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Volume 38 1984 Number 1
ISSN 0024-0966
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
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27 July 1984
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
LEE D. MILLER, President CHARLES V. COVELL, JR., KAROLIS BAGDONAS, Vice President Immediate Past President MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer
Members at large:
K. S. BROWN, JR. F. S. CHEW J. M. BURNS E. D. CASHATT G. J. HARJES F. W. PRESTON T. C. EMMEL E. H. METZLER N. E. STAMP
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Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- sion in the Sesiidae fascicle for the Moths of America North of Mexico, The dusky clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of willows and poplars.
JoURNAL OF Tue LEPIDOPTERISTS’ SOCIETY
Volume 88 1984 Number 1
Journal of the Lepidopterists’ Society 38(1), 1984, 1-12
THE LIFE HISTORY AND ECOLOGY OF EUPHYDRYAS GILLETTII BARNES (NYMPHALIDAE)
ERNEST H. WILLIAMS
Department of Biology, Hamilton College, Clinton, New York 13323
CHERYL E. HOLDREN AND PAUL R. EHRLICH
Department of Biological Sciences, Stanford University, Stanford, California 94305
ABSTRACT. Based on studies of several populations, the life stages of the montane butterfly Euphydryas gillettii and its natural history and ecology are described. E. gil- lettii shows unusual developmental flexibility in that it can diapause as second, third, or fourth instars, depending on climatic conditions; in addition, one population in a colder habitat is mostly biennial, while others are annual. In spite of this flexibility, the species has limited distribution in isolated populations over a narrow geographical range.
Euphydryas gillettii Barnes occurs in the middle Rocky Mountains, ranging from western Wyoming, through northern Idaho and western Montana, and into Alberta (Ferris and Brown, 1981). While much work has been published on other species of Euphydryas in the past 20 years (Ehrlich et al., 1975; Cullenward et al., 1979; Brown and Ehrlich, 1980; Stamp, 1982), little has been known about E. gillettii. Until very re- cently (Williams, 1981; Holdren and Ehrlich, 1981), the only report in the literature on the biology of this species was that of Comstock (1940), which describes the eggs and early instars. 3
We have studied E. gillettii in several locations recently and here report on its life history and ecology. Four populations have been observed extensively: natural populations in the Teton and Beartooth Mountains of Wyoming, and two populations introduced into Colorado from the Teton colony. In addition, several other populations have been visited.
2, JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Width of the head capsule, spine length, and body size for the different instars of Euphydryas gillettii.
Length of spines
Width of head a EO MI ETT Instar capsule (mm) Shaft (mm) Setae (mm) moving (mm) First 0.44 + 0.01 (30) 0.02 0.2-0.3 3-4 Second 0.61 + 0.03 (81) 0.30 + 0.04 (28) 0.2-0.4 4-6 Third 0.90 + 0.04 (46) 0.46 + 0.06 (43) 0.3-0.5 5-9 Fourth 1.17 + 0.18 (84) 0.69 + 0.10 (39) 0.5—-0.7 9-13 Fifth 1.47 + 0.18 (39) 0.74 + 0.10 (42) 0.6-0.9 12-18 Sixth DADE = OMe2 (a) 0.74 + 0.16 (27) 0.7-1.2 15-30
Study Sites
The Beartooth population lives along a small stream in a montane meadow of 2620 m (8600 ft) elevation. The butterflies fly in an elon- gate area, roughly 60 m by 240 m, which is surrounded by coniferous forest of primarily Picea engelmanii. The highest density occurs in an area of secondary growth, where trees are scattered sparsely through a moist bottom area near the stream.
The Teton population is the largest known for this species. The butterflies are widely scattered over an eastern facing slope at 2100 m (6900 ft) elevation, occurring in an area roughly 400 m by 1500 m of mostly herbaceous vegetation. Streams run down through this slope, and trees, mostly Picea and Populus tremuloides, grow along the stream beds. Adults are found throughout the slope.
The two Colorado sites are in Gunnison County. One, adjacent to the Rocky Mountain Biological Laboratory at Gothic (2900 m, 9500 ft), is similar to the Teton site. It consists of a moist meadow containing thick stands of willows on an east-facing slope, bounded by spruce forests, the East River, and the cliffs of Gothic Mountain. The second, Pioneer Resort (2700 m, 8800 ft), is less open than either the Gothic or Teton sites, but the flora is similar.
Description of Life Stages
Measurements of the head, spines, and body for the different instars are given in Table 1.
Egg. Nearly spherical; rounded base with sides sloping in to flattened top. Approxi- mately 22 longitudinal ridges which extend most of distance down from apex, with irregular pitting on base; horizontal striations between ridges (Comstock, 1940). Color yellow-green when first oviposited (see Egg Development for color changes). Diameter 0.78 + 0.02 mm (n = 11) and height 0.86 + 0.04 mm (n = 11) (eggmass shown in Fig. la).
First Instar Larva. Head blackish brown with few thin, colorless setae. Body pale greenish yellow with colorless setae arising from 12 longitudinal rows of brown papillae. Appearance is of a pale body spotted with brown. Spiracles brown. Anal prolegs darker than other prolegs, which are concolorous with body; true legs brown (Fig. 1b).
VOLUME 38, NUMBER 1 3
Fic. 1. Life stages of Euphydryas gillettii: a, egg mass; b, first and second instars (prediapause); ec, third instars (postdiapause); d, fourth instars (postdiapause); e, sixth instar prepupa; f, pupa.
4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Second Instar Larva. Head blackish brown with black setae. Body developing char- acteristic banding pattern of later instars: dorsal band pale yellow; dorsolateral band brown and irregular; lateral (stigmatal) band dull white; ventrolateral band light brown; and ventral band cream colored. Spiracles blackish brown. True legs brown. Crochets black; anal prolegs brown on outside. Branching spines develop from papillae and simple seta of first instar; shafts of spines light brown with black setae. Rows of spines as follows: one mid-dorsal in dorsal band; two in dorsolateral band, more dorsal row positioned caudal to second, and second on edge of next band; one row in lateral band; and two rows of small spines or tubercles adjacent to each other in ventrolateral band. Spines developed on all thoracic and abdominal segments, with exception of first and third rows, which are missing from thoracic segments (Fig. 1b).
Third Instar Larva. Head capsule black with black setae. Body has same banding pattern of previous instar, but with deeper colors. Ventral band with thin mid-ventral brown line. Prolegs yellow with black crochets; anal prolegs dark brown on outside. Spiracles black. Shafts of spines blackish brown on all rows except mid-dorsal row, in which shafts are yellow-brown (Fig. lc).
Fourth Instar Larva. Banding pattern further developed with greater contrast: dorsal band lemon yellow; dorsolateral band blackish brown with brown bases to spines; lateral band white with black spiracles, ventrolateral band brown; ventral band pale yellow with brown mid-ventral stripe. True legs black; prolegs yellow with brown bases and black crochets, and anal prolegs mostly black on outside. Shafts of all spines black, though with ring of lighter color at base of each, with yellow on light colored bands and brown on darker ones (Fig. 1d).
Fifth Instar Larva. Colors and patterns as in previous instar, with following exceptions: dorsal stripe bright lemon yellow, dorsolateral band black, spines and setae jet black, and all prolegs yellow but dark on outside.
Sixth Instar Larva. Continued development of previous banding pattern, with sharper contrast between bands. Midventral line blackish brown.
Prepupa. Slight discoloration of last instar, with some shortening and thickening (Fig. le).
Pupa. Ground color cream with black markings. Orange markings also occur except on wing cases; they are concentrated on abdominal segments, where there are seven orange warts per segment. Pupae average 16 mm long (Fig. 1f).
Adult. Head and thorax black; abdomen black above and somewhat lighter under- neath. Palpi and legs concolorous with distinctive brownish orange color of postmedian band (this color is closest to the reddish orange of color 7B7 in Kornerup and Wanscher, 1978; it is nearly identical to the orange-rufous, color II-11i, of Ridgway, 1912). Antennae black with thin white rings and with yellowish clubs. Dominant color of dorsal wing surface black; veins black; marginal band of orange and submarginal band of white much reduced, often disappearing in secondaries; postmedian band crossing both wings, 3-4 mm wide, and prominent; median spot band white and reduced, disappearing by anal margin; discal cell of primaries with four alternating spots of white and characteristic orange-rufous color, with another spot of each color in postcellular space; secondaries with three spots of each color in cell and postcellular space; basal area black. Underwings with same patterning as above, but black color reduced and spots expanded; this is especially true on secondaries in median to basal area, where there is great expansion of orange-rufous color and where black is limited to borders of spots. Males smaller than females, with forewing length 16.5 to 23 mm (mean = 20.9, n = 162); for females, fore- wing length 20.0 to 25.5 mm (mean = 23.7, n = 199) (Fig. 2b).
Ecology
Oviposition. As reported by Comstock (1940), the larval host is Lo- nicera involucrata (Rich.) Banks (Caprifoliaceae), a shrub 0.5 to 3 m tall that grows in moist soil in thickets and wooded areas throughout the geographical range of E. gillettii and far beyond (e.g., California,
VOLUME 38, NUMBER 1 5
Mexico, Alaska, and Quebec). The leaves are glabrous, short-petiolate, elliptic-oblong to elliptic-obovate in shape, and 5-14 cm long and 2- 8 cm wide (Hitchcock et al., 1959). Thus, the leaves are large enough to allow females to move completely to the underside of the leaves when ovipositing. Some authors (e.g., Tietz, 1972) have listed other larval foodplants, but eggs on or oviposition behavior near any plant other than L. involucrata is extremely rare. Of more than 600 egg masses seen in the Beartooth population, only four have been found on a plant other than L. involucrata; these occurred in 1982 on an unusually large and conspicuous specimen of Valeriane occidentalis Heller (Valerianaceae, a family related to the Caprifoliaceae). Post- diapause larvae may wander to other species of plants, however.
Female E. gillettii oviposit mostly in late morning. Prior to ovipo- sition they fly slowly above the shrub and herbaceous layer, fluttering near or touching branches that are among the most apparent (highest or densest). They do not appear to follow vegetational edges. While searching for oviposition sites, they occasionally touch plants other than Lonicera, but then they usually fly on within 2 sec.
Once a female does find L. involucrata, she flutters near the shrub, lands on a leaf, walks on it for a few seconds, and then flutters in the air, landing on the same or a different leaf. This process continues for 1 to 80 minutes, and even when she is blown or chased from the shrub, she returns to the same leaf or to one quite near it. She gradually increases the time spent on one leaf, walking up and down the dorsal surface near the leaf midrib, repeatedly opening and closing her wings, and occasionally moving entirely to the underside of the leaf. After the female backs over or flips sideways to the underside of the leaf, there is an initial quiescent period of a minute or two which generally precedes oviposition. Sometimes she may return to the upper surface after remaining quiescent for a brief time, walk around the leaf again, and perhaps even move to another leaf. When she finally begins ovi- positing, she remains motionless with the wings usually held open (Fig. 2b). Oviposition behavior of E. gillettii is quite similar to that described for E. phaeton (Stamp, 1982). Females appear to spend much time and effort assessing the potential oviposition site; individual females have been observed to spend more than two hours in the above be- haviors before actually beginning to oviposit.
The leaves chosen for oviposition are always large and near the top of a growing stem. In the Beartooth population, 51% of the egg masses were on the highest leaf pair and 36% on the next highest leaf pair (2 years, n = 453). Only one of 456 egg masses was found on the upper surface of a leaf, and only 7 of the 62 eggs from that egg mass hatched, while 30 were dislodged. The chosen leaves may or may not have other
6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fic. 2. Euphydryas gillettii: a, prediapause feeding web, which becomes the hiber- naculum; b, ovipositing female; ec, parasitized fifth instar prior to emergence of the parasitoid.
egg clusters already on them; in the Beartooth population, 44% (n = 456) of all egg masses were on leaves that had another egg mass on the same leaf (23% of all leaves with eggs, n = 332), resulting in a mean of 1.87 clusters per leaf. Egg masses are also clumped in E. phaeton (Stamp, 1982).
Approximately one-half of all egg clusters touch the leaf midrib. An ovipositing female faces the edge of the leaf and, while moving her abdomen back and forth, touches the lower leaf surface with the tip of the abdomen. If she then touches the midrib or another protruding leaf vein, she may use it as a guide in oviposition. Often she will use a previous egg mass as a guide. She lays the eggs row by row in both directions, and sometimes a second layer or more is oviposited upon the first. The far edge of the egg mass averages 2.0 cm from the edge of the leaf and the near edge 1.1 cm (n = 52), a distance which reflects the length of the body (roughly 1.6 cm).
In the Beartooth population, egg clusters have ranged in size from 23 to 310 eggs (n = 72), with a mean of 146 (Fig. la). In the Teton
~]
VOLUME 38, NUMBER 1
population, the average size over a three year period was 130 eggs per mass (n = 189), while in Colorado the average was 128 eggs per mass. In contrast, the egg masses of E. editha contain 45 eggs on average (Labine, 1968), while those of E. phaeton contain 274 (Stamp, 1982). Oviposition in E. gillettii proceeds at an average of 3.8 eggs per minute (n = 48 clusters), requiring 38 min to lay an average sized cluster; E. editha oviposits at a slower rate, needing 30 min to produce its smaller cluster (Labine, 1968). Based on observations of 150 marked female E. gillettii seen to display pre-oviposition behaviors or to oviposit at least once, none oviposited more frequently than every other day.
Egg Development. During the course of development in the Bear- tooth population, a mean of 13% of the eggs (n = 48 clusters) are lost from the egg mass due to dislodgement or detachment (19 eggs from a 146 egg average). Sometimes the edge of an egg mass peels away from a leaf, but most egg loss occurs where the eggs are more than one layer deep. The variance in egg loss per cluster is high, however, and most clusters lose few eggs. Presumably those eggs which detach from the leaves and fall to the moist, shady, predator-infested soil surface below do not hatch.
In Colorado, up to 30% of the egg masses are lost entirely during the developmental period due to heavy predation. Furthermore, few egg masses escape without some predation; losses of roughly 10 to 20% of the eggs in a mass are common. The predators are the same for the eggs as they are for the larvae: erythraid mites, myrid bugs, beetle larvae, and browsing mammals, the latter including moose and cattle.
The eggs change color during development from a pale straw-yellow when first oviposited, sometimes with a greenish tint, to a distinct gold, and then to darkening shades of red-brown. They become blue-gray about two days before hatching, a color which results from the for- mation of a dark head capsule beneath the white translucent egg shell.
The eggs hatch from July into September, while the snow begins falling in late August in these mountainous areas. Eggs hatch in 23 to 45 days in the Beartooth population, depending on the exposure of the oviposition site. In Colorado, the majority of the eggs hatch in 18-30 days, although those masses that are produced late in the season de- velop more slowly. Eggs at the center of the egg mass are the first to hatch, and most eggs hatch within a two day time span (Williams, 1981). A substantial fraction of the eggs, roughly 20% in the Beartooth population, hatch in early September after the leaves of L. involucrata have begun to wilt and turn yellow.
Prediapause Larvae. Newly emerged larvae feed partially on the egg shells, and within 24 hours they migrate to the upper surface of the leaf, where they begin forming a communal feeding web (Fig. 1b).
8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
The oviposition leaf is the first feeding site and is the base of the web; it curls inwards and is bound ever more tightly as time passes. Predia- pause larvae feed only on the epidermis and parenchyma of the leaves, leaving behind the patterned network of veins. Feeding occurs during the day; nocturnal feeding has not been observed. Gradually more leaves are added to the feeding web by binding lower leaf pairs to the first leaf. In this way the communal web grows larger, sometimes with the incidental binding of grasses and other leaves that are adjacent to the hostplant leaves. The “knots” (Scudder, 1889) thus formed are quite apparent in the field since they generally occur at the apices of the most conspicuous stems (Fig. 2a). Because different egg masses are often oviposited on the same or adjacent leaves, the larvae in a single feeding web may be the products of several different egg masses, even when these egg masses hatch on different dates.
Mortality is high during the prediapause period. Parasitic wasps identified as Benjaminia sp. (N. Stamp, pers. comm.) have been col- lected from the feeding webs, and the above-mentioned predators take a heavy toll. At least 80% of the larvae in the Beartooth population disappear before reaching winter diapause, while 50-60% of the Col- orado larvae die or disappear.
Most, if not all, of the larvae that result from a single egg mass remain in the same feeding web overwinter. Though these hibernacula are well attached to the woody stems of the shrubs, most are dislodged by winter snow.
Unlike the larvae of other well-studied Euphydryas, which diapause in the fourth instar, E. gillettii are apparently able to overwinter in response to environmental conditions as second, third, or fourth instars. For instance, the Beartooth colony, constrained by the rapid onset of winter at the end of the flight period, diapauses (first winter) in the second instar. In Colorado, where the two sites differ markedly in the length of both the larval and the food-plant growing season, the pop- ulations diapause at different instars even though they originated from the same parent colony in the Tetons. Like the original population, the larvae at 2440 m in Colorado reach the fourth instar, while at 2920 m they appear to overwinter successfully after the first molt but develop to the fourth instar given a sufficiently long summer (Holdren and Ehrlich, 1981). Overwintering larvae may pass through an extra molt before emergence, as occurs in E. editha (M. Singer, pers. comm.).
Postdiapause Larvae. In Colorado and Wyoming the larvae termi- nate diapause soon after the snow melts, which in most years is late May at 2440 m (8000 ft) and mid-June at 2920 m (9600 ft). The larvae feed on newly formed buds of L. involucrata, boring holes into the larger apical buds and consuming entirely the smaller axillary buds
VOLUME 38, NUMBER 1 9
(Fig. lc). By the time the larvae have molted into the fifth instar in the annual populations, the leaves are slightly expanded, measuring roughly 2 cm in length. In postdiapause fourth instars in the biennial population, the larvae may still feed in aggregations (Fig. 1d) on rel- atively large and well developed leaves. Although many postdiapause larvae feed on shrubs bearing the previous year’s webs, like other Eu- phydryas species, some disperse. Extensive, characteristic feeding dam- age as well as postdiapause larvae have been observed on isolated L. involucrata shrubs on which there had been no prediapause larvae. In the Beartooth population, some postdiapause larvae have been found feeding on Castilleja linariaefolia Benth. (Scrophulariaceae), Valeri- ana occidentalis Heller (Valerianaceae) and Pedicularis bracteosa Benth. (Scrophulariaceae). All of these plants have iridoid glycosides, secondary compounds known from the host plants of other Euphydryas (Bowers, 1981).
Diapause-related and postdiapause mortality appear to be quite high in Colorado. The number of postdiapause larvae found is consistently much smaller, by as much as two orders of magnitude, than the num- ber of large third instars observed shortly before diapause. Both post- diapause larvae and pupae may be parasitized, the latter in Colorado by the hymenopteran Ptermalus vanessae Howard, which oviposits into mature larvae or pupae. Parasitized larvae in the Beartooth pop- ulation cease feeding and movement in the fifth instar (Fig. 2c), and the Benjaminia parasitoid then emerges three to four weeks later.
Most, if not all, larvae in the Beartooth population return to diapause for a second winter before pupating; they spend the first winter in the second instar and the second in the fifth instar. The second diapause apparently is not obligate, but the shortness of the growing season in this habitat has led at least part of the population into a two-year life span; Williams (1981) has demonstrated another adaptation in this pop- ulation for the cold climate, that of ovipositing so that the eggs are warmed maximally by the sun. A biennial life cycle has also been reported for Euphydryas maturna (Forster and Wohlfahrt, 1955), a close, European relative of E. gillettii.
Larvae generally move away from the host shrubs for pupation (Figs. le & 1f), and the pupation sites are usually within 50 cm of the ground. While distinctive in color and pattern, the pupae are not easily found. Pupation requires about three weeks.
Adults. The adults fly during a four week period from June to mid- August. As is typical for butterflies (Wiklund and Fagerstrom, 1977), males are the earliest to emerge and show the greatest wing wear early in the season, and the male to female ratio declines gradually through the flight season (Williams, in prep.). Males also fly earlier in the morn-
10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
ing than females and in relatively greater numbers on cloudy days. Males are much stronger fliers; though smaller (in accord with Singer, 1982), they fly at faster speeds, are more difficult to catch, and are more difficult to manipulate when netted.
These butterflies spend much of the day sunning near the ends of branches high in coniferous trees, typically with the wings open slightly more than 180 degrees. Males fly back and forth through the habitat more than females, while females fly down to nectar more frequently. Occasional individuals puddle in the afternoon when other activity is reduced. Nights are spent in trees at heights of at least 3 m.
Mating is rarely observed because of the predilection of this species for the tops of nearby conifers. Chases of individuals near tree tops are common during the middle of the day, with males chasing both fe- males and other males. It remains curious, though, that males infre- quently chase females while females nectar in the herbaceous layer.
The butterflies do not have to move far to nectar. There is a pro- fusion of flowers in the E. gillettii habitat, largely because it is moist, and they feed readily at the available blossoms. The commonest nectar source for the Wyoming populations is a white geranium, Geranium richardsonii Fischer and Trautvetter (Geraniaceae), which is also used in Colorado where the most important source is probably Erigeron peregrinus (Pursh) Greene (Compositae). After senescence of the pri- mary nectar source, E. gillettii in Wyoming turns readily to yellow composites, mostly several tall Senecio which begin blooming as the Geranium cease. Given the abundance of flowers and the relatively limited time spent nectaring, adult food resources would not seem to be a major limiting factor in the population dynamics of this species.
DISCUSSION
Euphydryas gillettii was originally described and placed in the ge- nus Melitaea by Barnes (1897) from material collected in Yellowstone National Park, Wyoming; M. glacialis (Skinner, 1921) is a synonym. Gunder (1929), in his reorganization of North American Euphydryas, recognized the relationship of E. gillettii to the other Euphydryas species and pointed out that it is likely the most primitive of the North American species. L. G. Higgins (1978) then revised the genus Euphy- dryas and placed E. gillettii in a new genus, Hypodryas, along with the Palearctic species E. maturna, E. intermedia, E. eduna, and E. cynthia. Phenetically, E. gillettii seems most closely related with those species, although comparison of early stages and allozyme frequencies would clearly be desirable.
Following good taxonomic practice we have not accepted Hypodry- as as a genus; obligatory categories—genera, families, etc.—should be
VOLUME 38, NUMBER 1 ll
kept conservative to facilitate communication (Ehrlich and Murphy, 1982). Hypodryas could be considered as synonymous with “the ma- turna species group’ or, at most, a subgenus. Euphydryas is a phenet- ically quite uniform group. Because the genus is now so widely dis- cussed in the non-lepidopterological literature, we would not suggest any change in the widely accepted generic name.
Of current interest in the study of butterflies is whether or not the prior presence of eggs influences where a female lays her eggs. In several species—Battus philenor (Rausher, 1979), Pieris brassicae (Rothschild and Schoonhoven, 1977), and Anthocharis sara (Shapiro, 1980)—active egg load assessment is indicated, and in all of these cases females avoid ovipositing where eggs currently are or recently have been. Female E. gillettii rarely avoid leaves that already have eggs; moreover, the egg clusters are grouped together more than one would expect if they were distributed in the environment at random (Wil- liams, 1981). The same is apparently true of E. phaeton (Stamp, 1982). Though there has been no previous support for positive egg load as- sessment, the grouping of eggs or egg clusters together may further enhance survivorship of larvae if there is a selective reason, such as predator avoidance or thermoregulation, for grouping the eggs togeth- er initially. Stamp (1981, 1982) has considered reasons for such a group- ing, though in her experiments, E. phaeton suffered increased parasit- ism when the groupings were too large. Because the larvae from different clusters of E. gillettii eggs do mix freely in communal feeding webs, the contagious distribution of clusters may be adaptive.
E. gillettii displays sedentary behavior and occurs in localized col- onies with few populations known; these characteristics, along with the ease with which individuals may be caught, indicate that it could easily suffer from excessive human impact. How threatened the species may be is unknown, largely because it occurs in undisturbed mountain hab- itat, but much reduction in numbers in any one place could lead to the extinction of local colonies. Those who find a population in the field should exercise discretion when collecting, especially with fe- males.
ACKNOWLEDGMENTS
We thank Deane Bowers, Art Shapiro, and an anonymous reviewer for commenting on the manuscript. EHW was supported by grants from the Theodore Roosevelt Me- morial Fund of the American Museum of Natural History and from Wellesley College. The work of CEH and PRE was supported by a series of grants from the National Science Foundation, the most recent of which was DEB-8206961, and by a grant from the Koret Foundation of San Francisco. We thank the Brachman-Hoffman Foundation for support of publication.
LITERATURE CITED
BARNES, W. 1897. Some new species and varieties of Lepidoptera from the western U.S. Canad. Entomol. 29:39-42.
12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Bowers, M. D. 1981. Unpalatability as a defense strategy of western checkerspot but- terflies (Euphydryas Scudder, Nymphalidae). Evolution 35:367-375.
