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Karner blue

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117-566: The Karner blue ( Plebejus samuelis ) is an endangered species of small blue butterfly found in some Great Lakes states, small areas of New Jersey , the Capital District region of New York , and southern New Hampshire (where it is the official state butterfly) in the United States . The butterfly, whose life cycle depends on the wild blue lupine flower ( Lupinus perennis ), was classified as an endangered species in

234-637: A diapause (resting) stage, and the hatching may take place only in spring. Some temperate region butterflies, such as the Camberwell beauty , lay their eggs in the spring and have them hatch in the summer. Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time searching for and eating food. Although most caterpillars are herbivorous, a few species are predators : Spalgis epius eats scale insects , while lycaenids such as Liphyra brassolis are myrmecophilous , eating ant larvae. Some larvae, especially those of

351-520: A 590-foot (180 m) diameter. Maxwell recommends considering the amount of shaded wild lupine habitat within the dispersal distance of Karner blue butterfly when reducing overstory. Many investigators stress the importance of habitat patches being closely spaced. For instance, at sites in west-central Wisconsin, the average distance from an occupied wild lupine patch to another occupied patch was significantly (p=0.002) shorter (709 feet, 216 m) than to an unoccupied patch (1,155 feet, 352 m). Wild lupine patches in

468-963: A blade height >4 inches (10 cm) should be performed annually or biennially in the fall or winter and clippings should be left in place. Effects of mowing, burning, and other management techniques and land uses on the average number of Karner blue butterflies observed per survey hour are included in. Recommendations on the use of a wide range of management techniques such as thinning, rotational grazing , and planting of wild lupine and/or nectar species can be found in. According to reviews and general field observations, management activities that are typically harmful to Karner blue butterflies include management that increases deer and/or grouse populations, close-cropped grazing, frequent or poorly-timed mowing, plowing, use of herbicides that kill lupine or nectar plants, and use of pesticides that are detrimental to Karner blue butterflies, ants they associate with, or pollinators of species they use for nectar. Information on

585-493: A butterfly cannot fly until the wings are unfolded. A newly emerged butterfly needs to spend some time inflating its wings with hemolymph and letting them dry, during which time it is extremely vulnerable to predators. The colourful patterns on many butterfly wings tell potential predators that they are toxic. Hence, the genetic basis of wing pattern formation can illuminate both the evolution of butterflies as well as their developmental biology . The colour of butterfly wings

702-570: A butterfly through metamorphosis has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in

819-417: A common family. In some species, such as the great spangled fritillary , the eggs are deposited close to but not on the food plant. This most likely happens when the egg overwinters before hatching and where the host plant loses its leaves in winter, as do violets in this example. The egg stage lasts a few weeks in most butterflies, but eggs laid close to winter, especially in temperate regions, go through

936-627: A conspicuous, fluttering flight. The group comprises the superfamilies Hedyloidea (moth-butterflies in the Americas) and Papilionoidea (all others). The oldest butterfly fossils have been dated to the Paleocene , about 56 million years ago, though molecular likely originated in the Cretaceous . Butterflies have a four-stage life cycle , and like other holometabolous insects they undergo complete metamorphosis . Winged adults lay eggs on

1053-554: A feeding trial larval duration of Karner blue butterflies fed mildew-infected wild lupine was not significantly different from treatments that resulted in the shortest larval durations. Karner blue butterfly fed water-stressed wild lupine had significantly longer larval durations than many treatments including larvae fed flowering wild lupine, shade-grown wild lupine in seed, or mildew-infected wild lupine. Although Karner blue butterflies have been shown to benefit from their association with ants, wild lupine with Karner blue butterfly larvae in

1170-576: A few butterflies (e.g., harvesters ) eat harmful insects, and a few are predators of ants , while others live as mutualists in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts. The Smithsonian Institution says "butterflies are certainly one of the most appealing creatures in nature". The Oxford English Dictionary derives the word straightforwardly from Old English butorflēoge , butter-fly; similar names in Old Dutch and Old High German show that

1287-444: A few species. Some butterflies have organs of hearing and some species make stridulatory and clicking sounds. Many species of butterfly maintain territories and actively chase other species or individuals that may stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies are often characteristic and some species have courtship flight displays. Butterflies can only fly when their temperature

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1404-1058: A harbinger of more such events forthcoming owing to climate change . Among the recommendations were managed relocation of populations from the southern parts of its range into northward habitats with suitable conditions and supportive plants. The report supported this proposal by noting that stakeholders and managers "are growing more supportive of novel science-based interventions to save rare species from climate change—driven extirpation." Articles that address Karner blue butterfly sampling methodology include. Swengel found that surveys for Karner blue butterfly adults appeared more efficient than those for larvae. Evidence of different catchability and/or detectability of male and female Karner blue butterflies led King to suggest calculating male and female population sizes separately. General descriptions of Karner blue butterfly rearing methods and translocation/ reintroduction techniques are available. A wide variety of management techniques can be compatible with maintaining Karner blue butterfly populations when attention

