: 154-156.
1941. Obituaries of Hermann Schwarz, George W. Bock, August Knetzger, Charles L. Heink, and Vitae Kite. Ent. news 52: 118-120.
1948a. Brief biographies. 12. George Hazen French (1841-1935). Lepid. news 2: AQ.
1948b. Brief biographies. 14. George Duryea Hulst (1846-1900). Lepid. news 2: 6G.
1948c. Brief biographies. 15. Mary Esther Murtfeldt (1839-1913. Lepid. news 24593 SF
1948d. Book reviews and obituary note [Phil Rau]. Lepid. news 2: 43, 55, 62.
1949. A brief history of lepidopterology in Missouri. Lepid. news 3: 51-52.
1951. Harold I. O’Byrne (1898-1951) [Obituary]. Lepid. news 5: 11-12.
1953. The Languriide and Erotylide (Coleoptera) of Missouri with notes and keys. Journ. Kansas ent. soc. 26: 18-25. [With R. C. Froeschner.]
1956. Lepidoptera collecting at Ranken, Missouri. Lepid. news 10: 163-168.
P. SHELDON REMINGTON
12 Country Meadow Lane, Box 612, R.R.1, St. Louis 41, Mo., U. S. A.
76 Vol.16: no.1
RECENT LITERATURE ON LEPIDOPTERA
(Under the supervision of PETER F. BELLINGER)
Under this heading are included abstracts of papers and books of interest to lepidopterists. The world’s literature is searched systematically, and it is intended that every work on Lepidptera published after 1946 will be noticed here; omissions of papers more than 3 or 4 years old should be called to Dr. BELLINGER’s attention. New genera and higher categories are shown in CAPITALS, new species and sub- species are noted, with type localities if given in print. Larval foodplants are usually listed. Critical comments by abstractors may be made. Papers of only local interest and papers from this Journal are listed without abstract. Readers, not in North America, interested in assisting with this very large task, are invited to write Dr. BELLINGER (Dept. of Natural Sciences, San Fernando Valley State College, Northridge, Calif., U. S. A.). Abstractors’ initials are as follows:
[P.B.] —P. F. Bectuincer [W.H.] — W. Hackman [N.O.] — N. S. Osraztsov [..C.] —I. F. B. Common [T.].] — Taro Iwast [C.R.] — C. L. RemincTon [W.C.] — W. C. Coox [J.M.] —J. Moucua [J.T.] —J. W. Titpen [A.D.] — A. DrtaKonorFr [E.M.] — E. G. Munroe [P.V.] —P. BL. Views
B. SYSTEMATICS AND NOMENCLATURE
Marion, H., “Contribution a étude des Pyralide de Madagascar” [in French]. Mém. Inst. scient. Madagascar, sér. E, vol.5: pp.39-62, 1 pl, 14 figs. 1954. Contribution to the study of the Madagascar pyralids. Describes as new (from Ankaratra Mt. unless another type locality is given): Ancylolomia auripaleella; PRODUCTALIUS, & type P. tritzniellus; ANGUSTALIUS, & type A. ditzniellus; Tegulifera herbulotalis (Tananarive), T. sanguinalis (Tananarive); Obtusipalpis rubricostalis (Tananarive); EUPHYCIODES (type _ species: albotessulalis Mabille); Nacoleia pulveralis (Tananarive); Sylepta pauperalis; Diaphana viettealis (Tananarive), D. toulgoetalis (Tananarive), D. ankaratralis; Ischnurges paulianalis; Trigonuncus flavopunctalis; Pachyzancla ultratrinalis;s MEGATAR- SODES (type species: baltealis Mabille); Argyractis ambahonalis;s AMBAHONA, & type A. fusconebulis. [P. V.]
Marion, H., “Révision des Pyraustidze de la faune frangaise” [in French]. Rev. frane. Lépid., vol.14: pp.123-128, 1 pl; pp.181-188, 221-227, 16 figs.; vol.15: pp.41-58, 24 figs. 1954-55. Revision of the Pyraustide of the French fauna. The author begins with long comments on the history of the classification, the taxonomic categories and the morphological structures. The group of pyralids is divided by Marion into: Crambidze, Pyraustide, Acentropidz, Pyralide, Phycitide, and Galleriide. The subfamilies of the Pyraustide are Scopariina, Nymphuline, Schcenobiinze, Pyraustinze, Evergestinee. The author starts with the Acentropide; Schcenobiinzee and Scopariine (with two tribes: Scopariini and Cybalomiini) follow. Keys for families, subfamilies, tribes, genera, and species are given. Study to continue. [P. V.]
Marion, H., “Complement & la classification et nomenclature des Pyraustide d'Europe” [in French]. Entomologiste, vol.13: pp.129-130. “1957” [1958]. The two European spp. palustris Hbn. & nubilalis Hbn. should be placed in the genus Ostrinia Hbn. which has priority over Micractis W. Warren. [P. V.]
Marion, H., “Révision des Pyraustide de France” [in French]. Alexanor, vol.1: pp.15-22, 2 pls. 1959. Continuation of the revision of the French pyraustids. Here the genus Scoparia is begun. [P. V.]
Marion, H., “Classification et nomenclature des Pyraustide d’Enrope” [in French]. Entomologiste, vol.15: pp.44-45. 1959. Nomenclatorial note about the genus Epicorsia Hbn. and the new genus PARACORSIA (type species: repandalis Den. & ‘Schitt.)= URs Val
1962 Journal of the Lepidopterists’ Society a.
Marion, H., “Révision des Pyraustide de France (suite)” [in French]. Alexanor, vol.1: pp.103-110, 1 pl. 1959. Continuation. Description of the genus HYPERLAIS (Scopariine, Cybolomiini; type species: Hypolais siccalis Gn.). [P. V.]