Brown, I. L. & P. R. EHRLICH. 1980. Population biology of the checkerspot butterfly, Euphydryas chalcedona. Structure of the Jasper Ridge colony. Oecologia 47:239- 251.
Comstock, J. A. 1940. Notes on the early stages of Euphydryas gillettii Barnes. Bull. S. Calif. Acad. Sci. 39:111-118.
CULLENWABRD, M. J., P. R. EHRLICH, R. R. WHITE & C. E. HOLDREN. 1979. The ecology and population genetics of an alpine checkerspot butterfly, Euphydryas anicia. Oecologia 38:1-12.
EHRLICH, P. R. & D. D. MurpHy. 1982. Butterfly nomenclature: A critique. J. Res. Lepid. 20:1-11.
EHRLICH, P. R., R. R. WHITE, M. C. SINGER, S. W. MCKECHNIE & L. E. GILBERT. 1975. Checkerspot butterflies: A historical perspective. Science 188:221-228.
FERRIS, C. D. & F. M. BROWN. 1981. Butterflies of the Rocky Mountain States. Univ. Oklahoma Press, Norman.
Forster, W. & T. A. WOHLFAHRT. 1955. Die Schmetterlinge Mitteleuropas. Franck- h’sche Verlagshandlung, Stuttgart.
GUNDER, J. D. 1929. The genus Euphydryas Scud. of boreal America (Lepidoptera, Nymphalidae). Pan-Pac. Entomol. 6:1-8.
HiccIins, L. G. 1978. A revision of the genus Euphydryas Scudder (Lepidoptera: Nym- phalidae). Entomol. Gaz. 29:109-115.
HitcHcock, C. L., A. CRONQUIST, M. OWNBEY & J. W. THOMPSON. 1959. Vascular Plants of the Pacific Northwest. Part 4. Ericaceae Through Campanulaceae. Univ. Washington Press, Seattle.
HOLDREN, C. E. & P. R. EHRLICH. 1981. Long range dispersal in checkerspot butter- flies: Transplant experiments with Euphydryas gillettii. Oecologia 50:125-129. KORNERUP, A. & J. H. WANSCHER. 1978. Methuen Handbook of Colour. 3rd ed. Meth-
uen, New York.
LABINE, P. A. 1968. The population biology of the butterfly, Euphydryas editha. VIII. Oviposition and its relation to patterns of oviposition in other butterflies. Evolution 22:799-805.
RAUSHER, M. D. 1979. Egg recognition: Its advantages to a butterfly. Anim. Behav. 27: 1034-1040.
RipGway, R. 1912. Color Standards and Color Nomenclature. Publ. by the author; Washington, D.C.
ROTHSCHILD, M. & L. M. SCHOONHOVEN. 1977. Assessment of egg load by Pieris brassicae (Lepidoptera, Pieridae). Nature 266:352-355.
SCUDDER, S. 1889. The Butterflies of the Eastern United States and Canada. W. H. Wheeler, Cambridge.
SHAPIRO, A. M. 1980. Egg-load assessment and carryover diapause in Anthocharis (Pieridae). J. Lepid. Soc. 34:307-315.
SINGER, M. C. 1982. Sexual selection for small size in male butterflies. Am. Nat. 119: 440-443.
STAMP, N. E. 1981. Effect of group size on parasitism in a natural population_of the Baltimore checkerspot Euphydryas phaeton. Oecologia 49:201-—206.
STAMP. N. E. 1982. Selection of ovipositon sites by the Baltimore Checkerspot, Euphy- dryas phaeton (Nymphalidae). J. Lepid. Soc. 36:290-302.
TiETZ, H. M. 1972. An index to the described life histories, early stages and hosts of the macrolepidoptera of the continental United States and Canada. Vol. I. Allyn Museum of Entomology, Sarasota, Fla.
WIKLUND, C. & T. FAGERSTROM. 1977. Why do males emerge before females? Oeco- logia 31:153-158.
WiLuiAMs, E. H. 1981. Thermal influences on oviposition in the montane butterfly Euphydryas gillettii. Oecologia 50:342-346.
Journal of the Lepidopterists’ Society 38(1), 1984, 13-14
CORRECT NAME FOR THE NEOTROPICAL SQUASH-VINE BORER (SESIIDAE: MELITTIA)
VITOR O. BECKER! AND THOMAS D. EICHLIN?
ABSTRACT. The identity of the species of squash-vine borer occurring in Central and South America on cultivated Cucurbitaceae is established as Melittia pulchripes, not M. satyriniformis, which is a junior synonym of the Eastern squash-vine borer, M. cucurbitae. A lectotype is designated for M. riograndensis, a name which is then syn- onymized under M. pulchripes.
For more than a century the Melittia species whose larvae are com- monly found boring in stems of many cultivated species of Cucurbi- taceae in Central and South America has been referred to in the lit- erature as M. satyriniformis Hiibner. In a study by Duckworth and Eichlin (1978) it was found that in the Western Hemisphere these borers belong to a complex of three closely related species: cucurbitae (Harris), satyriniformis Hiibner, and a third which they described and named calabaza. According to these authors (1973:154), the three species of the complex are easily distinguished by their external features and genitalia. A fourth species, pauper LeCerf, apparently occurs only in the vicinity of Lima, Peru. Both cucurbitae and calabaza are restricted to the United States and Mexico and are sympatric in the southern part of their range. The species distributed from Guatemala through Cen- tral and South America was regarded by them as satyriniformis, fol- lowing the use of earlier authors.
Heppner and Duckworth (1981:26) established that satyriniformis is a junior synonym of cucurbitae. This was based mainly on the fact that Hiibner stated that the type locality of satyriniformis was “Geor- gia” and therefore, must be conspecific with cucurbitae, the only squash- vine borer from the region.
We have examined the syntypes of riograndensis Bréthes (1920:284) and found that they are the same species as the Central and South American species previously and currently misnamed as satyrinifor- mis. Two male syntypes of riograndensis were located in the Museo Argentino de Ciencias Naturalis (MACN), Buenos Aires, both bearing identical labels in Bréthes’ hand-writing: “17”; “E. Ronna, vi. 1919,
>> “<e >> <
Pelotas’’; ““Type’’; ‘“Melittia riograndensis Breth.”’; (red rectangle). The
‘Centro de Pesquisa Agropecuaria dos Cerrados, P.O. Box 70-0023, 73300-Planaltina, DF, Brazil. ? Division of Plant Industry, Insect Taxonomy Laboratory, California Department of Food and Agriculture, Sacra- mento 95814, USA.
14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
male specimen which had been dissected is here designated as the lectotype; the second male becomes a paralectotype. These two spec- imens are covered by mold, and their external features are somewhat obscured; however, the genitalia are identical to those of satyriniformis (sensu authors, including Duckworth and Eichlin, 19738:fig. 3c), bear- ing the peculiar quadrate expanded process at the center of the valva. They also agree with the genitalia of a specimen reared by the senior author from stems of Cucurbita sp. at Turrialba, Costa Rica. However, the oldest available name for this species is Melittia pul- chripes Walker (1856:67). Syntypes in the British Museum (Natural History) were examined and a lectotype designated (Duckworth and Eichlin, 1978:21). At this time it was determined to be conspecific with the Central and South American squash-vine borer. The previous ref- erences to satyriniformis for the Neotropical squash-vine borer should in fact be applied to pulchripes. Also, riograndensis now becomes a synonym of pulchripes (NEW SYNONYMY). The following is a summary of the species comprising the squash- vine borer complex: Melittia cucurbitae (Harris)—eastern half of United States, Gulf Coastal areas of Texas and Mexico to near Guatemala. Melittia calabaza Duckworth and Ejichlin—Arizona, central and western Texas, interior areas of Mexico to west coast. Melittia pulchripes Walker—Guatemala south throughout Central and South America to southern Brazil. Melittia pauper LeCerf—currently recorded only from Peru.
LITERATURE CITED
BRETHES, J. 1920. Insectos utiles y daninos del Rio Grande do Sul y de La Plata. Anales de la Sociedad Rural Argentiana 54:281-290. DuCKworTH, W. D. & T. D. EICHLIN. 1978. New species of clearwing moths (Lepi- doptera: Sesiidae) from North America. Proc. Entomol. Soc. Wash. 75:150-159. 1978. The type-material of Central and South American clearwing moths (Lep- idoptera: Sesiidae). Smithson. Contr. Zool. 261:1-28.
HEPPNER, J. B. & W. D. DuCKworTH. 1981. Classification of the superfamily Sesioidea (Lepidoptera: Ditrysia). Smithson. Contr. Zool. 314:1-144.
WALKER, F. 1856. List of the Specimens of Lepidopterous Insects in the British Mu- seum. Part 8, 271 pages. London: British Museum.
Journal of the Lepidopterists’ Society 88(1), 1984, 15-22
LIFE HISTORIES OF FOUR SPECIES OF PHILIRIS ROBER (LEPIDOPTERA: LYCAENIDAE) FROM PAPUA NEW GUINEA
MICHAEL PARSONS
Insect Farming and Trading Agency, Division of Wildlife, P.O. Box 129, Bulolo, Morobe Province, Papua New Guinea
ABSTRACT. The life histories of four species of Philiris, P. helena Snellen, P. agatha Grose-Smith, P. intensa Butler and P. ziska Grose-Smith, together with notes on their biologies, are described and illustrated.
Life histories of species of the genus Philiris Rober (Lycaenidae) from the Melanesian region have been little studied, but Forbes (1977) has recently detailed the life history of P. moira Grose-Smith from Papua New Guinea. Common and Waterhouse (1981) have briefly outlined the life history of P. innotata Miskin from Australia.
In the past, difficulty has been experienced with the placing of the correct females with the males of certain species of Philiris, but Sands (1979, 1980, 1981) has done much towards clarifying the taxonomy of the genus. The phyletic arrangement of all species, however, will re- main unclear until the biologies of further species are known. It is hoped that the addition to the literature of four new Philiris life his- tories, and information about the morphology of their early stages, will assist in such a study. I am also preparing to describe the life histories of Philiris harterti Grose-Smith, P. diana Waterhouse & Lyell, P. vi- oletta Rober, and P. praeclara Tite from Papua New Guinea, all of which feed on Litsea (Lauraceae).
Figs. 4 and 5 of the mature larva and pupa of P. moira are included here for ease of comparison. All life histories were studied from the Bulolo valley in the Morobe Province from approximately the center of the 10 km grid square reference DN50O. The early stages of all species show remarkable camouflage against their foodplants. The du- ration of the life cycle of each species was about one month from egg to adult.
Philiris helena Snellen
Egg. Diameter 0.75 mm; white, spherical when viewed from above, oval from the side, ventrally flattened; micropylar pit surrounded by six smaller pits; egg honeycombed with larger, regular, ovoid pits, the walls of which are produced into long, outward curving spicules which shorten gradually towards median line.
Larva. First instar. 1.55 mm in length, 0.5 mm in width; oval and elongate in dorsal profile; head pale brown; body pale green, edged with pale yellow; hirsute, fringed with fine white setae.
Second instar. 3.5 mm by 1 mm; anterior end slightly wider; head pale brown; body pale green with white middorsal line, broken at each segment.
16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Third instar. 5 mm by 2 mm; similar to second but white middorsal line bordered with tan brown.
Fourth instar. 10 mm by 4.5 mm; similar to third but middorsal line pinkish brown pattern encircled by cream; body color pink where larva has been feeding on small, young, pink leaves of foodplant, otherwise pale green; setal fringe 1.5 mm in length.
Fifth instar. 16 mm by 6.5 mm; similar to fourth but middorsal line broader (3 mm), forming a diffuse pattern of four lines of white dashes; laterally this pattern is continued but is fainter (Figs. 2 and 6).
Pupa. 11 mm in length, 5 mm in width; oval from above, ventrally flattened; hirsute, covered by fine pubescence of soft, white setae; dimorphic depending on substrate color being either pale green or pale brown; abdomen greenish yellow dorsally with two pale green dorsolateral lines either side of wider green middorsal line which continues onto thoracic segments. Supported by cremaster and fine silk girdle (Fig. 7). Duration, 12 days.
BIOLOGY. The foodplants are Macaranga aleuritoides F. Muell. and M. quadriglandulosa Warb. (Euphorbiaceae). Both species can grow up to 8 m in height. The leaves (Fig. 3) of both are broad and dark green. Those of M. quadriglandulosa are peltate, at the base (up to 35 mm in diameter) with a pointed tip and a serrated edge. Those of M. aleuritoides have five lobes are semipalmate and often grow to 70 mm in diameter. Both possess four to eight shiny, red, ovate glands on the upperside of the leaf bases which appear to be attractive to various. species of ants. Gressitt and Nadkarni (1978, p. 114) mention that these extrafloral nectaries are also attractive to several families of flies. M. quadriglandulosa has a coarse felt-like covering of hairs on its leaves. The sap of both is sticky, clear and gelatinous. The foodplants are common throughout the Lae-Wau region in regrowth areas from sea level to 1200 m.
Adults of P. helena can often be seen in abundance where the food- plant grows. They rest on the upper surface of the leaves and may be frequently seen drinking on damp sand.
Eggs are usually laid single on the leaf petiole and adhere to the long, felty hairs (Fig. 1). The larvae when young eat only the lower epidermis, creating windows in the leaf. Later they eat many full holes (Fig. 3), skeletonizing the leaf with no more than the veins to hold its former shape. Ants are almost always present on the Macaranga food- plants, but their association with the P. helena larvae is minimal, and they seem more preoccupied with the plant glands.
Pupae are always attached to the old leaf bracts at the base of the main stem. If the bracts are dry and brown, then the pupae tend to be brown also. If the bracts are still pale green, then the papae are of the same color.
From a final instar larva of P. helena which was collected wild, a small tachinid fly emerged. The species appears to be prone to attack by these parasitoids because numerous mummified skins of final instar
VOLUME 38, NUMBER 1 17
Fics. 1-3. Philiris early stages and feeding damage: 1, egg of P. helena on petiole of Macaranga aleuritoides; 2, eggs (above bract) and mature larva and empty pupal case of P. helena on bract of M. aleuritoides; 3, damage to leaves of a) M. aleuritoides, b) M. quadriglandulosa, c) M. involucrata, d) Ficus calopilina (by P. moira).
18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
larvae have been observed on the foodplants, each of which bore the ventral exit hole of a fly larva.
Philiris agatha Grose-Smith
Egg. Like that of P. helena.
Larva. The first three instars of P. agatha closely resemble those of P. helena but fourth instar pattern discernibly different. Larva of P. agatha slightly slimmer and more elongate than P. helena.
Fourth instar. 11] mm by 4 mm; head tan brown; body pale olive green; middorsal pattern bright white commencing from behind prothorax and composed of three parallel dashes on each segment which converge to form single broad line at anal end of larva; laterally, larva patterned with narrower parallel white lines; setal fringe 1.5 mm in length.
Fifth instar. 18 mm by 6 mm; similar to fourth (Fig. 8).
Pupa. 12 mm in length, 5 mm in width; hirsute, pale olive green, boldly patterned with white; prothorax with white, triangular markings above eyes; mesothorax with two broad white lines which curve around middorsal line; middorsal abdominal pattern com- posed of broad white dashes either side of midline; laterally abdomen bears single, broad, white, wavy line. Supported by cremaster and fine silk girdle (Fig. 9). Duration, 12 days.
BIoLoGy. The foodplant is Macaranga involucrata (Roxb.) Bail. (Euphorbiaceae), a tree which grows to 7 m in height. The leaves (Fig. 3) are variable in shape. They are either rounded or have three or five lobes, the central lobe being the longest. The leaves are covered with a fine white pubescence which makes them extremely soft and felt- like to the touch. They bear four small, often vestigial, glands on the leaf upperside near the petiole. The sap is clear, slightly sticky, and can have a strong camphor-like smell. It is a common plant of regrowth areas around the Bulolo valley.
Adults of P. agatha are less frequently seen than those of P. helena and may be classed as occasional in their habitat. They sometimes drink on damp sand and mud at creek margins and are very fast flying.
Eggs are usually laid on the underside of a small new leaf, or its petiole, at the apex of the foodplant. The larvae commence feeding on the young leaves. Then, as they grow, they move to feed on lower, older leaves. Leaf damage is shown in Fig. 3. No actual attendance of the larvae by ants was noted, although there were often brown tree ants on the foodplant.
The monomorphic pupae are invariably attached to the underside of a very young leaf (hardly larger than the pupa) at the apex of the foodplant. They match well the felty appearance and color of these leaves.
Philiris intensa Butler
Egg. Diameter 0.5 mm; white; hemispherical, with a regular covering of spicules.
Larva. First instar. 0.75 mm in length, 0.25 mm in width; oval in shape; head tan brown, lying well beneath prothorax; uniform greenish yellow; fringed with fine white setae.
VOLUME 38, NUMBER 1 19
Second instar. 3 mm by 1.5 mm; similar to first but with middorsal line patterned with reddish brown spots with dark green centers.
Third instar. 6 mm by 2.5 mm; similar to second but brown middorsal line broken centrally by two white spots.
Fourth instar. 8.5 mm by 3.5 mm; similar to third but spiracles white.
Fifth instar. 10 mm by 6.5 mm; similar to fourth but ground color darker green and matches that of leaf on which larva feeds; middorsal line of brown spots with white centers; setae fringe 1 mm in length, armed with prominent, outwardly directed barbs arranged in alternating rows along each hair (Fig. 10).
Pupa. 9 mm in length, 5 mm in width; smooth, not hirsute; apex of mesothorax with diffuse pattern of brown and white, ringed by olive green to edge ot pale yellow wings; abdomen pale lime-green with brown and white middorsal pattern on segments 1-5; spiracles white, those on first abdominal segment encircled by brown spots. Supported by cremaster and fine silk girdle (Fig. 11). Duration 10 days.
BIoLocy. The foodplant is Pipturus argenteus Willd. (Urticaceae), a common plant of creekside and regrowth areas, which grows to about 5 m tall. The leaves are pale to dark green and ovate with pointed tips and serrated edges. They average about 15-20 cm long and are felty to the touch. They are covered with minute white hairs. The small clusters of rounded, dimpled, opaque white fruit are gelatinous and are borne in alternating rows along fruit stalks.
Adults of P. intensa are commonly seen near the foodplant, and males are especially fond of drinking on damp sand.
Eggs are laid singly on the leaf underside, usually near the base. At all stages larvae eat the upper epidermis of the leaf and leave a char- acteristic long, narrow feeding trail of a meshwork of small veins. On occasion they were attended by small brown ants.
Pupation is always on the upperside of a large or small leaf of the foodplant and along the main vein just before it joins the petiole.
A small (8 mm long) orange ichneumonid wasp parasitoid was reared from a wild collected pupa.
Philiris ziska Grose-Smith
Egg. Diameter 0.55 mm; pale bluish-white, hemispherical with regular covering of spicules.
Larva. First instar. 0.75 mm in length, 0.25 mm in width; oval in shape; head tan brown, lying well beneath prothorax; uniform pale yellow; fringed with fine white setae.
Second instar. 3 mm by 1.5 mm; similar to first but color straw-yellow; dorsal vessel shows as dark green middorsal line; middorsal line white, not extending onto thoracic or anal segments; four orange-red spots laterally, two at center of body and two on penul- timate abdominal segment.
Third instar. 5 mm by 2.5 mm; similar to second but color greenish yellow with lateral spots brown; middorsal line extends along entire abdomen; setal fringe 0.55 mm in length.
Fourth instar. 8 mm by 4 mm, greatest width being at metathorax; dark green, paling to edges; middorsal line wholly white, or with pattern of brownish purple spots, three centrally and one on penultimate abdominal segment.
Fifth instar. 12 mm by 6 mm; dark green; middorsal line creamy white, along entire body length; setal fringe 2 mm in length (Fig. 12).
Pupa. 9 mm in length, 5 mm in width; similar to P. intensa; smooth, not hirsute; dark
20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 4-13. Philiris spp., larvae and pupae: 4 & 5, P. moira; 6 & 7, P. helena; 8 & 9, P. agatha; 10 & 11, P. intensa; 12 & 13, P. ziska.
green with middorsal pattern of chocolate-brown flecks on white on abdominal segments 1-4; same pattern repeated at apex of mesothorax and both areas surrounded by pale yellow; spiracles white. Supported by cremaster and fine silk girdle (Fig. 13). Duration, 12 days.
BioLocy. The foodplant is Malaisia scandens (Lour.) Bl. (Moraceae)
VOLUME 38, NUMBER 1 pas
which grows to about 5 m tall and is common along creeksides or road verges around Bulolo (altitude 700 m). Young leaves are soft and pale green. Old leaves are dark green and extremely brittle to the touch. They are ovate and may reach a length of 150 mm. The tree is a sprawling species which sends out long adventitious shoots. The small fruits are soft and red and are borne in many clusters along the branches.
Adults of P. ziska are often fairly common where the foodplant grows, especially alongside creeks.
Eggs are laid anywhere on the underside of new or old leaves, some- times five on a leaf. The newly emerged larva eats the top of the egg and leaves a white ring of shell which remains attached to the leaf. At all stages the larvae feed on the lower epidermis of the leaf and leave the upper epidermis as windows of tissue. They will not accept any other related plant species. The larval ground color is a perfect mimic of a vein. They were seen to be attended by small brown ants.
Pupation is always on the upperside of the leaf along the mid-vein just before it joins the petiole. Up to 12 pupae have been found dis- persed throughout one branch. The ground color is a perfect match of that of the leaf, and the brown markings edged with yellow resemble blemishes that are typically found on the leaves of various moraceous tree species.
P. ziska appears to be prone to attack by small, black chalcid wasp parasitoids. For example, one larva collected in its fourth instar ceased feeding and two 2 mm-long wasp larvae emerged to spin their cocoons beneath it before the larva died. Some pupae found had also died from what appeared to be a fungal disease.
DISCUSSION
A number of adults of each species have been reared and compared with material in the British Museum (Natural History) collection. In all cases males and females compared favorably with the pairings pre- sented in the collection (as figured by D’Abrera, 1977). Representatives of each species have been placed in the collection of the Insect Farming and Trading Agency in Bulolo.
Comparison of the early stages, especially the morphology of the pupae, shows that P. helena and P. agatha are closely related as are P. ziska and P. intensa. The pupa of P. moira resembles more the latter pair than those of P. helena or P. agatha but is very distinct in that it is prominently hirsute, not smooth as in P. ziska and P. intensa, and its maculation is different. It is very similar to that of P. innotata and also that of P. kapaura Tite. Pupae of the latter species I have
22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
found attached to saplings of a species of fig with large (500 mm diameter), hirsute leaves. Therefore, P. moira, P. innotata and P. ka- paura form a closely related group of species. The similarities between the larval morphologies and foodplant relations of P. ziska, P. intensa and P. moira (together with P. innotata and P. kapaura), however, suggest that these butterflies may also belong to the same species group. Females of P. moira are virtually indistinguishable from those of P. ziska.
It is noteworthy that I have found P. moira larvae feeding on Ficus semivestita Corner (Moraceae), near Bulolo, as well as Ficus calopilina Diels. The latter is the foodplant recorded for the species by Forbes (UST)
ACKNOWLEDGMENTS
My thanks to to Kenneth Airy Shaw of Kew Herbarium, England for his assistance in the identification of the foodplants and to Don Sands of C.S.I.R.O., Queensland, Australia for kindly sending me separates of his papers.
LITERATURE CITED
AirY SHAW, H. K. 1980. The Euphorbiaceae of New Guinea. Kew Bull. Additional series VIII. 243 pp.
COMMON, I. F. B. & D. F. WATERHOUSE. 1981. Butterflies of Australia. Second revised edition, Angus and Robertson, Sydney. 682 pp.
D’ABRERA, B. 1977. Butterflies of the Australian Region. 2nd edition. Landsdowne, Melbourne. 415 pp.
FORBES, G. R. 1977. The life history and polymorphic female of Philiris moira Grose- Smith (Lepidoptera: Lycaenidae) from Papua New Guinea. J. Aust. Entomol. Soc. 16:273-275.
GRESSITT, J. L. & N. NADKARNI. 1978. Guide to Mt. Kaindi: background to montane New Guinea ecology. Wau Ecology Institute Handbook No. 5. 135 pp.
SANDS, D. P. A. 1979. New species of Philiris Réber (Lepidoptera: Lycaenidae) from Papua New Guinea. J. Aust. Entomol. Soc. 18:127-183.
SANDS, D. P. A. 1980. The identity of Philiris nitens Grose-Smith (Lepidoptera: Ly- caenidae), with description of a new subspecies from Papua New Guinea. Aust. Entomol. Mag. 6:81-86.
SANDS, D. P. A. 1981. New species of Philiris Réber (Lepidoptera: Lycaenidae) from mainland New Guinea. J. Aust. Entomol. Soc. 20:89-96.