1521-606: A hard-ridged outer layer of shell, called the chorion . This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, called micropyles ; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly eggs vary greatly in size and shape between species, but are usually upright and finely sculptured. Some species lay eggs singly, others in batches. Many females produce between one hundred and two hundred eggs. Butterfly eggs are fixed to

1638-989: A large portion of its viable habitat. Remaining habitat is often fragmented, which is problematic for the lupine because it limits the range over which it can reproduce. Viable lupine habitat is often difficult to maintain because it flourishes after fires and other forms of disturbance. One reason this occurs is that lupine seed coats are so tough that only pressure changes due to rapid heating or abrasion are strong enough to allow water to penetrate and start germination . Moreover, fires, feeding by large ungulates , and mowing can improve habitat quality for established lupines by changing soil quality , vegetative structure, and leaf litter depth. Impacts of primary habitat loss have led to decreasing populations which are small and scattered which makes it harder for pollination to occur. Changes in land management such as prescribed burning, mowing, and mechanical thinning, would help protect and promote this plant and other plant diversity. Further research

1755-508: A leaf before eggs are laid on it. Many butterflies use chemical signals, pheromones ; some have specialized scent scales ( androconia ) or other structures ( coremata or "hair pencils" in the Danaidae). Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum. Many species show sexual dimorphism in the patterns of UV reflective patches. Colour vision may be widespread but has been demonstrated in only

1872-416: A leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue has been little researched but in the case of Pieris brassicae , it begins as a pale yellow granular secretion containing acidophilic proteins. This is viscous and darkens when exposed to air, becoming

1989-547: A longer lifespan of several months as adults. The thorax of the butterfly is devoted to locomotion. Each of the three thoracic segments has two legs (among nymphalids , the first pair is reduced and the insects walk on four legs). The second and third segments of the thorax bear the wings. The leading edges of the forewings have thick veins to strengthen them, and the hindwings are smaller and more rounded and have fewer stiffening veins. The forewings and hindwings are not hooked together ( as they are in moths ) but are coordinated by

2106-434: A maximum of 1,300 to 1,600 feet (400–500 m) wide to improve recolonization from neighboring areas. Depending on the available and surrounding habitat, managing for heterogeneity could mean implementing a wide range of techniques. Butterfly Butterflies are winged insects from the lepidopteran suborder Rhopalocera , characterized by large, often brightly coloured wings that often fold together when at rest, and

2223-452: A necessity for their successful establishment. Many butterflies, such as the painted lady , monarch, and several danaine migrate for long distances. These migrations take place over a number of generations and no single individual completes the whole trip. The eastern North American population of monarchs can travel thousands of miles south-west to overwintering sites in Mexico . There is

2340-612: A reason for lack of Karner blue butterfly presence on sites in New York. In addition, studies of Karner blue butterfly dispersal have led to recommended distance between patches of ≤980 feet (300 m) to allow for dispersal and management units no wider than 1,300 to 1,600 feet (400–500 m) to improve recolonization of treated areas. The presence of dispersal corridors may assist with Karner blue butterfly dispersal. Karner blue butterflies appear to disperse further in open habitats (see section "Timing of major life history events"). However, there

2457-658: A reverse migration in the spring. It has recently been shown that the British painted lady undertakes a 9,000-mile round trip in a series of steps by up to six successive generations, from tropical Africa to the Arctic Circle — almost double the length of the famous migrations undertaken by monarch. Spectacular large-scale migrations associated with the monsoon are seen in peninsular India. Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes . Butterflies navigate using

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2574-412: A segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular spinneret which is able to extrude silk. Caterpillars such as those in the genus Calpodes (family Hesperiidae) have a specialized tracheal system on the 8th segment that function as a primitive lung. Butterfly caterpillars have three pairs of true legs on the thoracic segments and up to six pairs of prolegs arising from

2691-454: A significant (p=0.009) positive correlation (r=0.2173) between patch area and number of disturbances (such as burning, vehicle traffic, and thinning and clearing), patches in a right-of-way in west-central Wisconsin occupied by Karner blue butterflies had a greater frequency of disturbance than unoccupied patches (p≤0.0125). In the same area, the abundance of the Karner blue butterfly spring brood

2808-435: A silken girdle may be spun to keep the pupa in a head-up position. Most of the tissues and cells of the larva are broken down inside the pupa, as the constituent material is rebuilt into the imago. The structure of the transforming insect is visible from the exterior, with the wings folded flat on the ventral surface and the two halves of the proboscis, with the antennae and the legs between them. The pupal transformation into

2925-665: A single generation, and a few in cold locations may take several years to pass through their entire life cycle. Butterflies are often polymorphic , and many species make use of camouflage , mimicry , and aposematism to evade their predators. Some, like the monarch and the painted lady , migrate over long distances. Many butterflies are attacked by parasites or parasitoids , including wasps , protozoans , flies , and other invertebrates, or are preyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents of pollination of some plants. Larvae of

3042-410: A time-compensated sun compass. They can see polarized light and therefore orient even in cloudy conditions. The polarized light near the ultraviolet spectrum appears to be particularly important. Many migratory butterflies live in semi-arid areas where breeding seasons are short. The life histories of their host plants also influence butterfly behaviour. Butterflies in their adult stage can live from