Marion, H., “Révision des Pyraustide de France (suite)” [in French]. Alexanor, vol.1: pp.175-182, 1 pl. 1960. Continuation; conclusion of Nymphuline, beginning of Evergestine, & complementary notes on Scopariine. Describes as new Scoparia cembre pfeifferi (Alps, Savoie, Peisey-Nancroix). [P. V.]
Marquant, R. M., “Une nouvelle race de Parnassius apollo Linné: P. apollo race ossalensis mihi” [in French]. Bull. mens. Soc. Linn. Lyon, vol.27: pp.51-52. 1959. New subspecies described from central Pyrenees, Col. d’Aubisque, alt. 1700m. PRA Vel
Mell, R., “Zur Geschichte der ostasiatischen Lepidopteren. I. Die Hebung Zentralasiens, das westchinesische Refugium zentralosiatischer Abkémmlinge und die Verbreitungsachse Sikkim/Kashiaberge—Zentralformosa (Achse V). Beitrage zur Fauna sinica XXV” [in German]. Deutsche ent. Zeitschr., N. F., vol.5: pp.185-213, 11 figs. 1958. Describes as new: (Sphingide) Clanis obscura (Kuatun); Rhagastis albomarginatus sauteri (Taihorin, Formosa); (Bombycidze ) THEOPHOBA (type not selected; genus includes Andraca albilunata), T. pendulans (Lungtaoshan, 800 m., N. Kuangtung); Ocinara diaphragma (Mahntsishan, N. Kuangtung), O. d. formosa (Shis, central Formosa); (Eupterotide) Ganisa longipennata (Kuatun, NW Fukien); (Notodontide) Andraca olivacens (Kuatun); (Acronictine ) Trisuloides klapperichii (Kuatun); (Saturniide) Antherzea yamamai titan (Kuatun). A very diffuse discussion of faunal origins in central Asia, illustrated principally by Sphingidze, Saturniide, and some other families of Lepidoptera. The paper is marred throughout by the presentation of probabilities or possibilities as certainties; for example, the author’s statements of the geographic origins of genera unknown as fossils, and designation of species known from single specimens as “double-brooded” or “obligatorily single-brooded”. [P. B.]
von Mentzer, Erik, “Erebia calcarius Lork. ssp. tridentina, ssp. nova (Lep., Satyridz )” lin German]. Ent. Tidskr., vol.79: pp.6-10, 5 figs. 1958. Described from Lago Nambino, 1800 m., Venezia Tridentina, Italy. Discusses systematic position of E. calcarius. [P. B.]
von Mentzer, Erik, “Ueber die Spezifizitat von Erebia neleus Frr. und Erebia aquitania Frhst. (Lep. Satyride)” [in German]. Ent. Tidskr., vol.81: pp. 77-90, 4 pls., 8 figs. 1960. Carefully distinguishes these 2 spp. on material from Mt. Blanc region, pointing out differences in habit & genitalia. Notes on application of names in this difficult group. & on E. tyndarus & E. cassioides (sensu Warren) from the same locality. [P. B.]
~Moucha, Josef, “Pieris napi L. und Pieris bryoniz O. im Vihorlat-Gebirge (Lep.
Pieride )” [in German; Czech summary]. Acta ent. Mus. nationalis Prage, vol.32: pp.91-100. 1958. The pooulation of P. bryoniz from Vihorlat Mts. (E. Slovakia ) is described. [J. M.]
Munroe, Eugene, “Hampson’s Schcenobiinz (Lepidoptera: Pyralide).” Proc. 10th internat. Congr. Ent., vol.l: pp.301-302. 1958. Redefines subfamily and _ lists included genera, those which must be transferred to other subfamilies, and some of uncertain position. Sinks Neobanepa aglossides to Pyralopsis peruviensis. Erects new subfamily MIDILINZ, for genera Midilo (=Singamia, Tetraphana), Cacographis, Gonothyris, Dismidilo; Hositea. Discusses relationships of several groups of Pyralidae. [P. B.]
Musgrave, A., “Some butterflies of Australia and the Pacific. Family Danaide— danaids II.” Austral. Mus. Mag., vol.9: pp.309-314, 9 figs. 1948. Discussion of Tellervo and species groups of Euplaa; some figures & descriptive notes. [P. B.]
Norman, T., “A new butterfly from Assam.” Journ. Bombay nat. Hist. Soc., vol.55: pp.180-181, 1 fig. 1958. Describes as new Ypthima cantliei (Margherita, upper Assam). [P. B.]
78 Recent Literature on Lepidoptera Vol.16: no.1
D. VARIATION AND GENETICS
Bernardi, G., “Le polymorphisme et le mimétisme de lHypolimnas dubia Palisot de Beauvois (Lep. Nymphalide)” [in French]. Ann. Soc. ent. France, vol.128: pp.141-158, 2 pls. 1959. Study on the polymorphism and mimicry of this nymphalid in Africa and Madagascar. [P.V.]
Boursin, Ch., “Note au sujet de l’article de M. Yves de Lajonquiére, ‘Deux formes nouvelles de lépidoptéres hétérocéres’” [in French]. Bull. mens. Soc. linn. Lyon, vol.28: p.151. 1959. 2 form of Scotia crassa named by de Lajonquiére was pre- viously named by Wagner. [P.V.]
Bowden, S. R., “Sexual mosaics in Pieris.” Lepid. News, vol.12: pp.7-13, 1 fig. 1958.
Bowden, S. R., “Yellow-green larve of Pieris napi L.” Entomologist, vol.91: pp.110- 111. 1958. Simple recessive mutant. [P.B.]
F. BIOLOGY AND IMMATURE STAGES
Kalshoven, L. G. E., “Twee soorten cacaomot in Nederland” [in Dutch; English summary]. Cacao, Chocolade en Suikerwerken, vol.22: no.10: 14 pp. 1954. Biology of Ephestia cautella & E. elutella in stored cacao. [P. B.]
