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Volume 24, Number 2

105

Bibliographie

Rothschild, W. 1914. Description of a New Troides. Nov. Zool. 21:275. Niepelt, W. 1916a. Lepidoptera Niepeltiana 2, p. 20-21, pi. 17, fig. 1-2. Niepelt, W. 1916b. Beitrag zur Kenntnis der Omithoptera alottei Rothschild. Ent.

Rundsch. 33:31. Peebless, H. M., W. SchmassMan. 1917. Description of the female of Troides al-

lottei Rothsch. Nov. Zool. 24:426-427. Zeuner, F. E. 1943. Studies in the Systematics of Troides Hiibner and its Allies.

Trans. Zool. Soc. 25. (3), 184 pp.

STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN

NEW ENGLAND. I. ABUNDANCE AND SEASONAL OCCURRENCE

OF THE SPECIES, 1961-1969

Theodore D. Sabgent

Department of Zoology, University of Massachusetts, Amherst, Massachusetts

and

Sidney A. Hessel

Entomology Section, Peabody Museum, Yale University, New Haven, Connecticut

The Catocala moths pose an interesting evolutionary enigma. This large genus, consisting of some 200 species in the broadest use of the generic name, is characterized by extensive sympatry throughout most North Temperate regions of the world, and its members occupy a rela tively narrow ecological niche, the larvae being oligophagous and the adults being cryptically marked when at rest on tree trunks. In New England alone, at least 52 Catocala species are known to occur (Forbes 1954), and we have taken as many as 21 species on a single night at onr location.1 Nevertheless, natural hybrids are unknown. This tremendous array of sympatric and rather closely related species immediately raises the question as to what sorts of mechanisms operate to limit natural hybridization within the genus. One long-range goal of these studies is to describe such mechanisms through detailed analyses of the behavior, ecology, and genetics of these moths.

Despite the popularity of the North American Catocala moths with collectors, relatively little of their total biology is known. They have been treated taxonomically by Grote (1872, 1873, 1876), Hulst (1880,

1 20 August 1961, Washington, Connecticut: C. epione, antinympha, badia, habilis, flehilis, ob-scura, residua, retecta, dejecta, palaeogama, subnata, neogama, ilia, parta, concumbens, sordida„ andromedae, ultronia, grynea, praeclara, and arnica.

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Journal of the Lepidopterists' Society

1884), Hampson (1910), McDunnough (1938), and Forbes (1954). Plates depicting most of the North American species are available in Holland (1903) and Barnes and McDunnough (1918). Present knowledge of the life histories and geographic ranges of the New England Catocala is summarized in Forbes (1954).

Information on the behavior and ecology of these moths is scanty and largely anecdotal. Some observations on the resting habits of adults have been recorded (e. g. Bunker, 1874; French, 1880; Johnson, 1882; Rowley and Berry, 1909; Kettlewell, 1958; Sargent and Keiper, 1969). Limited data on the movements of adults have been obtained from studies of color-marked individuals (Brower, 1930). Virtually nothing is known of courtship and mating behaviors—a fact that has precluded any detailed studies on the genetics of these moths.

The present report constitutes a first step in our study of the Catocala of southern New England: an account of the species that are present, based on daily counts of adults taken for several years in two localities. It is hoped that these data will provide a foundation for planning further investigations, will serve as a record for future comparative purposes, and will stimulate others to obtain comparable data from their localities.

Methods

The basic data utilized in this report are daily counts of adult Catocala taken in the vicinity of Amherst, Massachusetts (Area 1) for the years 1964 through 1969; and from Washington, Connecticut (Area 2) for the years 1961 through 1965, 1967, and 1969.

Area 1. Records here are based on collecting of 2009 specimens by T. D. Sargent and two graduate students at several localities in Franklin and Hampshire Counties in north-central Massachusetts, all localities lying within 10 miles of the center of Amherst. The vast majority of the records are from two sites, one in Pelham (1964-66) and another in Leverett (1967-69). Catocala taken at bait, a brown sugar-beer mixture painted onto tree trunks, comprise 84% of the records from this area, whereas Catocala taken at lights (several 150-watt incandescent bulbs, and one 15-watt black light fluorescent tube) comprise 12% of the records. The additional 4% of the records are based on the finding of resting moths in nature. Catocala were recorded on a daily basis from 1 July to 1 September (except for occasional 1 to 2 day absences, or days of extremely inclement weather), and more sporadically from 1 September to 1 November each year.