3159-541: A total of about 20,000 species. Traditionally, butterflies have been divided into the superfamilies Papilionoidea and the moth-like Hedyloidea . Recent work has discovered that Hedylidae, the only family within Hedyloidea, is nested within the Papilionoidea, meaning that Papilionoidea would be synonymous with Rhopalocera. The relationships between the rest of the 6 families are extremely well resolved, which

3276-470: A trend towards multivoltinism . Courtship is often aerial and often involves pheromones . Butterflies then land on the ground or on a perch to mate. Copulation takes place tail-to-tail and may last from minutes to hours. Simple photoreceptor cells located at the genitals are important for this and other adult behaviours. The male passes a spermatophore to the female; to reduce sperm competition, he may cover her with his scent, or in some species such as

3393-486: A tubular proboscis which is curled up at rest and expanded when needed to feed. The first and second maxillae bear palps which function as sensory organs. Some species have a reduced proboscis or maxillary palps and do not feed as adults. Many Heliconius butterflies also use their proboscis to feed on pollen; in these species only 20% of the amino acids used in reproduction come from larval feeding, which allow them to develop more quickly as caterpillars, and gives them

3510-745: A warmer year adults were 1st detected on 22 May. In addition, compared to the previous year the flight of the 2nd-brood during the hot year was shortened by 20 days. At Indiana Dunes National Park, evidence suggests that cool winters negatively impact 1st-brood populations and cool summers positively affect 2nd-brood populations. A review suggests that the cover provided by snowpack protects Karner blue butterfly eggs from hatching prematurely or being overheated by direct sunlight. Therefore, short periods of continuous snow cover due to site conditions or mild winters could result in decreased occurrence or smaller populations of Karner blue butterflies. Karner blue butterflies are dependent on heterogeneous habitat as it provides

3627-437: A water-insoluble, rubbery material which soon sets solid. Butterflies in the genus Agathymus do not fix their eggs to a leaf; instead, the newly laid eggs fall to the base of the plant. Eggs are almost invariably laid on plants. Each species of butterfly has its own host plant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of

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3744-422: A week to nearly a year depending on the species. Many species have long larval life stages while others can remain dormant in their pupal or egg stages and thereby survive winters. The Melissa Arctic ( Oeneis melissa ) overwinters twice as a caterpillar. Butterflies may have one or more broods per year. The number of generations per year varies from temperate to tropical regions with tropical regions showing

3861-587: A wide range of aspects, hydrology, microtopography as well canopy cover (0-90%) in conserving the Karner blue butterfly. However, Karner blue populations can persist in relatively homogenous habitats. For instance, a population has persisted at the Saratoga Airport in New York in open habitat maintained by regular mowing. More information on the habitat characteristics, environmental conditions, and/or geographic variability in habitat requirements that facilitate Karner blue butterfly's persistence in these areas

3978-450: Is above 27 °C (81 °F); when it is cool, they can position themselves to expose the underside of the wings to the sunlight to heat themselves up. If their body temperature reaches 40 °C (104 °F), they can orientate themselves with the folded wings edgewise to the sun. Basking is an activity which is more common in the cooler hours of the morning. Some species have evolved dark wingbases to help in gathering more heat and this

4095-570: Is commonly planted along roadsides. L. polyphyllus is not native to eastern North America, but has naturalized in areas in the upper Midwest and New England. L. polyphyllus has 11–17 leaflets that can reach 13 cm (5 in) in length, while L. perennis has 7–11 leaflets which only reach around 5 cm (2 in) in length. It is widespread in the eastern part of the USA (from Texas and Florida to Maine ) and Minnesota , Canada (southern Ontario , Newfoundland and Labrador ), and on

4212-458: Is derived from tiny structures called scales, each of which have their own pigments . In Heliconius butterflies, there are three types of scales: yellow/white, black, and red/orange/brown scales. Some mechanism of wing pattern formation are now being solved using genetic techniques. For instance, a gene called cortex determines the colour of scales: deleting cortex turned black and red scales yellow. Mutations, e.g. transposon insertions of

4329-619: Is discouraged due to their potential impacts on native food sources, especially wild lupine. Heterogeneity is an important feature of Karner blue butterfly habitat. Specific recommendations regarding the spatial arrangement of the various required habitat types and Karner blue butterfly habitat patches are discussed in Landscape effects. Creation and rotation of management units should incorporate this information as well as data on Karner blue butterfly movement (see section "Timing of major life history events"). Bidwell recommends management units be

4446-403: Is especially evident in alpine forms. As in many other insects, the lift generated by butterflies is more than can be accounted for by steady-state, non-transitory aerodynamics . Studies using Vanessa atalanta in a wind tunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture , vortices at the wing edge, rotational mechanisms and

4563-410: Is extruded and inserted into the female's vagina. A spermatophore is deposited in the female, following which the sperm make their way to a seminal receptacle where they are stored for later use. In both sexes, the genitalia are adorned with various spines, teeth, scales and bristles, which act to prevent the butterfly from mating with an insect of another species. After it emerges from its pupal stage,