Journal of the Lepidopterists’ Society 88(1), 1984, 23-31
COURTSHIP BEHAVIOR OF THE GULF FRITILLARY, AGRAULIS VANILLAE (NYMPHALIDAE)
RONALD L. RUTOWSKI AND JOHN SCHAEFER Department of Zoology, Arizona State University, Tempe, Arizona 85287
ABSTRACT. The courtship behavior of the Gulf Fritillary, Agraulis vanillae L., is described from motion picture records of successful and unsuccessful courtships between free-flying males and tethered virgin females. During most but not all courtships the male performs a previously undescribed wing clap display in which he alights next to the female and repeatedly claps his wings together, often catching the female’s antenna between his wings during each clap. In the discussion it is suggested that this display presents chemicals signals to the female and has evolved in response to female choice for males that clearly announce their species identity to the female.
In recent years substantial literature has developed concerning the behavior and ecology of the heliconiine butterflies (for review: Brown, 1981). Surprisingly, only two papers (Crane, 1955; Gilbert, 1976) have dealt specifically with the mating behavior of heliconiines in spite of a growing amount of information on the nutritious materials passed by males to females during copulation and the female’s use of these ma- terials in oogenesis (Boggs and Gilbert, 1979; Boggs, 1981). The only courtship description per se for a heliconiine is to be found in Crane’s paper on Heliconius erato Hewitson. Information on courtship behav- ior is essential in any attempt to evaluate the selective consequences of inter- and intra-specific variation in male’s abilities to produce ac- cessory gland secretions during copulation (Rutowski et al., 1983).
The following study expands our knowledge of the behavior of hel- iconiines by describing the courtship of a member of this subfamily that is common in the southern United States, the Gulf Fritillary (Agraulis vanillae Linnaeus). Particular attention during this study was directed at describing a previously unreported display performed by A. vanillae males and its role in successful courtship. The discussion will focus on the potential functions of this unique display.
METHODS
Observations were made on a population of A. vanillae in suburban Tempe, Arizona. Preliminary studies in the spring of 1980 led to in- tensive study from March to June in 1981. From about 0800 h to 1600 h, adults of A. vanillae frequently visit Lantana spp. (adult nectar source) and Passiflora spp. (larval foodplant). Virgin females were ob- tained from larvae and eggs collected on Passiflora spp. either in the field or in cages in the laboratory. Larvae were reared to adulthood in translucent plastic shoeboxes (9 x 16 x 30 cm) on cuttings of Passi- flora. The shoeboxes were kept near a window which exposed the
24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
larvae to a normal light-dark regime. Temperature and humidity were not controlled.
To observe courtship, newly emerged, virgin females no more than four to five days old (most were one day old) were tethered (for tech- nique: Rutowski, 1978) and placed on a conspicuous perch on a Pas- siflora plant. Usuaully within a few minutes, passing males approached and courted the female. Courtships, both successful and unsuccessful, were filmed at 24 and 70 frames per sec with a Beaulieu 4008 ZM 2 super-8 movie camera.
RESULTS Successful Courtship
By separating males and females immediately after coupling, twen- ty-five successful courtships (leading to copulation) were filmed using ten females. No single female was used to film more than three court- ships. Of these 25 courtships, one involved two males and so, was disregarded, leaving 24 for the detailed analysis that gave rise to the description that follows.
Courtship began when either the male alit next to the female or the female began a flutter response as the male approached. Both were often preceded by a brief period during which the male hovered about 15 cm over the female before descending and initiating physical con- tact. Once the male alit he positioned himself with his wings open, his head close to that of the female, and the long axis of his body forming about a 45 degree angle with that of the female (Fig. 1). Once in this position the male began what will hereafter be referred to as the wing clap display. During this display the male’s body remained in position but he repeatedly clapped his wings shut and then quickly reopened them. Between claps the wings were opened to about 90 degrees rel- ative to one another. It was typical that in the position assumed by the male, the female’s antenna on the side next to the male was laid back between the male’s wings and was caught between them during each clap. During wing clapping and in some instances even before the male alit, the male’s claspers were visibly spread. Just as, or before, the male ceased clapping his wings he began probing by curling his ab- domen toward the female’s hindwings and attempting to insert its tip up between her closed wings. Coupling occurred between the female’s hindwings where its occurrence could not be closely monitored. How- ever, it is assumed that either when it occurred or shortly thereafter the male became still briefly before he began slowly waving his wings and moving to a position facing away from the female while still
VOLUME 38, NUMBER 1] 25
Yeon ~
Fic. 1. Above: an A. vanillae male (left) as he appears when courting a female (right) with a wing clap display. Below: the same pair except that the male has now begun probing. In both figures, note that the female’s antenna on the side next to the male is laid back between his wings.
26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Temporal structure of successful courtship in Agraulis vanillae.
Time event occurs before end of courtship
Courtships with wing clap display Courtships without wing clap display
Event x + SD (s) range (sec) n x + SD (s) range (sec) n
First contact by 6 10.9 = 5.1 PPro) UT 8.04 + 3.67 5438-153 6
Last contact by 6 Chil ae BOX AAVaIS.j3 — IT Oj) as 2/7 1.8-7.46 6 6 begins wing clap
display ees) 22 2 1.92-15.8 17 — —-
Ol
6 begins probing 3.538 + 3.18 0.5-13.0 14 2.97 + 1.46 1.46-5.53 6 ends wing clap
display Wess) ae Pho) OS f—alPAish 7 — = a 6 stops moving 0 0
coupled. Wing waving often evolved directly into strong wing flapping by the male in an effort to initiate a post-nuptial flight.
This sequence of events was observed in 14 of the 24 courtships filmed. In another three records the male broke contact with the perch and alit again at least once but no more than twice. In one case the male performed a bout of probing while perched the first time. After alighting a final time, all of these males performed wing claps before probing.
A more striking variant of the courtships described above was ob- served in seven courtships in which the wing clap display was com- pletely omitted. In these cases the male simply began probing imme- diately after alighting for the last time as they did in some of the courtships with wing claps but in these cases the males were successful. Hence, the wing clap display is not a requisite part of successful court- ship.
Females opened and closed or fluttered their wings in only eight of the 24 courtships. In six of these the female performed a single flutter before or at the time of first contact by the male. If the male made repeated contact each contact often elicited a single flutter. Twice flutters were observed during the wing clap display and twice after the male had become still. Fluttering in immediate response to male contact was observed in three of the seven courtships in which males did not wing clap.
The temporal structure of courtships in which the male performed a wing clap display is shown in Table 1. In these summaries, there are some courtships in which the first contact was also the last; hence, some data points were used in the summaries for both events. Also, for convenience and summary purposes, we regarded the last time the male initiated probing as the time when the male began probing. This was not true of three courtships. The sample size of the summary of
VOLUME 38, NUMBER 1 ; PLT)
30
Frequency (°%o)
O — O ICO> 2007300; "400 >400
Duration (msec)
Fic. 2. The frequency distribution of duration for 132 wing claps. See text for details.
this event is less than that for the other events because in three cases the angle of filming did not permit viewing of the beginning of prob- ing.
Courtships with the wing clap display have an average duration of about 11 sec and wing clapping begins within a second after the male alights for the last time. Probing usually began in the second before the display ended and slow wing waving followed coupling within 2 sec.
Table 1 also summarizes the temporal structure of courtships in which the male did not perform a wing clap display. The sample size for some events is less than seven, either because the film record began after the event occurred or because the filming angle prohibited view- ing the event. Statistical comparisons of the time of occurrence of events held in common between the courtships with and without the wing clap display revealed that none of the differences were signifi- cant. In particular, neither the time of first contact nor the time of last contact differed from one type of courtship to the next. Factors con- tributing to this lack of differences were the high variances associated with the times of occurrence and the small sample sizes for courtships without the wing clap display.
The average duration of the wing clap display was 4.73 + 3.34 sec (n = 17). Fig. 2 shows the average duration of each clap from the time the wings start to close from their spread position until they start to close for the next clap. The data are taken from 132 wing claps per-
28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
formed in five displays of 6, 12, 16, 41, and 57 claps, respectively, and filmed at 70 frames per sec. The histogram shows a mild but distinct bimodality. This reflects the fact that during the display the temporal patterning of the claps may take one of three forms: (1) a series of long claps, (2) a series of short claps, and (3) a series of alternating long and short claps. From the films we also measured that, regardless of overall clap duration, the wings were closed an average of 26.4 + 9.37 msec (n = 127). Hence, the difference in duration between long and short claps is due primarily to a difference between them in the time the wings are open.
Most courtships were filmed at 24 frames per sec which made count- ing of individual wing claps difficult; thus, we were unable to obtain a good average for the number of wing claps per courtship. However, using a mean wing clap display duration of 4.73 sec and an average of 165 msec per clap, it may be estimated that in the typical courtship the male performs about 30 wing claps.
Unsuccessful Courtships
Courtships that did not lead to copulation were studied in less detail than those that lead to copulation. This was because unsuccessful court- ships were more variable and difficult to characterize. In addition many of the film records of unsuccessful courtship were incomplete. How- ever, 20 complete records were obtained and are summarized here. In five of these courtships the male contacted the female and hovered over her briefly before departing. In the other 15 the male alit next to the female at least once during the courtship. Of the males that alit, four departed without performing a wing clap display while the other 11 performed the display at some point in the courtship. Two males actually probed before wing clapping while three did not begin prob- ing until they had wing clapped for some time. The other six males left the female after wing clapping without probing. During these courtships the females either did nothing (10 cases), fluttered the wings (8 cases), or assumed a posture like the pierid mate refusal posture (Obara, 1964) with the wings spread and the abdomen held perpen- dicular to the plane of the wings (2 cases). When females fluttered or spread their wings it could be seen that the pair of glands associated with the tergites at the end of the abdomen were periodically everted. Whether or not this occurs when the wings are closed was not deter- mined.
In summary, courtship terminated before coupling either because the male left before an attempt (males did not probe in 16 cases) or because the female did not behave in a way that permitted coupling (4 cases).
VOLUME 38, NUMBER 1 29
DISCUSSION
The wing clap display performed by males of Agraulis vanillae is clearly an important part of successful courtship. Its form is reminis- cent of displays performed by males of some other nymphalids. The male of the grayling (Hipparchia (=~Eumenis) semele Linnaeus) after alighting next to a perched female moves so that he is face to face with the female and, by rocking forward with the wings open, catches the female’s antennae between his forewings. The male then closes his wings and rocks gently back, drawing the female’s antennae across patches of scent scales on the male’s forewings (Tinbergen et al., 1942). During the courtship of the great spangled fritillary (Speyeria cybele Fabricius), the male assumes a position perched next to the female like that seen in A. vanillae, and “‘at intervals he would suddenly open and close his wings’ (Clark, 1932:110). The interval between these open- ings and closings becomes less as the courtship progresses. Magnus (1950) reports that males of the fritillary (Argynnis paphia Linnaeus) when perched alongside and facing a female, clap their wings in such a way that the female’s head and antennae are caught between them. In Heliconius erato, the closest relative of A. vanillae that has been carefully studied, the male persists in flapping his wings after alighting and while moving into position for coupling (Crane, 1955). Perhaps this behavior was the one from which the wing clapping display of A. vanillae was evolutionarily derived.
Indications are that the wing clapping display is involved in pre- senting chemical signals to the female. Obviously, the male’s position behind and to the side of the female is poor for presentation of visual stimuli, and the fact that the female’s antenna is between the male’s forewings during the display suggests that the display is designed to deliver a chemical signal. This conclusion is reinforced by the obser- vation that androconia that are likely to be scent-producing are found on the dorsal surface of the male’s forewings along several of the veins (M,, M,, M,, Cu,, Cu,, 2A; Muller, 1877). In addition there are large apparently glandular structures associated with the internal faces of the claspers that might produce a chemical signal. The claspers are often spread or opened during most of the courtship. Experiments are planned to determine the roles of these structures in the courtship behavior of A. vanillae.
Rutowski (1983) has recently discussed the selection pressures that have led to the evolution of species-specific male courtship displays in butterflies. A similar analysis of the courtship of A. vanillae suggests that selection for signals that announce the species identity of the male to the female has been of particular importance. The nutrient invest- ment of A. vanillae males and the general ecology and behavior of
30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
this species is not so different from that of the other species of butter- flies as to suggest that they led to the evolution of the wing clapping display (Rutowski et al., 1983). In addition, although A. vanillae females may occasionally approach and chase males as observed in other species (Crane, 1955; Rutowski, 1980; Rutowski et al., 1981; Wik- lund, 1982), it does not seem that problems of sexual discrimination for females are any more severe than they are in Heliconius erato, which like A. vanillae, is essentially sexually monomorphic with re- spect to visual characters but whose courtship lacks the wing clap display (Crane, 1955).
Species discrimination would seem, however, to pose a potential problem for A. vanillae females. A field guide to the butterflies of North America (Pyle, 1981) reveals that there are at least eight but- terflies sympatric with A. vanillae that are very similar in color and markings. These species include Danaus gilippus Cramer, D. plexip- pus Linnaeus, Dryas julia Fabricius, Marpesia petreus Cramer, Li- menitis archippus Cramer, Dione moneta Hiibner, Anaea floridalis Johnson and Comstock, and A. andria Scudder. Given that an A. va- nillae female is likely to be approached and courted by males of any of these species, selection might favor those females that prefer con- specific males that perform a display or offer a signal clearly indicative of their species identity. While it is known that the danaids have their own unique male courtship behavior (hairpencilling: Brower et al., 1965; Pliske, 1975) the other species are not behaviorally well-known enough to assess the extent to which their courtships are different from that of A. vanillae.
A final question about the wing clap display concerns the fact that it is not performed in all successful courtships like the wing spread display sometimes performed by males of the pierid, Nathalis iole Boisduval (Rutowski, 1981(83)). Why do some females accept a male without the performance of the display? Obviously, females vary in receptivity, and some will accept males without the display. These may be males with such potent chemical signals that they are able to sat- isfactorily stimulate the female without the display. In any event it seems from both successful and unsuccessful courtships that males will attempt copulation without wing clapping, which in turn suggests that the display has certain costs for males. Just what these costs are or might be is currently not clear (although, see: Burk, 1982).
ACKNOWLEDGMENTS Thanks to Michael Boppré for showing us the location of androconia in A. vanillae,
to Mark Newton for assistance in filming courtship, and to the National Science Foun- dation for financial support (Grant No. BNS 80-14120).
VOLUME 38, NUMBER 1 ; 3]
LITERATURE CITED
Bocecs, C. L. 1981. Selection pressures affecting nutrient investment at mating in heliconiine butterflies. Evolution 35:931-940.
Boccs, C. L. & L. E. GILBERT. 1979. Male contribution to egg production in butterflies: evidence for transfer of nutrients at mating. Science 206:83-84.
BROWER, L. P., J. V. Z. BROWER & F. P. CRANSTON. 1965. Courtship behaviour of the queen butterfly, Danaus gilippus berenice (Cramer). Zoologica 50:1-39.
Brown, K. S., JR. 1981. The biology of Heliconius and related genera. Ann. Rev. Entomol. 26:427-—456. ;
BuRK, T. 1982. Evolutionary significance of predation on sexually signalling males. Florida Entomol. 65:90-104.
CLARK, A. H. 1932. The butterflies of the District of Columbia and vicinity. Smithson- ian Institution, U.S. National Museum Bulletin 157. U.S. Government Printing Office, Washington.
CRANE, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara Hewitson, with special reference to the social use of color. Zoologica 40:167-196.
GILBERT, L. E. 1976. Postmating female odor in Heliconius butterflies: a male-con- tributed antiaphrodisiac? Science 193:419—420.
Macnus, D. B. E. 1950. Beobachtungen zur Balz und Eiablage des Kaisermantels Argynnis paphia L. (Lep., Nymphalidae). Zeit. Tierpsych. 7:435-449.
MULLER, F. 1877. The scent-scales of the male of Dione vanillae. Kosmos 2:38-41. (Translation in: Longstaff, G. B. 1912. Butterfly Hunting in Many Lands. Longmans, Green, and Co., London.)
OparRA, Y. 1964. Mating behavior of the cabbage white Pieris rapae crucivora. II. The ‘mate refusal posture’ of the female. Dobut. Zasshi 73:175-178.
PLISKE, T. 1975. Courtship behavior of the monarch butterfly, Danaus plexippus L. Ann. Entomol. Soc. Amer. 69:143-151.
PYLE, R. M. 1981. The Audubon Society Field Guide to the North American Butterflies. A. A. Knopf, Inc., New York. 916 pp.
RUTOWSKI, R. L. 1978. The form and function of ascending flights in Colias butterflies. Behav. Ecol. Sociobiol. 3:163-172.
RUTOWSKI, R. L. 1980. Courtship solicitation by females of the checkered white but- terfly, Pieris protodice. Behav. Ecol. Sociobiol. 7:113-117.
RUTOWSKI, R. L. 1981(83). Courtship behavior of the dainty sulfur butterfly, Nathalis iole, with a description of a new facultative male display. J. Res. Lepid. 20:161- 169.
RUTOWSKI, R. L. 1983. The wing waving display of Eurema daira males (Lepidoptera, Pieridae): its structure and role in successful courtship. Anim. Behav. 31:985—989.
RUTOWSKI, R. L., C. E. Lonc, L. D. MARSHALL & R. S. VETTER. 1981. Courtship solicitation by Colias females (Lepidoptera: Pieridae). Amer. Midl. Nat. 105:334- 340.
RUTOWSKI, R. L., M. NEWTON & J. SCHAEFER. 1988. Interspecific variation in the size of the nutrient investment made by male butterflies during copulation. Evolution 34:708-713.
TINBERGEN, N., B. J. D. MEEUSE, L. K. BOEREMA & W. W. VAROSSIEAU. 1942. Die Balz des Samtfalters, Euwmenis (=Satyrus) semele (L.). Zeits. Tierpsych. 5:182-226.
WIKLUND, C. 1982. Behavioural shift from courtship solicitation to mate avoidance in female ringlet butterflies (Aphantopus hyperanthus) after copulation. Anim. Behav. 30:790-793.
Journal of the Lepidopterists’ Society 38(1), 1984, 32-39
CHECKLIST OF MANITOBA BUTTERFLIES (RHOPALOCERA)
PAUL KLASSEN Box 212, Elm Creek, Manitoba
ABSTRACT. A list of butterflies (Rhopalocera) occurring in Manitoba is compiled from records of resident and non-resident collectors, published literature, museums, uni- versity collections and the author’s collection.
It has been forty years since the last published checklist of Manitoba butterflies (Rhopalocera) by G. Shirley Brooks in “A Revised Check List of the Butterflies of Manitoba” (1942). Since that list is out-dated and not readily available, the present list has been prepared, including a number of species not previously recorded.
Many parts of Manitoba have been collected very sparingly, and I am afraid the habitat will be destroyed before these areas have been studied. There is very little virgin prairie left in this province, and some of that is not accessible to collectors. Some species in this habitat are threatened. Most of the province, however, is largely undeveloped, and there are large tracts of virgin forests, marshes, bogs, taiga and tundra untouched by the bulldozer. This will pigea lly remain so for a long time.
It is hoped that this checklist will encourage more study of the fascinating butterfly fauna of Manitoba. Any comments and criticism of this list and the notes following it will be appreciated.
The sequence of taxa follows the order of the Miller and Brown Catalogue/Checklist (1981), and the species are numbered accord-
ingly. DISTRIBUTION
Most of Manitoba is covered by boreal forest including many lakes, rivers and bogs. The southern part, especially toward the west, consists of grasslands changing to an aspen parkland region farther north. The area bordering the coast of Hudson Bay contains some tundra.
For practical reasons the following definitions are used:
FN = Far North. An area just southwest of Hudson Bay. Here Churchill and vicinity have been collected quite intensively and most far north records are from here.
N = North. Northern third of the province excepting the far north. This area consists of boreal forest with make lakes, rivers and bogs. Not much collecting has been done in this area.
NW = Northwest. The western half of “N”’.
NE = Northeast. The eastern half of “N”.
C = Central. The middle third of the province running north and south. Geographi- cally this area is like the north. Very little collecting has been done here.
VOLUME 38, NUMBER 1 a
WC = West Central. The western half of “C”.
EC = East Central. The eastern half of “C”’.
S = South. The southern third of the province.
SW = Southwest. The western half of “S”. This area consists of dry prairie in the southwest turning to moist prairie farther north and east. A large part of this is in the parkland or transition zone and is broken up by the Turtle Mountain in the extreme south and the Riding and Duck Mountains to the north. Lake Manitoba is east of Riding Mountain. Most of this area is agricultural land with very little virgin prairie left.
SE = Southeast. The eastern half of “S’’. This area consists of boreal forest in the north, mixed forest farther south with moist prairie along the Red River valley. The southern end of Lake Winnipeg is included in this area. Most of the prairie is now in agriculture.
G = General Distribution. Covers the whole province.
Note: Only those areas for which there are actual records of butterflies have been listed. Some species probably cover a much larger area than is indicated in the checklist below.
CHECKLIST OF MANITOBA BUTTERFLIES (RHOPALOCERA)
Hesperiidae Latreille Epargyreus Hiibner
clarus clarus (Cramer)—S, C Ta. Thorybes Scudder
pylades (Scudder)—G (except FN) 48. Erynnis Schrank
icelus (Scudder & Burgess)—G (except FN) 83.
brizo brizo (Boisduval & Leconte)—S, C 84a.
juvenalis juvenalis (Fabricius)—S 85a.
martialis (Scudder)—SE 92.
lucilius (Scudder & Burgess)—S 96.
persius persius (Scudder)—SW, NE, FN 99a. Pyrgus Hiibner
centaureae freija (Warren)—SE, WC, N, FN 100a.
communis (Grote)—S 104. Pholisora Scudder
catullus (Fabricius)—S eS: Carterocephalus Lederer
palaemon mandan (Edwards)—S, C 120a. Ancyloxypha
numitor (Fabricius)—S, C 142. Oarisma Scudder
poweshiek (Parker)—S 144.
garita (Reakirt)—S 145. Thymelicus Hiibner
lineola (Ochsenheimer)—SE 150. Hesperia Fabricius
uncas uncas Edwards—SW 156a.
comma assiniboia (Lyman)—S 158b.
c. borealis Lindsey—NE, FN 158d.
ottoe Edwards—S 160.
leonardus Harris—SE 161.
pawnee Dodge—SW 162.
dacotae (Skinner)—S 169.
sassacus manitoboides (Fletcher)—SE 171b.
nevada (Scudder)—SW £78.
34
Polites Scudder
coras (Cramer)—S
themistocles (Latreille)—S
mystic dacotah (Edwards)—S, WC Atrytone Scudder
logan lagus (Edwards)—SW Poanes Scudder
hobomok (Harris)—S Euphyes Scudder
ruricola metacomet (Harris)—S, C Atrytonopsis Godman
hianna hianna (Scudder)—S Amblyscirtes Scudder
hegon (Scudder)—SE
vialis (Edwards)—S
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
174.
179. 181b. 189b.
1973 217b. 219a.
235. 245.
Papilionidae Latreille
Papilio Linnaeus polyxenes asterius Stoll—SE bairdii Edwards—SW kahli F. & R. Chermock—SW
machaon hudsonianus Clark—SW, WC, N
cresphontes Cramer—S
glaucus canadensis Rothchild & Jordan—G
troilus troilus Linnaeus—SE(?)
308a. 308. 306. 310b. 314. 320b. 325a.
Pieridae Duponchel
Pieris Schrank protodice Boisduval & Leconte—S
occidentalis occidentalis Reakirt—S, FN
napi oleracea Harris—G rapae (Linnaeus)—S, FN Euchloe Hiibner
ausonides mayi F. & R. Chermock—G
olympia (Edwards)—SW
Colias Fabricius philodice philodice Godart—S, C eurytheme Boisduval—S, FN
alexandra christina Edwards—SW, WC
hecla hela Strecker—FN
boothii Curtis—FN
nastes moina Strecker—FN gigantea gigantea Strecker—FN g. mayi F. & R. Chermock—SW
pelidne pelidne Boisduval & Leconte—FN
interior interior Scudder—S, WC palaeno chippewa Edwards—FN cesonia (Stoll) —SW Eurema Hiibner mexicana (Boisduval)—SW Nathalis Boisduval iole Boisduval—S
334. 335a. 336d.
338.
84le. 344.
351a.
on2. 355e. 357b.
358. 360c. 362a. 362c. 368a. 364a. 365a. 368a.
380.
389.
Lycaenidae Leach
Feniseca Grote
tarquinius tarquinius (Fabricius)—S
Lycaena Fabricius xanthoides dione (Scudder)—S
39 la.
395b.