Area 2. Records here are based on collecting of 5806 specimens by S. A. Hessel at one site in Washington, Litchfield County, in west-central

Volume 24, Number 2

107

Connecticut; this site lies approximately 65 air-miles to the southwest of Amherst, Massachusetts.2 All records of Catocala from this area are based on specimens taken at lights, most at one Robinson mercury vapor light-trap, and some at one 15-watt black light fluorescent tube. These lights were in operation from mid-March to mid-November each year (except for occasional 1 to 3 day absences).

Several important differences in collecting procedures between the two areas should be stressed:

(1)     Captures in Area 1 were predominantly at bait; captures in Area 2 were exclusively at lights.

(2)     Collecting in Area 1 was generally terminated by midnight; in Area 2 the Robinson trap was operating continuously from dusk to dawn.

(3)     Several collecting sites are included in Area 1; Area 2 includes only one collecting site.

(4)     Collecting in Area 1 was sporadic after September 1, but continued unabated in Area 2.

Because of these differences, the records for the two areas will be presented separately.

It should also be noted that some Catocala individuals may have been recorded on more than one occasion, as the majority of specimens in both areas were released after examination. However, studies of color-marked Catocala in Area 1 (Sargent, in prep.) indicate that very few individuals are captured on more than one occasion.

The species of Catocala were identified as keyed and described in Forbes (1954), and the species names used throughout this report are from that source. It should finally be noted that specific distinctions were not always made between gracilis and sordida in both areas, and among crataegi, blandula, and mira in Area 1. It is known from mounted specimens, however, that all of these species occurred in both areas.

In area 1, the sex of Catocala specimens was determined, and the precise time of their capture noted. These data are not included in the present report, but will be treated in subsequent papers.

Results and Discussion Abundance. The Catocala taken each year in Areas 1 and 2 are enumer-

2 This site lies at the bottom of a very narrow north-south valley through which an all-season stream flows southward. The lower end of the valley, only one mile distant, opens onto terrain which rolls gently southward to the Connecticut coast, while northward, the Litchfield Hills, of which it is part, become the Berkshires of Massachusetts. The surrounding hills are largely mixed deciduous woodlands, but include several serai stages resulting from the abandonment of farms, pastures, and woodlots of the Colonial period. Climatically, as demonstrated by meteorological maps, the site is to be included with territory considerably more northern than with the Connecticut and Hudson River valleys of its latitude.

108                                                          Journal of the Lepidopterists' Society

TABLE 1. Catocala spp. recorded in the Amherst, Mass. area, 1964-69.

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109

TABLE 1. Continued.

* records of blandula and mira included here

ated in Tables 1 and 2 respectively. In both cases, the species are listed in a descending order of overall abundance. Examination of these tables reveals some differences between results from the two areas. Catocala numbers, in terms of both species and individuals, are greater from Area 2. In part, this difference must be due to the longer daily and seasonal collecting periods in Area 2, and perhaps to a greater efficiency of the Robinson trap, when compared to bait, as a collecting device. However, further consideration indicates that those species that are markedly more abundant in Area 2 are almost invariably hickory and walnut (Juglan-daceae) feeders. Accordingly, the data were reanalysed with reference to the known foodplants of the various Catocala species (Forbes, 1954), and this procedure revealed that remarkably more hickory-walnut feeders, both in terms of species and individuals, were taken in Area 2 (Table 3).

This disparity in records of feeders on the Juglandaceae seems to reflect something more than the previously listed differences in collecting procedures between the two areas. It seems more likely that differences in the frequency of hickories and walnuts are important. In this regard, the presence of Butternut (Juglans cinerea L.) as a common tree in the woodlands near the collecting site in Washington, and its virtual absence in all collecting areas near Amherst, seems most suggestive. It could be that

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Volume 24, Number 2

111

TABLE 2. Continued.

Totals and Percent of Numbers of individuals                                , *} r

& (Percent of Yearly Total for All Species)            T°tal for

--------_--------------------------------------------------------------------- Entire

1961 1962 1963 1964 1965 1967 1969 Period

Species

piatrix

cerogama

* During the period 1952—60, one additional species was noted in this area—-C. robinsonii, 15 September 1956, one specimen.

112

Journal of the Lepidopterists' Society

TABLE 3. Foodplants of the Catocala spp. from two localities.

