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Showing papers in "Journal of The Lepidopterists Society in 1986"


Journal Article
TL;DR: For example, Calvert et al. as mentioned in this paper studied the migration behavior of monarch butterflies in the Transvolcanic Belt of central Mexico and found that during November and December the numerous small groups consolidate into a few large compact aggregations and move downward into more protected positions closer to water.
Abstract: Each year monarch butterflies migrate from breeding grounds in the United States and Canada to the Transvolcanic Belt of central Mexico. Here, within the montane fir forests, they initially aggregate in small groups of loose clusters scattered along high ridge crests. During November and December the numerous small groups consolidate into a few large compact aggregations and move downward into more protected positions closer to water. Butterfly activity increases in the last half of February due to seasonal warming. The consolidation and compaction processes that marked the beginning of the season reverse, and the colonies spread out and often split into two or more parts. After mid-March, colony size decreases as the butterflies begin to remigrate northward. Several characteristics of the climate and physiography of the Transvolcanic Belt, including moisture, altitude, and slope exposure and inclination, are important to the overwintering biology of the monarch butterfly. The forests of the zone playa major role in satisfying the overwintering monarchs' microclimatic requirements by moderating temperature extremes and conserving moisture. By colonizing this high altitude area in the tropics, the butterflies appear to satisfy microclimatic requirements that include temperatures low enough to keep activity, metabolism, and lipid expenditure to a minimum, but not so cold as to cause freezing; sufficient solar input to allow thermoregulatory basking and consequent flight; and sources of moisture and nectar. Each autumn, millions of monarch butterflies (Danaus plexippus L.) migrate southwest or south (Urquhart & Urquhart 1978, Schmidt-Koenig 1979) from breeding grounds in eastern and central United States and southern Canada to overwintering sites in Mexico. Funneling through Texas, they cross into Mexico and encounter the southern extension of the Rocky Mountains, the Sierra Madre Oriental. Here they change their southwesterly course and follow the ranges to the southeast, eventually cross them, and continue to the Transvolcanic Belt, the volcanic mountains that extend across the southern end of Mexico's Central Plateau (Altiplanicie Mexicana) between 19° and 200N latitude. At a few isolated places within the high altitude coniferous forests, which are scattered through this belt of mountains (Fig. 1), monarchs spend the winter in aggregations estimated to be in the tens of millions (Brower et al. 1977, Calvert, in prep.). Monarchs migrate south in the fall to avoid winter cold and survive in cool, moist places where they can conserve fuel reserves in a state of reproductive inactivity until making the return trip north in the spring. Yet weather in the overwintering areas does not ideally meet monarch requirements. Not only do temperatures occasionally fall into the lethal range (Calvert et al. 1983), but also intense insolation on VOLUME 40, NUMBER 3 165 clear and partly cloudy days stimulates butterfly activity to an extent that appears to contradict their need to conserve fuel. In an attempt to resolve these apparent contradictions, and to understand better why the monarchs choose these particular areas in Mexico, we here describe characteristics of the annual overwintering cycle and ecological features of several overwintering areas that we studied for nine seasons (December 1976 through spring 1985). PHYSIOGRAPHIC FEATURES, CLIMATE AND VEGETATION Volcanic cones and ranges dominate the terrain of the Transvolcanic Belt, which has an area of 60,000 km2, and measures approximately 640 km across by 95 km wide (Moore 1945). To the north it is bounded by the high Mexican plateau, and on the south by the large Balsas River drainage (Rzedowski 1978). Its eastern portion averages 2200 m elevation with numerous peaks rising above 3600 m, including the highest mountains in North America south of Alaska (Goldman 1951). The western portion contains fewer high peaks, and declines in elevation towards the Pacific. The central area where the monarch colonies are located (Fig. 1) is drained to the north and east by the Rio Lerma and to the south and west by the Balsas-Mezcala river system (Arbingast et al. 1975). Classic wet-dry season weather patterns prevail through most of the Transvolcanic Belt. Precipitation and heavy clouding is frequent from May until October, especially in the mountains, but winters are dry, and arid conditions prevail on the interior plains (Goldman & Moore 1945). However, winter and early spring storms occasionally occur in the area, and the higher elevations are subjected to high winds, heavy rains, snow, and ice storms (Mosina-Aleman & Garcia 1974). While potentially lethal to the overwintering butterflies (Calvert et al. 1983), these storms are also beneficial because they reduce the severity of the winter drought in the high-elevation overwintering areas. Because of the wide range of altitudes and climatic conditions, vegetation within the Transvolcanic biotic province is extremely varied. High interior plains and valleys consist largely of grasslands intermixed with patches of small trees, shrubs, yuccas, agaves and cacti. On mountainous slopes, forests dominated by oaks and pines give way to firs at about 2750 m (Goldman 1951), but in more humid areas, the firs commence as low as 2400 m (Rzedowski 1978). On the highest peaks, firs give way to alders and other species of pine and eventually to grassland and tundra (Goldman 1951, Goldman & Moore 1945). As is true of the lower limits, vegetational transitions depend on moisture and exposure, and the altitudinal limits of the fir zone may be influ-

75 citations


Journal Article
TL;DR: The responses of pipevine swallowtail caterpillars to simulated attacks of invertebrate enemies and to actual attack by coccinel\id larvae were examined and it was shown that they were more reactive to the simulated attack of a biting predator than to the simulate touch by an insect enemy.