Kasy, F., “Die Raupe von Gnophos stevenaria Bsd. (Lepidoptera, Geometride )” [in German; Serbocroatian summary]. Fragm. balcanica, vol.1: pp.115-116. 1954. Describes larva, reared on Prunus spinosa. [P. B.]
Kasy, F., “Kleinschmetterlingsraupen als Bewohner der von der Weidenholzgall- miuicke (Helicomyia saliciperda Duf.) befallenen Weidendste und -stammchen” [in German]. Ent. Nachrichtenbl. osterr. und schweizer Ent., vol.6 (Sonderheft 1954): pp.1-4. 1955. Notes on biology of Euzophera bigella, Laspeyresia legum- inana, L. corollana, Stenolechia albiceps, Chrysoclysta bimaculella, Trochilium vespiforme; all reared from Salix branches stimulated to luxuriant growth by attack of a gall midge. [P. B.]
Kasy, F., Uber die Jugendstadium und Bionomie von Athetis lepigone Moeschl. (Lep., Agrot.)” [in German]. Zeitschr. wiener ent. Ges., vol.41: pp.289-297, 17 figs.; vol.42: pp.4-5. 1956, 1957. Describes early stages & phenology; larva poly- phagous (numerous plants tested & preferences recorded); young larve refused fresh leaves of Taraxacum but ate dead leaves. [P. B.]
Kasy, F., “Neues tiber Chondrosoma fiduciarium Anker” [in German]. Ent. Nach- richtenbl., Vienna, vol.4, no.4: pp.1-2. 1957. Describes habitat; notes on biology. Pupa susceptible to dessication, but can also take up water from damp soil. Larvz fed on Taraxacum, Lactuca, Solidago, & Achillea. [P. B.]
Kato, Akira, “Hibernation of Colias hyale poliographus Motschulsky ( Pieridz )” [in Japanese]. Shin Konchu, vol.4, no.13: pp.18-19. 1951. Pupal hibernation sug- gested. [T. I.]
Kauffman, Guido, “Beobachtungen tiber die ersten stande einiger alpiner Pyrginze” [in German]. Zeitschr. wiener ent. Ges., vol.65: pp.23-28. 1954. Notes on early stages of Pyrgus cacalizx, P. carling, P. malvoides, & P. alvenus. [P. B.]
Kawada, Akira, “Key to the pupz of Saturnide” [in Japanese]. Shin Konchu, vol.5, no.2: pp.23-26, 13 figs. 1952.
Kawada, Akira, “Key to the pupx of Sphingide” [in Japanese]. Shin Konchu, vol.5, no.o: pp.39-47, 83 figs. 1952.
Kawazoe, Akito, “On the eggs and the young caterpillars of Melitza diamina pro- tomedia Ménétries” [in Japanese; English summary]. Trans. lep. Soc. Japan, vol.7: pp.26-28, 4 figs. 1956. Describes & figures these stages; foodplant Patrinia scabio- sefolia. [P. B.]
Keji, Joseph A., “Oviposition observations.’ Lepid. News, vol.5: p.69. 1951.
Kelsey, J. M., “Imsects attacking tussock.” New Zealand Journ. Sci. Tech. (A), vol.38: pp.638-643. 1957. Of the fourteen species of Lepidoptera listed as pests
eS
1962 Journal of the Lepidopterists’ Society 79
of New Zealand Tussock grasses (Festuca and Poa spp.) at altitudes of 1,500 to 4,500 ft., species of Crambus, Persectania, & Leucania are stated to be the most important. [I. C.]
Kelsey, J. M., “Virus sprays for control of Pieris rape L.” New Zealand Journ. Sci. Tech. (A), vol.38: pp.644-646. 1957. Experiments to control P. rape in New Zea- land by spraying larve with suspensions of undetermined virus described. [I .C.]
Kershaw, R. C., “Notes on the Army Worm.” Victorian Nat., vol.73: pp.76-77. 1956. Records larve of Persectania ewingii (Hadenine) attacking pear fruits on trees in West Tamar, Tasmania. [I. C.]
Kershaw, S. H., “Phragmatobia fuliginosa L. 6 6 attracted by scent of 2 Panaxia dominula L.” Ent. Rec. & Journ. Var., vol.65: pp.219-220. 1953.
Kettlewell, H. B. D., “A further case of similar assembling scents.” Entomologist, vol.88: p.19. 1955. Xanthorhoe montanata 6 4 to Q Biston betularia [P. B.]
Kettlewell, H. B. D., “The breeding of Coscinia cribraria Linn.” Ent. Rec. & Journ. Var., vol.64: pp.106-107. 1952.
Kettlewell, H. B. D., “The pupation habits of Gonimabrasia tyrrhea Cram. (Lep. Saturniide ).” Ent. Rec. & Journ. Var., vol.65: pp.118-119. 1953. Larva of this South African sp. burrows into sand & pupates within larval skin. [P. B.]
Kettlewell, H. B. D., “Brood size and dispersal in Biston betularia L. (Lepidoptera ).” Entomologist, vol.88: p.50. 1955. Reports 1100 pupe reared from single mating. Suggests that young larve disperse by silk ballooning, avoiding concentration; larve cryptic, not social. [P. B.]
Kettlewell, H. B. D., “Xanthorhoe montanata and X. spadicearia assembling to females of Biston betularia.” Entomologist, vol.89: p.130. 1956. Suggests similarity in 2 scent. [P. B.] :
Kettlewell, H. B. D., “Lithophane lapidea Hiibner (Lep. Noctuide) breeding in Britain.” Entomologist, vol.90: pp.1-8, 1 pl. 1957. Summarizes knowledge of early stages (foodplant Cupressus); discusses distribution & method of arrival in Britain (probable recent immigrant ).