VOLUME 38, NUMBER 1
hyllus (Cramer)—S epixanthe michiganensis Rawson—SE dorcas dorcas Kirby—G helloides (Boisduval)—S Harkenclenus dos Passos titus titus (Fabricius)—S Satyrium Scudder acadica acadica (Edwards)—SE a. watrini (Dufrane)—SW edwardsii (Grote & Robinson)—S_ - calanus falacer (Godart)—S liparops fletcheri (Mitchener & dos Passos)—S Incisalia Scudder augustus augustus (Kirby)—S, C, N polios polios Cook & Watson—S, C, N henrici henrici (Grote & Robinson) —SE niphon clarki Freeman—S, C eryphon eryphon (Boisduval)—N Strymon Hiibner melinus humuli (Harris)—S Everes Hiibner comyntas comyntas (Godart)—S, C amyntula aibrighti Clench—SW, WC, N, FN Celastrina Tutt ladon lucia (Kirby)—G (except FN) l. argentata (Fletcher)—SW Glaucopsyche Scudder lygdamus couperi Grote—G (except FN) l. afra (Edwards)—-SW Plebejus Kluk
Argyrognomon scudderii (Edwards)—SW, WC, FN
a. nabokovi Masters—SE
melissa melissa (Edwards)—-SW
m. samuelis Nabokov—SE
saepiolus amica (Edwards)—G optilete yukona (Holland)—C, N, FN franklinii franklinii (Curtis)—FN
f. lacustris (Freeman)—C, N
f. rustica (Edwards)—S
Heliconiidae Swainson
Agraulis Boisduval & Leconte vanillae incarnata (Riley)—SW
Nymphalidae Swainson
Euptoieta Doubleday claudia (Cramer)—S Speyeria Scudder
cybele pseudocarpenteri (F. & R. Chermock)—S
aphrodite aphrodite (Fabricius)—SE
a. manitoba (F. & R. Chermock)—S idalia (Drury)—S
edwardsii (Reakirt)—SW
callippe calgariana (McDunnough)—S atlantis atlantis (Edwards)—SE
a. hollandi (F. & R. Chermock)—S, WC
565d. 566a. 566d.
567.
569. o72p. o7 4a. o74c.
36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
a. dennisi dos Passos & Grey—SW 574u. mormonia eurynome (Edwards)—SW 576i. Boloria Moore eunomia dawsoni (Barnes & McDunnough)—G 578c. selene atrocostalis (Huard)—S, WC, FN 579f. bellona bellona (Fabricius)—S, WC, N 580a. frigga saga (Staudinger)—-G 58 la. improba improba (Butler)—FN 582a. polaris stellata Masters—FN 585b. freija freija (Thunberg)—G 586a. titania boisduvalii (Duponchel)—FN 589a. t. grandis (Barnes & McDunnough)—G (except FN) 589c. chariclea arctica (Zetterstedt)—N 590a. Chlosyne Butler gorgone carlota (Reakirt)—S 605b. nycteis nycteis (Doubleday)—S 606a. n. reversa (F. & R. Chermock)—SW 606c. harrisii harrisii (Scudder)—S 607a. h. hanhami (Fletcher)—S, WC 607c. Phyciodes Hiibner tharos tharos (Drury)—G 623b. batesii (Reakirt)—S, C 624. Euphydryas Scudder phaeton phaeton (Drury)—SE 635a. Polygonia Hiibner interrogationis (Fabricius)—S 636. comma (Harris)—S 637. satyrus neomarsayas dos Passos—S 638b. faunus faunus (Edwards)—S, WC 639a. gracilis (Grote & Robinson)—N, FN 643. progne (Cramer)—S, N, FN 645. Nymphalis Kluk vau-album j-album (Boisduval & Leconte)—S 646a. californica californica (Boisduval)—S 647a. antiopa antiopa (Linnaeus)—G 648a. milberti milberti (Godart)—S, WC, FN 649b. Vanessa Fabricius virginiensis (Drury)—S, FN 650. cardui (Linnaeus)—S, WC, FN Gal. atalanta rubria (Fruhstorfer)—S, C, N 653a. Junonia Hiibner coenia Hiibner—S 656. Limenitis Fabricius arthemis arthemis (Drury)—SE 663a. s. rubrofasciata (Barnes & McDunnough)—S, C 663b. archippus archippus (Cramer)—S, WC 664a.
Satyridae Boisduval Lethe Hiibner
anthedon Clark—S TU
eurydice eurydice (Johansson)—S 718a. Euptychia Hiibner
cymela cymela (Cramer)—S, C 723a. Coenonympha Hiibner
inornata inornata Edwards—SW, C 728d.
i. benjamini McDunnough—S 728e.
VOLUME 38, NUMBER 1] OU
Cercyonis Scudder
pegala olympus (Edwards)—S 732e. Erebia Dalman rossi ornata Leussler—FN 737a. disa mancinus Doubleday & Hewitson—G 738a. discoidalis discoidalis (Kirby)—G 7Ala. theano sofia Strecker—FN 742a. epipsodea freemani Ehrlich—SW, WC 744b. Neominois Scudder ridingsii ridingsii (Edwards)—SW . 748a. Oeneis Hiibner macounii (Edwards)—S (ile chryxus calais (Scudder)—C 752b. uhleri varuna (Edwards)—-SW 753a. alberta alberta Elwes—S 754a. bore ssp.—FN 756. jutta ascerta Masters & Sorensen—SE 757b. j. ridingiana F. & R. Chermock—SW, WC Tne: j. harperi Chermock—N, FN Told. melissa semplei Holland—FN 758c. polixenes polixenes (Fabricius)—FN 759a.
Danaidae Duponchel
Danaus Kluk plexippus (Linnaeus)—S, WC 760.
NOTES
C. palaemon mandan—Type-locality—“Lake Winnipeg’, restricted to Pine Ridge by F. M. Brown and L. Miller, is common in most wooded areas of southern Manitoba.
T. lineola, first recorded from Manitoba in the early 1970’s, is now firmly established in Winnipeg and east of there (Preston and Westwood, 1981).
H. comma borealis from Churchill, should perhaps have another subspecific name.
P. asterius polyxenes is rare in southeastern Manitoba.
P. kahli—Type-locality—“Riding Mtns., Man.”, is found mostly in the Riding Moun- tain and Duck Mountain area, but some are found as far east as the Red River. There seems to be some intergradation between this and the latter species and P. machaon.
E. ausonides mayi—Type-locality—“Riding Mtns., Manitoba”.
C. hecla hela—Type-locality—“‘above Fort Churchill”.
C. nastes moina—Type-locality— ‘above Fort Churchill”.
C. g. giganiea—Type-locality—‘“west coast of Hudson Bay above Fort York”’.
C. g. mayi—Type-locality— “Riding Mtns., Manitoba”’.
L. d. dorcas—Type-locality— ‘Lat. 54°”, restricted to The Pas, Manitoba, by Ferris.
H. t. titus—In southwestern Manitoba some specimens perhaps belong to immaculo- sus.
S. a. acadica flies in the southeast and watrini in the southwest.
S. liparops fletcheri—Type-locality—““Manitoba’”’.
C. ladon argentata—Type-locality—‘“Cartwright, Manitoba’, flies in southwestern Manitoba, while lucia is found in most of the rest of the province.
G. lygdamus afra—Type-locality— “Deer River country ’’, restricted to vic. Brandon, Man. by F. M. Brown, flies in southwestern Manitoba with couperi in the rest of the province. They are quite hard to tell apart as they are variable in size and in the ventral spots and color.
P. argyrognomon scudderii—Type-locality—“Lake Winnipeg, Manitoba’, flies in western Manitoba. The bands of submarginal orange lunules, both ventral and dorsal, are on the average more complete than in nabokovi which is found in the southeastern part of the province. The subspecies are very variable and so difficult to tell apart.
38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
P. m. melissa flies in south and central Manitoba with samuelis in the southeastern corner.
P. f. franklinii is found in the Churchill area.
P. f. lacustris—Type-locality—“Norway House’, is in central Manitoba.
P. f. rustica occurs in southern Manitoba.
S. aphrodite manitoba—Type-locality—“Sand Ridge’, which is east of Riding Moun- tain. These formerly went under the name of mayae and occupy most of southern Manitoba. S. a. aphrodite is in the extreme southeast.
S. a. atlantis is sometimes found in the extreme southeast.
S. a. hollandi—Type-locality— ‘Riding Mtns., Manitoba’, flies in most of southern Manitoba.
S. a. dennisi—Type-locality—““Beulah, Manitoba’, closely resembles lais from Sas- katchewan. It was known by that name for some time. It probably intergrades with that subspecies. In the Riding Mountain area, hollandi flies in the wetter areas and dennisi in the drier, more open areas, but adults feed at flowers in the same places. Should these belong to different species?
B. polaris stellata—Type-locality—“Churchill, Manitoba’, flies in the Churchill area in odd-numbered years.
B. titania grandis flies in southern and central Manitoba.
B. t. boisduvalii from the north is abundant at Churchill.
C. nycteis reversa—Type-locality—“Riding Mountains, Manitoba’, refers to most specimens from Manitoba, however some are like subspecies nycteis in facies.
C. harrisii hanhami—Type-locality— ‘Bird Hill, near Winnipeg, Manitoba’, flies in southern Manitoba. There seems to be some intergradation with the subspecies harrisii, as some specimens are like the ones from Ontario in facies.
L. arthemis rubrofasciata—Type-locality— “Manitoba, Saskatchewan, Alberta’, is common in southern Manitoba with some arthemis found in the eastern part, where they intergrade.
L. eurydice, formerly known as transmontana in this area, is quite distinct as the ground color very pale, almost white in some specimens, as compared to the dark spec- imens found in eastern Ontario.
C. i. inornata—Type-locality—“Lake Winnipeg’, emended to “Saskatchewan River between Lake Winnipeg and The Pas, Man.”, by F. M. Brown, flies mostly in the parklands area and benjamini on the prairies in southern Manitoba. C. ochracea probably does not fly in the province.
E. rossii ornata—Type-locality—“Churchill”, is abundant in the Churchill area most years.
E. theano sofia—Type-locality—“Fort Churchill, Manitoba” was formerly known as canadensis, is locally common at Churchill most years.
O. bore ssp., flies at Churchill in even-numbered years. It is quite rare most years, but locally more common, in some. It is quite variable and has a darker ground color than bore hanburyi from Baker Lake, N. W. Territories, Canada.
O. jutta ascerta flies in eastern Manitoba. It is dark and the orange bands are less developed or even lacking in some males. It is found in the odd-numbered years with the rare exception.
O. j. ridingiana—Type-locality—“Riding Mountains, Manitoba’’, is found mostly in even-numbered years in western Manitoba, but some fly every year. The orange bands are well developed.
O. j. harperi—Type-locality—“Gillam, Manitoba’, is a little smaller than the two preceding subspecies. It is quite variable with the orange bands in some females well developed to faint in others. It resembles alaskensis. It is common at Churchill every year.
Some butterflies have, over a number of years, been taken very rarely in Manitoba. The following are probably strays from the south: P. cresphontes, P. t. troilus, C. cesonia, E. mexicana, N. iole, A. vanillae, S. idalia and N. californica.
There are also some species that, although rare, apparently breed in the province. Some of these may be seen to be more common after the areas have been more exten-
VOLUME 38, NUMBER 1 39
sively collected. Here is a list of these: P. catullus, O. poweshiek, H. ottoe, H. dacotae, H. nevada, A. logan, P. bairdii, E. olympia, L. epixanthe, I. eryphon, S. melinus, S. edwardsii, S. callippe, E. phaeton, P. gracilis, J]. coenia and N. ridinsii.
A small number have been included that maybe should be deleted from the list. Papilio bairdii is included based on records from Beulah and Birtle and records of bairdii ore- gonia from Beulah. The author suspects that these may be misidentified specimens of machaon or kahli. The latter is quite variable. C. boothii, C. pelidne and B. improba have been recorded from “north Manitoba”. C. boothii and B. improba could occur northwest of Churchill and C. pelidne could be found east of there. B. chariclea is recorded from Kettle Rapids. Formerly Boloria titania from Manitoba were called chari- clea titania. As there is no proven reason to the contrary, the above specimens are all included in the checklist.
The following species, included in older lists, have been deleted: H. comma manitoba, no records for Manitoba.
P. zelicaon probably does not occur in the province. The records possibly refer to machaon or forms of kahli.
E. ausonides coloradensis is supposed to fly in southeastern Manitoba. I cannot see any difference between the mayi, type-locality, “Riding Mtns., Man.’’, and the ausonides from the rest of the province.
P. zephyrus recorded from Aweme and Beulah probably were misidentified Polygonia.
S. cypris = ethene and S. a. columbia included in older lists probably are S. a. mani- toba, which they closely resemble.
S. lais, included in old lists flies in Saskatchewan and Alberta and intergrades with dennisi in Manitoba.
S. calanus calanus recorded as calanus is deleted as the subspecies that flies in the province is falacer.
S. heathii, also omited, because it is an aberration of the latter.
S. liparops strigosa does not occur in the province. Although some specimens of fletch- eri from Manitoba closely resemble strigosa with no orange spots on the fore-wings, these occur in the same populations together with specimens having orange patches covering one-half of the front wings. This subspecies is very variable.
Mitoura spinetorum probably does not fly in Manitoba.
ACKNOWLEDGMENTS
Many thanks go out to all the following, who sent in data and helped in other ways: George T. Austin, Patrick J. Conway, Richard E. Gray, W. W. Gregory, R. J. Heron, Ronald R. Hooper, Brian McKillop and William B. Preston of the Manitoba Museum of Man and Nature, David Parshall, James D. Reist, Oakley Shields and Jim Troubridge, and to the personnel of the University of Manitoba and the Canada Agriculture Research Station.
LITERATURE CITED
BROOKS, G. SHIRLEY. 1942. A check list of the butterflies of Manitoba. Can. Entomol. 74:31-36.
MILLER, LEE D. & F. MARTIN BROWN. 1981. A Catalogue/Checklist of the Butterflies of America North of Mexico. The Lepid. Soc. Memoir No. 2. 280 pp.
PRESTON, W. B. & A. R. WESTWOOD. 1981. The European Skipper, Thymelicus lineola (Lepidoptera: Hesperiidae), in Manitoba and Northwestern Ontario. Can. Entomol. 113:1123-1124.
WALLIS, J. B. 1927. A Colour Key to the Manitoban Butterflies. Nat. Hist. Soc. Man. 31 pp.
Journal of the Lepidopterists’ Society 38(1), 1984, 40-46
THE LIFE HISTORY AND BEHAVIOR OF EPIMARTYRIA PARDELLA (MICROPTERIGIDAE)
PAUL M. TUSKES 1444 Henry St., Berkeley, California 94709!
AND
NORMAN J. SMITH 2192 Jenni Ave., Sanger, California 93657
ABSTRACT. Adults of Epimartyria pardella (Walsm.) are rather sessile and exhibit a clumped distributional pattern. Moths are active during the day and usually closely associated with liverworts. Larvae from eggs deposited in the lab feed on liverworts. There are three larval instars and in captivity 1.75 years were spent in the larval stage. Collection of wild larvae suggest that 2 years are also required to complete development under natural conditions.
The family Micropterigidae is recognized as the most primitive group of Lepidoptera known. The adult moths are the only Lepidoptera with functional mandibles which they use for feeding on pollen. Microp- terigidae are aglossate, jugate moths whose closest relatives are believed to be the Heterobathmiidae. Chapman (1917) and Hinton (1946) placed the Micropterigidae in their own order, the Zeugloptera, because of the primitive characters the larvae express, but Common (1970), Kris- tensen (1971) and Richards and Davies (1959) treated the Zeugloptera as a suborder of Lepidoptera. Fossil micropterigids in lower Cretaceous amber indicate that relatively little change has occurred in the group during the last 185 million years (Whalley, 1977, 1978).
In the United States this unique suborder is represented by the new world genus Epimartyria Walsm. (1898) that consists of two species. A great deal of work has been done on the systematics and evolutionary status of the micropterygids (Hinton, 1958; Common, 1975; Heath, 1976; Whalley, 1978; Kristensen and Nielsen, 1979), but observations dealing with their behavior and habitat are for the most part lacking. In this paper, information is presented on the biology and habitat of Epimartyria pardella (Walsm.).
The type series of E. pardella consists of five specimens which were collected near the coast in southern Oregon during early June 1872. The description that Walsingham (1880) published is brief and accom- panied by a color illustration. The moth (Fig. 1) has a wingspan of 10 to 11 mm. The forewing is metallic brown with three distinctive gold spots, while the hind wings are only metallic brown. The fringe of both
1 Present address: 7900 Cambridge 141G, Houston, Texas 77054.
VOLUME 38, NUMBER 1 Al
fore and hindwings is yellow and brown. The abdomen and thorax are gray-brown; the legs and a portion of the head are golden yellow. From the head to the posterior tip of the abdomen the moth measures just under 3.5 mm.
Last Instar Larva
Head. Length 0.5 mm, diameter 0.27 mm. Brown. Antennae prominent, trisegmented and situated on small tubercles located on dorsal lateral portion of head (Fig. 3). Stem- mata with 5 facets and located at the base of the antenna. Labrum simple with a pair of trisegmented palpi. Mandibles simple and dark brown.
Body. Length 4.3 to 4.6 mm; width 1.4 mm; height 1.2 mm. The body tapering at both ends with highest and widest point at abdominal segment 4. Dorsal and lateral surface brown to dark brown, ventral surface light brown. Prothoracic shield with 10 peg-like setae, 8 on the anterior and lateral border and 2 dorsally. Prothorax distinctly narrower than mesothorax. Mesothorax with 8 setae, 6 on dorsal and lateral anterior portion of gray brown pigmented area, and 2 just ventral to this pigmented area. Setae of metathoracic segment similar to those of mesothorax except subdorsal seta is greatly reduced in size. All thoracic segments have additional small micro-seta just dorsal to each true leg. True legs brown, with 3 segments and simple claw. Abdominal segments (A) Al to A8 (and T2 and T3) with sawtooth-shaped knobs which form a dorsal and lateral ridge, areas between ridges concave. The middorsal area concave and small dark depres- sion occuring on posterior of segments T2 to A8. Segments Al to A8 each with one dorsal seta (0.18 mm) atop dorsal ridge. Segments Al to A8 with reduced, almost microscopic subdorsal seta (0.04 mm) and prominent lateral seta (0.12 mm) on lateral ridge. Dorsal, subdorsal and lateral setae occur in brown pigmented area which has rough and wrinkled appearance. Dorsal and lateral intersegmental area constricted and may contain series of 8 to 20 microscopic dots. Ventral to lateral ridge, cuticle smooth and light brown. Series of brown dots form pattern around fixture that usually support a small seta. Conical ventral “prolegs” occur on segments Al to A8 and small sclerotized protuberance appears on ventral surface of each. Segments A9 and A10 fused and with enlarged simple sucker. Spiracles posterior and ventral to lateral setae. Head diameter of first and second instar larvae 0.11 and 0.22 mm, respectively.
Habitat
Observations were made in Prairie Creek State Redwood Park, Humboldt County, California. All locations where adults were ob- served or captured were within a few km of the ocean and at relatively low elevations. Although some moths were found along creeks and moist hillsides in the redwood-fir forest, the preferred habitat appears to be steep-walled, moist canyons near the coast which are dominated by ferns and bryophytes (Fig. 6). The prominent bryophytes that are associated with the adults and larvae are, Conecephalum conium, Pel- lia sp., Hookeria lucens, and Atrichum undulatum. Other plants in the immediate area include: bracken fern, Pteridium aquilinum; sword fern, Polystichum minitum; deer fern, Blechnum spicans; and five- finger fern, Adiantum pedatum.
Climatic conditions in this area are moderate and stable. Weather records from Prairie Creek Campground, located about 4 km east of the beach at an elevation of 160 m indicated the mean daily temper-
42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 1-5. 1, Adult male E. pardella (14x); 2, ova on underside of liverwort thallus (8x); 3, last instar larva (9x); 4, cocoon (5x); 5, pupa (9x).
ature for January (9.5°C) and July (15.5°C) of 1981 differed by only 6°C. Although the summer months (June to September) are relatively dry, approximately 140 cm of rain falls between October and May. During the 1980-81 rainy season (October to May) there were 12 days when the temperature dropped below O0°C (32°F); the lowest temper- ature recorded during that time was —1.5°C (28°F). Barbour et al. (1973) suggested that seasonal temperature fluctuation reaches a min-
VOLUME 38, NUMBER 1 43
Fic. 6. Habitat of E. pardella in Northern California.
imum in this area because of off shore upwelling. They indicated that the mean monthly air temperature normally changes only a few de- grees between the coldest and warmest months, and the ocean tem- perature changes very little.
Adult Behavior
The flight season begins in late May and continues to early or mid- July, with the peak adult density in June. The moths are active during the day, generally between 0900 and 1930 h, but this is influenced by temperature, humidity, and light intensity. When abundant, adults may be observed perched on vegetation; at low densities the best means of locating a colony is by sweeping suitable habitat with an aerial net. Behavioral notes were made on the activities (in situ) of individual moths that were observed from one to seven hours.
In areas protected from wind, adults frequently perched on the upper surface of fern fronds or other plants near patches of liverworts growing on canyon walls or beside creeks. Adults exhibit a clumped distributional pattern and, where common, densities reached 6 moths/ m?. When windy, or if the humidity is low, adults find shelter among the moist bryophytes with which they are always closely associated. The antennae are held at a 45 to 60 degree angle above the midline of the body while the moths are perched (Fig. 1). As they walk, the
44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
antennae wave up and down often touching the substrate. The hind legs of pardella are almost equal in length to that of the entire moth, and are occasionally used to jump or hop a few centimeters. Adults may remain motionless for hours and then walk or fly a few centi- meters and perch again. During a single two and one half hour obser- vation, a moth traveled 25 cm in a sporadic pattern and came to rest for the evening less than 5 cm from where it was first observed. Another moth less than 50 cm away walked less than 15 cm during this time. Most moths in 1981 were observed from one to three hours and trav- eled less than 30 cm. In 1982, five moths were carefully observed for a total of 29.2 hours. Again, the adults were extremely sessile, often remaining for hours in the same position. During the 29.2 hours, 16 flights were observed with an average distance of 21 cm per flight; they walked an average of 17 cm. Moths changed positions to perch in sunny locations, to avoid predators, and in the case of females, to oviposit. Adults are the prey of various small predators. One moth was captured in a spider web and another chased by a small hunting spider of the genus Theridion. A third moth was stalked but not captured by a small Olympic salamander (Rhyacotriton olympicus). Moths fly when disturbed but normally flight is infrequent and brief; the flight pattern is fluttery and weak but usually direct.
Adult micropterigids of other genera are reported to feed on pollen rather than nectar and have unique mouthparts. The mandibles are well developed, and the hypopharynx is concave on the upper surface. As pollen grains are ingested they are ground by the action of the mandibles against the hypopharyngeal spines and then digested (Till- yard, 1923; Hannermann, 1956). European species have been collected at the blooms of many plant species, including: Compositae, Acer, Carex, Scrophulariaceae, Quercus, and Ranunculus (Heath, 1960). Al- though various Ranunculus, Compositae, and Scrophulariaceae were near by and in bloom, no moths were observed at the flowers. Adults were frequently observed drinking water. Since they lack a proboscis they lower their head to the droplet of water by extending their meso- thoracic legs to the side of their body. This lowers the head and raises the abdomen, allowing the moth to drink. If deprived of moisture moths die in less than two days, but when provided with water, they survived in captivity from nine to 18 days, and females deposited ova.
The only mating pair of moths was found just prior to 1000 h. In captivity females laid an average of 8.2 eggs per day. Ova were de- posited on the underside of the liverwort thalli singly or in small clus- ters containing up to five ova (Fig. 2). The females generally remained on the upper surface and would simply swing their abdomen under the edge of the thallus to oviposit.
VOLUME 38, NUMBER 1 45
Immature Stages
The ova are flattened, circular and smooth when first deposited but become spherical in a short time and are covered with a series of small white projections (Fig. 2). The ova are white and measure 0.40 x 0.44 mm. At 22°C the eggs hatch in 21 days. The first instar larvae emerge from the side of the egg and are about 0.75 mm long. They vary from light brown to light gray and appear to have the same setal pattern and shape as mature larvae but have the ability to flatten themselves when at rest.
Larvae were reared in either a terrarium or petri dishes. Although both species of liverwort (Conocephalum and Pellia) were available, the larvae showed a marked preference for Pellia, the smaller of the two species. Mature larvae are active primarily at night but early instar larvae may be active at any time. While feeding, the margin of the liverwort is not damaged, rather the underside of the living thallus is eaten away but not through. Many micropterigid species feed on bryo- phytes, but the work of Luff (1964) and Lorenz (1961) indicates that some species do not.
In captivity the larvae are rather inactive, avoid intense light, and are usually found on the underside of the thalli during the day. In the field larvae were also found under living thalli during the day. Their coloration and size allowed them to blend well with the dead thalli which occur under the living growth. As the larva walks the true legs grasp the substrate; from above it appears to glide across the surface as the rhythmic undulations of the ventral surface are not apparent. When disturbed or inactive the head may be withdrawn so that only the prothoracic shield is visible; when extended the antennae which are located above the eyes are prominent (Fig. 3).
Unlike the European species which have a one year life cycle (Heath, 1976; Lorenz, 1961), pardella appears to have a two year cycle. In captivity eggs deposited in June 1981 became adults in June 1983. In the field, second instar larvae were commonly collected each year during the adult flight period. These larvae must represent the off- spring from ova deposited the previous year, as reared larvae one year old were also in the second instar. Davis (pers. comm.) observed that E. auricrinella (Walsm.) from the eastern United States also has a two year life cycle.
