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Journal of The Lepidopterists' Society

Volume 22 1968 Number 3

SOME ASPECTS OF MATING BEHAVIOR IN BUTTERFLIES

Lee D. Miller

Catholic University of America, Washington, D. C. 20017

AND

Harry K. Clench

Carnegie Museum, Pittsburgh, Pennsylvania 15213

Mating butterflies tend to remain nearly motionless. As a result we witness the activity infrequently and our knowledge of it remains extremely limited. Two kinds of mating data, however, are reasonably accessible to the general observer and give promise of increasing value as more records accumulate: the time of day when mating occurs; and the sex of the active (flying) partner. The first requires simply a notation of the time of day. The second is not so simple. In many cases mated pairs must be disturbed deliberately to cause them to fly. The determination of the sex of the one that flies, however, is by no means easy in those species where the sexes closely resemble one another.

For several years each of us has been noting this information as opportunity presented during the course of collecting. In January, 1966, when we went to Mexico as the Carnegie Museum-Catholic University of America Expedition, gathering additional records was on our agenda. Our accumulated observations are given in the present paper. We have added such published records as we have been able to find, but we have made no intensive literature search.

The absolute necessity for accurate reporting (which includes admission of doubt or deletion of a questionable datum) is underscored by the curious paper of Pronin (1964) on this subject. We cite his records, but we consider all of them uncertain and several, v/hich we indicate, are highly questionable. The latter (cf. Aporia crataegi, Pieridae; Danaus plexippus, Danaidae) contradict several to many others, and cast doubt on the rest of his records. This opinion is reinforced by his numerous unwarranted conclusions and unsupported categorical statements. For

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example, he gives (p. 40) a table of the mating times of ten species of butterflies, all periods of an hour or less (and no indication of how many pairs of each species were observed). Immediately following the table he states, "Thus, each species of butterfly . . . has its own definite mating time." The conclusion is certainly not legitimately derivable from his data and is, furthermore, incorrect (see records below of Fieris rapae, Euptychia cymela, Erynnis juvenalls).

In the following list we use these abbreviations: DC, District of Columbia and vicinity; PNR, Powdermill Nature Reserve (9 miles south of Ligonier, Westmoreland County, Pennsylvania); and CM-CUA, Carnegie Museum-Catholic University of America Expedition to Eastern Mexico, 1966. We include in the list only those records in which either the time of day, or the sex of the active partner, or both, is recorded, and add the data of the observation. To save space, and because the precise locality is not of critical importance, we usually give only the state or country in which the observation was made.

Papilionidae

There are only Pronin's (1964) records of Papilio machaon Linnaeus (Europe, 1-2 P.M., ? flying) and Papilio multicaudatus Kirby (California, 2-3 P.M.).

Pieridae

Records are particularly numerous in this family. Mating of pierids apparently takes place in the late morning to early afternoon (but note early morning record of protodice). The male is the active partner in nearly all records.

Fieris rapae ( Linnaeus)

Pennsylvania (PNR), 6.V.1964, 11:35 A.M. and another 3:35 P.M. (HKC).

Iowa, $ flying (LDM).

DC, $ flying (Clark, 1932)

Europe, $ flying (Pronin, 1964) Fieris napi ( Linnaeus)

Europe, $ flying (Pronin, 1964) Fieris virginiensis Edwards

Pennsylvania (PNR), 23.IV.1964, 1:20 P.M., S flying; and another 1:30 P.M., $ flying (HKC). Fieris protodice Boisduval & Le Conte

New Mexico, 21.VII.1963, 9:30 A.M. (HKC). Aporia crataegi ( Linnaeus )

Europe, 9 flying (Pronin, 1964) [record questionable] Appias d. drusilla (Cramer)

Tamaulipas (Mexico), 9.1.1966, 2:25 P.M., ^flying (CM-CUA). Catasticta n. nimbice (Boisduval)

Hidalgo (Mexico), 13.1.1966, 12:30 P.M., $ flying (CM-CUA).

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Colias eurytheme Boisduval Iowa, $ flying (LDM). DC, $ flying (Clark, 1932). Colias hyale (Linnaeus)

Europe, $ flying (Pronin, 1964). Colias croceus Fourcroy

England, August 1964, ca. 2:30 P.M. Disturbed on four occasions, the $ flying each time (M. Clifton). Eurema daira (Godart)

Veracruz (Mexico), 16.1.1966, 2:40 P.M. (CM-CUA). Eurema lisa Boisduval & Le Conte

Veracruz (Mexico), 16.1.1966, 1:45 P.M., $ flying (CM-CUA). Tamaulipas (Mexico), 26.1.1966, 12:55 P.M., $ flying (CM-CUA). Georgia, 9.IX.1966: A mated pair was repeatedly disturbed. On the first two occasions the male flew; on the third, the female; on the fourth to sixth, the male. When the female took the lead the cover may have been so dense that the male could not fly (LDM). Florida, 22.111.1967, 10:55 A.M., $ flying (LDM).