Abstract: The responses of pipevine swallowtail caterpillars (Battus philenor: Papilionidae) to simulated attacks of invertebrate enemies and to actual attack by coccinel\\id larvae (Hippodamia convergens: Coccinellidae) were examined. The caterpillars were more reactive to the simulated attack of a biting predator than to the simulated touch by an insect enemy. Active fifth instars reached around to the posterior or walked away in response to stimuli, whereas prepupal fifth instars were more likely to extrude the osmeterium and never moved away from the stimuli. Caterpillars that were larger than the coccinellid predators were attacked but seldom eaten. In contrast, larvae that were the same size or smaller than the coccinellids were killed more frequently. When the caterpillars were attacked posteriorly, they defended a limited area by reaching around while the prolegs remained attached. The area defended depends on cuticular stretch, number of attached prolegs, current physiological state, and type and degree of stimulation. A common view of insects as prey is that behaviorally they are rather defenseless. Dixon (1973) stated, \"The general impression conveyed by the literature is that aphids and related small insects are helpless, sedentary and thin-skinned creatures that invite the attention of any predator that comes along.\" Generally, that same view is held for caterpillars and other immature insects. Yet caterpillars can and will defend themselves under certain circumstances, such as when attacked by insect predators and parasitoids. However, as I show here, pipevine swallowtail caterpillars (Battus philenor (L.): Papilionidae) have important constraints that limit the effectiveness of defensive behavior. Most six-legged insects can turn up to 3600 in their own defense. For instance, aphids can move forward or backward rapidly, and they can kick their adversaries (Banks 1957). Aphids can escape just before contact or immediately afterward by simply moving away quickly; many invertebrate predators must be within a centimeter of, or bump into, prey before detecting them (Russel 1972). Furthermore, before or after contact by predators, aphids can swivel on their stylets up to 1800 and thus continue feeding while avoiding their enemies (Russel 1972, Brown 1974). In contrast, caterpillars have a cylindrical body with short thoracic legs anteriorly and short prolegs posteriorly. This means that they differ greatly in their maneuverability from six-legged insects, especially aphids which are pear-shaped with relatively long legs. As a consequence of the mobility of six-legged insects, most invertebrate predators are more mobile than caterpillars and can outmaneuver them. 192 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Thus, caterpillars may benefit by escape, often dropping off their host plant (Myers & Campbell 1976). However, leaving the host plant can be risky. Caterpillars (and other soft-bodied insects) may starve before they locate a host plant, or die from desiccation or ground predation (Dethier 1959, Roitberg & Myers 1978, Rausher 1979). Larvae may drop on a thread of silk, which they can later climb, returning safely and quickly to their host plant (Dempster 1971). But predatory ants may climb down silk threads to capture larvae (Allen et al. 1970). Some caterpillars feed in hiding, a strategy that appears to be especially effective for early instars (Allen et al. 1970, Lopez et al. 1976). Web-making caterpillars may disappear into their webs when disturbed (Fitzgerald 1980). But webs do not deter some wasps and pentatomids from pursuing caterpillars within (Morris 1972, Schaefer 1977). Carabid beetles may tear open webs (Langston 1957). Chrysopid larvae with their long, sicklelike mandibles, and pentatomid and reduviid bugs with their long beaks, can attack prey through cloth and webbing (Fleschner 1950, Bornemissza 1966, Allen et al. 1970, Berisford & Tsao 1975). Furthermore, some predatory pentatomids and spiders live in webs of caterpillars (Morris 1972, E. W . Evans, pers. comm.). Therefore, caterpillars may benefit by vigorous defense when escape is less effective or more risky, such as when an insect predator initiates contact but cannot overwhelm the caterpillar. Typically, a defensive caterpillar attaches firmly to the substrate with the prolegs, lifts the thoracic legs and swings the anterior of the body toward the attacker, especially when approached from the side or rear by a predator. Caterpillars may use their bodies to hit and their mandibles to grasp an attacker (Morris 1963, McFadden 1968, Iwao & Wellington 1970, Frank 1971, Heinrich 1979, Suzuki et al. 1980, Stamp 1982). Unlike vertebrates and adult insects, caterpillars do not use their legs defensively . Instead, they may regurgitate or wipe offensive glands on attackers (Eisner & Meinwald 1965, Feltwell 1982). The questions posed in this study were: 1) when does a caterpillar opt to escape or for defense? 2) how does it defend itself? and 3) how effective is it in defending itself, or when are insect enemies successful in countering a caterpillar's defense?

