A MIGRATION PROBLEM - VANESSA CARDUI (NYMPHALID.tE), THE PAINTED LADY BUTTERFLY. by CHARLES H. ABBOTT

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1 1962 Journal of the Lepidopterists' Society 229 A MIGRATION PROBLEM - VANESSA CARDUI (NYMPHALID.tE), THE PAINTED LADY BUTTERFLY by CHARLES H. ABBOTT The subject of insect migration, and particularly of butterfly migration, has reached the point where certain laws of migration are beginning to emerge. We are also learning that exact information is lacking on many points which we have taken for granted, and that when we secure this information we shah be in a better position to formulate laws. As a first basis we must admit, as WILLIAMS (1958) has so many times pointed out, we cannot arbitrarily divide migration into rhythmic migration and one-way migration (also called emigration or, better, unidirectional migration with no return). Only one butterfly, the Monarch, Danaus pzexippus (L.) is usually considered to have a typical rhythmic migration, in which the same individuals make a two-way flight; while WILLIAMS has collected data showing return flights by successive generations in an increasing number of species. Therefore, if two-way and one-way insect migrations are essentially different, we do not have enough knowledge to divide the butterflies between them. Our problem is to find some underlying principles or laws, and there will be time enough to determine how wide can be their application. As has been previously pointed out, the Painted Lady, Vanessa cardui ( Linmcus) was selected for this study, because in migration years it is so abundant that there is limitless material on which to work, and because its study has been neglected. The conclusions to date may be found in the author's 1950, 1951, and 1959 papers, and may be very briefly summarized as follows: The species is usually found, so far as North America is concerned, only south of the United States, or possibly, in southernmost Imperial Valley; and comes north only in migration years, which are few in number and at irregular intervals (with possible minor exceptions to be noted later). If one could observe the start of a migration at the point of origin, probably large numbers would be noted all at once, but at points in California very small numbers appear at first with a gradual build-up. The direction of migration is consistent, year after year, in most California localities toward the north-northwest. The geographical extent of the migration in anyone year appears correlated with the numbers involved, the larger the numbers the greater the distance covered by the species. It was determined in 1958, when the migration was traced eventu-

2 230 ABBOTT: migration Vo1.l6: noa ally from the Mexican border to the Cascades of Washington, that several generations were involved, probably four in California, and the number farther north undetermined. This makes one wonder if each generation found an optimum of conditions in the area which it reached and where it laid eggs, and that such a combination of optima over the entire Pacific Coast area rarely occurs. I believe we have already the most complete picture available (of course far from complete) of the migration of a single North American butterfly, except probably the MoriarcliTDanaus plexippus); and from what WILLIAMS says in his latest book Insect Migration (1958), 1 do not believe as complete a picture is available of any species in Europe. What I want to outline now is particular observations which are needed to fill in gaps in our knowledge, which are practicable for field lepidopterists. As a reason for doing it, I want to quote a paragraph from page 2 of WILLIAMS'S Insect Migration (1958): "We can ask for example why Painted Ladies migratc; this is a problem of the origin and evolution of the habit. But we have also to consider the much more detailed problem of how or why a particular individual butterfly is migrating in a particular directign at a particular time. In my opinion it is by finding answers to the second that we may be able to solve the first of these questions." l. What is the distribution of this species in western North America in a non-migration year? WILLIAMS (1938, 1958) has rcported, on the basis of published reports from this part of the country, that the species is found north of Mexico only in years of extensive migrations. There seems to be little doubt that the source of our great migrations is south of the Mexican border (Abbott, 1959), although little is known of the exact location or nature of the breeding grounds. One in Baja California, discovered by GARTH and DAWSON in 1949, has been described (Abbott, 1951: p.159). The above statement appears to be open to exceptions, as there are several unpublished reports by competent lepidopterists of small resident populations in California and Nevada which have been observed year after year. There is a difference of opinion whether these populations take part in a general migration. It is important that information on these resident populations should be published, particularly as to the number of generations per year, the length of each stage of metamorphosis, and what stage is inactive in a carry-over from one year to the next. As a possible explanation of this last paragraph, it was suggested by P. F. BELLINGER (personal communication) that V. cardui might have