4680-451: Is good, especially in some species in the blue/violet range. The antennae are composed of many segments and have clubbed tips (unlike moths that have tapering or feathery antennae). The sensory receptors are concentrated in the tips and can detect odours. Taste receptors are located on the palps and on the feet. The mouthparts are adapted to sucking and the mandibles are usually reduced in size or absent. The first maxillae are elongated into

4797-612: Is home to the world's largest population of Karner blues, which benefit from its vast area of savanna and extensive lupine. In 2003, the Canadian Species at Risk Act listed the Karner blue as being extirpated from Canada. In 2012, after an unusually hot and dry year, the Karner blue was also extirpated in the Indiana Dunes National Park . The male and female of this small (wingspan of about one inch) butterfly are different in appearance. The topside of

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4914-871: Is likely to influence Karner blue butterfly abundance. In a right-of-way in west-central Wisconsin, the frequency that Karner blue butterflies stayed in an area between recaptures was significantly (p<0.05) positively related to percent cover of nectar flowers such as Canadian horseweed (Conyza canadensis var. canadensis) and spotted beebalm ( Monarda punctata ). Preliminary results from a study in southeastern Minnesota suggest that openings where Karner blue butterflies were detected tended to have more flowering nectar species. In Michigan, more Karner blue butterflies were observed on sites with more nectaring plants, especially butterfly milkweed ( Asclepias tuberosa ). As discussed in Canopy cover, nectar species increase in open habitats. For instance, at Fort McCoy in west-central Wisconsin, all

5031-581: Is native to the Americas, but in the nineteenth century or before, spread across the world, and is now found in Australia, New Zealand, other parts of Oceania, and the Iberian Peninsula . It is not clear how it dispersed; adults may have been blown by the wind or larvae or pupae may have been accidentally transported by humans, but the presence of suitable host plants in their new environment was

5148-585: Is needed. Size of habitat patches can also influence Karner blue butterfly abundance (see section "Wild lupine"). At the Allegan State Game Area, wild lupine patches occupied by Karner blue butterflies were larger than unoccupied patches. This was also the case at Fort McCoy in Wisconsin. In addition to the effect of amount of wild lupine, a review suggests that it is easier to maintain Karner blue butterfly habitat in larger patches. Lane notes

5265-458: Is oval with a light hilum . The germination of this plant isn't too selective over day and night as both still occur equivalently. Researchers found that the seeds of L. perennis require scarification to germinate and ideal temperatures range from 24–29 °C (75–84 °F). Lupinus perennis is commonly mistaken for the Western species Lupinus polyphyllus (large-leaved lupine), which

5382-663: Is paid to implementing them at appropriate times and at intensities, scales, and frequencies that Karner blue butterflies can tolerate. For example, mowing can maintain open areas with little to no detrimental effect on Karner blue butterflies on sites where burning is impractical or in areas too small to support populations of Karner blue butterflies likely to survive a burn. On restored oak savanna sites in south-central Wisconsin, no significant (p=0.924) differences were detected between Karner blue butterfly densities on sites burned in summer, sites mowed in August, and control sites. Mowing with

5499-465: Is restricted to the males, and studies have suggested that the nutrients collected may be provided as a nuptial gift , along with the spermatophore, during mating. In hilltopping , males of some species seek hilltops and ridge tops, which they patrol in search for females. Since it usually occurs in species with low population density, it is assumed these landscape points are used as meeting places to find mates. Butterflies use their antennae to sense

5616-491: Is silky, without bractlets ; its upper labium with a protuberant basis, is integral or weakly emarginate, the lower one is integral, almost twice longer than upper. Floral bracts are styliform, shorter than the calyx, early falling. The corolla is three times longer than the calyx. The vexillum is shorter than the wings. The carina is weakly ciliate. Pods are yellow-grayish-brown, with straight lines, necklace-shaped, short and closely hirsute, easy shattered, with 5–6 seeds. Seed

5733-698: Is summarized in the below cladogram. Papilionidae [REDACTED] Hedylidae [REDACTED] Hesperiidae [REDACTED] Pieridae [REDACTED] Nymphalidae [REDACTED] Lycaenidae [REDACTED] Riodinidae [REDACTED] Butterfly adults are characterized by their four scale-covered wings, which give the Lepidoptera their name ( Ancient Greek λεπίς lepís, scale + πτερόν pterón, wing). These scales give butterfly wings their colour: they are pigmented with melanins that give them blacks and browns, as well as uric acid derivatives and flavones that give them yellows, but many of

5850-405: Is taken up by the gut, but there may also be large silk glands, and special glands which secrete distasteful or toxic substances. The developing wings are present in later stage instars and the gonads start development in the egg stage. When the larva is fully grown, hormones such as prothoracicotropic hormone (PTTH) are produced. At this point the larva stops feeding, and begins "wandering" in

5967-782: Is uncertainty regarding what constitutes a corridor or a barrier to dispersal. Creation of corridors with both lupine for larvae and nectar species for adults may be useful in connecting habitat patches. Karner blue butterflies have two broods per year, following wild lupine ( Lupinus perennis ) phenology quite closely. Eggs laid by Karner blue butterflies in late summer overwinter and hatch in mid- to late April. Development from egg through four larval instars and pupation takes from 25 to 60 days. The average lifespan of adult Karner blue butterflies has been reported at between 3 and 5 days. The first Karner blue butterfly flight generally occurs sometime between mid-May and mid-June, with males typically appearing earlier than females. First flight females lay