Catocala species included:

1 relicta, parta, briseis, unijuga, car a, concumbens, amatrix

2 antinympha, hadia

3 piatrix, epione, habilis, serena, Judith, flebilis, obscura, residua, retecta, dejecta, vidua, palaeo-gama, subnata, neogama

* ilia, coccinata, similis, micronympha, arnica

5 ultronia, crataegi, mira, blandula, grynea

0 gracilis, sordida, andromedae

" innubens (Gleditsia), cerogama (Tilia), praeclara (Quercus ?, Crataegus ?).

Area 1 is near or beyond the northern limit of some of these Catocala species, due perhaps to a sparse distribution of certain foodplants, or to climatic marginality for the insects themselves.

The presence and absence of a foodplant seems to explain the single remaining difference in species between Areas 1 and 2 (i.e. bayberry, Myrica pensylvanica Loisel, the foodplant of C. badia, is present only in Area 2).

It is also possible, of course, that some differences in records from the two areas are due to differences in collecting procedures, which reflect, in turn, behavioral differences among the Catocala species. For example, C. ilia has been taken much more often at bait than at lights (96% of 371 captures in Leverett, 1967-69), and this fact may well explain its apparently higher numbers in Area 1 (and the consequent higher percentage of feeders on the Fagaceae). In contrast, C. antinympha has been more prevalent at lights than at bait (88% of 74 captures in Leverett, 1967-69),

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113

TABLE 4. Early occurrence of several common Catocala species in two localities.

and this could account for its apparently higher frequency in Area 2. Differences of this sort, as well as differences in the time of flight of the various species during the night, are being carefully studied, and will form the basis of further papers in this series.

Changes in Abundance. The records presented here do not cover a sufficient number of years to allow extended analyses of changes in abundance. However, the variations and trends in annual numbers of certain species warrant brief comment.

Several species exhibited wide fluctuations in annual abundance (e.g. C. ilia and C. ultronia in Area 1, C palaeogama and C. habilis in Area 2, and C. antinympha in both areas). Two species in Area 1, C. amatrix and C. residua, were never recorded until 1969, and then both were relatively common. These fluctuations indicate that detecting long-term trends in Catocala abundance may often require longer series of annual records than those considered here.

The records from Area 2, together with notes of S. A. Hessel extending back to 1952, do indicate that two species, C. serena and C. Judith, have recently become remarkably more common in that area. Although neither of these species was collected in Area 2 prior to 1960, since then 104 serena and 35 Judith have been collected. These results could be due to erratic long-term oscillations in the abundance of these species, or might represent their recent range extension from more southern regions into Area 2. If these two species are presently extending their ranges, then we might eventually expect to take specimens in Area 1, where neither has yet been taken.

In all of the records from both areas, only one species, C. subnata, seems to be showing any evidence of a recent decline in numbers. This species is known as one which exhibits long-term fluctuations in abundance (Forbes, 1954).

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Journal of the Lepidopterists' Society

TABLE 5. Seasonal occurrence of Caiocala.

1 Species with less than 10 records not included.

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115

TABLE 5. Continued.

serena (103) residua

(898) concumhens

(385)

retecta (401)

cerogama

(14) porta

(34)

obscura

(130)

cava

(126)

flebilis (19) neogama

(339)

amatrix (38)

habilis

(661)

10

36

37

30

16

18

11

10

26

29

30

36

42

71

35

32

36

58

34

18

15

24 19 21 24 14 21 39 29 11 32 55 36

13

32

17

21

19

41

1 Species with less than 10 records not included.

The wide annual variation in total Catocala individuals taken at any one locality seems largely due to dramatic fluctuations in the abundance of a very few common species (e.g. C. ilia and C. ultronia in Area 1; C. pa-laeogama, C. residua, and C. habilis in Area 2). Weather is another factor which undoubtedly influences records of annual abundance. It is well known among collectors that Catocala are more frequently taken on warm, humid nights; and thus the number of such nights during a summer will affect the total number of Catocala recorded. Data on temperature and humidity with respect to Catocala abundance will be obtained in our areas in subsequent years.

Seasonal Occurrence. The seasonal appearance of adult Catocala in Areas 1 and 2 seemed virtually identical. For example, of the 30 species common to both areas, 17 had earliest records of capture from Washington, and 12 had such records from Amherst (in one species the earliest re-

116

Journal of the Lepidopterists' Society

corded date was the same in both areas). Data on the early occurrence of the six species for which there are 50 or more records from each area are summarized in Table 4. (Late occurrences of the species could not be as meaningfully compared across areas because late season collecting was much less complete in Area 1 than in Area 2.)