36 citations




Journal Article
TL;DR: The hypothesis that butterflies as a group are primarily nectar thieves, rather than pollinators, of many flowers that they visit was tested by observing skippers and quantifying their pollen loads, and data do not reject the above hypothesis.
Abstract: The hypothesis that butterflies as a group are primarily nectar thieves, rather than pollinators, of many flowers that they visit was tested by observing skippers and quantifying their pollen loads. Two species of skippers, Atalopedes campestris and Epargyreus clarus, were studied. Adult A. campestris visited 23 flower species and Epargyreus clarus visited 27 flower species. Fifty-nine male and female E. clarus carried a mean of 45.1, and 283 male and female A. campestris carried a mean of 68.4 pollen grains from eight species of very frequently visited flowers. Skippers carried most of the pollen in their facial cavities and on their probOScides. At least one skipper of each species carried pollen from each of these flowers in its genital cavity, a newly documented pollen-carrying structure for butterflies. The skippers may have occasionally pollinated their nectar flowers, because they were constant to particular species during foraging bouts; they transported pollen; and they contacted stigmas with their pollen-bearing proboscides. Nevertheless, the skippers evidently functioned mainly as nectar thieves. They foraged mostly on asteriads rather than other kinds of flowers, primarily probing innermost (male-stage) disk florets, and they tended not to contact the outermost (female-stage) florets with their more pollen-laden parts. Moreover, they carried pollen loads that were too small to make them significant pollinators. Thus, our skipper data do not reject the above hypothesis. Many butterfly species visit flowers from which they imbibe nectar (Faegri & van der Pijl, 1966; Shields, 1972; Barrows, 1976, 1979; Schemske, 1976; Wiklund et aI., 1979; Schemske & Horwitz, 1984). For example, 197 butterfly species found in eastern North America use at least 5.9 ± 0.55 SE (1-15) genera of flowers as nectar sources (Opler & Krizek, 1984). Butterflies undoubtedly pollinate some flower species (Grant & Grant, 1965; Levin, 1972; Levin & Berube, 1972; Barrows, 1979; Cruden & Hermann-Parker, 1979), and they are definitely nectar thieves of others (Spears, 1983; Schemske & Horvitz, 1984). An individual nectar thief is an animal that takes nectar through a natural orifice of a flower without pollinating it (Inouye, 1980). Further, if an animal species thieves nectar during more than 50% of its visits to a particular flower species, the entire animal species could be classified as a thief species with regard to this plant species. Delpino (1874) suggested that male butterflies are likely cross pollinators of their nectar plants, but decades later Robertson (1924: 100101) stated that butterfly \"relations to flowers are often that of nectar thieves.\" Subsequently, Wiklund et aI. (1979) studied the flower visiting of the pierid Leptidea sinapsis L. in Sweden. From this species they generalized that, \"Butterflies as a group may have evolved to occupy a parasitic mode of life as adults, feeding on the nectar of flowers without pollinating them,\" but they did not refer to Delpino's 300 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY or Robertson's assertions. All in all, however, pollination effectiveness and efficiency of butterflies is little known (Gilbert & Singer, 1975; Kevan & Baker, 1983; Spears, 1983). In an attempt to test further the butterflies-as-nectar-thieves hypothesis, we studied foraging behavior of two common skippers, Atalopedes campestris (Boisduval) and Epargyreus clarus (Cramer), in Washington, D.C. The identities and relevant characteristics of the skippers' nectar flowers, skipper foraging behavior, and the locations and amounts of pollen that skippers carried were examined to test the hypothesis. Both skipper species that we studied are native to the Washington, D.C., area, where they have three broods per season (Clark, 1932). Atalopedes campestris fly in the garden from mid-July through September; E. clarus, from mid-June to early August. A future paper will discuss whether butterflies, in general, are nectar thieves or pollinators. MATERIALS AND METHODS In our study, we define \"foraging bout\" as a skipper's feeding activity on one or more flower species, starting when it was first discovered on a flower until it could no longer be followed due to its flying out of sight. A \"visit\" is a skipper's alighting upon or near a flower, extending its proboscis into it for at least 1 sec, and presumably feeding. An \"infrequently