Kettlewell, H. B. D., “Lithophane lapidea Hubner (Lep.: Noctuide) breeding in Britain. A foreword.” Entomologist, vol.90: pp.285-286, 1 pl. 1957. Breeding notes & figures of larve. [P. B.]
Kettlewell, H. B. D., E. A. Cockayne, & C. N. Hawkins, “Thalera fimbrialis Scopoli in England.” Ent. Rec. & Journ. Var., vol.65: pp.305-308. 1953. Description of larva & pupa & account of larval habits; reared on Achillea millefolium. [P. B.]
Kettlewell, H. B. D., & G. Haggett, “The life history of Hydracia hucherardi Mabille (Lep., Agrotide ).” Entomologist, vol.88: pp.217-223, 2 pls., 1 fig. 1955. Describes rearing in captivity; discovery of larvae & pupz in nature; & full-grown larva & pupa. Foodplant Althza officinalis. [P. B.]
Khan, Mohammad Quadiruddin, “Note on life history and habits of castor-semi- loopers.” Proc. 33rd Indian Sci. Congr.: p.122. 1947. Abstract; on Achza janata, Paralellia algira.
G. PHYSIOLOGY AND BEHAVIOR
Berger, Richard, “Fang der Birkeneule Brephos parthenias’ [in German]. Ent. Zeitschr., vol.69: pp.14-15. 1959. Notes on adult behavior. [P.B.]
Bergerard, J., Ph. Dreux, & J. Fiszer, “Influence des ions alcalins et alcalino-terreux sur l’automatisme cardiaque de deux insectes: Gryllulus domesticus L. (adulte) et Galleria mellonella L. (Jarve)” {in French]. Arch. Sci. physiol., vol.4: pp.225-240, 5 figs. 1950.
Bittner, Alfred, “Besonderheiten der Bewegungsformen der Wachsmotten als Beispiel fiir die Untersuchung von Insektenlaufspuren” [in German]. Naturwissenschaften, vol.46: p.119, 3 figs. 1959. Suggests that erratic running of Galleria & Achroia
80 Recent Literature on Lepidoptera Vol.16: no.J
mimics dance of host bees and so protects moths. Movement studied by recording tracks. [P.B.]
Blaustein, Mordecai P., & Howard A. Schneiderman, “The effect of barbituric acid and its analogues on the adult development of giant silkworms.” Anat. Rec., vol.132: p.415. 1958. Abstract; Callosamia & Antherea.
Blaustein, Mordecai P., & Howard A. Schneiderman, “The effect of certain antimetabolites and hydrolytic enzymes on the adult development of giant silk- worms.” Anat. Rec., vol.132: p.416. 1958. Abstract; Callosamia promethea.
Blest, A. D., “The evolution of protective displays in the Saturnioidea and Sphingide (Lepidoptera )” [in Englisiu; German summary]. Behaviour, vol.11: pp.257-307, 22 figs. 1957. Study of behavioral response to stimulation in adults of 35 satumiids and 6 sphingids. Behavior is classified and correlated with type of coloration. It is suggested that displays (but not cryptic behavior) evolved from preparatory flight movement. [P.B.]
Blest, A. D., “Central control of interactions between behaviour patterns in a hemileucine moth.” Nature, vol.184: pp.1164-1165. In Automeris aurantiaca the duration of a rocking response to stimulation is proportional to the duration of a preceding flight. This relationship is not affected by suppression of afferent impulses (removal of parts of the body, destruction of sense organs), and must be controlled in the central nervous system. [P.B.]
Bonnemaison, L., “Elimination de la diapause chez la noctuelle du chou (Mamestra Brassice L.) (Lep. Noctuide)” [in French]. Bull. Soc. ent. France, vol.65: pp.73- 78, 4 figs. 1960. Reports experiments on the effects of specific low temperatures at specific durations in breaking diapause of pupz. [P.B.]
I. MISCELLANY
Brady, Philip, “Boarding house for butterflies.” Nature Mag., vol.51: pp.188-190, 3 figs. 1958. Report on establishment of Massachusetts Audubon Society at South Lincoln, for rearing and studying butterflies. [P.B.]
Bruce, W. G., “Bible references to insects and other arthropods.” Bull. ent. Soc. Amer., vol.4: pp.75-78. 1958. Includes references to caterpillars and moths. [P.B.]
Christ], Ott, “Entomologische Forschungsarbeit in Linz’ [in German]. Zeitschr. wiener ent. Ges., vol.43: pp.187-206. 1958. Brief history of entomology of Linz, Austria, from time of Schiffermiller to present; list of publications of entomologists of Upper Austria since 1945. [P.B.]
Collins, D. L., & D. P. Connola, “Some recent research on forest insects in New York State.” Proc. 10th internat. Congr. Ent., vol.4: pp.335-340, 1 map. 1958. Includes notes on research on lepidopterous pests, especially Malacosoma disstria. [P.B.]
Dexter, Ralph W., “Packard’s final arrangement for publication of his monograph on the bombycine moths.” Lepid. News, vol.11: p.206. 1958.
Fontes, Elena Martinez, & Juan Jose Parodis, Guia de Naturalista Sudamericanos. 138 pp., Buenos Aires. 1949. See notice in Lepid. News, vol.12: p.56.
Frederick, A. C., “John Howley Cook, 1878-1958.” Lepid. News, vol.12: p.207. 1959.
Gibson, William W., & José Luis Carillo S., “Lista de insectos en la colleccion entomologica de la Oficina de Estudios Especiales, S.A.G.” [in Spanish & English]. Folleto misc. Sec. Agr. Ganad., Mexico, no.9: 254 pp. 1959. Records all specimens, with brief data; numerous Lepidoptera. Collection in Mexio City. [P.B.]