Danaidae

With the exception of Pronin's doubtful record, males are the active partner exclusively in danaids. The observations of Brower et al. (1965) are particularly significant, as they were part of an extended study of D. berenice courtship behavior and many pairs were observed.

Danaus plexippus (Linnaeus)

Iowa, VII, VIII, afternoon, several pairs, $ flying (LDM).

California, ? flying (Pronin, 1964) [record questionable].

No Locality, £ always flying (Urquhart, 1960: 52). Danaus gilippus strigosus (Bates)

Arizona, VIII. 1958, afternoon, at least 2 pairs, £ flying (LDM).

Veracruz (Mexico), 16.1.1966, 3:30 P.M., $ flying (CM-CUA). Danaus gilippus berenice (Cramer)

Florida, betw. 22,VII-11.VIII.1960 and betw. 2.VII-11.VIII.1961, betw. 2^5 P.M. (EST), many records, $ always flying (Brower et ah, 1965).

Satyridae Another well represented family. Females, without exception in the records, are the active partners, and matings take place from late morning to mid-afternoon, with a distinct predominance of records in early afternoon.

Cercyonis pegala maritima (Edwards)

Massachusetts, 27.VII.1964, 1:00 P.M. (HKC). Cercyonis pegala (Fabricius ), subspecies

Pennsylvania, 23.VII.1961, 1:30 P.M., $ flying (LDM). Euptychia cymela ( Cramer)

Pennsylvania (PNR), 28.V.1964, 12:45 P.M., $ flying; 5.VI.1964, 2:00 P.M.; and 3.VI.1965, 1:30 P.M., $ flying (HKC). Euptychia hermes (Fabricius ) complex

Veracruz (Mexico), 18.1.1966, 1:30 P.M., $ flying (CM-CUA). Maniola jurtina ( Linnaeus )

England, 5.VII.1964, 1:30-3:00 P.M., 2 pairs, $ flying (LDM).

Europe, $ flying (Pronin, 1964).

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Melanargia galathea (Linnaeus), Erebia medusa Scliiffermuller, Pararge aegeria (Linnaeus), Aphantopus hyperanthus (Linnaeus): all Europe, all 9 flying (Pronin, 1964). Pierella I. luna (Fabricius)

Costa Rica, 18.VIII.1963, 11:00 A.M., 9 flying (LDM). Pierella, helvina incanescens Godman & Salvin

Costa Rica, 18.VIII.1963, 11:00 A.M., $ flying (LDM).

Nymphalidae

There is much diversity in this family in the sex of the active partner. Females clearly predominate, but note the exceptions: Speyeria nokomis (sometimes) and Anartia fatima (but not Anartia jatrophae). Mating time may vary from group to group. The records suggest (hardly forcefully at this point) that Phyciodes and Anartia may mate in late morning, Speyeria in early afternoon, Nymphalis in mid aflernoon and Vanessa in late afternoon.

Euptoieta claudia ( Cramer)

Nuevo Leon (Mexico), 2.VII.1966, 10:30 A.M., $ flying (C. J. McCoy, Jr.). Speyeria cybele ( Fabricius)

Iowa, 27.VIII.1956, 9 flying (LDM). Speyeria aphrodite ( Fabricius )

Pennsylvania (PNR), 8.VII.1965, 12:30 P.M., 9 flying (HKC). Speyeria idalia ( Drury)

'DC, 9 flying (Clark, 1932). Speyeria nokomis nokomis (Edwards)

Colorado, 2.IX.1965, 2:30 P.M., 3 flying (F. M. Brown). Mr. Brown has since observed a number of mated pairs of this species. On some occasions the male took the lead, on others, the female. Argynnis paphia (Linnaeus)

Europe, VII, 9 flying (Pronin, 1964). Phyciodes tharos (Drury)

DC, 9 flying (Clark, 1932).

Pennsylvania (PNR), 25.V.1961, 12:00 noon (HKC).

New York, 3.VIII.1966, 1-2 P.M. (Sister M. Celestine). Phyciodes phaon (Edwards)

Texas, 8.VII.1963, 10:00 A.M. (HKC). Asterocampa leilia (Edwards)

Tamaulipas (Mexico), 8.1.1966, 11:00 A.M., 9 flying (CM-CUA). Anartia jatrophae luteipicta Fruhstorfer

Tamaulipas (Mexico), 8.1.1966, 11:00 A.M., 9 flying (CM-CUA). Anartia fatima ( Fabricius)

Costa Rica, 6.VIII.1963, 10:00-12:00 A.M., $ flying (LDM). Limenitis hredoivii (Geyer)

California, VIII, 2-3 P.M., 9 flying (Pronin, 1964) Vanessa atalanta (Linnaeus)

Russia, V, 4:15 P.M. (Pronin, 1964). Vanessa cardui (Linnaeus)

Russia, VII, 6:00 P.M. (Pronin, 1964). Nymphalis antiopa (Linnaeus), N. polychloros (Linnaeus), N. io (Linnaeus), N. urticae (Linnaeus), Polygonia c-alhum (Linnaeus): all Russia, IV, 2-3 P.M. (Pronin, 1964).