visited flower species (IVFS)\" is a flower that we saw only one individual skipper visit during only one of the ten 2-week observation periods of our study. A \"frequently visited flower species (FVFS)\" is a flower that we saw two to four conspecific skippers visit, and a \"very frequently visited flower species (VFVFS)\" is a flower that we saw five to hundreds of skippers visit during two or more of the 2-week observation periods. A \"clear day\" is one over 75°F, with no rain, and with less than 20% cloud cover. A \"facial cavity\" is a concavity into which a skipper's proboscis coils; a \"genital cavity,\" one at the end of a skipper's abdomen, formed in a female by scales surrounding her papilla anal is above and lamella antevaginalis below and in a male by scales surrounding his uncus above and valvae below. Skippers were studied from May through October 1982 in the 0.9ha vegetable and flower garden where Lazri and Barrows (1984) investigated flower visiting in Pieris rapae L. The garden is a community garden used in 1982 by about 146 gardeners, and it contains about 265 species of entomophilous plants, including vegetables, ornamentals, herbs, wildflowers, and weeds. Flowers visited by the skippers and the relative numbers of skippers present at each species were noted during a total of 12 30-min meandering walks made through the garden twice each month in June, July, August, and September. The walks were made once every 2 weeks on VOLUME 39, NUMBER 4 301 a clear day, every hour on the hour, from 0800 to 2000 h (EDT). At each skipper-frequented flowering plant or group of such plants, we made short (10 sec) counts to standardize the amount of time spent at a plant or group of plants. A total of 564 skippers of both species was counted during the entire census. To measure flower corolla lengths, we collected flowers in plastic bags and kept them moist until they could be examined. Dial calipers, accurate to 0.01 mm, were used to measure corollas (Lazri & Barrows, 1984). We made a pollen reference collection from pollen collected in the study area. In studying possible flower constancy, frequency of flower use, and pollen deposition of skippers, we observed 22 foraging A. campestris and 60 foraging E. clarus. A stopwatch and tape recorder were used when needed. To discriminate focal individuals from other nearby skippers when they were common, we marked forewings of focal individuals with small spots of enamel paint, which did not appear to affect their behavior. Forty additional skippers were each observed for 10 min as they foraged at asteriad disk and ray florets. In examining possible pollen transport and deposition, we collected 285 A. campestris and 77 E. clarus; 3 to 23 males and 5 to 25 females were taken from each VFVFS. Before it was captured, each skipper was followed as it visited two consecutive flower heads, extending its proboscis into a flower in each head for at least 1 sec. After it was netted, a skipper was paralyzed by carefully pinching the sides of its thorax between a thumb and forefinger and then placed into a glassine envelope on which relevant data were recorded. The enveloped skipper was immediately put into an insulated bag filled with frozen cold packs. Within the hour, all skippers in the bag were put into a cooler filled with more frozen cold packs. At the end of a collecting day, the skippers were put into a freezer until they could be examined for pollen (Turnock et aI., 1978). In searching for pollen on a skipper, we removed its legs and proboscis, placed them on a clean glass slide, and covered them with a drop of Permount® and a coverslip. The rest of the skipper was placed on a watchglass. Its proboscis, legs, body, glassine envelope, slide, and watchglass were examined for pollen under a compound microscope (up to 400 power), a dissecting microscope (up to 30 power), or both. Pollen adhering to the skipper's labial palpi were included in its facial cavity count. Free floating pollen grains on the slide and watch glass and in the envelope were also counted. Adult skipper age was estimated to be young, middle-aged, or old, based on the amount of scale loss and wing tattering that was present on a skipper's wings and body. A young skipper was one that was almost totally intact; a middle-aged 302 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY one had slight wing tattering and a few scales missing; and an old one had very tattered wings and many scales missing. Quantitative analyses were made with the Statistical Analysis System (SAS) computer package (Ray, 1982a, b). Pollen count and corolla depth values were log transformed to obtain homoscedastic data for the Duncan's multiple range test (DMRT). Possible differences between groups were analyzed with the t-test (TT) or paired t-test (PTT) corrected for heteroscedasticity when necessary, the Fisher exact probability test (FEPT), and the Chi-square test (CST) . Kendall's rank correlation coefficient (KRCC) was used to test for significant correlations. RESULTS AND DISCUSSION