Glen, Robert, et. al., “Entomology in Canada up to 1956. A review of developments and accomplishments.” Canad. Ent., vol.88: pp.290-371. 1956. History of entomology, especially applied entomology; this summary was prepared for the 10th International Congress of Entomology by various government and university entomologists in Canada. [P.B.]
Henne, Christopher, “Charles Henry Ingham (1904-1957).” Lepid. News, vol.’ pp.169-170, 1 fig. 1958.
EDITORIAL BOARD OF THE JOURNAL
Editor-in-Chief: Cuar.es L. REMINcTON Associate Editor
(Literature Abstracting) : PETER F. BELLINGER Associate Editor
(“Especially for Collectors”): FreD T, THORNE
Harry K. Crenco —N. S. Osraztsov — EuceNE G. MunRoE
Editor, News of the Lepidopterists’ Society J. W. TrmpENn
Manager of the Memoirs: Smpney A. HEssEL
NOTICE TO CONTRIBUTORS TO THE JOURNAL
Contributions to the Journal of the Lepidopterists’ Society may be on any aspect of the study and collection of Lepidoptera in any part of the world. Papers of more than twenty pages will not normally be accepted, but if they are, authors may be required to pay for overage.
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Titles must be kept as short as possible; Latin names of genera and species will be italicized, and authors of such Latin names WILL NOT APPEAR IN THE TITLE of any paper but may appear once in the text. The title should indicate the family of the subject. The style should conform to that used in recent issues of the Journal. PLEASE NOTE EXACT STYLE FOR REFERENCES. Footnotes should be kept at a minimum. The editors reserve the right to adjust style to fit standards of uniformity.
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Printed in U. S. A. By Tue S. Z. Fretp Co., Inc. New Haven, Connecticut
1962 Journal of the Lepidopterists’ Society Vol.16: no.1
TABLE OF CONTENTS — SOMMAIRE — INHALT
Taxonomic notes on Erora (Lycenidz)
DY, KMLIAN ROEVER 660.005.5056. yececetescesslecarcseadeet a er | 1-4 Evolutionary mechanisms of the Pieris napi — bryonie group [first part]
By. Ze LORROVIG oles ccsscacscasnc ewveleiinacendenn 5-19 A biting midge ectoparasitic on Arizona lycenids
by (PAUL Re vBBRLICH .0...eccsccjendepestiseocsstsikiiinscel oie ee 20-22, Ecological studies of Rhopalocera at Donner Pass, California. JI.
by. Tuomas C. & Joan F. EMMEG ooo... ccc cescie ses cnee 23-44 Authorship of names of Papilio credited to Boisduval or Lucas
by; Cyan. F. ps PASSOS ©.0....6..5:5.cccccccassessedeneeseseeestet sie oe 45-46 Studies of Neotropical Lepidoptera. I. Life history of Dyops
by BoP. Wan rsHme: ooo... isecelessdocstitnotses eile 47-54 Preliminary notes on the life history of Callophrys (Sandia) macfarlandi
by D. B. Srauuincs, J. R. TuRNER, & P. R. ESRLICH ...............0..0000 55-57
REVIEWS: Collins & Weast, Wild silk moths of the United States, by C. L. Hocus; Hayward, Guia para el entomologo principiante, Dy) Cx Ly RENGINGTON oo. loisescee Ob ccesedec csc docddevscervanswduet ceed e 22, 58
FIELD NOTES Satyrium behrii (Lycenidz) in Oregon
DY REO RS CENCE 6.5005. Solce oes isc sake co! anceuee ssddetaseeasbbencedsatusteenace een 44 Colias nastes in the U. S. A.
| 2) atin BD G GRKS) = 1) of 01 0 LAR OS a RPE UO OC AUMSRSMIR NE RCEICN 46 Albino Lycena phleas in Connecticut
BV PDR soe oo. Sc ub nesian cczeu hheoetasedcssgu0s ane nae ear 59 Whitish Lycena phieas in Ohio
Vat teal MENTE NG Oy er ee a aaa) Aomisésstes en 59-60 Larval notes on Chlosyne iacinia and californica
DY. Pe ROBIE os oss bk clos eoteeesccs sav ovsacceat iaancoadas shila rr 61 Foodplants of Southwestern Saturniide
BS eRe RS NVR Ee Leh aslak cous cane cecck owns cateaes nace ene ee cr 61-62 Adopea lineola in northern Ontario
DY Ps Ba NET a oes oo ias Uns esezeennsiekoapeoesonedtbeess nen 62 Migration of Nymphalis californica in Washington
By UWUG EAs RGGI or) ocak cs nue ciel cepsisnaneae da each un teary aci eae ear 63
ESPECIALLY FOR FIELD COLLECTORS Collecting Gneis, etc. on Vancouver Island, with a theory on hilltopping
by FRIGHARD GGUPEY bios ncce recs canes ac econtva gene dence ecesbactseed uate eee 64-66 Collecting is still good in the Northwest By (Bla Je INWCOMER ios os cea eetaatonadecnde este sade tenses 67-70 Biographical obituary of Edwin P. Meiners (1893-1960) by Py Sermon | JRENEINGTON 246....5.....42. sere dsavctvcandseitesdteeed ee 71-7 CO Wer Pa TEISE i Re ee ae ERLE a UE CO, oe sae clu Sa ae Aa ee Ae 66 RECENT: LITERATURE. ON LEPIDOPTERA: 0.000.000. cecpeestca es ene 76-80
Volume 16 — 1962 Number 2
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
In This Issue
OVERLAPPING SUBSPECIES IN SPEYERIA ZERENE OVIPOSITION ON TOXIC PLANTS ANALYSIS OF PIERIS NAPI & BRYONLZ IN EUROPE NOTES ON PAPILIO INDRA MINORI
(Complete contents on back cover)
5 December 1962
THE LEPIDOPTERISTS’ SOCIETY 1962 OFFICERS
President: Joun A. Comstock (Del Mar, Calif., U. S. A.) Ist Vice President: H. B. D. KETTLEWELL (Oxford, England) Vice President: Himosut INoveE (Fujisawa, Japan) Vice President: TARSICIO ESCALANTE (Mexico City, Mexico) Treasurer: GrorcE EHLE (Lancaster, Penna., U. S. A.) Asst. Treasurer: SmwNEY A. HEsseu (Washington, Conn., U. S. A.) Secretary: Paux R. Euruicu (Stanford, Calif., U. S. A.) Asst. Secretary: FLoyp W. Preston (Lawrence, Kansas, U. S. A.) EXECUTIVE COUNCIL Terms expire Dec. 1962: Harry Krocerus ( Helsingfors, Finland ) F. M. Brown (Colorado Springs, Colo., U. S. A.) Terms expire Dec. 1963: J. C. E. Rrotre (Toronto, Canada) ‘Daxrpor Povotny (Brno, Czechoslovakia) Terms expire Dec. 1964: H. A. FREEMAN (Garland, Texas, U. S. A.)