Pupation occurs close to the ground among vegetation. The brown cocoon, which measures 5.5 x 4.5 mm, is oval, thin walled and tightly woven (Fig. 4). Strands of coarse silk attach the cocoon to vegetation. The exarate pupa is white to light brown (Fig. 5).
Based on the illustration of Micropterix calthella (L.) larvae by Lo- renz (1961), the larvae of E. pardella exhibit a number of differences.
46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
The setae of calthella are club-shaped and apparently uniform in length. The larvae of pardella have peg-shaped setae which vary in length according to their location. In addition, the distribution of the larval setae and pupal setal patterns also differ. A preserved pupa, cocoon, and larvae were deposited in the collection of the California Academy of Sciences, San Francisco.
ACKNOWLEDGMENT
The research conducted at Prairie Creek Redwood State Park was done under a permit from the California Department of Parks and Recreation. We wish to thank the park service for their cooperation. We also wish to thank Ann McGowan-Tuskes for two years of field assistance and for reviewing the manuscript.
LITERATURE CITED
BaRrBour, M. G., R. B. Craic, F. R. DRUSDALE & M. T. GHISELIN. 1973. Coastal Ecology: Bodega Head. Univ. of Calif. Press, Berkeley.
CHAPMAN, T. A. 1917. Micropteryx entitled to ordinal rank; Order Zeugloptera. Trans. Entomol. Soc. London 1916:310-314.
Common, I. F. B. 1970. Lepidoptera. In insects of Australia. Melbourne. Melbourne Univ. Press. Pp. 765-866.
1975. Evolution and classification of the Lepidoptera. — Rev. Entomol. 20: 183-203. ‘
HANNERMANN, H. J. 1956. Die Kopfmuskulatur von Micropteryx calthella L. Mor- phologie und funktion. Zool. Jahrb. Anat. 75:177-206.
HEATH, J. 1960. The foodplants of adult micropterygids. Entomol. Mon. Mag. 95:188.
1962. The eggs of Micropteryx. Ibid. 97:179-180.
1976. The moths and butterflies of Great Britain and Ireland. Vol. 1. Pp. 151-
155.
HINTON, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of lepidoptera. Trans. Roy. Entomol. Soc. London 97:1-87.
1958. The phylogeny of the oaneneatd orders. Ann. Rev. Entomol. 3:181—206.
KRISTENSEN, N. P. 1971. The systematic position of the Zeugloptera in the light of recent anatomical investigations. Proc. XIII Int. Cong. Entomol. 1:261.
KRISTENSEN, N. P. & E. S. NIELSEN. 1979. A new subfamily of micropterigid moths from South America. A contribution to the morphology and phylogeny of the Mi- cropterigidae, with a generic catalogue of the family (Lepidoptera: Zeugloptera). Steenstrupia 5(7):69-147.
LORENZ, R. E. 1961. Biologie und morphologie von Micropterix calthella (L.). Dt. Ent. Z. (N.F.) 8:1-28.
Lurr, M. L. 1964. Larvae of Micropteryx [sic] (Lepidoptera; Micropterygidae). Proc. R. Entomol. Soc. Lond. (C) 29:6.
RICHARDS, O. W. & R. G. DaAvigs. 1957. In a general textbook of entomology. A. D. Imms. London, Methuen. 9th ed. 886 pp.
TILYARD, R. J. 1923. On the mouth parts of the Micropterygoidea (Lepidoptera). Trans. Roy. Entomol. Soc. London 181-206.
WHALLEY, P. E. S. 1977. Lower Cretaceous Lepidoptera. Nature 266:526.
1978. New taxa of fossil and recent Micropterygidae with a discussion of their evolution and a comment on the evolution of Lepidoptera. Ann. Transvaal Mus. 31: 71-86.
WALSINGHAM, T. 1880. On some new and little known species of Tineidae. Proc. Zool. Soc. London 83-84.
1898. Description of a new micropterygid genus and species and a new erio-
craniad species from N. America. Entomol. Rec. J. Var. 10:161-163.
Journal of the Lepidopterists’ Society 88(1), 1984, 47-50
A NEW ACANTHOPTEROCTETES FROM THE NORTHWESTERN UNITED STATES (ACANTHOPTEROCTETIDAE)
DONALD R. DAVIS
Department of Entomology, Smithsonian Institution, Washington, D.C. 20560
ABSTRACT. Acanthopteroctetes aurulenta Davis, new species, is described from Oregon and Utah. Both male and female are illustrated.
Recent collecting in central Utah by Ronald W. Hodges resulted in the discovery of the male of an undescribed species of Acanthopter- octetes previously mentioned in the literature (Davis, 1978:96, 129) but not named. The availability of both sexes of this species now en- ables me to name this insect, which constitutes only the fourth species described for the family.
Acanthopteroctetes aurulenta, new species
Length of forewings. 6, 7.4 mm; 8, 5.1 mm (Fig. 1). Head. Vestiture rough, pale yellowish brown to nearly white. Antennae with 43 seg- ments; vestiture of scape extremely rough with prominent pecten of more than dozen
Fic. 1. Acanthopteroctetes aurulenta, new species. Holotype 6, wing expanse 15 mm.
48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 2-5. Acanthopteroctetes aurulenta, new species, male genitalia: 2, lateral view,
(J = juxta; Tr = transtilla); 3, ventral view; 4, valva, mesal view; 5, aedoeagus. Scale = 0.5 mm.
long whitish hairs extending over eye; flagellum smooth, uniformly banded with white and pale brown scales. Haustellum naked except for scattered, fine setae. Maxillary palpi greatly lengthened, 5-segmented, geniculate; vestiture white. Labial palpi considerably shorter than maxillary palpi, covered with whitish scales.
Thorax. Pronotum covered with smooth, golden brown scales; central tuft of approx- imately one dozen elongate golden hairs present. Forewings uniformly pale golden brown,
VOLUME 388, NUMBER 1 49
Fics. 6-8. Acanthopteroctetes aurulenta, new species, female genitalia: 6, ventral view (CO = common oviduct, U = utriculus, V = vesicle); 7, dorsal view; 8, vestibulum and bursa copulatrix, dorsal view. Scale = 0.5 mm.
slightly lustrous; R, slightly variable, either connate with R,,;+M, or shortly stalked. Hindwings more thinly scaled, uniformly pale gray. Venter of thorax white. Legs mostly white; epiphysis absent.
Abdomen. Sparsely covered with pale golden brown scales above, more whitish be- neath. External glands absent. Caudal margin of eighth segment in female with encir- cling ring of elongate sensory setae; median setae longest with setae decreasing in length ventrally.
50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Male genitalia (Figs. 2-5). Uncus slender, minutely bifid, with 5 minute, subapical serrations along ventral margin. Ninth segment relatively long cylinder, about twice length of uncus, without lateral separation between tegumen and vinculum. Both anterior and posterior margins of vinculum deeply excavated. Median process of transtilla with 3 pairs of ventral serrations. Juxta elongate, length over 2.5 its width; basal half darkly sclerotized. Valvae slender, greatest width (at base) 0.25 its length; saccate membrane arising from elongate pouch along distal half of cucullus. Aedoeagus elongate, exceeding genital capsule in length; prominent cluster of approximately 6 elongate cornuti present.
Female genitalia (Figs. 6-8). Apex of ovipositor broad, depressed, triangular in outline, with approximately 15-17 serrations bordering lateral margins. Posterior apophyses stou- ter than anterior pair. Vestibulum enlarged, extremely irregular in outline, and with highly folded, thickened walls. Spermatheca with minute spherical vesicle at posterior end of elongate, slightly inflated utriculus; spermathecal papillae not sclerotized. Corpus - bursae reduced in size, membranous.
Types. Holotype 6: Head Ephraim Canyon, 10,000-10,300 ft [3049-3140 m], Senpete Co., UTAH, 1 Aug 1981, R. W. Hodges, blacklight, USNM 100671. Paratype: Baker, Oregon, Spring Creek, 12, 8 Jul 1966. J. H. Baker (USNM).
Distribution. Northwestern Oregon and central Utah.
Remarks
The uniformly light golden brown forewings of A. aurulenta easily distinguishes it from the other darker, banded-wing species in the fam- ily. This characteristic color pattern has suggested the specific name, derived from the Latin aurulentus (golden, ornamented with gold). The valvae of A. aurulenta are also unusual in possessing a very dis- tinct, thinly sclerotized pocket from which arises the peculiar saccate membrane found in all members of the genus.
The Spring Creek, Oregon habitat can be characterized as a pine- sagebrush association with Ceanothus (the host of A. unifascia Davis (Davis and Frack, in press)) occurring nearby. The type locality in the Wasatch Mountains of central Utah, which has been heavily grazed in recent times (D. C. Ferguson, pers. comm.), is an open, subalpine plateau.
ACKNOWLEDGMENTS
I wish to thank my assistant, Ms. Biruta Akerbergs Hansen, for preparing the illustra- tions for this paper, and Dr. Ronald Hodges of the Systematic Entomology Laboratory, USDA, for his efforts in collecting this species.
LITERATURE CITED
Davis, D. R. 1978. A revision of the North American Moths of the superfamily Erio- cranioidea with the proposal of a new family, Acanthopteroctetidae (Lepidoptera). Smithsonian Contr. Zool., No. 251, 131 pages, 344 figs.
Journal of the Lepidopterists’ Society 38(1), 1984, 51-56
TWO INTERESTING ARTIFICIAL HYBRID CROSSES IN THE GENERA HEMILEUCA AND ANISOTA (SATURNIIDAE)
RICHARD STEVEN PEIGLER! 303 Shannon Drive, Greenville, South Carolina 29615
AND
BENJAMIN D. WILLIAMS The Lawrence Academy, Groton, Massachusetts 01450
ABSTRACT. Two crosses were reared to the adult stage with saturniid moths from different areas of the United States. These were Hemileuca lucina 6 x H. nevadensis 2 reared in Massachusetts and Texas on Salix, and Anisota senatoria 6 x A. oslari 2 reared in Connecticut on Quercus coccinea. Larvae and adults of both crosses were interme- diate. Descriptions and figures of the hybrids are given. Several isolating mechanisms between the parent species were tested and are discussed.
Dozens of artificial crosses in the Saturniidae have been successfully reared since the previous century, but virtually all of these have in- volved species of the subfamily Saturniinae. This paper deals with two remarkable crosses obtained by the junior author utilizing small satur- niid moths belonging to the subfamilies Hemileucinae and Ceratocam- pinae.? In both crosses, species native to the Southwest were reared in the Northeast and females from those rearings attracted congeneric diurnal males native to the Northeast. The species involved were Hemileuca lucina Henry Edwards, H. nevadensis Stretch, Anisota senatoria (J. E. Smith) and A. oslari W. Rothschild. For information on the adult morphology, wing pattern, immature stages, hostplants, reproductive behavior, and geographical distributions of these four parent species, the reader is referred to works by Ferguson (1971) and Riotte and Peigler (1981).
Hemileuca lucina 6 x H. nevadensis 2
In mid-September 1977 two virgin females of H. nevadensis (stock from Escondido, San Diego Co., California) were placed on twigs of Salix gracilis Anderess at the edge of a wet meadow in Groton, Mid- dlesex Co., Massachusetts, which supports a sizable population of H. lucina. The emergence time of the reared H. nevadensis in Groton coincides with the flight time of H. lucina, i.e., mid-September through early October. The females emitted pheromone, and males of H. lu- cina were attracted. We assumed that pheromone from wild females
‘Museum Associate in Entomology, Los Angeles County Museum of Natural History.
52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
of H. lucina was also present in the air, and therefore we concluded that males of H. lucina may not discriminate between pheromones of the two species. No behavioral isolation was observed as is often the case in achieving cross-matings of Saturniidae; there was no hesitation by the males nor resistance to them by the females.
Each female produced an egg ring around a twig, each ring con- sisting of ca. 100 eggs. The egg is the overwintering stage in most species of this genus. One egg ring was sent to the senior author. The following spring eclosion of both egg rings was near 100 percent, and both authors reared broods successfully to the adult stage. In Groton the larvae were reared on Salix gracilis under a cloth bag, surviving an unseasonal 25 cm snowfall on 10 May while in the second instar. In Brazos County, Texas, the senior author reared his brood on Salix sp. (probably nigra Marshall) under cloth bags. Adult emergence in 1978 in Texas and Massachusetts differed, probably as a result of dif- ferences in photoperiod between the two regions where the pupae were kept. In Texas males emerged 21 July through 24 August peaking in the middle of August; females appeared during the second half of August and early September. The hybrid brood in Massachusetts yield- ed males from 6 September to 8 October and females mostly during the second week of October. A brood of pure H. nevadensis (Escon- dido, California) reared alongside the hybrids in Texas produced adults of both sexes in September, too late to permit attempts to backcross the hybrids, but coinciding with the emergence pattern of pure H. nevadensis in Massachusetts as mentioned above.
Hybrid females of both broods were sterile, based on the fact that their abdomens appeared to contain few or no ova. In both broods most adults expanded their wings normally after emerging, but some specimens, especially among females, failed to spread their wings par- tially or totally. This problem is encountered in several species of the genus with reared material and is not considered to indicate reduced viability resulting from hybridization.
Only the final instar larva is described, this one showing greatest divergence from parent species, but H. nevadensis and H. lucina hard- ly differ structurally. Pupal differences were difficult to find between the parent species also, so that those given below may not be reliable. Adults of this group do not exhibit sexual dimorphism; thus, the sexes are not described separately.
Description Mature larva. Integument intermediate: black with numerous dull white oval flecks,*
many converging but all individually distinct except around spiracles; in H. lucina flecks smaller and more widely separated; in H. nevadensis flecks larger and converging to
VOLUME 38, NUMBER 1 53
Fics. 1-8. 1 & 2, hybrid pair of Hemileuca lucina 6 x H. nevadensis 2; 3 & 4, pair of Anisota senatoria from Pomfret, Connecticut; 5 & 6, hybrid pair of Anisota senatoria 6x A. oslari 2; 7 & 8, pair of Anisota oslari from Santa Cruz Co., Arizona.
o4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
form yellowish white areas on integument, especially dorsally. Two dorsal rows of tufted- spine scoli stramineous with black tips as in both parents. Subdorsal and subspiracular branched scoli black with whitish tips in hybrid and both species.
Pupa. Anterior rim of each abdominal segment wider as apparently for H. nevadensis. Cremaster with stouter curved spines as in H. lucina; H. nevadensis apparently with thinner, straighter spines, and possibly fewer than in H. lucina. Head and thoracic characters of both parent species and hybrid indistinguishable.
Adult (Figs. 1 & 2). Elongated white scales on meso- and metathorax sparsely distrib- uted among black scales, these white scales more numerous than in H. lucina but much less numerous than in H. nevadensis. Whitish band of forewing agreeing with that of H. lucina by being wider on ventral side than on dorsal side, but more like H. nevadensis by having crenulate outer margin curving parallel with outer wing margin. Discal mark in hindwing containing white slit as in father species, this mark often solid black in H. nevadensis. Black portions of wings more opaque and coal-black than either parent species.
Anisota senatoria 8 x A. oslari 2
A freshly emerged female of A. oslari was permitted to emit pher- omone in an exposed location at Pomfret, Windham County, Con- necticut, during a clear, cool, and windy day in mid-July 1978. The undersized moth had been reared on scarlet oak (Quercus coccinea Muenchh.) from eggs received from Madera Canyon, Arizona the pre- vious year. Anisota senatoria flies in southern New England from mid- June through mid-July, whereas reared specimens of A. oslari have emerged from mid-July through mid-August. Males of A. senatoria seek females from ca. 1130 to 1530 h EST, and the circadian flight time of A. oslari is also known to be during midday hours. A male of A. senatoria arrived but had considerable difficulty locating the female due to gusty wind. He persisted for ca. 1 h before making physical contact, at which time copulation readily occurred. Attempts to obtain this cross the previous year had apparently failed due to the normally larger size of the females of A. oslari, which prevented the males of A. senatoria from achieving copulation.
After mating, the female of A. oslari oviposited freely. Eclosion of the eggs was virtually 100 percent. The hybrid larvae were vigorous, and several were reared to maturity under cloth bags on scarlet oak. The following year the female hybrids emerged during the last few days of May, whereas their male siblings appeared from 9 June through 19 July. Females were apparently sterile, having shrunken abdomens as mentioned under the previous cross.
Description
Mature larva. Head brown with bold black markings on each side (head of A. senatoria solid black; head of A. oslari solid brown; the two-colored head of hybrids remarkable because all known species of Anisota have solid colored heads in all instars). Prothoracic tergite black as in A. senatoria. Body color black with bold orange stripes on sides and two broken orange stripes on top. Anal plate and anal prolegs orange with black markings (solid black in A. senatoria, solid brown in A. oslari). Pattern of spines on body more as
VOLUME 38, NUMBER Il 55
in A. senatoria but size and arrangement of spines on anal plate intermediate between parent species. Median caudal spine long as in A. oslari.
Male (Fig. 5). Overall appearance strikingly intermediate. Wingshape as in A. sena- toria but large size as in A. oslari. Ground color dark purplish brown, forewings having brownish orange overtones. Postmedian line weak; barely discernable transparent patch in forewing (absent in A. oslari, well-developed in A. senatoria). White discal mark large. Forewing with sparse sprinkling of dark spots. Outer margins of hindwings straight.
Female (Fig. 6). Intermediate in most characters. Wingshape closer to A. oslari. Ground color light brownish orange with pinkish suffusion in postmedian area as in father species and on hindwing as in mother species. Postmedian line weak in forewing, very faint in hindwing. White discal mark surrounded by purple as in A. senatoria. Forewing with a few dark spots.
DISCUSSION
Hybridization experiments such as these provide data on isolating mechanisms and degree of phylogenetic divergence. Aside from the obvious one of allopatry, other isolating mechanisms tested by these crosses include mechanical, behavioral, viability of immature stages, and fertility of adult hybrids. Remarks on each of these were given above for both crosses. The differing emergence times between the sexes of an individual hybrid brood were proposed by Peigler (1981) as an isolating mechanism, because this reduces frequency of F, or backcross matings when hybrid broods are produced in nature (when primary isolating mechanisms fail). This phenomenon, now widely rec- ognized in hybrid Lepidoptera, is well illustrated in the two present crosses. We use the term “isolating mechanism”’ in the traditional sense as did Solignac (1981), notwithstanding the valid arguments put forth by Key (1981) that several independent principles are encompassed by the term.
Genetic compatibility between two taxa, which is to some extent correlated with phylogenetic divergence, falls along a continuum. The pairs of species in the present study are demonstrated to have an in- termediate affinity when compared to the following two extremes. Minimal compatibility of parent species would be seen if eggs fail to eclose or larvae die in the first instar. This was demonstrated by the cross Hemileuca nuttalli (Strecker) ¢ x H. eglanterina (Boisduval) in the studies of Collins and Tuskes (1979), which might be expected because the parent species are sympatric. On the other hand, what appears to be total genetic compatibility in Saturniidae is illustrated by crosses (both reciprocals) between the Indian Antheraea roylei Moore and the Chinese A. pernyi (Guérin-Méneville). The parent species have chromosome numbers of n = 30 and n = 49 respectively, and the hy- brid (n = 30) has been reared through more than 20 generations, main- taining its increased vigor over the parent species (Jolly, 1974, 1981). Most known crosses of Lepidoptera result in more or less vigorous F, hybrid adults with reduced fertility, especially in females.
56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
It is our hope that this paper will encourage lepidopterists to exploit every opportunity to achieve interspecific matings of species that they rear. When fertile eggs and viable larvae result, records and descrip- tions should be kept, results published, and material deposited in mu- seums.
ACKNOWLEDGMENTS
We are grateful to J. Steve McElfresh of San Diego, California, for supplying eggs of H. nevadensis and A. oslari. Our figures were made by Thomas Marion Hill of Green- ville, South Carolina. Drs. W. D. Winter, Jr. and Joseph E. Eger, Jr. made color photo- graphs of living larvae of the hybrids and/or pure species which aided formulation of the larval descriptions. Material supplied by Earll M. Brown of San Diego also was useful in this study. Specimens of both crosses, including the four hybrids figured, have been deposited in the Los Angeles County Museum of Natural History, and a pair of the Hemileuca cross is in the American Museum of Natural History.
LITERATURE CITED
CoLLins, M. M. & P. M. TuskEs. 1979. Reproductive isolation in sympatric species of dayflying moths (Hemileuca: Saturniidae). Evolution 33:728-7383.
FERGUSON, D. C. 1971. Bombycoidea, Saturniidae (in part), in R. B. Dominick et al., The moths of America north of Mexico, fasc. 20:2A:153 pp., 11 col. pls., E. W. Classey, London.
Jotty, M. S. 1974. Discovery of new field of tasar on oak and its impact on national economy. Central Tasar Res. Sta., Ranchi, Bihar, India. 4 pp.
1981. Distribution and differentiation in Antheraea species (Saturniidae: Lep- idoptera), pp. 1-14 in S. Sakate & H. Yamada eds., Study and utilization of non- mulberry silkworms. Symposium in 16th Internat. Congr. Entomol., August 1980, Kyoto, Japan. (12) + 78 pp.
Key, K. H. L. 1981. Species, parapatry, and the morabine grasshoppers. Syst. Zool. 30: 425-458.
LEMAIRE, C. 1978. Les Attacidae americains ... The Attacidae of America (=Satur- niidae), Attacinae. C. Lemaire, Neuilly. 238 pp., 49 pls.
PEIGLER, R. S. 1981. Demonstration of reproductive isolating mechanisms in Callosa- mia (Saturniidae) by artificial hybridization. J. Res. Lepid. 19:72-81.
RIOTTE, J. C. E. & R. S. PEIGLER. 1981. A revision of the American genus Anisota (Saturniidae). J. Res. Lepid. 19:101-180.
SOLIGNAC, M. 1981. Isolating mechanisms and modalities of speciation in the Jaera albifrons species complex (Crustacea, Isopoda). Syst. Zool. 30:387—405.
TUSKES, P. M. 1976. A key to the last instar larvae of West Coast Saturniidae. J. Lepid. Soc. 30:272-276.
* Lemaire (1978:23) explained in detail why the name Ceratocampinae is to be used instead of Citheroniinae. * Tuskes (1976) stated that these flecks are circular, but in all material we have seen, consisting of several species of the genus, these are distinctly oval.
Journal of the Lepidopterists’ Society 38(1), 1984, 57-59
SPERMATOPHORE PERSISTENCE AND MATING DETERMINATION IN THE GYPSY MOTH (LYMANTRIIDAE)!
CYNTHIA R. LOERCH AND E. ALAN CAMERON
Department of Entomology, The Pennsylvania State University, University Park, Pennsylvania 16802
ABSTRACT. Spermatophores were detectable in all female gypsy moths dissected within 1.5 h following inception of copulation. After 1.5 h, the percentage of detectable spermatophores decreased with time; by 4.5 h, no spermatophore could be detected in any mated female moth. The percentages of detectable spermatophores did not differ significantly among three gypsy moth populations (laboratory-reared, high and moderate density natural populations) for intervals timed from inception of copulation. Examina- tion of the bursa copulatrix for the presence of a spermatophore can be useful for rapid determination of female gypsy moth mating success.
The spermatophore of the gypsy moth, Lymantria dispar (L.), is formed within the female bursa copulatrix during the first 10 min of copulation (Klatt, 1920; Leonard, 1981). It consists of an oval sperm sac with a tapered neck that extends into the ductus bursae and a proteinaceous mass secreted by the male accessory glands. Proteolytic enzymes produced by the female begin to dissolve the spermatophore shortly after its formation (Chapman, 1971; Engelmann, 1970).
However, little is known of the fate of the gypsy moth spermato- phore between formation and disintegration. Taylor (1967) reported that the spermatophore disintegrates within one or two hours of cop- ulation but did not state whether this is time accrued from inception or termination of copulation. The distinction is essential since copula- tion averages 60-73 min (range = 20-198 min) (Forbush and Fernald, 1896; Doane, 1968; Waldvogel et al., 1981). Because the gypsy moth spermatophore is not persistent, determination of female mating suc- cess relies on examining eggs for embryonation several weeks after deposition or examining the female reproductive system for the pres- ence of sperm (Stark et al., 1974). This paper presents, for the first time, data on the persistence of the gypsy moth spermatophore, with implications for rapid determination of female mating success.
MATERIALS AND METHODS
Laboratory-reared virgin gypsy moths were mated, uninterrupted, in arenas described by Waldvogel et al. (1981). The time in copula was recorded for each pair. To obtain data on the persistence of the
‘ Authorized for publication as Paper Number 6368 in The Journal Series of The Pennsylvania Agricultural Experi- ment Station. This work was conducted under Experiment Station Project No. 2044, and supported in part by Regional Research Project NE-84 (Revised).
58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Percentages of spermatophores detectable at intervals timed from inception of copulation for three gypsy moth populations: laboratory-reared, and high and mod- erate density natural populations.