14 citations




Journal Article
TL;DR: The biology and life history of Parnassius clodius Menetries is examined in the Pacific Northwest, where the species was formerly found in the Portland and Seattle metropolitan areas and is still quite abundant in the low foothills surrounding the Willamette Valley and the Puget Sound trough.
Abstract: This paper examines the biology and life history of Parnassius clodius Menetries in the Pacific Northwest. Habitats used by the species include subalpine meadows high in the mountains and lowland rain-forests west of the Cascade Range. The primary larval foodplants belong to the genera Dicentra and Corydalis of the family Fumariaceae. Larvae in alpine habitats often display a gray-brown camouflage pattern that blends with the rocks of the habitat. However, larvae in lowland rain-forests display a conspicuous black and yellow-spotted pattern that appears to mimic the warning colors of polydesmid millipedes. Larval development in lowland habitats is completed within a single year, and pupation takes place inside a strong, well-formed silken cocoon. Male butterflies display a "rape" type of mating, with no evidence of courtship behavior or sexual pheromones. Tough, tear-resistant wings and a large female sphragis may be related to this sexual behavior. Parnassius clodius Menetries belongs to a genus that is considered to be relatively primitive within the Papilionidae (Tyler, 1975). These are the only butterflies that have a moth-like pupa enclosed within a silken cocoon. Because of the putatively "primitive" nature of these butterflies, their life history and ecology is of considerable interest. Of the three species of Parnassius found in North America, only P. clodius is uniquely endemic to this continent and is widely distributed in the western mountains from southern Alaska to central California, western Wyoming, and northern Utah (Ferris, 1976). Some details of the life history and ecology of this species are outlined by Edwards (1885), Tyler (1975), and Dornfeld (1980). During the past twenty years, the present authors have studied various aspects of P. clodius biology in Oregon, Washington, and western Wyoming, resulting in much additional information. Ecology and Life History In terms of ecology, P. clodius occupies two distinctly different types of habitat. One consists of open subalpine meadows and rocky slopes above timberline at high elevations in the mountains. We have observed the species in subalpine meadows throughout western Oregon and Washington, and in Yellowstone National Park of Wyoming. We Volume 39, Number 3 157 also observed the species on alpine talus slopes above timberline at Harts Pass, Okanogan County, Washington. However, the most frequent habitat of P. clodius in the Pacific Northwest is the lowland rain-forests extending from the western slope of the Cascade Range west to the Pacific Ocean. Although typically found in moist riparian habitats along forest streams and mountain valleys, the species was formerly found in the Portland and Seattle metropolitan areas and is still quite abundant in the low foothills surrounding the Willamette Valley in Oregon and the Puget Sound trough in Washington. This forest habitat extends from the 4000 ft. (1200 m) elevation down to sea level near the ocean. The primary larval foodplant in these coastal rain-forests is the wild bleeding heart Dicentra formosa Andr., which is very abundant in moist forest habitats along the West Coast. A second probable foodplant is Corydalis scouleri Hook., a relatively uncommon species. We have not yet observed P. clodius larvae on this plant in the field, but they accept it readily in the laboratory. At high elevations in the alpine habitat and east of the Cascades, Dicentra uni flora Kell. is a likely foodplant. This species is a known foodplant of P. clodius in northern California (John F. Emmel, pers. coram.). All of these plants belong to the family Fumariaceae, and it is probable that related species such as Dicentra cucullaria L. and Corydalis aurea Willd. would also provide acceptable food plants. The female butterflies oviposit on and near the Dicentra plants. However, we have also observed females ovipositing on shrubs up to four feet above the Dicentra beds. Evidently a specific chemical emanating from the foodplant is sufficient