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6084-479: The Lycaenidae , form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions . Large blue ( Phengaris arion ) caterpillars trick Myrmica ants into taking them back to the ant colony where they feed on

6201-474: The Weis-Fogh ' clap-and-fling ' mechanism. Butterflies are able to change from one mode to another rapidly. Butterflies are threatened in their early stages by parasitoids and in all stages by predators, diseases and environmental factors. Braconid and other parasitic wasps lay their eggs in lepidopteran eggs or larvae and the wasps' parasitoid larvae devour their hosts, usually pupating inside or outside

6318-455: The ZW sex-determination system where females are the heterogametic sex (ZW) and males homogametic (ZZ). Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species. Of these, 775 are Nearctic ; 7,700 Neotropical ; 1,575 Palearctic ; 3,650 Afrotropical ; and 4,800 are distributed across the combined Oriental and Australian / Oceania regions. The monarch butterfly

6435-600: The non-coding DNA around the cortex gene can turn a black-winged butterfly into a butterfly with a yellow wing band. When the butterfly Bicyclus anynana is subjected to repeated inbreeding in the laboratory, there is a dramatic decrease in egg hatching. This severe inbreeding depression is considered to be likely due to a relatively high mutation rate to recessive alleles with substantial damaging effects and infrequent episodes of inbreeding in nature that might otherwise purge such mutations. Although B. anynana experiences inbreeding depression when forcibly inbred in

6552-1166: The Allegan State Game Area in Michigan were not detectably closer to ant hills than wild lupine without caterpillars. Due to effects of environmental variation and differing requirements among life stages, broods, and sexes, Karner blue butterflies require a mosaic of sun and shade. Adult Karner blue butterflies' preference for open, sunny areas has been well documented. Increased lupine and nectar abundance, higher temperatures allowing for longer activity periods, and ease of finding mates have been suggested as possible reasons for adult preference of open areas. Adult Karner blue butterfly females are more likely to use shaded habitats than males. Avoiding harassment by males and compromising between greater amounts of wild lupine in open areas and better quality of wild lupine in shaded areas (see below) have been suggested as possible reasons for increased occurrence of females in shade. Differences between broods have also been observed. In west-central Wisconsin, abundance of spring adults positively correlated with decreasing canopy cover, while correlation with summer adults

6669-540: The Apollos ( Parnassius ) plugs her genital opening to prevent her from mating again. The vast majority of butterflies have a four-stage life cycle: egg , larva (caterpillar), pupa (chrysalis) and imago (adult). In the genera Colias , Erebia , Euchloe , and Parnassius , a small number of species are known that reproduce semi-parthenogenetically ; when the female dies, a partially developed larva emerges from her abdomen. Butterfly eggs are protected by

6786-607: The United States in 1992. First considered a subspecies of Plebejus melissa , it was first identified and described by novelist Vladimir Nabokov . The name originates from Karner , New York (located half-way between Albany and Schenectady ) in the Albany Pine Bush , where it was first discovered. In the novel Pnin , Nabokov describes a score of Karner blues without naming them. Lupine blooms in late May. There are two generations of Karner blues per year,

6903-430: The abdomen, generally with short prolegs on segments 3–6 and 10; the three pairs of true legs on the thorax have five segments each. Many are well camouflaged; others are aposematic with bright colours and bristly projections containing toxic chemicals obtained from their food plants. The pupa or chrysalis, unlike that of moths, is not wrapped in a cocoon. Many butterflies are sexually dimorphic . Most butterflies have

7020-405: The abdominal segments. These prolegs have rings of tiny hooks called crochets that are engaged hydrostatically and help the caterpillar grip the substrate. The epidermis bears tufts of setae , the position and number of which help in identifying the species. There is also decoration in the form of hairs, wart-like protuberances, horn-like protuberances and spines. Internally, most of the body cavity

7137-502: The adult colour pattern are marked by changes in the expression of particular transcription factors in the early pupa. The reproductive stage of the insect is the winged adult or imago . The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a single epidermal cell. The head is small and dominated by the two large compound eyes . These are capable of distinguishing flower shapes or motion but cannot view distant objects clearly. Colour perception

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7254-467: The air for wind and scents. The antennae come in various shapes and colours; the hesperiids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensory organs known as sensillae . A butterfly's sense of taste is coordinated by chemoreceptors on the tarsi , or feet, which work only on contact, and are used to determine whether an egg-laying insect's offspring will be able to feed on

7371-520: The ant Formica nitidiventris was only seen in open (≤15% cover) areas, while another ant, Dolichoderus plagiatus was only observed in areas with a dense canopy (≥76% cover). Parasitoids in the genera Phaeogenes and Orthostigma were all seen in closed habitats, while 90% of damsel bugs ( Nabicula subcoleoptrata ), a potential predator, were observed in open areas. Closed habitats had insignificantly (p=0.116) more parasitoids on average than partial (16%–75% cover) and closed habitats. Nectar availability