Hours
following % spermatophores detectable
inception of
copulation Laboratory-reared High density Moderate density Total Le 100.0 (11) 100.0 (9) 100.0 (10) 100.0 (30) 2.0 81.8 (11) 66.6 (9) 70.0 (10) 73.8 (80) ao 70.8 (24) 50.0 (10) 40.0 (10) 59.1 (44) 3.0 16.7 (24) 44.4 (9) 30.0 (10) 25.6 (48) 3.5 0.0 (19) - 1 (9) 10.0 (10) 5.3 (38) 4.0 =) 0 (9) 10.0 (10) 5.3 (19) 4.5 — (0) — (0) 0.0 (9) 0.0 (9)
* Values in parentheses are numbers of mated female moths dissected. Percentages did not differ significantly (Chi- square test, Fisher’s exact test; P > 0.05) among populations at each time interval.
spermatophore, females were dissected under a microscope at 30x magnification, at intervals timed from inception of copulation. A me- dial incision through the abdominal terga provided access to the bursa copulatrix. The bursa copulatrix was then dissected in situ and its contents compared with those of an unmated female. All matings and dissections were performed at room temperature.. These procedures were repeated with virgin moths that emerged from pupae collected from moderate density (ca. 83000 egg masses/ha) and high density (ca. 70,000 egg masses/ha) natural populations in Clearfield County, Penn- sylvania. Egg mass densities were estimated by the method of Wilson and Fontaine (1978).
RESULTS AND DISCUSSION
Duration of copulation averaged 87 + 2.3 min for all mated pairs (n = 213, range = 22-218 min). The percentages of spermatophores that remained detectable at intervals timed from inception of copu- lation are presented in Table 1. For each time interval, the percentages of detectable spermatophores did not differ significantly among pop- ulations (Chi-square test, Fisher’s exact test; P > 0.05). Within 1.5 h following inception of copulation, 100% of the spermatophores in all populations could be detected. During this period, the shiny white spermatophore was visible through the wall of the bursa copulatrix. After 1.5 h, the percentage of detectable spermatophores decreased with time; the spermatophore was rarely visible through the bursa copulatrix wall, and dissection was necessary to determine its presence. At 3.5 h following inception of copulation, the spermatophore was detectable in less than 12% of the moths examined from any popula- tion. By 4.5 h, the contents of the bursa copulatrix of all mated females were indistinguishable from those of an unmated moth.
VOLUME 38, NUMBER 1 59
These data eliminate the ambiguity arising from Taylor’s (1967) report. His observations, if timed from termination of copulation, roughly agree with our findings. In other species of Lepidoptera, where the spermatophore may persist for several days or more, the bursa copulatrix can be examined for the presence of a spermatophore to determine whether a female has mated (Burns, 1968; Snow and Car- lysle, 1967; Taylor, 1967). Although the gypsy moth spermatophore is not persistent, it can be useful for rapid determination of female mat- ing success, which may be required in some precopulatory behavioral studies. Examination of the bursa copulatrix for a spermatophore is highly reliable within 1.5 h following inception of copulation. The presence of a spermatophore indicates female mating success and es- tablishes that mating occurred less than 4.5 h prior to examination. Unfortunately, the absence of a spermatophore does not establish that the female gypsy moth is unmated. When no spermatophore is de- tectable, the most immediate recourse is examination of the sperma- theca for the presence of sperm (Stark et al., 1974).
ACKNOWLEDGMENTS
We thank W. Metterhouse and R. Chianese of the New Jersey Department of Agri- culture, Division of Plant Industry, for providing laboratory-reared pupae, and S. J. Brumbaugh for assisting with mating observation. We also wish to thank P. H. Adler and R. O. Mumma, Department of Entomology, The Pennsylvania State University, for their helpful criticisms of the manuscript.
LITERATURE CITED
BuRNS, J. M. 1968. Mating frequency in natural populations of skippers and butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci. U.S.A. 61:852-859.
CHAPMAN, R. F. 1982. The insects: structure and function. 3rd Ed. American Elsevier Publishing Co., Inc., New York. 992 pp.
DOANE, C. C. 1968. Aspects of mating behavior of the gypsy moth. Ann. Entomol. Soc. Am. 61:768-773.
ENGELMANN, F. 1970. The physiology of insect reproduction. Pergamon Press Inc., New York. 807 pp.
ForsusH, E. H. & C. H. FERNALD. 1896. The gypsy moth, Porthetria dispar (Linn.). Wright and Potter Printing Co., Boston. 495 pp.
KuLaTT, B. 1920. Beitrage zur Sexualphysiologie des Schwammspinners. Biol. Zentralbl. 40:539-558.
LEONARD, D. E. 1981. Bioecology of the gypsy moth, in The gypsy moth: research toward integrated pest management, Doane, C. C. & M. L. McManus, eds., U.S. Dep. Agric., Tech. Bull. 1584. pp. 9-29.
SNow, J. W. & T. C. CARLYSLE. 1967. A characteristic indicating the mating status of male fall armyworm moths. Ann. Entomol. Soc. Am. 60:1071-1074.
STARK, R. S., E. A. CAMERON & J. V. RICHERSON. 1974. Determination of mating and fertility of female gypsy moths. J. Econ. Entomol. 67:296-297.
TAYLOR, O. R., JR. 1967. Relationship of multiple mating to fertility in Atteva punc- tella (Lepidoptera: Yponomeutidae). Ann. Entomol. Soc. Am. 60:583-590.
WALDVOGEL, M. G., C. H. COLLISON & E. A. CAMERON. 1981. Durations of pre- copulatory periods of laboratory-reared irradiated and non-irradiated male gypsy moths. Environ. Entomol. 10:388-389.
WILSON, R. W., JR. & G. A. FONTAINE. 1978. Gypsy moth egg-mass sampling with fixed- and variable-radius plots. U.S. Dep. Agric., Agric. Handbk. 523.
Journal of the Lepidopterists’ Society 38(1), 1984, 60-61
GENERAL NOTES
INSECT PARASITES AND PREDATORS OF HACKBERRY BUTTERFLIES (NYMPHALIDAE: ASTEROCAMPA)
During the course of collecting and rearing immature stages of hackberry butterflies (Nymphalidae: Asterocampa) over the past five years, a number of arthropod parasites and predators were encountered. These arthropods have been preserved or their behay- iors recorded in hopes of understanding some of the selective pressures which might affect the courses of evolution for Asterocampa species. This note is a report of insect species which have a greater or lesser effect on survival of the various stages of the butterflies.
Identifications were made by the author with the aid of the cited references and the reference collection at Texas A&M University. Help in the collection or identification of specimens, or review of the manuscript was provided by L. G. Friedlander, P. Davis, D. and D. Paschley, and Drs. H. R. Burke, J. C. Schaffner, and R. Wharton.
The most frequently encountered parasites of hackberry butterflies are the scelionid egg parasites, which occur in all Asterocampa observed. Stink bugs, such as the one figured by Langlois and Langlois (1964, Ohio J. Sci. 64:1-11, fig. 11), are the most common predators. Only one other insect (at the generic level) has been positively re- ported to attack Asterocampa, the larval parasite, Hyposoter fugitivus (Say) (Hym.: Ichneumonidae) (Townes, 1945, Mem. Amer. Entomol. Soc. No. 11, Pt. I, pp. 479-925).
Parasites Reared from Eggs
1. Hym.: Eulophidae: Tetrastichus spp. (Boucek, 1977, Bull. Entomol. Res. 67:17-80): A. clyton (Boisduval & Leconte) egg masses (TEXAS: Brazos Co., 14-VII-79; Menard Co., 20-VI-79).
2. Hym.: Scelionidae: Telenomus spp. (Masner, 1976, Mem. Entomol. Soc. Canada No. 97, 87 pp.): A. argus (Bates) egg mass (MEXICO: Oaxaca, 11-VII-81); A. celtis (Boisduval & Leconte) eggs (TEXAS: Hidalgo Co., 4-VI-81); A. clyton egg masses (AR- IZONA: Pima Co., 23-VIII-80; TEXAS: Brazos Co., 14-VII-79; Menard Co., 20-VI-79; San Patricio Co., 3-VI-81; Travis Co., 14-X-77; Waller Co., 8-VII-79; VIRGINIA: West- moreland Co., 22-VI-80); A. leilia (Edwards) eggs (TEXAS: Starr Co., 6-VI-81).
Parasites Reared from Larvae
1. Dip.: Tachinidae: Euphorocera prob. floridensis Townsend (Aldrich and Webber, 1924, Proc. U.S. Natl. Mus. 63:1-90; Cole, 1969, The flies of western North America, Univ. Calif. Press, Berkeley and Los Angeles, 693 pp.): A. celtis last stage larva (TEXAS: Austin Co., 6-VIII-79).
2. Dip.: Tachinidae: Lespesia prob. aletiae (Riley) (Beneway, 1963, Univ. Kansas Sci. Bull. 44:627-686; Cole, 1969, loc. cit.): A. clyton late stage larvae (TEXAS: Gonzales Co., 30-IX-79).
3. Hym.: Braconidae: Cotesia spp. (Mason, 1981, Mem. Entomol. Soc. Canada No. 115, 147 pp.): A. clyton third stage larvae (TEXAS: Gonzales Co., 21-IX-79; Hidalgo Co., 138-XI-77; Jeff Davis Co., 15-VIII-81; Uvalde Co., 23-IX-79).
4. Hym.: Braconidae: Meteorus spp. (Tobias, 1966, Entomol. Rev. 45:348-358): A. clyton larvae! (TEXAS: Goliad Co., 6-VI-81; Travis Co., 29-V-78, 20-VII-79).
5. Hym.: Eulophidae: Elachertus sp. (Peck et al., 1964, Mem. Entomol. Soc. Canada No. 34, 120 pp.): A. celtis last stage larva (TEXAS: Travis Co., 21-VI-78); A. clyton middle stage larvae (TEXAS: Brazos Co., 14-VII-79; Travis Co., 28-X-77).
6. Hym.: Ichneumonidae: Microcharops tibialis (Cresson) (Townes, 1969, Mem. Amer. Entomol. Inst. No. 18, 307 pp.; Townes and Townes, 1966, Mem. Amer. Entomol. Inst. No. 8, 367 pp.): A. clyton third stage larva (LOUISIANA: St. Tammany Parish, 30-III- 82).
VOLUME 38, NUMBER | 61
Parasites Reared from Pupae
1. Hym.; Chalcidiae: Brachymeria sp. (Howard, 1885, U.S. Dept. Agric., Bur. Ento- mol., Bull. No. 5, 47 pp.): A. clyton pupa (TEXAS: Gonzales Co., 15-X-77).
2. Hym.: Ichneumonidae: Itoplectis conquisitor (Say): A. clyton pupa (TEXAS: Dim- mit Co., 21-IV-79).
Predators
1. Hem.: Pentatomidae: Apateticus cynicus Say (Slater and Baranowski, 1978, How to know the true bugs (Hemiptera-Héteroptera), Wm. C. Brown Co. Publ., Dubuque, Iowa, 256 pp.): A. clyton early stage larvae (TEXAS: Travis Co., 26-III, 24-V, 18-X, 31- X-77).
2. Hem.: Pentatomidae: Apateticus lineolatus (Herrick-Schaeffer) (det. J. Eger): A. clyton larvae (TEXAS: Cameron Co., 13-III-79).
3. Hem.: Pentatomidae: Podisus maculiventris (Say) (Slater and Baranowski, 1978, loc. cit.): A. clyton early stage larvae (TEXAS: Travis Co., 24-V, 1-VI, 23-28-X-77, 23- V-78).
4. Hem.: Reduviidae: Sinea prob. sanguisuga Stal: A. clyton second stage larva (TEX- AS: Travis Co., 29-V-78).
5. Hem.: Reduviidae: Sinea spinipes (Herrick-Schaeffer) (Slater and Baranowski, 1978, loc. cit.): A. clyton early stage larvae (TEXAS: Travis Co., 28-X-77).
6. Hym.: Vespidae: Polistes exclamans Viereck: A. celtis fifth stage larva (TEXAS: Travis Co., 24-IV-78); A. clyton third stage larvae (TEXAS: Travis Co., 25-X-77).
7. Hym.: Vespidae: Vespula sp.: A. clyton third stage larvae (TEXAS: Travis Co., 31- X-77).
TIMOTHY P. FRIEDLANDER, Department of Entomology, Texas A&M University, College Station, Texas 77843.
‘None were reared from larvae. One female Meteorus was observed to oviposit in A. clyton larvae. One female hyperparasite of Meteorus was observed to oviposit in larvae of the same species. One of these hyperparasites was reared from Meteorus cocoons taken in close association with A. clyton larvae.
Journal of the Lepidopterists’ Society 38(1), 1984, 61-63
ITHOMIINE BUTTERFLIES ASSOCIATED WITH NON-ANTBIRD DROPPINGS IN COSTA RICAN TROPICAL RAIN FOREST
Adult females of Mechanitis and the allied genus Melinaea (Brown, 1977, Syst. Ento- mol. 2:161-197) feed on the fresh droppings of birds (primarily antbirds) that follow swarms of army ants through tropical rain forest in Costa Rica (e.g., Ray and Andrews, 1980, Science 210:1147-1148). These authors conclude that bird droppings resulting from birds following army ant swarms provide a predictable nutrient resource for these female butterflies, and that the exploitation of this resource may be related, in some yet to be studied way, to egg production. In this note I extend the findings of Ray and Andrews (op. cit.) to the association of female ithomiines of various genera to fresh droppings of bird species not associated with army ant swarms in Costa Rican tropical rain forest. I conclude that fresh bird droppings of any kind in such a habitat provide a resource exploited by ithomiines on an opportunistic basis.
Between 1972 and 1980, I conducted several studies of various butterfly species in a small parcel of relatively undisturbed mixed primary and secondary-growth tropical rain forest (premontane tropical wet forest) at “Finca La Tigra’, near La Virgen (220 m elev.), Heredia Province, Costa Rica. The site is about 20 km from the “Finca La Selva”
62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fic. 1. Female Godyris zavelata caesiopicta feeding at fresh bird dropping at a light gap in the forest habitat at Finca La Tigra in northeastern Costa Rica.
study site of Ray and Andrews (op. cit.). During this lengthy period, I observed ithomiine butterflies feeding on fresh bird droppings splashed on leaves of understory plants, par- ticularly along foot paths and light gaps in the forest. This was not a deliberate search for butterflies, but rather accidental encounters within an approximately 500-square meter area usually visited three or four months each year. Mechanitis spp. and Hypothy- ris euclea leucania (Bates) were the most frequently observed ithomiines exhibiting this behavior. These ithomiines are very abundant, relative to others, at this locality (Young, 1976, Pan Pacific Entomol. 53:104-113; Young, 1979, J. Lepid. Soc. 33:68-69; Young and Moffett, 1979a, Amer. Midl. Natur. 101:309-319; 1979b, Deutschen Ent. Zeitschr. 26:21-38). A less numerous ithomiine, Godyris zavelata caesiopicta (Niepett) was also observed feeding on fresh bird droppings at various times in the same period (Fig. 1). In my experience, encounters of such behavior consisted of usually one or two butterflies, either both on the same dropping or on separate droppings in the case of two or more. Large aggregates of ithomiines on bird droppings were not encountered. At the same times, however, I did not notice any swarms of army ants in the same areas, or in adjacent open areas such as a cacao plantation forming the border to the forest study site. In one instance with Godyris (10 July 1982 at 1600 h) I noticed a single butterfly feeding at a dropping for close to forty minutes but with frequent interruptions by several flies (Dip- tera) that chased it away temporarily. Godyris zavelata females are easily distinguished from males by wing colors (Young, 1974, Entomol. News 85:227-238). It is by no means as abundant locally (in this area) as Mechanitis and Hypothyris. Several other bluish clear-wing ithomiines (undetermined) also visited fresh bird droppings in the same forest patch.
Based upon these preliminary observations made at irregular intervals over several years at the same forest patch in northeastern Costa Rica, I suggest that the females of several genera of ithomiine butterflies routinely exploit, on an opportunistic basis, fresh
VOLUME 38, NUMBER 1 63
bird droppings splashed on understory vegetation. Areas of tropical rain forest with disruptions in the canopy, such as light gaps and foot paths, are particularly attractive gathering places for various species of birds, perhaps because many insects, potential prey, and other arthropods are also found in these microhabitats. In turn, bird droppings occur there frequently, although perhaps in an unpredictable fashion, selecting for op- portunistic foraging by female ithomiines. When large concentrations of bird droppings become available, such ithomiines, at least Mechanitis and Melinaea, may exhibit delib- erate orientation to such food resources and become abundant there, as reported else- where (Ray and Andrews, op. cit.).
I thank Luis Poveda for identification of the Godyris larval food plant, and Dr. J. Robert Hunter for allowing access to Finca La Tigra.
ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- waukee, Wisconsin 53233.
Journal of the Lepidopterists’ Society 38(1), 1984, 63-64
SATYRIUM KINGI (LYCAENIDAE) TAKEN IN MARYLAND
At 1600 h on 22 July 1982, after spending a discouraging time collecting in three areas in Wicomico and Worcester Counties in Maryland, I caught a Satyrium kingi (Klots and Clench) near Millville, Worcester County. This capture represents a significant northward extension of the known range of this species on the coastal plain.
The orange cap on the blue spot on the hindwing ventrum showed the identity of this rare find. Its abdomen was thin, and its long tails were gone, but the slight roundness of its wings and the fact that it landed on a sweetgum sapling at about 5-6 feet above the ground corresponded with the description of Gatrelle (1974, J. Lepid. Soc. 28:33-37) of the flight habits of females. Its flight was slow, due possibly to its age, the lateness of the hour, or the deep shade in the area, but it does agree with the “sluggish” adjective used by Covell and Straley (1973, J. Lepid. Soc. 27:144-154). The very late date and the condition of the specimen (Fig. 1) indicated that this was possibly the last survivor of the season’s brood.
Fic. 1. Left: S. kingi, male. Suffolk, Nansemond County, Virginia, July 1, 1974, lower aspect; Right: S. kingi, female, Millville, Worcester County, Maryland, July 22, 1982, lower aspect.
64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
The specimen was taken along a damp trail from a sandy road, where the vegetation consisted primarily of sweetgum (Liquidambar styraciflua L.), red maple (Acer rubrum L.), white cedar (Chamaecyparis thyoides (L.), loblolly pine (Pinus taeda L.), sweet bay (Magnolia virginiana L.), tassel-white (Itea virginica L.), blueberry (Vaccinium sp.), and sweet pepperbush (Clethra alnifolia L.), which was just coming into bloom. This habitat resembles in some respects that designated as Group A for kingi by Gatrelle (1974).
Only three worn Megisto cymela (Cramer) and one fresh male Wallengrenia otho (J. E. Smith) were seen in the same area on the date of capture. Incisalia henrici (Grote & Robinson) was common and I. augustus (Kirby) rare in the previous spring, the only other time I had collected there.
WILLIAM A. ANDERSEN (M.D.), 220 Melanchton Avenue, Lutherville, Maryland 21098.
Journal of the Lepidopterists’ Society 38(1), 1984, 64
THE IDENTITY OF WING HAIRS IN MEGALOPYGIDAE
The wings of Megalopygidae were described as being ee with long, wrinkled or wavy hairs that gave them a wooly appearance.
By making transparent impressions of the upper surface of the front wings of both male and female Megalopyge opercularis (J. E. Smith), using the replica method de- scribed by Khalaf (1980, Fla. Entomol. 63(3):307-340), it became clear that the wings were covered with scales that were deeply divided (Figs. 1 & 2); the apices were atten- uate; and the branches formed the so-called “hairs”. The base of the scales was cuneate (attenuate) as in other moths.
This investigation received support from the Academic Grant Fund of Loyola Uni- versity.
KAMEL T. KHALAF, Loyola University, New Orleans, Louisiana 70118.
Fics. 1 & 2. Light micrograph of replica of the front wing of Megalopyge opercularis (J. E. Smith), showing deeply divided scales: 1, female; 2, male.
VOLUME 38, NUMBER 1 65
Journal of the Lepidopterists’ Society 38(1), 1984, 65
POPULATION OUTBREAK OF PANDORA MOTHS (COLORADIA PANDORA BLAKE) ON THE KAIBAB PLATEAU, ARIZONA (SATURNIIDAE)
The pandora moth (Coloradia pandora Blake) is fairly widespread in the pine forests of the Rocky Mountains, and occasionally exhibits large population outbreaks as noted by Ferguson (1971. Moths of America North of Mexico, Fascicle 20.2A, E. W. Classey, Ltd., London). Such an impressive outbreak was noted on a visit to the Kaibab Plateau of northern Arizona in August 1982. During a field trip to the plateau, thousands of adult pandora moths were observed flying about or landed upon tree trunks in yellow pine (Pinus ponderosa) forest in the daytime hours. While driving a northsouth transect the full length of the Kaibab Plateau on 15 August 1982, the greatest concentrations of pandora moths were noted within a two-three mile zone surrounding the Jacob Lake Junction, on State Highway 89 (Alt.) and Highway 67. Hundreds of adult moths (many freshly emerged) and thousands of eggs were noted on the buildings and tree trunks at Jacob Lake, especially near outside lights that were kept on at night.
Adult males and females were active in large numbers nocturnally as well as diurnally, because “black lighting” at night produced heavy catches near the North Rim of the Grand Canyon on 16 August. Wygant (1941. Jour. Econ. Entomol. 34(5):697-702) noted in Colorado that the peak emergence of adults was in July, every-other-year, because of a two-year life cycle, and the primary food plant was lodgepole pine (Pinus contorta). In another area, Oregon, yellow pine was reported to be the principal food plant of the pandora moth by Packard (1914. Mem. Nat'l. Acad. Sci. 12:1—276). Since the yellow pine predominates on the Kaibab where pandora moths were observed to be most abundant in August 1982, this pine is probably the most important food plant there.
Several hundred eggs were oviposited by freshly collected females placed in glassine envelopes. The ova were glossy blue-green spheres which hatched in early September three to four weeks after oviposition. This fits with Ferguson’s notation that the young larvae overwinter, mostly in the second instar, on the pine branches at the base of needles. Attempts to rear the larvae on Pinus palustris (which was available to the author) failed.
Adult pandora moths are clearly strong flyers, since one was observed flying across a barren desert landscape some 45 miles west of the edge of the Kaibab Plateau and the nearest pine trees. Undoubtedly, during large population outbreaks, some individuals wander great distances in search of suitable food plants to oviposit upon.
LARRY N. BROWN, Department of Biology, University of South Florida, Tampa, Florida 33612.
Journal of the Lepidopterists’ Society 38(1), 1984, 65-66
TWO LARGE COLLECTIONS OF MACROLEPIDOPTERA TO THE MILWAUKEE PUBLIC MUSEUM
The Milwaukee Public Museum in recent years has received two major Lepidoptera collections, the William E. Sieker Collection of Sphingidae and the James R. Neidhoefer Collection of Macrolepidoptera of several families.
A donation from the wife of the late Mr. Sieker and daughter Marie, the Sieker Collection was acquired by the Milwaukee Public Museum in September 1982. Amassed
66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
over almost fifty years, this outstanding collection totals over 9000 prepared Sphingidae representing some 150 genera, 1000 specific and subspecific taxa, and includes some type material. All major faunistic regions are well represented. Mr. Sieker acquired a major portion of the collection through exchange and donations from biologists conducting field research in different parts of the world. The collection includes all of the known Wis- consin sphingid species, the result of Mr. Sieker’s own collecting and his strong association with other naturalists in the state over the years. There are also several hundred prepared Catocala moths (Noctuidae), an additional several thousand papered specimens, associ- ated field notes, several hundred reprints of important works on sphingids and other groups, and reference books.
William E. Sieker was born in 1912 in Milwaukee, Wisconsin and died in Madison, Wisconsin in January 1982 at age 70. Although a tax attorney by profession, he pursued a second career of collecting and studying sphinx moths, particularly in the northern reaches of Wisconsin. His interest in Sphingidae was fused with a dedication of helping conservation efforts, particularly in Wisconsin. Mr. Sieker was a founder of the Wisconsin Entomological Society and a past president of that organization as well as of the Madison Audubon Society. He was also legal counsel for the Wisconsin Chapter of Nature Con- servancy and helped that organization acquire the Ridges Sanctuary at Baileys Harbor in Door County.
The museum acquired the James R. Neidhoefer Collection in January 1976, a collec- tion which includes about 95,000 specimens of Macrolepidoptera (approx. 45,000 pre- pared and 50,000 papered specimens) of which approximately 1200 are gynandromorphs, sexual mosaics and aberrations (structural and color). The collection is particularly strong in Papilionidae, Nymphalidae, Heliconiidae, Ithomiidae, Morphidae, Pieridae, Saturni- idae and Sphingidae. All of the major faunistic regions are represented, with particular strengths in the Neotropical and Indo-Australian Regions. Mr. Neidhoefer’s donation also included 182 insect storage cabinets with 105 drawers, and an extensive library of rare books, reprints, and monographs on the Lepidoptera. At the time the collection was donated to the museum, Mr. Neidhoefer also financed the renovation of a collection storage room in the Invertebrate Zoology Section (which includes entomology).