S. L. DE La Torre y CALLejas (Havana, Cuba) and ex-officio: the above six elected Officers and the Editor-in-Chief
The object of The Lepidopterists’ Society, which was formed in May, 1947, and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches,...... to issue a periodical and other publications on Lepidoptera; to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed toward these aims (Constitution, Art. II). A special goal is to encourage free interchange among the lepidopterists of all countries.
Membership in the Society is open to all persons interested in any aspect of lepidopterology. All members in good standing receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer the full dues for the current year, together with their full name, address, and special lepidopterological interests. All other correspondence concerning membership and general Society business should be addressed to the Secretary. Remittance in dollars should be made payable to The Lepidopterists’ Society. There are three paying classes of membership:
Active Members — annual dues $6.00 (U. S. A.) Sustaining Members — annual dues $15.00 (U. S. A.) Life Members — single sum $75.00 (U.S. A.)
Dues may be paid in Europe to our official agent: E. W. Classey, 353 Hanworth Road, Hampton, Middx., England.
In alternate years a list of members of the Society is issued, with addresses and special interests. All members are expected to vote for officers when mail ballots are distributed by the Secretary annually. There are four numbers in each volume of the Journal, scheduled for February, May, August, November, and eight numbers of the News each year.
The Lepidopterists’ Society is a non-profit, scientific organization. The office | of publication is New Haven, Connecticut (see address inside back cover). Application for Second-class mail privileges has been approved at New Haven, Connecticut.
oun? oF
Tue Lepiporprerists’ SOCIETY
Volume 16 1962 Number 2
NOTES ON OVERLAPPING SUBSPECIES. I. AN EXAMPLE IN SPEYERIA ZERENE (NYMPHALID:)
by L. P. Grey and A. H. Moeckx
During the 1959 and 1960 seasons Moecx collected series of Speyeria zerene (Boisduval) in and near the Warner Mountains, Modoc County, California, demonstrating a geographical overlap of two radically dif- ferent subspecies and thus reinforcing a hypothesis of relationship hitherto deduced more indirectly.
For background, illustrative material appended includes a resume (table 1) of subspeciation accredited to zerene, and a chart (map 1) indicating geographical distribution of the subspecific characters; these should enable orientation of the forms to be discussed herein. Topography of the particular areas mentioned is shown by map 2; specimens selected to depict normal and extreme variation in spot localities are illustrated in figures 1 and 2. The field notes on these series will be found appended, in a concluding section.
Figure 1 is of a series of males (in ventral aspect) from an unnamed ridge about 16 miles east of Eagleville, California, lying in Washoe County, Nevada. Normal variation in that locality is represented in rows 2 and 3 of figure 1, and is referable to the subspecies of zerene termed cynna. [Note: names considered lower-than-species are used throughout as catchwords; extent of blending discourages formal cate- gorical applications. |
In a population of this sort, the upper singleton in figure 1 is remarkably abnormal, as would be more evident if seen in full color; it has well- developed ruddiness. The lower singleton has a reddish tinge; the specimens immediately above it (row 4) are largely unsilvered. Redness,
81
82 Grey & Moecx: Speyeria zerene Vol.16: no.2
and lack of silver—these are characters unheard-of in cynna throughout the large area which that subspecies preempts in central and northern Nevada, in southern Idaho and southeastern Oregon. These two par- ticular characters are, however, quite the normal thing in another zerene subspecies occurring just across the street, so to speak, from this Nevada colony.
Turning to figure 2, the three examples in the bottom row (M, N, O) will convey a good idea of normal zerene variation in the nearby Warner Mountains, i.e., running a bit darker but otherwise quite like the nomino- typical Sierran forms. The remainder of the series in figure 2, namely, the sixteen specimens A through L in the four upper rows, are from the Patterson Meadows, at the southern end of the Warners in the south- eastern corner of Modoc County. All were taken within less than a two-mile radius in substantially identical ecology and at about the same elevations (7300-7500 ft. ).
Typical zerene predominated (samples G, H, I, L) but cynna also was procured here (samples A, B, J, K). Some individuals were strikingly intermediate (samples C, D, E, F) and all the more interesting since two (D, E) were in a “congregation of cynnas” and one (F) was in association with zerene. The greatest extremes (J and L) were netted from the same flower bed although in different years.
Attempts to secure large samplings were unsuccessful, being hampered by the aridity and lack of roads. The Patterson Meadows locality is the only one as yet discovered along the east rim of the Warners where this blending is widespread and obvious although occasional singleton in- termediates have been taken in those mountains. Closer search was clued and encouraged by the gene recombinations noticed in the Washoe County series.