7488-434: The ant eggs and larvae in a parasitic relationship. Caterpillars mature through a series of developmental stages known as instars . Near the end of each stage, the larva undergoes a process called apolysis , mediated by the release of a series of neurohormones . During this phase, the cuticle , a tough outer layer made of a mixture of chitin and specialized proteins , is released from the softer epidermis beneath, and

7605-658: The blues, greens, reds and iridescent colours are created by structural coloration produced by the micro-structures of the scales and hairs. As in all insects, the body is divided into three sections: the head, thorax , and abdomen . The thorax is composed of three segments, each with a pair of legs. In most families of butterfly the antennae are clubbed, unlike those of moths which may be threadlike or feathery. The long proboscis can be coiled when not in use for sipping nectar from flowers. Nearly all butterflies are diurnal , have relatively bright colours, and hold their wings vertically above their bodies when at rest, unlike

7722-472: The coasts of the Arctic Ocean, where it grows in sandy areas such as dunes and savannas . Lupinus perennis is used as foodplants by the caterpillars of several Lepidoptera . Among these are the clouded sulphur , eastern tailed blue , gray hairstreak , silvery blue , wild indigo duskywing , frosted elfin ( Callophrys irus ), the eastern Persius duskywing ( Erynnis persius persius ), and

7839-481: The density of wild lupine. Wild lupine are typically more abundant in open areas than in shade. Mildew infection of wild lupine may be increased in denser wild lupine patches. Lower mildew infection rates in shadier areas have been reported. However the implications of mildew infection on Karner blue butterfly are uncertain (see section "Wild lupine"). It has also been suggested that the low density of wild lupine in shadier habitats could provide better larval habitat due to

7956-569: The dependence of larvae, especially 2nd-brood larvae, on shaded habitat is the possibility of early senescence of wild lupine in open areas resulting in a lack of larval food. Shade-grown wild lupine being more nutritious, possibly due to nitrogen content limiting photosynthesis to a greater extent in open areas, was one of several explanations. The size of wild lupine has been positively associated with Karner blue butterfly larval length and amount of feeding damage. In addition, there may be shade-related effects on Karner blue butterflies that are related to

8073-441: The desiccated husk. Most wasps are very specific about their host species and some have been used as biological controls of pest butterflies like the large white butterfly . When the small cabbage white was accidentally introduced to New Zealand, it had no natural enemies. In order to control it, some pupae that had been parasitised by a chalcid wasp were imported, and natural control was thus regained. Some flies lay their eggs on

8190-545: The direct impact of varying environmental conditions over the course of the season on Karner blue butterflies (see section "Temperature") and the associated effects on wild lupine (see below). Larvae in shaded habitat apparently have an advantage over those in open areas. The increase in larvae in shaded habitats is likely due to effects of shade on wild lupine. Shade-grown wild lupine has been shown to provide higher quality larval resource than sun-grown lupine. Several reasons for this have been suggested. An often cited reason for

8307-474: The dominants included grasses (22.7%), northern dewberry ( Rubus flagellaris , 5.0%), other blackberries (4.8%), and sheep sorrel (4.3%). Several studies have found a positive relationship between measures of wild lupine and Karner blue butterfly abundance. However, available evidence suggests that senescent wild lupine is a poor larval food source. The effects of other phenological stages are more ambiguous. In west-central Wisconsin, Karner blue butterfly abundance

8424-419: The epidermis begins to form a new cuticle. At the end of each instar, the larva moults , the old cuticle splits and the new cuticle expands, rapidly hardening and developing pigment. Development of butterfly wing patterns begins by the last larval instar. Caterpillars have short antennae and several simple eyes . The mouthparts are adapted for chewing with powerful mandibles and a pair of maxillae, each with

8541-512: The family Fabaceae . The leaves are palmately compound with 7–11 leaflets arranged radially. Their stalks are numerous, erect, striated, and slightly pubescent . The leaflets are obovate , with a blunted apex or pointed spear, and sparsely pubescent. Petioles are longer than leaflets; stipules are very small. The inflorescence is long, sparsely flowered, sometimes almost verticillate. Flowers color can be white, blue, purple, or pink, but are most often blue or bluish purple. The calyx

8658-741: The family Hesperiidae (skippers). Molecular clock estimates suggest that butterflies originated sometime in the Late Cretaceous , but only significantly diversified during the Cenozoic, with one study suggesting a North American origin for the group. The oldest American butterfly is the Late Eocene Prodryas persephone from the Florissant Fossil Beds , approximately 34   million years old. Butterflies are divided into seven families that contain

8775-420: The first in late May to mid June, the second from mid-July to mid-August. Local conservation efforts, concentrating on replanting large areas of blue lupine which have been lost to development (and to fire suppression, which destroys the open, sandy habitat required by blue lupine), are having modest success at encouraging the butterfly's repopulation. The Necedah National Wildlife Refuge in central Wisconsin

8892-403: The food plant on which their larvae , known as caterpillars , will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in a chrysalis . When metamorphosis is complete, the pupal skin splits, the adult insect climbs out, expands its wings to dry, and flies off. Some butterflies, especially in the tropics, have several generations in a year, while others have