Mr. Neidhoefer acquired his collection over forty years, through buying and exchang- ing specimens with collectors all over the world, and through field expeditions (i.e., Brazil) financed by him and for the purpose of collecting. With the cooperation of a former curator of the Milwaukee Public Museum, Kenneth MacArthur, Mr. Neidhoefer was instrumental in acquiring other Lepidoptera collections for the museum, most notably the George Berg Collection (which includes a good series of Nicaraguan Rhopalocera), the P. Gagarin Collection (Brazil), and others.
James R. Neidhoefer was born in Milwaukee in 1917 and became an avid naturalist at an early age. He received an undergraduate degree in zoology from Marquette Uni- versity and did a thesis on the freshwater sponges of Wisconsin. He took over the family carpet business in Milwaukee but continued to pursue his interests in natural history by collecting Lepidoptera and teaching his 12 children about insects. Prior to moving to Miami, Florida in 1981 to pursue a new retirement job as president of a wholesale pet distributorship, Mr. Neidhoefer was very active in local nature organizations and the Boy Scouts, as well as the Milwaukee Public Museum. As an honorary curator for the museum, he now collects Lepidoptera and other invertebrates in Florida during his spare time.
With the acquisition of the Neidhoefer Collection, and through grants from the Insti- tute of Museum Services and the Friends of the Milwaukee Public Museum, a major collection reorganization and upgrading of facilities was initiated by the museum’s full time curators in the Lepidoptera area, myself and Susan S. Borkin. With the acquisition of the Sieker Collection, and combined with further collecting efforts in Wisconsin and also from ecological studies in the Neotropical Region, one of our goals is to make these outstanding collections of use to curators, systematists, and biologists working on groups represented in them.
ALLEN M. YOUNG, Curator and Head, Invertebrate Zoology Section, Milwaukee Pub- lic Museum, Milwaukee, Wisconsin 532338.
VOLUME 38, NUMBER 1 67
Journal of the Lepidopterists’ Society 38(1), 1984, 67
ARE CHAIN-LINK FENCES BARRIERS TO BUTTERFLIES?
During the summers of 1982 and 1988, I regularly collected European cabbage but- terflies, Pieris rapae Linnaeus from the Fenway Victory Gardens, Boston, Massachusetts, and Dunback Meadows, Lexington, Massachusetts. From 24 June to 18 August 1982, and 2 June to 1 August 1983, I observed 27 confrontations between free flying P. rapae and chain-link fences. On each occasion the butterfly flew within 5-10 cm of the fence, back and forth over a 1 to 1.5 meter area, and then added a vertical movement of equal distance. Three times P. rapae succeeded in flying over the fence. Once a male flew to the end of the fence and around if, and once a butterfly proceeded after a 2-3 second delay to pass through the fence after I tried unsuccessfully to capture it. On 21 occasions P. rapae changed their flight direction nearly 180° after confronting chain-link fences. On one occasion an alfalfa butterfly, Colias eurytheme Bdy. was observed to change direction approximately 90° after confronting a fence. A 90° change was also observed once for a P. rapae after physically striking a fence. The openings in a chain-link fence measure approximately 7 cm in height and width. The mean wing spread of P. rapae is only 3.8 cm. On several occasions I have seen individual P. rapae squeeze their folded wings through 1.8 cm wire screening of a flight cage in the laboratory; and in the field, I have observed individuals fly without hesitation through thin wire fences with openings of 12-15 cm. Even though chain-link fences have openings through which a P. rapae could physically pass without contact, the butterfly rarely does so. Perhaps P. rapae can not accurately judge the opening size; it may appear small and likely to damage wing tips; or perhaps the thick shiny wire on all sides of the butterfly may be distorted by the butterfly’s visual system and perceived as a nearly solid barrier.
Chain-link fencing is used widely to keep would-be intruders out of areas or keep in desired objects. Mountain alpine areas are under increasing pressures from human visitors each summer. Some parks have posted personnel to keep visitors on established trails, others have begun to rope off areas. Chain-link fences have been proposed as a means to save badly trampled alpine areas.
The construction of chain-link fences and other obstacles may have a variety of effects on butterfly populations depending on the species involved and the habitat. Williams (1930. The migration of Butterflies, Oliver and Boyd, London. 473 pp.) states that Be- lenois severina and Vanessa cardui usually fly over obstacles with little or no lateral deviation from their line of flight. Feltwell (1982. Large White Butterfly The Biology, Biochemistry, and Physiology of Pieris brassicae (Linnaeus), Dr. W. Junk Publishers, The Hague, 535 pp.) reports that P. brassicae typically flies over obstacles rather than around them. However, Andronymus neander predominantly flies laterally with little or no vertical rise when confronted by an obstacle in its flight path (Williams, 1930. ibid). Generally, alpine lepidoptera fly very low to the ground to avoid winds. If fences are encountered, movement may be hindered, adding an additional energetic pressure on mountain butterfly populations which are often already low in number. Therefore, there may be serious deleterious effects on alpine butterfly populations if chain-link fences are built in these areas.
These observations are limited in number and species involved. Perhaps a more quan- tified investigation is merited. Such an investigation should be concerned with the height and opening sizes of fences, with a look at a number of different species in various habitats to determine if the observations reported here can be generalized.
MARK K. WourMs, Department of Biology, Boston University, 2 Cummington Street, Boston, Massachusetts 02215.
Journal of the Lepidopterists’ Society 38(1), 1984, 68
BOOK REVIEW
CATALOGO SISTEMATICO DE LOS LEPIDOPTEROS IBERICA. (I) MACROLEPIDOPTERA, by M. R. Gomez-Bustillo and M. Arroyo-Varela. 1981. Inst. Nac. Invest. Agrarias, Ministerio de Agricultura y Pesca, Madrid. 499 pp., 6 col. pls. (1200 Pta. [=$9.40)).
Recent catalogs and checklists, including those of Bradley et al. (1972) for England, Karsholt and Nielsen (1976) for Denmark, and Leraut (1980) for France and Belgium, have almost covered the entire Lepidoptera fauna of the most western parts of Europe with up-to-date checklists. The new catalog by Gomez-Bustillo and Arroyo-Varela closes the gap by covering the fauna of Spain and Portugal. Their work is the first of two volumes; the second volume is to cover the Microlepidoptera. The catalog initially strikes one as very different from most catalogs, since the cover has a large color photograph of the pierid Aporia crataegi (L.), not what one usually finds on catalog covers. Additionally, there are six color plates near the back of the book with photographs from nature of a representative species of each family in the book. The text is also untraditional inasmuch as bibliographic references are included for each family in terms of literature on the species of the Iberian Peninsula. There is an initial brief summary of the classification adopted for the catalog, generally following recent classifications like that of Common (1970, Insects of Australia), followed by a short introduction on the origins and evolution of Iberian Lepidoptera. The main text treats 39 families of so-called Macrolepidoptera, grouping many primitive families together with the normal macros. This arrangement produces an artificial and utilitarian arrangement for the catalog designed to conform to the older concepts, whereby large-sized moths were placed in “Bombyces’’. This is not altogether detrimental, since a phylogenetic chart of families is included, but it does maintain the myth that these “Bombyces” are somehow related more than they really are, and it also detracts from a strictly systematic treatment of families from primitive to more advanced. Nonetheless, the catalog is a welcome addition to the works listing the European fauna.
The authors follow a family usage that splits families too much, in my view, but does follow the practice of many European specialists. Thus, such groups as Syssphingidae, Riodinidae, Danaidae, Thaumetopoeidae, Dilobidae, and Ctenuchidae, which many con- sider only of subfamily status, are here raised to family level. I did not make any detailed checks of nomenclature. In Sesiidae, however, not all synonymies are included for each species, only a few of the major ones. Each family name is provided with authorship and dates, as well as for other higher categories. The specific and generic checklist then follows, with a discussion section and reference list for each family. The species are all listed with their dates of authorship, with parentheses added when the names of species have been recombined. The names of subspecies and forms, however, are not provided with dates. Each species is also given a notation as to its place in a European faunal district; thus, statements are made such as “endemic to Iberia” or “supramediterranean. ”
The catalog is in spanish, but since the main text involves a checklist of taxa of the Iberian Peninsula, it is easily used by anyone. It is a welcome addition to the growing rank of faunal catalogs and checklists. One can only hope that now, in lieu of a new checklist of the entire Palearctic region, others will follow the lead of the authors and provide additional regional catalogs (e.g., the Balkans, Russia, the Far East), so that in this way we may in time have new lists of all the areas within the Palearctic region.
J. B. HEPPNER, Department of Entomology, Smithsonian Institution, Washington, D.C. 20560.
Date of Issue (Vol. 38, No. 1): 27 July 1984
EDITORIAL STAFF OF THE JOURNAL THOMAS D. EICHLIN, Editor
% Insect Taxonomy Laboratory 1220 N Street Sacramento, California 95814 U.S.A.
MacpDa R. Papp, Editorial Assistant Douc.Las C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor NOTICE TO CONTRIBUTORS
Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instruc- tions.
Abstract: A brief abstract should precede the text of all articles.
Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first mention of a plant or animal in the text should include the full scientific name, with authors of zoological names. Insect measurements should be given in metric units; times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where italics are intended. References to footnotes should be numbered consecutively, and the footnotes typed on a separate sheet.
Literature Cited: References in the text of articles should be given as, Sheppard (1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading LITERATURE CITED, in the following format:
SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 209 pp.
196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10: 165-216.
In the case of general notes, references should be given in the text as, Sheppard (1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30).
Illustrations: All photographs and drawings should be mounted on stiff, white back- ing, arranged in the desired format, allowing (with particular regard to lettering) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced photographically to that size or smaller. The author's name, figure numbers as cited in the text, and an indication of the article's title should be printed on the back of each mounted plate. Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illustrations.
Tables: Tables should be numbered consecutively in Arabic numerals. Headings for tables should not be capitalized. Tabular material should be kept to a minimum and must be typed on separate sheets, and placed following the main text, with the approx- imate desired position indicated in the text. Vertical rules should be avoided.
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CONTENTS
THE LIFE HISTORY AND ECOLOGY OF EUPHYDRYAS GILLETTII BARNES (NYMPHALIDAE). Ernest H. Williams, Cheryl E. Holdren & Paul R. Ehrlich ee
CORRECT NAME FOR THE NEOTROPICAL SQUASH-VINE BORER (SE- SUDAE: MELITTIA). Vitor O. Becker & Thomas D. Eichlin
LIFE HISTORIES OF FOUR SPECIES OF PHILIRIS ROBER (LEPI- DOPTERA: LYCAENIDAE) FROM PAPUA NEW GUINEA. Michael Parsons 22 ee
COURTSHIP BEHAVIOR OF THE GULF FRITILLARY, AGRAULIS VA- NILLAE (NYMPHALIDAE). Ronald L. Rutowski G John Schaefer 00 Ee
CHECKLIST OF MANITOBA BUTTERFLIES (RHOPALOCERA). Paul Klassen 2.00 ee ee ee
THE LIFE HISTORY AND BEHAVIOR OF EPIMARTYRIA PARDELLA (MICROPTERIGIDAE). Paul M. Tuskes & Norman J. Smith
A NEW ACANTHOPTEROCTETES FROM THE NORTHWESTERN UNITED STATES (ACANTHOPTEROCTETIDAE). Donald R. DGbis a ee
Two INTERESTING ARTIFICIAL HYBRID CROSSES IN THE GENERA HEMILEUCA AND ANISOTA (SATURNIIDAE). Richard Steven Peigler & Benjamin D; Williams _.... ee
SPERMATOPHORE PERSISTENCE AND MATING DETERMINATION IN THE Gypsy MOTH (LYMANTRIIDAE). Cynthia R. Loerch & E. Alan: Camerore 2c
GENERAL NOTES
Insect parasites and predators of hackberry butterflies (Nymphalidae: Aster- ocampa).. Timothy P. Friedlander: 2.0304 2) J
Ithomiine butterflies associated with non-antbird droppings in Costa Rican tropical rain forest. Allen M. Young 201000)»
Satyrium kingi (Lycaenidae) taken in Maryland. William A. Andersen .. The identity of wing hairs in Megalopygidae. Kamel T. Khalaf |... Population outbreak of pandora moths (Coloradia pandora Blake) on the
Kaibab Plateau, Arizona (Saturniidae). Larry N. Brow? .-cccccccsccceeneeneee
Two large collections of Macrolepidoptera to the Milwaukee Public Mu- seum. Allen M. Young
Are chain-link fences barriers to butterflies? Mark K. Wourms
BOOK REVIEW 09.2500 Cl eS
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1984 Number 2
Volume 38
ISSN 0024-0966
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Published quarterly
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
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16 August 1984
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
LEE D. MILLER, President CHARLES V. COVELL, JR., KAROLIS BAGDONAS, Vice President Immediate Past President MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer
Members at large:
K. S. BROWN, JR. F. S. CHEW J. M. BURNS E. D. CASHATT G. J. HARJES F. W. PRESTON T. C. EMMEL E. H. METZLER N. E. STAMP
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Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- ume, and recent issues of the NEWS are available from the Publications Coordinator. The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire to Publications Coordinator.
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044.
Cover illustration: Head (antennae mostly missing) of Paranthrene tabaniformis (Rot- temburg). This drawing was prepared by George Venable, Smithsonian artist, for inclu- sion in the Sesiidae fascicle for the Moths of America North of Mexico. The dusky clearwing, a Holarctic species, is a borer in the exposed roots, stems and branches of willows and poplars.
JOURNAL OF
Tue LeprporreRrists’ SOCIETY
Volume 38 ~ 1984 Number 2
Journal of the Lepidopterists’ Society 38(2), 1984, 69-84
LIFE HISTORIES OF TAENARIS (NYMPHALIDAE) FROM PAPUA NEW GUINEA
MICHAEL PARSONS
Insect Farming & Trading Agency, Division of Wildlife, P.O. Box 129, Bulolo, Morobe Province, Papua New Guinea
ABSTRACT. Descriptions and illustrations of the early stages and ecology of Taen- aris onolaus Kirsch and Taenaris catops Westwood are given with a brief description and illustrations of the early stages of Taenaris myops Felder. Adults of both T. onolaus and T. catops were frequently seen imbibing cycad juices which probably enhances their assumed distastefulness to predators. Their foodplant specializations and aposematic at- tributes are discussed together with the mimetic relationships of Taenaris.
The genus Taenaris Hiibner in Papua New Guinea numbers 18 species. Together with three species of the genus Morphopsis Oberthiir and the monotypic genera Hyantis Hewitson and Morphotenaris Fruhstorfer, these are the only representatives of the Morphinae to be found in the country. A further six species of Taenaris and one of Morphopsis are known from Irian Jaya. Torres Strait marks the bound- ary of the distribution of these few closely related genera and species in the Melanesian region. They do not occur on the Australian main- land.
The Morphinae occur widely throughout the Indo-Australian region and number about 100 species. The morphology of the early stages and of the adults indicate their close affinity with the Satyrinae. For example, adults of Morphopsis albertisi Oberthiir in Papua New Guinea superficially resemble the smaller satyrine Tisiphone helena Ollift. from north Queensland, Australia, where no mimicry could be in- volved.
Adults exhibit little sexual dimorphism, but males tend to be smaller than the females, have a more concave inner margin of the forewing and bear sub-basal androconial tufts on the hindwing.
Little was known of the biology of the Morphinae in the Melanesian region. Rosier (1940) gave some details of the biology of Taenaris
70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
horsfieldii Swains. from Java and D’Abrera (1977) mentioned briefly the early stages and foodplants of Taenaris catops Westwood and Taenaris phorcas Westwood. D’Abrera stated that a paper describing the life history of M. albertisi was in preparation, but until now there has been no detailed study of any species of this subfamily published for the region.
Taenaris onolaus Kirsch
D’Abrera (1977) lists four races of this species, two occurring in Irian Jaya. The description of the subspecies ida Honrath fits the butterfly described here and this, therefore, represents an extension of its range from the known type localities for the subspecies in the Huon Penin- sula. The 10 km grid square reference in which the subspecies has been found in Bulolo is DN50 at approximately 700 m. The present study was made from October to December 1979.
Egg (Fig. 2). 1.5 mm in diameter; pearly white when laid, changing within two days through cream to deep pink; almost spherical, but slightly tapered towards flattened apex; chorion covered with evenly spaced, shallow dimples. Duration, 14 days.
Larva. First instar. Length 4 mm on hatching, 5 mm at end of instar; head jet black, shiny with fine white setae; thorax and abdomen with fine white setae up to 1 mm in length, initially cream, gradually changing to yellowish green, then orange-red; prothorax with dorsolateral black spots. Duration, 2 days, and a further 5 days of inactivity during pre-ecdysis and ecdysis.
Second instar. Length 10 mm at end of instar; head jet black, shiny, 1 mm in diameter with setae 3.5 mm in length and pair of truncate, slightly forwardly curved horns 0.75 mm in length, each horn with 3 strong spines; thoracic setae 4 mm, abdomen with setae 3.5 mm in length; thorax and abdomen deep pink, abdomen with a dorsal black spot on anal segment. Duration, 4 days, plus 3 days of inactivity during pre-ecdysis and ecdysis.
Third instar. Length 22 mm at end of instar; similar to second but head 2 mm in diameter, horns 1 mm in length, each with 5 spines; thoracic setae 6 mm; thorax and abdomen pink with 4 indistinct, but continuous orange-yellow lines, 2 dorsolateral and 2 lateral. Duration, 6 days, plus 2 days of inactivity during pre-ecdysis and ecdysis.
Fourth instar. Length 35 mm at end of instar; head 3 mm in diameter, horns 1.5 mm, each with 6 spines; thorax and abdomen wine-red, orange-yellow lines slightly more prominent; body setae up to 9 mm in length; below these a layer of strong, sharp, black setae 1.55 mm in length. Duration, 5 days, plus 2 days of inactivity during pre-ecdysis and ecdysis.
Fifth instar (Fig. 4). Length 60 mm at end of instar; similar to fourth but head 5 mm in diameter, horns 2.5 mm with 6 strong spines (Fig. 20c); body setae up to 10 mm; lower black setae 3 mm. Duration, 8 days, plus 2-3 days spent wandering.
Prepupa. Larval color changes from wine-red to yellow after suspension prior to pupation so that lower black setae and black spots of prothoracic and anal segments become very prominent. Duration, about 1 day of hanging before larva to pupa ecdysis.
Pupa (Figs. 6 & 7). Length 30 mm; ovate, smooth, translucent creamy white; cremaster black; anal rise with 2 black tubercles; apical margin of front bifid, forming 2 short, conical horns above each eye 1 mm in length. Duration, 17-20 days.
Ecological Observations
Foodplant and habitat. The foodplant is, unusually, a gymnosperm, Cycas circinalis (L.) Laut. & K. Sch. of the order Cycadales. This,
VOLUME 388, NUMBER 2 Teak
Fics. 1-9. Taenaris onolaus: 1, female ovipositing on cycad; 2, eggs; 3, second instar larvae; 4, mature larva; 5, mature larvae at rest in the leaf litter near the base of their foodplant; 6, dorsal profile of the pupa; 7, lateral profile of the pupa; 8, upperside of male; 9, underside of female.
2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
however, is not a unique specialization among the butterflies. Taenaris butleri Oberthiir is known from the same foodplant (T. Fenner, pers. comm.) and also Luthrodes cleotas Guérin of the Lycaenidae (Szent- Ivany et al., 1956). Another lycaenid, Theclinesthes onycha Hewitson, is also known from Cycas in Australia (Sibatani & Grund, 1978).
The foodplant exhibits a well defined distribution in the study area, being restricted to a well drained ridge alongside a gravel road behind the Bulolo Forestry College. The cycads cover about 3 acres which makes the area ideal for the study of a defined population of T. ono- laus.
Cycas circinalis occurs locally at a quite high density (in places up to six plants per 10 square meters), mainly along the top of the ridge and under a 15 year old Pinus plantation. The plantation provides a fairly open habitat with only semi-shading by the thin pine canopy. Saplings of other trees occur sporadically throughout the plantation. These conditions appear to be ideal for the growth of the cycads and may explain why the plant is not found locally outside the area where the scrub becomes thicker. A number of plants were fruiting prolifi- cally during visits to the area in October, November and December 1979, and there were many cycad nuts on the ground.
Oviposition and phenology. Eggs are laid by females in batches ranging in number from 20 to 40 with an average batch size of about 30. The highest number recorded in a single batch was 77. They are deposited close together, but not touching, on the undersides of one (or sometimes two) leaves of the older, tougher, dark green fronds. They are always placed about one third of the way down from the tip of the frond. Occasionally (seen in at least seven batches), there are one or two unfertilized eggs which remain white after the others have changed to pink.
The plants on which the females choose to oviposit are all of about the same height, approximately 1.5 m tall, and with usually 5-15 fronds. Cycads are extremely slow growing, and these plants are esti- mated to be from 5-6 years old (possibly older). As yet they have little or no trunk, and the fronds of most of them arise directly from the ground. No eggs or larvae were found on the younger plants with only two or three fronds and of smaller overall size at the beginning of the study period.
From observations of two females made late one afternoon in De- cember from 1735 h onwards, it appears that T. onolaus only oviposits during the period of about two hours before complete darkness which is at 1900 h, dusk (or half-light) coming at about 1830 h. (This was suggested later by two further observations of females ovipositing at
VOLUME 388, NUMBER 2 73
dusk.) One female was discovered at 1745 h below a cycad frond, having laid about 30 eggs. Approximately every two minutes she de- posited another egg in the row of four across the cycad leaf. Having completed a row she then moved slowly forward and positioned herself to begin a new row, from side to side. It is estimated, therefore, that a whole batch of about 50 eggs (this particular female had gone by the next morning but laid 45 eggs) would take approximately two hours to lay.
A second female was seen at the same time flying around another cycad, repeatedly settling on the upperside of a frond and then crawl- ing beneath it. She then flew behind some vegetation, which obscured the other half of the same cycad, and settled out of sight. Soon after she was re-located sitting on a batch of about 25 newly laid eggs ready to recommence egg laying. It appears, therefore, that some females take periods of rest away from the cycads on which they are ovipositing then return to lay their eggs at intervals. Both females were still ovi- positing in near darkness at 1850 h.
It is possible that females are able to lay further batches of eggs. However, it appears that their ovaries produce a certain number of eggs that are laid as a single batch in a short period of time. They are probably fairly short lived once they have paired and have finished ovipositing. It is also evident that females can detect the presence of eggs or larvae (probably visually) that are already present on the cycad, because when the area was studied in mid-November, no suitable look- ing plant was found to have more than one batch of eggs or larvae on HU
Eggs of T. onolaus were first discovered at the beginning of October 1979 which marked the end of an extremely dry dry-season. This lasted from the end of June for three months. During this time there was no rain recorded for the Bulolo Valley. In October, however, there were a few batches of T. onolaus larvae to be found, mainly fourth and fifth instars. Two egg batches were located at this time, which indicates that, even though the climate can be seasonally extremely dry in the area, generations can be continuous throughout the year because of the hardiness of the cycads; their foliage remains constant all the year round. (During the 1979 dry season in the Bulolo Valley, many angio- sperms, especially vines, even in fairly dense forest areas, began to wilt and/or ceased new leaf growth. Often the dry season is hardly appar- ent.)
When the area was again revisited at the end of the first week in November, heavy rains had recommenced during the previous four weeks. At this time all but about eight suitable-looking plants had eggs
74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
or larvae, and the approximate census was as follows: 18 batches of eggs, 7 groups of first instar larvae, 5 groups of second instar larvae and 2 of third instar larvae.
During the latter half of December females were still ovipositing, and a number of cycads, even the smaller, single-frond plants, were seen with newly laid egg batches. Some hosted up to three age classes of larvae, all of which fed together. This suggested that the area was now almost saturated with early stages due to the build up in numbers of adults and that intraspecific competition can occur where the species has limited, or finite, food resources. The few suitable cycads with no larvae at this time implied that any eggs laid on them may have been the subject of predation. Although no predators have been seen taking early stages, certain fresh egg batches were often found to have been eaten by the next day. Tetigoniid grasshoppers and predatory bugs are likely to be responsible.
In general, it may be concluded that as T. onolaus is a cycad feeder of tropical distribution, it is subject to little periodicity, i.e., that it is continuously brooded all year round, but that any large fluctuation in population numbers is reciprocal of extremes of wet and dry weather. Prolonged dry periods appear to produce aestivative (diapause) pupae and may also retard growth of new cycad fronds, so that the result at the onset of new rains is a large buildup of adults and early stages which compete intraspecifically for foodplants in areas with limited distribution of cycads.