to induce oviposition anywhere in the general vicinity of the foodplant. The larvae develop within the egg shell but do not emerge from the egg until the following spring. Eggs deposited on shrubs usually reach the Dicentra beds when the shrubs drop their leaves in the fall. Foodplant records such as Viola and Rubus mentioned by Ackery (1975) are almost certainly in error and may be due to this indiscriminate oviposition by the females. Early instar larvae have small tubercles, but later instars are mostly smooth with fine hairs. The larvae stay hidden in debris at the base of the foodplant most of the time. Feeding takes place very rapidly, so the larvae are exposed from cover only briefly. Nevertheless, P. clodius is frequently parasitized by tachinid flies in many localities. Osmeteria are poorly developed in Parnassius larvae and are not as important for defense against predators compared to Papilio larvae. Parnassius clodius larvae display two very distinct color morphs. One form is black with a lateral row of bright yellow spots on each side of the body (Fig. 1). The form of these spots is highly variable. 158 Journal of the Lepidopterists' Society Figs. 1-3. Left (1), larva of P. clodius, black form, Benton Co., Ore. Middle (2), larva of P. phoebus, Yakima Co., Wash. Right (3), larva of P. clodius, gray-brown form, Castle Lake, Siskiyou Co., Calif. Figs. 4-6. Left (4), Harpaphe haydeniana, Polk Co., Ore. Middle (5), open net cocoon and pupa of P. phoebus (behind thick Sedum stems in lower center). Right (6), well-formed cocoon of P. clodius cut open to reveal pupa ready to eclose. ranging from large round spots to long slender bars, or may be divided into several smaller spots. This color pattern is very similar to that of P. phoebus Fabr. (Fig. 2) and the Eurasian P. apollo L. (illustrated by Stanek, 1969). However, P. phoebus differs in having a second, more dorsal row of yellow spots on each side of the body. The second color form in P. clodius is gray-brown or pinkish gray with creamy yellow lateral spots and dorsal rows of narrow chevron markings equivalent to the dorsal row of spots seen in P. phoebus (Fig. 3). In our experience, Volume 39, Number 3 159 Table 1. Sequence of experiment testing the mimicry-model system of Parnassius clodius larvae and the millipede Harpaphe haydeniana as protection against the grasshopper mouse Onychomys leucogaster. 1. Clodius larvae given to mouse—larvae eaten. 2. Millipedes given to mouse—millipedes bitten, producing defense odor detectable to observer, mouse then rejected millipedes. 3. Meal worms given to mouse—worms eaten. 4. Clodius larvae given to mouse—larvae sniffed and rejected. 5. Adult meal worm beetles given to mouse—beetles eaten. 6. Clodius larvae given to mouse—larvae sniffed, handled, finally eaten after long delay. 7. Millipedes given to mouse—millipedes sniffed and rejected. 8. Meal worms given to mouse—worms eaten. the gray-brown form is dominant in alpine populations of P. clodius, for example at Harts Pass in Okanogan County, Washington and at Donner Pass in Nevada County, California. This morph appears to be a camouflage pattern that blends with the rocks in the alpine habitat. By sharp contrast, the black and yellow-spotted form is very conspicuous, is dominant in the lowland rain-forest populations of P. clodius, and appears to mimic the warning colors of polydesmid millipedes such as Harpaphe haydeniana Wood (Fig. 4). These millipedes are very abundant in the moist, riparian habitats used by P. clodius larvae. Some populations of P. clodius are polymorphic for both larval color forms. For example, larvae sent to us by John F. Emmel from Castle Lake in Siskiyou County, California displayed both color forms. Likewise, an adult female butterfly collected at Chinook Pass near Mt. Rainier National Park, Washington produced ten larvae, five of the black form and five of the pinkish gray form. In these, the black larvae retained the narrow yellowish dorsal chevrons of the gray larvae, a trait absent in most lowland black larvae. This ratio between the black and gray forms is suggestive of a simple Mendelian inheritance for these color morphs. However, the chevron markings are apparently controlled by a separate set of gene loci. In 1973, one of the present authors (McCorkle) conducted an experiment to test the predator protection of the mimicry-model system that apparently exists between lowland P. clodius larvae and the millipede Harpaphe haydeniana. Grasshopper mice (Onychomys leucogaster Max.) from eastern Oregon were used as predators in this experiment, since these insectivorous rodents do not occur within the ranges of the butterfly or millipede and