The rough balance of segregation vis-a-vis blending occurring in the vicinity of Patterson Meadows lends credence to a rather commonly held hypothesis, which is that the colonial and individual “pale zerene” turning up along the Sierran-Basin tension zone (the malcolmi near the Mammoth Lakes, the mixtures in the Tahoe passes, etc.) are resultants from contacts between the red Sierran and the yellow Great Basin forms. The relationships originally were deduced from cumulative gene replacements observable around a two-thousand-mile horseshoe (Moeck, 1957); present records serve to confirm opinions of the phylogeny as based on directions taken by variation in geographically arranged ma- terial. The extreme divergence of the end-products discouraged hope of ever finding intermediates; the segregation accompanying the blending is not at all surprising.
1962 Journal of the Lepidopterists’ Society
83
Fig. 1: Samples of S. zerene ssp. from Washoe Co., Nev.
data in record section, comments in text.
See locality on map 2,
84 Grey & Morck: Speyeria zerene Vol.16: no.2
Fig. 2: Samples of S. zerene ssp. from Modoc Co., Calif. See localities on map
2, data in record section, comments in text.
1962 Journal of the Lepidopterists’ Society 85
Nore: One record from Haines Alaska about as far trom Vancouver as Vancouver is from Sanfrancisco. Further records from the Transition-Canadian ¥orest coast zone may be expected.
— Cline Code ——
eetorezcscsa
Z fi i i !
| | | |
VAN Tra
cy > 0 CY- Pye PL PL oy
on ag Cree =a | e i i pi | Sj |
Map 1: Distribution of S. zerene. See table 1 for key to symbols. The shaded rectangle denotes area shown in map 2.
DIscUSSION
Mention of other geographically correlated peculiarities of zerene seems due, but is abbreviated because of a desire to expand some re- marks on problems at which this and succeeding articles are aimed.
86 Grey & MorEck: Speyeria zerene Vol. 16: no2
Washoe Co.
S iS) Ss
cS iS) ~
fle
California MOULD, =
|
Se
_ Lassen Co
Map 2: Topography of a section of Modoc Co., Calif., and Washoe Co., Nev., where S. zerene ssp. illustrated in figs. 1 & 2 were taken. See record section and text comments.
For that matter, students familiar with Western topography and with zerene probably will find all of their observations and conjectures implicit on map 1. Appearing there, for examples, are (1) the gross correlations of the four large divergencies with the four physiographic provinces; (2) the zones of stability and instability — accordingly as clines converge or overlap, and with individual variation lessening
1962 Journal of the Lepidopterists’ Society 87
toward the middle and south portions of each province; (3) the hints of refugia, viz: (a) that the Coast Range set is incongruously largely divergent, possibly from an earlier archipelago, (b) that platina likely is nearest to the earlier “heartland” interior form, garretti a recent offspring wave northerly, sinope a Wyoming Basin isolate, cynna a relatively undisturbed Snake River Plateau resident, (c) that the southern distribution (as always) is the most obscure—is carolz a relict, parent to Sierran forms, or a recent emigrant therefrom? — and why the hiatus in the Greenhorns and Tehachapi, the rarity in the Plateau country? — this insect demonstrates successes and tolerations to warrant some expectations which are disappointed southerly, so the failures probably have special significence; (4) the area of greatest plasticity, in the Pacific Northwest, as is inevitably predicted by the cornering of the four zones in that region and from the ideas of glacial retreat which indicate a recency of the contacts. Now, to the Modoc-Washoe County material: this, almost surely, exemplifies intraphyletic segregation. Herein, and in following papers, the writers will point to instances, mostly in Speyeria, of apparent partial or total segregations occurring within the distributions of varieties presumed related to the extent of belonging to single phyletic lines —or, if defined in a quibbling spirit, belonging within a single “species” as best that category can be applied with present knowledge. Overlaps of this nature are engaging attention of researchers in a number of fields of evolutionary inquiry, for it is believed that such collisions of isolates are likely to provide rich data for genetical, cytological, paleogeographical and other studies. Questions raised by these occurrences seem almost endless; some will be noted in later papers; in this discussion attention is drawn to three aspects, to wit: (1) the taxonomic, (2) the geographical (correlations with environment and barriers) and (3) the paleogeographical — which belongs closely bracketed with (2) because of uncertainties as to when the divergencies within population structures have been acquired. Notations of overlaps such as these are obligations falling largely on taxonomists. It is they who presume to name and index species, from which listings the studies of evolution take their directions. Some of the conceptual modifications brought on by discoveries of taxonomists ( é.g., intraphyletic segregations — wherein species fragments differ to the extent of appearing locally discrete, and siblings — wherein convergence is uppermost, so that specific distinctness may be thoroughly masked. lying more in biology and behavior than in facies ) obviously have affected evolutionary thinking. It is quite certain that more local anomalies of such caliber remain to be discovered, and that existing classifications will
Vol.16: no.2
Speyeria zerene
Grey & MOECK:
88
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90 Grey & Morck: Speyeria zerene Vol.16: no.2
be set up by them. Evolutionary conceptions draw their warrants from the prevailing editions of “species”; taxonomy provides the impetus.
The cynna - zerene overlap falls within boundaries of credibility as established in the 1945 dos Passos and Grey catalogue of Speyeria, wherein cynna is listed as a subspecies of zerene. A regrettable error in that index is corrected hereby: the carole population in southern Nevada pretty surely belongs within zerene and not under coronis (Behr) where it was described originally.
Present taxonomic worries therefore seem unnecessary, except as all such relegations have to continue passing the yearly tests of new dis- coveries and changing concepts. The Speyeria classification mentioned was framed deliberately to reduce species to the utter minimum allowable against sympatrisms, allopatrisms and intergradings as then known. Future revisions probably will be more of concept than of groupings: it will appear that those who have catalogued Limenitis, for one example, make more species than the present speyerian philosophy would admit, and so it goes; in various genera some will lump, while others, familiar with the same data, will claim that lumping is not the best answer.