Larval behavior. Larvae of T. onolaus are gregarious throughout their feeding period. In the fourth and fifth instars, however, the dis- tance that separates each larva is increased, and they may be found feeding singly, or in sub-groups of up to five. First to third instar larvae spin an almost invisible mat of extremely fine silk on which they rest below the frond of the cycad, so that when the plant is viewed from above they are completely obscured from the observer.
When feeding, early instar larvae begin at the tips of the leaves of the cycad frond and eat each leaf back separately to the base of the main stem. The group will then begin to feed again on the next leaf and progress gradually downwards. They often defoliate a whole frond as they grow. The smaller larvae form very orderly rows when resting or as they feed on the edge of the leaf lamina (Fig. 3). Final instar larvae tend to be cannibalistic on soft, newly formed pupae if many are caged together. One particular batch of about 12 fourth instar larvae were found resting during the day at the base of a frond of one cycad and were thus hardly visible beneath the leaf litter trapped there (Fig. 5). This does not, however, appear to be typical behavior. They were not undergoing ecdysis, and it is possible that these larvae were
VOLUME 38, NUMBER 2 ‘ko
feeding at night and seeking shelter from predators during the day. All instars have been found feeding at various times during the day with no specific feeding or resting times.
In general, all instars are fairly slow in their movements. When touched they sometimes react by thrashing the head from side to side. This appears to be an effective means of warding off insect predators. Fifth instar larvae tend to curl up and fall off the cycad fronds if handled, behavior which enhances their very moth-like appearance.
Adult behavior and abundance. Females appeared to be most fre- quent in the study area and were seen at all times flying randomly throughout the pine plantation. Specimens were seen on every visit during the study period although never in great abundance at any one time. Numbers ranged on average from 1-4 (flying in close proximity) seen per hour.
At one time in mid-December a sample census resulted in the sight- ings during one hour of three males flying in a restricted gully on the border of the study area (one feeding on damp mud), and four females. Two of these females hung inertly beneath a cycad frond (late after- noon) and did not react to rapid hand movements nearby. One was picked up and promptly flew off when released. Invariably, however, adults are very wary and do not allow one to approach to within less than 2 meters if they are at rest and alert on the upperside of broad- leaved foliage.
There are no succulent fruit trees in the pine plantation, and none of the saplings which produce small berries that fall to the ground have proven attractive to adults. However, on a number of occasions both sexes have been seen feeding on the fermenting skins of cycad nuts that have fallen to the ground when brown and ripe.
Competition. Intraspecific competition has been mentioned. How- ever, there also exists in the area, interspecific competition for Cycas circinalis between T. onolaus and a chrysomelid beetle of the subfam- ily Criocerinae. The small 1 cm long, orange beetle which is probably Crioceris clarkii B. Baly (based on the discussion in Szent-Ivany et al., 1956), feeds as a cream colored larva on the cycads. It has a definite preference for the soft, new, light-green cycad fronds. Therefore, by selecting only the older tougher fronds on which to oviposit, T. onolaus probably avoids competition for individual plants. Nevertheless, the beetle does cause much damage to the cycads in the area and can be classed as a successful competitor with T. onolaus.
The feeding damage caused by the beetle larvae is very character- istic. Even for a long time after a cycad has been eaten back by either herbivore it is possible to determine whether it was fed on by beetle or butterfly. Whereas T. onolaus eats the whole leaf of a frond, the
76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
chrysomelid eats only the underside of a lamina and leaves the top waxy cuticle as a window. This soon dries, turns yellow, and is left trailing, still attached to the frond.
Many new, recently unfurled, cycad fronds were the subject of at- tack by the beetle in mid-December 1979. These beetles also appear to play a significant réle in controlling the population size of the but- terfly. It is possible that they cause a final crash in the numbers of a cohort of T. onolaus because, if there are sufficient numbers of the beetle, then growth of cycads in the area may be halted completely. There will not, therefore, be enough fronds which reach maturity for the benefit of T. onolaus.
Taenaris catops Westwood
D’Abrera (1977) lists 21 races of this species. As emphasized by Brooks (1950) the named subspecies of T. catops may be very artificial as the species is widely distributed in New Guinea, of common status, and exhibits a great phenotypic variability both locally and regionally. Considering for example the supposed subspecies mylaecha from Sud- est Island which is described by D’Abrera as an “albinotic extreme” (i.e., very white), the same form is now recorded widely from the Western Highlands Province of the mainland (Fig. 22). The other ex- treme is an extremely dark form of T. catops, in which black and dark grey have replaced almost all the white. Supposed subspecies of T. catops should, therefore, be accepted with caution and are more likely the result of clinal variation or Miillerian mimetic associations within their genus.
The life history of this species was also recorded from the T. onolaus study area in March and April 1980. The egg and first two instars cannot be described as only the third instar onwards were available.
Third instar (Figs. 10 & 11). Larvae grew extremely rapidly from 8 to 25 mm in 3 days; head jet black, shiny, 2 mm in diameter, covered with fine white setae, horns similar to those of T. onolaus, 1.5 mm in length; body covered with soft, white setae, longest (5 mm) on the prothoracic and anal abdominal segments, decreasing to 4 mm at body center; thorax and abdomen dark grey with 2 dorsolateral and 2 lateral white lines; spiracles encircled with yellowish orange; claspers laterally yellowish orange, dorsally with black patch surmounted by two short (0.55 mm) pointed tubercles. Duration, 3 days, plus a day of inactivity spent during pre-ecdysis and ecdysis.
Fourth instar. Length 32 mm at end of instar; similar to third but body laterally black with middorsal black line bordered with grey. Duration, 7 days, plus 1% days of inactivity spent during pre-ecdysis and ecdysis.
Fifth instar (Fig. 12). Length at end of instar 57 mm; similar to fourth but head 4.5 mm in diameter, horns with 8 long, thin spines (Fig. 20a); body jet black but for 2 dorsolateral white lines and 2 lateral yellow lines; spiracles black encircled with orange; soft white setae 7 mm longest; strong, sharp lower setae tan brown, 2.5 mm in length. Duration, 9 days.
Pupa (Figs. 13 & 14). Length 31 mm; smooth ovate, pale green; cremaster pale yellow,
VOLUME 38, NUMBER 2 eT.
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Fics. 10-15. Taenaris catops: 10, third instar larvae at rest; 11, third instar larvae feeding; 12, mature larva; 13, ventral profile of pupa; 14, lateral profile of pupa; 15, adult female imbibing the juices of a damaged cycad nut.
tipped with black. In shape pupa like that of T. onolaus but frontal horns slightly longer, more pointed, tipped with yellow and below this is a ring of pale brown; tubercles of anal rise not as prominent as those of T. onolaus, only faintly tipped with brown. Du- ration, 12 days.
78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Ecological Observations
Foodplant and habitat. The early stages of T. catops were discov- ered at the center of the T. onolaus study area previously described. The foodplant is a new record for T. catops. It is a 1.5 m tall ground orchid with large, predominantly white flowers, Phaius tancarvilleae (Banks in L’Herit) Bl. The plant has been found in the Bulolo Valley at 800 m growing under Pinus in the plantation. This may not, how- ever, be the usual foodplant for the species as D’Abrera (1977) states that T. catops feeds on the Black Palm (Caryota rumpha: Palmae), Betel-nut palm (Areca catechu: Palmae) and Banana (Musa: Musa- ceae). Pyle and Hughes (1978) list T. catops from Cordyline terminalis (Liliaceae) which is used in hedges in many highland areas of the mainland.
Larval behavior. Like T. onolaus, the larvae are gregarious and remain so up to the final instar. They feed in line from the tip of the leaf lamina and eat the blade gradually downwards to halfway or a little less (Fig. 11). The larvae pause at intervals and then move slightly back up the blade to rest.
Adult behavior and abundance. Females of. catops, like those of T. onolaus, were more often encountered in the area than males. They were also slightly more abundant than those of T. onolaus. Occasion- ally, up to five at one time were seen in one area.
T. catops has also been found just before dusk hanging inertly be- neath foliage. Only at this time can they be approached because they are otherwise always alert and wary when feeding or resting on the uppersides of leaves. In forest areas they prefer to fly in shade. The species has been observed in many localities on the mainland flying just above the leaf litter in search of fermenting fruits on which to feed or probing moist leaf litter.
In spite of their preference for shady habitats, both sexes of T. catops can commonly be seen flying through gardens in Bulolo and in straight lines across any open grassland areas in the Bulolo Valley. In sharp contrast, T. onolaus has never been observed outside the study area.
It is interesting to note that both T. catops and T. onolaus were fond of visiting the fermenting husks of cycad nuts on the ground (Fig. 15). At one time a female of T. onolaus was seen feeding between two T. catops females. At another time five T. catops were flushed from beneath two close-growing cycads on which the chrysomelid beetle larvae were feeding. They were seen to probe the fresh green frass of the beetle larvae where it had fallen to the ground. On numerous occasions the cut ends of cycad fronds on the ground which had exuded sap were seen to be extremely attractive to T. catops—this is discussed further below.
VOLUME 38, NUMBER 2 79
Fics. 16-19. Taenaris myops: 16, mature larva; 17, prepupal larva; 18, latero- ventral profile of pupae; 19, underside of male.
Taenaris myops Felder
D’Abrera (1977) lists 13 races of this species. Its full life history was studied from a batch of 37 eggs. These were collected from the un- derside of the leaf of a monocotyledon, Tapenochilus sp., of the Cos- taceae found in November 1980, growing on a creekside near Eilogo Falls (Port Moresby, Central Province, 10 km grid square EK45). This represents a new foodplant record. However, as the author, after lo- cating the eggs and watching them hatch, had other commitments, the early stages of T. myops were reared and photographed by Peter Clark. He noted that, in general, the whole life history was similar to that of T. catops.
Egg. Slightly lighter pink but otherwise similar to that of T. onolaus.
Larva. First instar. 4 mm long on hatching; head jet black, shiny, covered with fine white setae; body with long fine white setae, up to 1 mm in length; thorax and abdomen Opaque creamy white, gut from behind head to last 4 abdominal segments shows as pinkish red line, anal segments with traces of pinkish red.
Second to fourth instars. Larvae at each instar exhibited similar growth rates and maximum sizes as those of T. catops. They grew steadily darker so that by fourth instar
they were brownish black. Fifth instar. Length 59 mm at end of instar; head horns with 6 spines (Fig. 20b);
80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
20
Fic. 20. Frontal profile of left horn and lateral profiles of Taenaris final instar larval head capsules: a, T. catops; b, T. myops; e, T. onolaus.
prothoracic segment wine-red, remainder of thorax and abdomen jet black, not lined as larvae of T. onolaus and T. catops; spiracles black, encircled with wine-red; body setae soft, long, white, laterally 4 mm, dorsally 10 mm in length; lower strong, black setae 4 mm longest.
Prepupa (Fig. 17). Larval color changed to dark grey dorsally and pale green ventrally after larvae had suspended themselves prior to pupation. Pupation took place 40-42 days after larvae hatched.
Pupa (Fig. 18). 30 mm in length; shape like that of T. onolaus but color like that of T. catops. Duration, 18 days.
The overall duration from the time that the eggs hatched to the emergence of the adults was 54 days. T. myops has been previously recorded in Papua New Guinea feeding on coconut (Cocos nucifera) and oil palm (Elaeis guineensis) both of the Palmae (Dept. Primary Industry, unpublished).
DISCUSSION
Cycads are known to be toxic and often lethal to cattle. Whiting (1963) discussed the toxicity of cycads in general, and Yang and Mick- elsen (1968) have shown that the husk of Cycas circinalis is toxic to rats. It is quite probable, therefore, that the larvae of T. onolaus, like many “pharmacophagous”’ butterflies (the Aristolochia-feeding swal- lowtails, for example), can sequester, and store, certain compounds (such as bitter alkaloids) which render them distasteful to birds and other predators. Their bright wine-red color suggests this. The larvae
VOLUME 38, NUMBER 2 81
of T. butleri, which also feeds on cycads, are also wine-red (T. Fenner, pers. comm.).
It is possible that the larvae of T. catops and T. myops are more palatable to their predators, because their foodplants are not known to have toxic properties. Other species and their foodplants, which have not yet been mentioned but which are relevant to this discussion, in- clude Taenaris artemis Vollenhoven on coconut (Cocos nucifera: Pal- mae) and T. phorcas on tanget (Cordyline: Liliaceae) (T. Fenner, pers. comm.). Rosier (1960) has found the wine-red larvae of T. horsfieldii on Smilax (Smilacaceae) and, according to Corbet and Pendlebury (1978), the closely related genus Faunis in Malaysia feeds on Smilax (Smilacaceae), Musa (Musaceae) and Pandanus (Pandanaceae). Recent records of other Taenaris foodplants sent into the Insect Farming and Trading Agency include Taenaris dimona Hewitson on banana (Musa: Musaceae) and Taenaris gorgo Kirsch on Black Palm (Caryota rum- pha: Palmae). Both records were from the Maprik area, East Sepik Province. I have recorded the life history of Taenaris artemis on Pan- danus (Pandanaceae) in the Western Province. The larvae were pre- dominantly yellow marked with black.
Although Taenaris larvae do not appear to advertise their presence, all species nevertheless feed gregariously, which is behavior character- istic of distasteful Lepidoptera. However, on some foodplants the lar- vae of certain Taenaris species may be unable to store adequate sec- ondary plant compounds for their effective protection. If T. catops obtains no such protection by feeding on ground orchids, then this may explain why adults were seen to imbibe cycad juices and consequently were so common in the study area. A similar conclusion was reached by Edgar et al. (1976) for danaine butterflies that enhanced their un- palatability by visiting the withered leaves of plants which produced pyrrolizidine alkaloids. The observation that T. onolaus, even as an adult, imbibed cycad juices strongly supports the hypothesis that Taen- aris is a distasteful group of butterflies and that some species enhance this as adults. It may be added that the fermenting skins of cycad nuts have an extremely nauseating smell.
All species of Taenaris so far studied in the field have exhibited great wariness and are quick to avoid capture. This, together with their eyespots and the protective hairs and bristles of their larvae, may be considered to be secondary lines of defence if they have been retained from an ancestral form that was more cryptically colored and in which these characters were of primary protective function. Such an ancestor may have looked like the small, dull, species of Fawnis found in Ma- laysia today. The general trend to enhance the aposematic attributes
82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
of Taenaris appears to have been for the butterflies to increase in size, to become lighter, and for the eyespots to become enlarged and high- lighted with broader orange borders. Of the cycad feeding species so far studied, the ground color is predominantly black, and the extent of the orange has been greatly increased so that it is highlighted as a warning color. It is interesting to note that the same also appears to be true of the cycad feeding lycaenid Luthrodes cleotas from Papua New Guinea which has large patches of orange on the upper and underside of its hindwings in both sexes.
It is possible that the ability of certain Taenaris species to feed on cycads as larvae is a recent evolutionary advance. J. Holloway (pers. comm. in discussion) suggested that the initial transfer to these prim- itive gymnosperms may have been a result of the similarity in the appearance of cycad fronds and those of coconut palms, for example, so that some Taenaris females began to oviposit on them by mistake. Alternatively, the transition from angiosperms, such as palms, to the cycad gymnosperms may have been through other angiosperms (such as Cordyline or Tapenochilus), that acted as “bridges,” i.e., they con- tained secondary substances that were common (or similar) to both. These may have acted either as oviposition cues to the females or phagostimulants to the larvae. The fact that adults of T. catops imbibe cycad juices could be taken to imply a closer link of this species with cycads in the past; however, it is also indicative of a chemical similarity between its normal foodplants and cycads.
T. catops exhibits a wide range of geographical forms and it is probable that it is a Millerian mimic of its close relatives. A recent sampling of Taenaris in the Cape Rodney area, Central Province, revealed what appears to be a Miillerian mimicry complex involving four species of morphines (pers. obs., Dec. 1979). These were Hyantis hodeva Hewitson, Taenaris mailua Grose-Smith, T. catops and T. myops. They were all extremely alike, and in particular, T. catops was more heavily marked than usual with extended black margins to the apices of the fore and hind wings. T. mailua differed in the area from the form of the nominate race and was slightly less heavily marked with black. It appears, therefore, that there was a convergence of the phenotypes of all the species in the area. All four species looked iden- tical on the wing.
Miillerian mimicry within the Morphinae appears to be a wide- spread phenomenon throughout New Guinea in general, and another good example has been recorded from Minj in the Western Highlands Province between H. hodeva and T. catops where the extremely white form of T. catops is predominant (Figs. 22 & 24). H. hodeva in the area is almost white, lacks its usual heavy black apical margins and has
VOLUME 38, NUMBER 2 83
Fics. 21-24. Miillerian mimicry in female morphines: 21, normal Taenaris catops from Bulolo; 22, albinotic T. catops from Minj; 23, normal Hyantis hodeva from Bulolo; 24, albinotic H. hodeva from Minj. (Males from the two localities are like their females. )
reduced eyespots. In and around the Bulolo Valley the same species are also alike, but in this locality they are heavily marked (Figs. 21 & 23) and conform to the more normal and widespread phenotypes. Other butterflies are probably Batesian mimics of Taenaris. For example, the female of Mycalesis drusillodes Oberthiir is thought to be mimetic of H. hodeva (Vane-Wright, 1971). Both model and mimic have been collected from the Torricelli Mountains near Maprik in the East Sepik Province (P. Clark, pers. comm.) and at Mt. Bosavi in the Southern Highlands Province (pers. obs., April 1980). The satyrine genus Elymnias is apparently mimetic of certain species of Euploea (Danainae), and Elymnias agondas Boisduval females are extremely good mimics of Taenaris bioculatus Guérin and T. catops where the models and mimics occur sympatrically. Hypolimnas deois Hewitson (Nymphalidae), in color and pattern, is very Taenaris-like and may be mimetic of T. onolaus in the Bulolo Valley. It is also assumed that the female form onesimus Hewitson of Papilio aegeus Donovan (Pa-
84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
pilionidae) mimics T. catops, and this form is also commonly seen around Bulolo. If this is so, then the female form amanga Boisduval of this swallowtail is a good candidate to be a mimic of T. onolaus. It has been seen frequently in the study area and strongly resembles T. onolaus in flight.
A more detailed study of the foodplant relations and mimetic asso- ciations of these butterflies will prove most interesting as further life- histories and the foodplants of other species of Taenaris are discovered.
ACKNOWLEDGMENTS
I am most grateful to Ted Fenner (Department of Primary Industry, Darwin, Austra- lia), Dick Vane-Wright and Jeremy Holloway (British Museum of Natural History, Lon- don), all of whom have improved this manuscript by their most helpful suggestions. Also to Peter Clark (Insect Farming and Trading Agency, Bulolo, PNG) for his assistance in completing the rearing and the photography of T. myops.
LITERATURE CITED
BROOKS, C. J. 1950. A revision of the genus Tenaris Hiibner. Trans. R. Entomol. Soc. Lond. 101:179-238. j
CorBET, A. S. & H. M. PENDLEBURY. 1978. The Butterflies of the Malay Peninsula. 3rd edition revised by J. N. Eliot. Malayan Nature Society. 578 pp.
D’ABRERA, B. L. 1977. Butterflies of the Australian Region. Landsdowne. 2nd edition, 415 pp.
EDGAR, J. A., P. A. COCKRUM & J. L. FRAHN. 1976. Pyrrolizidine alkaloids in Danaus plexippus L. and Danaus chrysippus L. Experientia 32:1535-1537.
PyLe, R. M. & S. HUGHES. 1978. Conservation and utilization of the insect resources of Papua New Guinea. 157 pp. Unpublished mimeo. report, Wildlife Division, Papua New Guinea.
Rosier, J. P. 1940. Aateekeningen over ontwikkelingsstadia van eenige javaansche vlinders. Entomol. Med. Ned.-Indie. 6:61-64.
SIBATANI, A. & R. GRUND. 1978. A revision of the Theclinesthes onycha complex (Lepidoptera: Lycaenidae). Trans. Lepid. Soc. Jap. 29:1-34.
SZENT-IVANY, J. J. H., J. S. WOMERSLEY & J. H. ARDLEY. 1956. Some insects of Cycas in New Guinea. P & NG. Ag. J. 11:1-4.
VANE-WRIGHT, R. I. 1971. The systematics of Drusillopsis Oberthiir (Satyrinae) and the supposed Amathusiid Bigaena van Eecke (Lepidoptera: Nymphalidae), with some observations on Batesian mimicry. Trans. R. Entomol. Soc. Lond. 123:97-123.
WHITING, M. G. 19638. Toxicity of cycads. Econ. Bot. 17:271-302.
YANG, M. G. & O. MICKELSEN. 1968. Cycad husk from Guam: Its toxicity to rats. Econ. Bot. 22:149-154.
Journal of the Lepidopterists’ Society 38(2), 1984, 85-87
A NEW SPECIES OF SIMILIPEPSIS AND TAXONOMIC PLACEMENT OF THE GENUS (SESIIDAE)
PING YUAN WANG!
~ Research Entomologist, Institute of Zoology, Academia Sinica, Peking, China
ABSTRACT. A new species of the wasp-like sesiid of the genus Similipepsis is de- scribed, and the taxonomic placement of this genus into the subfamily Tinthiinae is proposed.
The Section of Entomology of the Carnegie Museum of Natural History (CMNH) maintains a large collection of insects that has been vastly underutilized by systematists. The collection is rich in all insect groups but butterflies and moths are particularly abundant. The di- versity of taxa is particularly evident among the collection of unsorted moths in which I found a sesiid specimen with remarkable ichneu- monoid resemblance.
Further study of this wasp-like moth revealed that it belongs in the genus Similipepsis, a genus described by LeCerf (1911) and heretofore taxonomically unaligned in the Sesiidae hierarchy. Heppner and Duck- worth (1981:44) in their recent work made no study of this genus. They listed Similipepsis among other “‘unassigned” sesiid genera, leav- ing this problem for further research.
My studies of the genus revealed that Similipepsis species are char- acterized by having the abdomen constricted to a slender pedicel at the base, the proboscis normal, labial palpus oblique with the second joint of long hairs, forewing veins R, and R; stalked and M, missing, hindwing with vein Cu, from just before angle of cell and widely separated from Cu,, hind leg wasp-like. The genus is further recogniz- able by the absence of the scale tuft on the tip of the antennae. Ac- cording to recent classification (Naumann, 1971; Duckworth & Eichlin, 1977), these two characteristics suggest that Similipepsis has affinities and should be placed with genera of the subfamily Tinthiinae.
To date, there are only four known species of Similipepsis, S. aurea Gaede, S. lasiocera Hampson, S. typica Strand and S. violaceus Le- Cerf. The genus is paleotropical in origin and is confined geographi- cally to the Ethiopian and Oriental regions. After reviewing specimens and literature of known species (Strand, 1913; Hampson, 1919; Gaede, 1929), I determined that the aforementioned specimen in the Carnegie
‘Resident Museum Specialist, Section of Entomology, Carnegie Museum of Natural History, USA.
86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fic. 1. Adult male (holotype) of Similipepsis ekisi Wang, new species.
collection collected from the Cameroons was quite distinctly different and not conspecific with the known species.
I am indebted to Dr. Ginter Ekis for offering me the opportunity to study in the Section of Entomology. I would also like to express my appreciation to Dr. Chen Wen Young, the Collection Manager of the Section, for various courtesies during my six month research visit. I am indebted to Anna Tauber and Pat Vachino for literature and clerical assistance, and also to Vincent Abromitis, Section of Exhibits, for pho- tographic assistance. Dr. Craig Black, Director of the Carnegie Mu- seum of Natural History, provided the financial assistance that made it possible for me to come to the United States.
Similipepsis ekisi, new species
Holotype: Male Metet (Adamaoua), Cameroon (Republic of Cameroon), 15 August 1919. A. I. Good. Carn. Mus. Acc. 6552 (deposited in CMNH, Holotype number 775).
The holotype is associated with the following items: sex label (white, machine print); locality label (white, machine print); collection date label (white, machine and hand print); accession label (white, machine and hand print); CMNH repository label (yellow, machine print); holotype label (red, machine and hand print).
VOLUME 38, NUMBER 2 87
Male. Head: vertex black; frons brown; occipital fringe greyish white; vertex laterally with fringe greyish white mixed with black; labial palpi upturned, first and second segments brown and covered with extended long bushy scales, second segment less ex- panded than first, with brown scales on both sides and white erect scales on inside border, third segment white and sharply upturned above vertex; antenna brown and bipectinate, devoid of apical scale tuft; proboscis present. Thorax dark brown, tegula brown; meta- thorax with minute, slender brown and white hairy scales extended from base of hind- wing. Abdomen dark brown, first segment expanded slightly, second extremely narrowed and extended into a long stalk, third slightly expanded, fourth and fifth greatly expanded, sixth and seventh narrowed; anal tuft covered with setaceous, V-shaped brown scales; underside of third abdominal segment with ring of V-shaped white scales. Forewing transparent, except on costal margin; stem of R vein covered with dark brown scales, dark brown scales scattered in region of cell; cilia brown. Hindwing hyaline, with few scattered scales; veins and margins brown; cilia brown. Foreleg: front of femur, with long row of