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Lycaenidae

All the reliable records indicate that the female is the active partner in this family. There are, however, several possible exceptions: those of Pronin (1964) which are uncertain; and several, not listed, observed by HKC with sex determination doubtful. Mating time is exceedingly varied, from mid-morning to early evening.

The possible restriction of mating in some species (crysalus, acadica, augustinus iroides) to the hours of early evening is noteworthy and so far known only in this family.

Hypaurotis crysalus (Edwards)

Colorado, 11.VIII.1962, 7 P.M. (Chambers, 1963). Satyrium acadica (Edwards)

Michigan, 22.VII.1951, 6-7 P.M., several pairs; and 24.VII.1951, 6-8 P.M., several pairs ( HKC ).

Connecticut, evening (teste C. L. Remington, Chambers, 1963). Chrysophanus titus ( Fabricius)

Pennsylvania, 22.VII.1961, betw. 12:00-2:00 P.M., 9 flying (LDM). Callophrys (Incisalia) henrici ( Crote & Robinson)

New York, 7.IV.1906, 10:30 A.M. (Cook, 1907). Callophrys (Incisalia) augustinus iroides (Boisduval)

California (Contra Costa Co.), 24.V.1963, 5:30 P.M. (PDT); 26.V.1963,

5:20-7:45 P.M., 2 pairs (sunset); 30.V.1963, 5:15 P.M. "Although . . .

individuals often perched on the tree during midday hours, none were

seen mating earlier than 5:00 P.M. (4:00 P.M. P.S.T.)." (Powell, 1964).

Lycaena phlaeas americana Harris

Pennsylvania (PNR), 16.IX.1965, 1:15 P.M. (HKC). Lycaeides melissa (Edwards)

New Mexico, 23.VII.1963, 2:00 P.M. (HKC). Plebeius argus (Linnaeus) and Polyommatus icarus (Rottemburg)

Europe, VII, £ flying (Pronin, 1964). Maculinea arion (Linnaeus)

Europe, VIII, $ flying (Pronin, 1964). Everes c. cornyntas (Godart)

Pennsylvania, 14.VIII. 1952, 6:30 P.M. (HKC). Everes comyntas texanus Chermock

Costa Rica, 11.VIII.1963, betw. 11:00 A.M.-L00 P.M., 2 pairs, $ flying (LDM).

Veracruz (Mexico), 16.1.1966, 3:05 P.M., $ flying (CM-CUA). Celastrina pseudargiolus (Boisduval & Le Conte)

Michigan, 29.IV.1951, betw. 11:30 A.M.-L30 P.M. (HKC).

Hesperiidae

As far as records indicate, only the female is the active partner in hesperiids. There is a marked preponderance of records in mid to late afternoon.

Polythrix asine ( Hewitson)

Costa Rica, 3.VIII.1963, betw. 11-12 A.M., $ flying (LDM). Cogia calchas ( Herrich-Schaffer)

Costa Rica, 2.VIII.1963, 3:30 P.M., $ flying (LDM).

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Erynnis juvenalis (Fabrieius)

Pennsylvania (PNR), 25.V.1961, 2:00 P.M. (HKC).

Pennsylvania, 10.V.1964, 1:20 P.M., 9 flying (LDM, HKC).

Maryland, 11.V.1965, 4:00 P.M., $ flying (HKC). Adopaea s. sylvestris (Poda)

England, 5.VII.1964, betw. 1:30-3:00 P.M., $ flying, 2 pairs (LDM). Polites peckius (Kirby)

Pennsylvania (PNR), 3.VI.1965, 4:20 P.M., $ flying; and 5.VIII.1965, 3:20 P.M., 9 flying (HKC). Euphyes conspicua conspicua (Edwards)

DC, 9 flying (Clark, 1932). Hesperia Columbia (Scudder)

[California], 2:15 P.M. (MacNeill, 1964:30). Hesperia juba (Scndder)

Western United States, $ flying, 2 pairs (MacNeill, 1964:30). Hylephila phyleus (Drury)

Florida, IX.1962, ca. 4 P.M., $ flying (J. N. L. Stibick).