9009-409: The friction of their overlapping parts. The front two segments have a pair of spiracles which are used in respiration. The abdomen consists of ten segments and contains the gut and genital organs. The front eight segments have spiracles and the terminal segment is modified for reproduction. The male has a pair of clasping organs attached to a ring structure, and during copulation, a tubular structure

9126-555: The grass was growing. The earliest Lepidoptera fossils date to the Triassic - Jurassic boundary, around 200   million years ago. Butterflies evolved from moths, so while the butterflies are monophyletic (forming a single clade ), the moths are not. The oldest known butterfly is Protocoeliades kristenseni from the Palaeocene aged Fur Formation of Denmark, approximately 55   million years old, which belongs to

9243-545: The height of adjacent trees. According to a review, subpopulations in habitat patches of less than 0.25 ha (0.62 acres) are vulnerable to extirpation. Given their typically short dispersal distance, the spatial arrangement of habitat is important to the conservation of the Karner blue butterfly. Distance between the various types of habitat required by Karner blue butterflies is likely an influential landscape attribute. Lane suggests that suitable Karner blue butterfly habitat occurs in areas where open and closed canopies occur within

9360-440: The impacts of an insecticide on Karner blue butterflies and some herbicides on Karner blue butterflies as well as lupine and nectar species are available. According to the Karner blue butterfly recovery plan, management activities that can have a detrimental effect should be timed to allow at least 2 generations between repeat treatments and, if possible, critical subpopulations should be divided into discrete management units. Given

9477-402: The importance of shaded habitat led to a recommendation that 20% to 40% of Karner blue butterfly habitat provide shaded lupine and that 1/3 have >60% canopy cover with succession in these dense areas allowed to progress to complete closure. Maxwell also suggests avoiding any management that would result in habitat homogenization. The Karner blue butterfly recovery team mentions the importance of

9594-418: The increased search effort required by predators. This trade-off between lupine quality and quantity is another reason heterogeneous habitat is important for Karner blue butterfly. On sites in Wisconsin and Minnesota, canopy cover did not have a significant (p>0.06) effect on total numbers of ants, parasitoids, or predators. However, certain species did show trends across canopy cover categories. For instance

9711-1030: The laboratory it recovers within a few generation when allowed to breed freely. During mate selection, adult females do not innately avoid or learn to avoid siblings, implying that such detection may not be critical to reproductive fitness. Inbreeding may persist in B anynana because the probability of encountering close relatives is rare in nature; that is, movement ecology may mask the deleterious effect of inbreeding resulting in relaxation of selection for active inbreeding avoidance behaviors. Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen , tree sap, rotting fruit, dung, decaying flesh, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants. In general, they do not carry as much pollen load as bees , but they are capable of moving pollen over greater distances. Flower constancy has been observed for at least one species of butterfly. Adult butterflies consume only liquids, ingested through

9828-400: The major predators of Karner blue butterflies. The seven-spotted lady beetle is one of the few confirmed predators of Karner blue butterfly larvae. Paper wasps ( Polistes spp. ), spined soldier bugs , and ants ( Formica spp. ) have been observed removing larvae, and the ant Monomorium emarginatum has been seen chewing on Karner blue butterfly eggs. There is currently no explanation for

9945-506: The majority of moths which fly by night, are often cryptically coloured (well camouflaged), and either hold their wings flat (touching the surface on which the moth is standing) or fold them closely over their bodies. Some day-flying moths, such as the hummingbird hawk-moth , are exceptions to these rules. Butterfly larvae , caterpillars , have a hard ( sclerotised ) head with strong mandibles used for cutting their food, most often leaves. They have cylindrical bodies, with ten segments to

10062-1391: The male is silvery or dark blue with narrow black margins. The female is grayish brown, especially on the outer portions of the wings, to blue on the topside, with irregular bands of orange crescents inside the narrow black border. The underside of both sexes is gray with a continuous band of orange crescents along the edges of both wings and with scattered black spots circled with white. The Karner blue butterfly occurs in portions of eastern Minnesota, Wisconsin, Michigan, and New York. Reintroductions have been initiated in Ohio, Indiana, Illinois and New Hampshire. The Karner blue butterfly appears extirpated from Iowa, Pennsylvania, Massachusetts, Maine, and Ontario in Canada. Although Karner blue butterflies are characteristic of oak savannas ( Quercus spp. ) and pine barrens ( Pinus spp. ) habitats, they also occur in frequently disturbed areas such as rights-of-way, old fields, and road margins. In east-central New York, Karner blue butterflies occurred in 3 rights-of-way habitat types: wild lily-of-the-valley-starflower (Maianthemum canadensis-Trientalis borealis), sweetfern-whorled yellow loosestrife ( Comptonia peregrina-Lysimachia quadrifolia ), and blackberry-sheep sorrel ( Rubus spp. Rumex acetosella ). An index of Karner blue population size