3 1962 Journal of the Lepidopterists' Society 231 phases such as those described for locusts, a migratory phase, associated with overcrowding, and a non-migratory phase, called solitary in locusts. (For discussion of UVAROV'S Phase Theory, see Williams, 1958: pp This discussion also reports similar phases in certain moths.) It seems to be more difficult to determine if there is an inactive period in the annual life history of the species in its home in Mexico whence the migrations periodically come. The key point which 'needs to be determined, and is also the most difficult to find out, is what combination of conditions starts a species migrating. Is it a set of conditions which work directly on the adult insect, or is it a combination of favorable conditions resulting in a larger number of eggs laid, a larger number of larvre developing to the pupa stage, or a larger number of pupre surviving to emerge as adults? 2. The only known set of external conditions which appears to favor a very large generation of V. cardui in the desert or semi-desert south of the border is the right combination of rains and temperature to give an unusual crop of desert vegetation, chiefly annual wild flowcrs. This serves as food for the larva.~, and could result in the next generation of egg-laying and potentially migrating adults being much larger. This makes extremely important the recording of all possible data on rainfall, temperature, and vegetation in subsequent migration years; also facing the fact of the great variation in conditions in different parts of the desert. The past data on this point are rather scarce and inconclusive. The question which needs to be answered is: Is a heavy migration aliways correlated with heavy winter rains in the desert at the right season to produce a large crop of desert annual wild flowers? Data available at present make a "yes" answer to this question doubtful, but the data are inconclusive (Abbott, 1951: p.162.) If this is a correct conclusion, it is probable that large crops of annuals at points somewhat farther north from the Mexican border would help to explain the very large second and third generations such as were noted in But what determines the large first generation in a migration year? Is it a combination of conditions having a sudden effect on some stage of development? Or does a migration result only when there has been cumulative build-up of nnmbers during preceding generatiom, perhaps since the last preceding migration? 3. An alternative theory, which deserves investigation, is that the instinct to migrate is always present in this species or at least in the migratory phase of this species. In non-migration years they are too few and fly too short distances to be observed by any except trained

4 232 ABBOTT: migration Vo1.l6: no.4 observers who happen to be in exactly the right localities. This would explain the report by R. C. DICKSON that a small migration occurs in Imperial Valley every year. The following is a suggested pattern for a non-migration ycar. At the beginning of the season only a small number of imagoes transform from puplb, and being few in number they do not have to fly far to find vegetation on which to lay eggs; or, owing to a very short growing season of plants, only a very small number of eggs from the last preceding generation develop as far as adults of the first generation of the new season. Parasites in the usual numbers would also be more effective in controlling a small generation. Predators are usually at least one generation behind their prey in a fluctuation in numbers (Elton, 1927: p.141 ), and this may be true of the so-called parasitic Hymenoptera and Diptera, which are really midway between predators and parasites. If the instinct to migrate is always present, it would explain why in a migration year the succeeding generations, even if many miles from the base habitat and perhaps from the original cause, continue to migrate until adverse conditions stop the migration. 4. Hew long do female V. cardui live after egg-laying, and do the individuals continue migrating after egg-laying? Do males migrate the same distance as females? 5. Is the migration of a particular season stopped by any or all of the following? a. Egg-laying, causing adults to stop flying. b. Failure of new generation to develop in egg, larval, or pupal stage. Might be in larval stage, due to drying up of food plant, or in any stage, due to parasitism. We also do not know if unfavorable weather conditions may have a direct effect on any stage. c. If the adults have died after egg-laying and the next generation has failed to develop, it would explain why there is no over-wintering in the regions reached by migration, or no return flight, if there is any instinct for return flight. d. If there is ever any evidence discovered for over-wintering following migration, in a locality where this species is normally absent, it should be reported immediately. It is even more important to report any evidence for a return flight to the south. In the case of the Africa ~ Europe flyway of the same spec.ies, WILLIAMS (19.58) has found no evidence of over-wintering in the north, but several instances of southward flight in autumn. Of course these were not the same individuals which flew north, but a subsequent generation.

5 1962 Journal of the Lepidopterists' Society Is there any authentic evidence in the Pacific states for the migration of any other species of Vanessa often associated with V. cardui, or closely resembling it? I refer, particularly, to V. carye, V. virginiensis, and V. atalanta. I have found V. carye and V. atalanta feeding on the samc bush with migratory V. cardui, but have never seen either species flyaway from a bush and join a migratory swarm. Occasionally I have seen a single V. atalanta flying with migratory V. cardui. V. carye are not easily distinguished in flight. V. carye is regularly described as resident and non-migratory. WILLIAMS, et az. (1942) give two reports each for migrations of V. virginiensis and V. atalanta in the eastern United States. This paper is based on one presented before the Pacific Slope Section of the Lepidopterists' Society at Santa Barbara, California, August 26, It has outlined points on which exact information would be of help in interpreting the migration of Vanessa carciui, the Painted Lady Butterfly, and perhaps of Lepidoptera in general. Its purpose is a request for information. If any observers of the migration of this species have any past data on these points or acquire any in the near future which they do not wish to publish separately, it would greatly assist the writer if they would send them to him. Literature Cited Abbott, C. H., Twenty-five years of migration of the Painted Lady Butterfly, Vanessa cardui, in southern California. Pan-Pac. ent. 26: , A quantitative study of the migration of the Painted Lady Butterfly, Vanessa cardui L. Ecology 32: , The 1958 migration of the Painted Lady Butterfly, Vanessa cardui (Linnreus) in California. Pan-Pac. ent. 35: Elton, C., Animal ecology. 207 pp. Macmillan Co., New York. vvilliams, C. B., Recent progress in the study of some North American migrant butterflies. Ann. ent. soc. America 31: , Insect migration. 235 pp. Macmillan Co., New York. Williams, C. B., G. F. Cockhill, M. E. Gibbs, & J. A. Downes, Studies in the migration of Lepidoptera. Trans. my. ent. soc. London 92: Bay View Drive, Hermosa Beach, Calif., U. S. A.