A summary of the seasonal occurrence of the Catocala species is presented in Table 5. These data were obtained by summing across years and areas, a procedure which tends to extend the apparent season of each species to some extent. Nonetheless, it is clear that seasonal isolation alone is insufficient to separate most of the Catocala species from one another (all of the species, for example, may occur during the latter half of August). Some species may occur over the entire summer (e.g. C. unijuga, with extreme dates of 8 July and 9 October), and these certainly must be isolated from other species by factors other than seasonal occurrence. However, it seems equally clear that seasonal offset may coact with other factors in effectively isolating certain closely related pairs of species (e.g. hlandula and mira, serena and habilis, dejecta and retecta, subnata and neogama).

Summary

Daily records of adult Catocala have been kept over several years at two localities in southern New England. For the period 1961-1969, nearly 8,000 individuals of 39 species were recorded. These data have been summarized here in an attempt to establish the abundance, and fluctuations in abundance, as well as the seasonal limits, of the species. In addition, suggestions relating to the differences in species composition at the two localities have been advanced.

Acknowledgments

This report was written while the first author was a visiting fellow in the laboratory of Dr. Charles L. Remington at Yale University. The stimulation of discussions with Dr. Remington, and his suggestions regarding this manuscript, are gratefully acknowledged.

Some of the Catocala records from the Amherst area were contributed by Dr. Ronald R. Keiper (1967-68) and Charles G. Kellogg (1969), and their cooperation and assistance is greatly appreciated.

Literature Cited

Barnes, W. and J. McDxjnnough. 1918. Illustrations of the North American species of the genus Catocala. Mem. Amer. Mus. Nat. Hist. vol. 3, part 1.

Brower, A. E. 1930. An experiment in marking moths and finding them again (Lepid.: Noctuidae). Entomol. News 41: 10-15.

Bunker, R. 1874. Notes on collecting Catocalas. Canad. Entomol. 6: 25-26.

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117

Forbes, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III. Noc-

tuidae. Cornell Univ. Agri. Exp. St. Memoir 329. French, G. H. 1880. Notes on Catocala hunting. Canacl. Entomol. 12: 241-242. Grote, A. R. 1872. On the North American species of Catocala. Trans. Amer. Entomol.

Soc. 4: 1-20.

---------- 1873. On the genus Catocala. Canad. Entomol. 5: 161-164.

---------- 1876. On species of Catocala. Canad. Entomol. 8: 229-232.

Hampson, G. F. 1910. Catalogue of the Lepidoptera Phalaenae in the British Museum. Vol. 9. London.

Holland, W. J. 1903, The Moth Book. New York. (Reprinted 1968, with annotations by A. E. Brower).

Hulst, G. D. 1880. Remarks upon the genus Catocala with a catalogue of species and accompanying notes. Bull. Brooklyn Entomol. Soc. 3: 2-13.

---------- 1884. The genus Catocala. Bull. Brooklyn Entomol. Soc. 7: 13-56.

Johnson, J. S. 1882. Catocalae taken in the vicinity of Frankford, Pennsylvania.

Canad. Entomol. 14: 59-60.

Kettlewell, H. B. D. 1958. The importance of the microenvironment to evolutionary trends in the Lepidoptera. Entomologist 91: 214-224.

McDunnough, J. 1938. Checklist of the Lepidoptera of Canada and the United States of America. Part 1. Macrolepidoptera. Mem. Southern Calif. Acad. Sci. Vol. 1.

Rowley, R. R. and L. Berry. 1909. Notes on the study of some Iowa Catocalae. Entomol. News 20: 12-18.

Sargent, T. D. and R. R. Keiper. 1969. Behavioral adaptations of cryptic moths. I. Preliminary studies on bark-like species. J. Lepid. Soc. 23: 1-9.

COMMUNAL ROOSTING IN COLIAS AND PHOEBIS (PIERIDAE)

Harry K. Clench

Carnegie Museum, Pittsburgh, Penna.

Two instances of communal roosting in coliadine pierids have recently come to my attention. They have to do with different genera, of considerably different body size, and occurring in different major environments, but there are several striking similarities and I believe a common explanation may satisfy both. 1

Colias eurytheme Boisduval

In an open meadow near Pittsburgh, Pennsylvania, at about 5:30 PM, EST, on 18 September 1969, I saw Colias eurytheme in fairly large numbers preparing for the night. A few of them were still active, but most had already sought roosts. The sun was within 15-20° of the true horizon, but was destined to disappear a little prematurely behind a low, tree-covered hill, and the field was already partly shaded.