would have no prior experience with these arthropods. The sequence of this experiment is shown in Table 1. This experiment appears to demonstrate that the mimicry color pattern of lowland P. clodius larvae can give them a degree of protection 160 Journal of the Lepidopterists' Society against predators, although predators may with sufficient experience learn to distinguish the larvae from millipedes. In nature, however, the millipedes are commonly exposed in the open, while P. clodius larvae are usually hidden and only briefly exposed during feeding. Thus, the mimicry may work quite well in nature, since predators would be expected to have abundant experience with the millipedes and little experience with the larvae. In lowland populations of P. clodius, development is completed in a single year. The larvae emerge from the egg shells during March and start to feed on the young Dicentra plants. Full larval development is reached usually by late April or May, followed by pupal development of several weeks, and adult butterfly emergence in June and July depending upon elevation. The pupa is short and rounded, dark brown in color, and quite similar to a saturniid moth pupa. It is enclosed within a strong, well-formed silken cocoon (Fig. 6). By contrast, the cocoon of

8 citations





Journal Article
TL;DR: The development of a hardware-cloth trap that will work for years without repair and that promises, with specified improvements, to catch more than 70% of the migrants that encounter it is described.
Abstract: Three models of a flight trap made principally of hardware cloth were tested at Gainesville, Florida. All models had a 6 m long central barrier of lh inch mesh hardware cloth. Butterflies encountering opposite sides of the barrier were trapped separately, allowing calculation of net movement up or down the Florida peninsula. The most efficient model has a barrier 3.7 m high and a two-stage trapping superstructure of v.. inch hardware cloth. It catches 22-70% of migrant Phoebis sennae, Agraulis vanillae, and Urbanus proteus. Migrating butterflies characteristically fly in a straight line a few meters above the ground and rise and fly over obstacles rather than deviating laterally (Williams, 1930). Beginning in 1975, I have used stationary flight traps that intercept and trap migrant butterflies at Gainesville, Florida (Walker, 1978, 1980; Walker & Riordan, 1981). My first traps were made of polyester, which ripped in strong winds and deteriorated in sunlight. They consequently required frequent repair and annual replacement. Furthermore, they lost about 90% of the migrants they intercepted. In this paper I describe the development of a hardware-cloth trap that will work for years without repair and that promises, with specified improvements, to catch more than 70% of the migrants that encounter it.









Journal Article
TL;DR: The following three new species of olethreutine moths are described to facilitate completion of a catalogue-checklist of the moths of Texas, a project originated by Andre Blanchard.
Abstract: Three new species are described; the male imago and male and female genitalia of each are figured. Eucosma rosaocellana is described from eight specimens from northwest Texas, and contrasted with E. salaciana Blanchard & Knudson. Dichrorampha hroui is described from 37 specimens from southeast Louisiana and northeast Texas, and is contrasted with D. leopardana (Busck) and D. incanana (Clemens). Pammene medioalbana is described from nine specimens from central Texas, and is contrasted with Cydia latiuscula (Heinrich) and C. gallaesalaciana (Riley). The following three new species of olethreutine moths are described to facilitate completion of a catalogue-checklist of the moths of Texas, a project originated by Andre Blanchard. Many other undescribed species of Texas Lepidoptera remain or are likely to be discovered in the Tortricidae and other families. In many such cases, the families or their subdivisions are in such great need of revision that isolated descriptions of new species could confuse the situation. Since the last comprehensive revision of Eucosma (Heinrich 1923), numerous new Eucosma species have been described, but the one described below is quite distinctive, and is closely related to E. salaciana Blanchard & Knudson (1981). Heppner (1981) described two new species of Dichrorampha from Florida, and Miller (1983) reduced five Dichrorampha species names to synonyms, and summarized other recent work in this genus. No new species of Pammene have been described from North America since Heinrich's (1926) revision. The holotypes of all species described below are in the U.S. National Museum of Natural