Discussion

Not surprisingly, more questions are raised than answered by the foregoing data. Even so, a few generalizations are possible and a few thoughts are suggested by the data.

1. Mating time of day. We give (fig. 1) frequency polygons of the observed flight times for each of the major families. They include even rough records, and are not corrected for Daylight Saving Time, so they are crude representations. We intend them to show only (a) that much variety exists; (b) that the Hesperiidae appear to mate somewhat later, on the average, than others; and (c) that the evening mating of some Lycaenidae is a significant departure from the norm.

The polygon of combined records (bottom figure) shows a marked peak between the hours of 1:00 and 3:00 P.M., the hottest part of the average day. This, together with the great variability in observed times of some species, suggests that clock, or solar, time may be of less significance in a particular mating than the ambient or antecedent weather. Despite the rarity of observations of mated pairs in the field, it is remarkable how many of the above records consist of two or more pairs seen on the same day, even the same hour. This may be additional indication that appropriate weather conditions are important.

Early evening mating in some lycaenids is well established; and late afternoon mating in some Hesperiidae also seems to be indicated. The significance of these facts we cannot even guess; nor can we explain the difference between such close relatives as Callophrys (Incisalia) henrici (morning mating) and C. (7.) augustinus iroides (evening mating).

2. Mating date. The calendar date on which mating occurs means little by itself; but taken in conjunction with the local flight period of

1968 Journal of the Lepidopterists' Society 131

8 noon 8

Explanation of Figure

Fig. 1. Frequency polygons of observed mating times in various families. P = Pieridae; S = Satyridae; N = Nymphalidae; L — Lycaenidae; H = Hesperiidae, "all" = sum of all these, plus the Papilionidae and Danaidae. Note: (1) the somewhat later average mating time in Hesperiidae; (2) the bimodality of the Lycaenidae, with a significant secondary peak in early evening; (3) the peak of all observations falling between 1-3 P.M., the warmest part of the average day.

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the species it may be quite significant. In Euptychia cymela, the Powder-mill records are all very early in the flight period: mating apparently occurs shortly after eclosion of the tardier sex. This appears to be true also in Pieris rapae, P. virginiensis, and Phyciodes tharos. On the other hand, in Satyrium acadica, Chrysophanus titus, and possibly Hypaurotis cry solus and Cercyonis pegala, the records indicate that mating occurs much later in the flight period, perhaps even towards its end. In pegala it may coincide approximately with the first eclosion of the very tardy females, but this is not true in the lycaenids.

3. Sex of the active partner. Some families seem to be quite consistent: Pieridae (males), Danaidae (males), Satyridae (females), Hesperiidae (females). The Lycaenidae also may be consistent (females), but data are too few to be sure. In the Nymphalidae, although females predominate, there are several inconsistencies. Brown's observations that in Speyeria nokomis either sex may take the lead may untimately be found true of many other nymphalids. The behavior of Eurema lisa (Pieridae) suggests that even in consistent families the normally passive partner may take the lead in some stress situations.

Literature Cited

Brower, L. P., J. V. Brower, & F. P. Cranston, 1965. Courtship behavior of the Queen butterfly, Danaus gilippus berenice (Cramer). Zoologica, 50: 1-39, 11 figs., 7 pis.

Chambers, D. S., 1963. Evening mating in Hypaurotis crysalus (Lycaenidae) in Colorado. J. Lepid. Soc, 16: 200.

Clark, A. H., 1932. The butterflies of the District of Columbia and vicinity. U. S. Natl. Mus., Bull. 157, ix + 337 pp., 64 pis.

Cook, J. H., 1907. Studies in the genus Incisalia. III. Incisalia henrici. Canad. Ent., 39: 181-187, pi. 4.

MacNeill, C. D., 3964. The skippers of the genus Hesperia in western North America, with special reference to California (Lepidoptera: Hesperiidae). Univ. Calif. Publ. Ent., 35: 1-221, 28 figs., 8 pis.

Powell, J. A., 1964. Mating behavior of Incisalia iroides (Boisduval) (Lepidoptera: Lycaenidae). Pan-Pacific Ent., 40: 100.

Pronin, G., 1964. The mating time of Lepidoptera. J. Lepid. Soc, 18: 35-41.

Urquhart, F. A., 1960. The monarch butterfly. Univ. Toronto Press, xxiv + 361 pp.

PACIFIC SLOPE SECTION—1968 MEETING

The fifteenth annual meeting of the Pacific Slope Section of the Lepidopterists' Society will be held September 6-8, 1968, at the University of California, Berkeley. The program will include a field trip and open house at the museum Friday, September 6; presentation of papers Saturday and Sunday; and a banquet Saturday evening. Collections of the California Insect Survey and library facilities of the Department of Entomology will be available for study. Details of the program will be mailed to all Pacific slope members and to others who request them, in August.