Some valuable taxonomic moralizings probably can be drawn from study of intraphyletic segregations within Speyeria; the situations de- scribed assuredly will be found duplicated, substituting places and species, in genera such as Papilio, Euphydryas, Cercyonis, etc., which seem now on the verge of long-awaited syntheses. It may be helpful to emphasize, as is the intention, that overlaps can be obscure in many ways and in sophisticated fashions, almost sure to be overlooked unless the fund of distributional knowledge is at once broad and detailed. Some hybrid zones shout their presence and are matters of common knowledge, as in Northeastern Limenitis and Colias — even then they may baffle taxonomists intent on pouring all of the new wine into the old skins of Linnaean concepts — but others may be entirely unsuspected because with data as skimpy as ordinarily is available the early stage of hypothesis would have to be made in defiance of the very cornerstone tenets of the species-makers, for these reasons:
The usual picture of subspecies is of geographically oriented divergencies which blend with other intraspecific forms along sharp or ill-defined boundary lines. But divergence in plastic genera can be extreme to the point of incredibility. Intergradings which would clue relationships may be slightly or not at all in evidence, perhaps confined to poorly explored areas or even lost by extinction. What, then, if two great extremes work into contact? Local samplings are likely to come up with two very different forms, each apparently discrete. By con-
1962 Journal of the Lepidopterists’ Society 91
ventional standards these indeed are species. Local naturalists must be allowed their word of truth: sympatry plus evident biogenetic differentia- tion spells out “species” quite positively. But students of population structures also have a right to be heard: if locally discrete sympatres display a gradual convergence when traced geographically into other regions, ending in an area or areas where blending is conspicuous or total, then a phyletic wholeness is manifested, which it is incorrect to claim for any of the parts.
Arguments and rebuttals of this sort will be heard increasingly as distributions are studied more intensively. Even now, the quirks of speciation and subspeciation known in butterflies compare favorably with those recorded for other groups of animals, and they will be scanned even closer in the future by students of evolution. Systematic taxonomy of butterflies therefore is in need of continued scrutiny; there are new frontiers; they lie, however, where they always have, i.e., with segregations and overlaps. The concepts and tools need refining to deal with complexity hitherto suspected and now being confirmed.
A second arresting feature of intraphyletic overlaps, in addition to their challenges to taxonomy, are their morphological dissimilarities which so obviously flaunt the hypotheses built around ideas of variation re- sponding to local ecogeographies. There has been a great deal of research along the latter lines, prompted by convergencies and parallels seen not only within closely related genera but also among forms known to be well apart phyletically. As an example of explanations which have been sought in particular factors of a shared environment the work by Hovanitz (1941) is a classic of its kind.
Parallels within Speyeria, as noted by Hovanrrtz, lend themselves irresistably to evolutionary speculations, being numerous to the point of endlessness, often involving very different species (and thus throwing heavier burdens on taxonomists quite sufficiently beset with anatomical likenesses and ecological overlaps ). Hovanrtz’s correlations of melanism with insolation stand unanswerable. He makes no further special claim that environment has the paramount role in determining variation, as some others have done with less excuse, or have so implied by pointing out the many geographically correlated parallels in various genera.
Recently (1959), Van SoMEREN and Jackson have summarized some ways in which parallels are supposed to evolve consequent from various kinds of mimicry, including arithmetic mimicry which certainly ought to apply in Speyeria to the advantage of locally scarce species; in these cases the convergence results from a “dwelling together” but has nothing directly to do with geography. :
92 Grey & MoeEck: Speyeria zerene Vol.16: no.2
Such theories have the attraction of offering plausible rationalizations of wide applicability. They have a drawback of seeming to finalize “causes’ which almost surely are incredibly multi-factorial however plainly one element may appear to stand out.
Dissimilarities evidenced in intraphyletic segregations give salutary check to environmental postulates. For example, cynna and zerene are at the very antipodes in coloration, and yet they face one another in similar ecologies across narrow barriers — and now turn out to fly in the same meadow. In such instances, it may be inferred that properties which are intrinsic (incompatabilities based in physiology) outweigh all extrinsic things such as geographical propinquity and identity of ecologi- cal background.
In a later paper attention will be directed to a segregation which really may have resulted in large part from differential adaptability to micro-environmental factors, as perhaps can be found upon close analysis in the other overlaps of isolates. Still and all, it hardly seems tenable to suppose that segregations such as in Appalachian Limenitis can be explained as being due to any sharp breaks in climate or ecology. There, and most likely in the great majority of overlaps, the better rationaliza- tions seem to lie with genetics, with time, and with paleogeography.
The time element undoubtedly is the great enigma underlying the questions with which evolutionarily oriented taxonomy grapples. How long does it take, what particulars of environment are needed to cause species to diverge? Once established, how stable then are the gene systems thus integrated harmoniously to place, time and species? As the years bring changes, as populations migrate with climates, and as balances are struck between selection pressures and mutation rates, what then does local variation reflect of immediate situation and of earlier history?
A sharp dichotomy runs through the whole literature and theory of evolution: On the one hand, a multiplicity of “causes” are adduced which logically ought to lead only to chaos, the supposed mechanisms and modifiers being varied and acting at random; on the other hand, the natural order reflects crystallization into definite patterns down to such trivialities as that certain subspecies of butterflies occupy territory to the exclusion of others. Can it be that the “causes” by their very infinity give statistical expectation of orderliness in resultants, as in the modern “laws” of physics?
This touch of paradox is delightfully exposed in Forp’s (1954) Butter- flies, where mention is made of a strain of Euphydryas which was observed to subspeciate markedly in less than three decades—followed
1962 Journal of the Lepidopterists’ Society 93
later by a chapter discussing Pleistocene population movements as