10179-758: The most commonly reported parasitoids of Karner blue butterflies. A tachinid fly, Aplomya theclarum , has also been listed as a Karner blue butterfly parasite. Two wasps, one from the family Trichogrammatidae and another tentatively identified as a member of the family Eulophidae , are suspected to parasitize Karner blue butterfly eggs. Karner blue butterfly larvae benefit from a facultative , mutualistic relationship with several ant species. In pitch pine - bear oak ( Pinus rigida - Quercus ilicifolia ) habitat in New York , significantly more larvae tended by ants survived (67%) than untended larvae (38%). The 19 ant species tending Karner blue butterfly larvae were from

10296-401: The name is ancient, but modern Dutch and German use different words ( vlinder and Schmetterling ) and the common name often varies substantially between otherwise closely related languages. A possible source of the name is the bright yellow male of the brimstone ( Gonepteryx rhamni ); another is that butterflies were on the wing in meadows during the spring and summer butter season while

10413-413: The nectar of 41 different species in a single study in west-central Wisconsin. Broods differ significantly in the species used for nectaring, likely due to differences in their phenology. Several blackberries have been documented as food sources for the spring brood, while spotted beebalm, white sweetclover ( Melilotus alba ), and flowering spurge ( Euphorbia corollata ), are widely cited sources of nectar in

10530-463: The nectar species listed for Karner blue butterflies require either full or partial sun. In addition, treatments such as cutting with or without herbicides, herbicide application alone, or mowing at varying intervals resulted in a significant (p<0.01) increase in the number and cover of nectar species. See section "Food habits" for species that Karner blue butterflies use as nectar sources. Other habitat characteristics: Although Schweitzer asserts that

10647-424: The need to find a balance between having patches of different required habitats within the activity range of Karner blue butterflies and having open areas large enough that they do not become shaded too quickly. Canopy openings with diameters of at least 82 feet (25 m) were recommended based on research at Indiana Dunes National Park, and research in Wisconsin and Minnesota led to a recommended opening size of 1.5 times

10764-402: The occurrence of Karner blue butterflies in habitats of varying canopy cover. The lower temperatures occurring in partial and shaded habitats of Wisconsin and Minnesota meant that 1st-flight females only had access to these areas for a few hours a day. In contrast, these females could access open habitats an average of 10.5 hours a day. The percent of adults in habitats of varying canopy categories

10881-411: The outside of caterpillars and the newly hatched fly larvae bore their way through the skin and feed in a similar way to the parasitoid wasp larvae. Predators of butterflies include ants, spiders, wasps, and birds. Lupinus perennis Lupinus perennis (also wild perennial lupine , wild lupine , sundial lupine , blue lupine , Indian beet , or old maid's bonnets ) is a flowering plant in

10998-567: The presence of litter is important to Karner blue butterflies in some years, abundance in a right-of-way in west-central Wisconsin was negatively (p<0.05) related with average litter cover. In west-central Wisconsin, the amount of Karner blue butterfly larval feeding damage increased with grass cover. A review suggests that grass cover may provide roosting sites for Karner blue butterflies and that 5% cover of tall grass would most likely meet this need. Karner blue butterflies may be associated with disturbance. Although results may have been biased by

11115-664: The primary causes of the decline of the Karner blue butterfly. The main threats to Lupinus perennis are thought to be habitat loss, habitat fragmentation , and poor management. Currently it is considered "rare" in Pennsylvania , a species of special concern in Rhode Island , threatened in Iowa , Maryland , and New Hampshire ; it is endangered in Vermont , and is extirpated from Maine . Human development has eliminated

11232-529: The proboscis. They sip water from damp patches for hydration and feed on nectar from flowers, from which they obtain sugars for energy, and sodium and other minerals vital for reproduction. Several species of butterflies need more sodium than that provided by nectar and are attracted by sodium in salt; they sometimes land on people, attracted by the salt in human sweat. Some butterflies also visit dung and scavenge rotting fruit or carcasses to obtain minerals and nutrients. In many species, this mud-puddling behaviour

11349-431: The quest for a suitable pupation site, often the underside of a leaf or other concealed location. There it spins a button of silk which it uses to fasten its body to the surface and moults for a final time. While some caterpillars spin a cocoon to protect the pupa, most species do not. The naked pupa, often known as a chrysalis, usually hangs head down from the cremaster, a spiny pad at the posterior end, but in some species

11466-539: The rare and endangered Karner blue ( Plebejus samuelis ), whose caterpillars feed only on the lupine leaves. Leaves that have been fed on by Karner blues have distinctive transparent areas where the larvae have selectively eaten only the green, fleshy parts. The lupine has been declining in number and range since the Industrial Revolution . It is estimated that it has declined in number by about 90% since 1900. This decline has in turn been deemed one of

11583-660: The removal of larvae or chewing of eggs by some of the same ant species that tend larvae. Other potential predators of Karner blue butterfly larvae include green lacewings , soldier beetles , and damsel bugs . Predators of adult Karner blue butterflies include dragonflies , robber flies , ambush bugs (Phymatidae) and spiders, such as crab spiders . Assassin bugs (Reduviidae) are also likely predators of adult Karner blue butterflies. Other likely predators include white-tailed deer and birds. Incidental predation of Karner blue butterfly eggs, larvae, and pupae by white-tailed deer grazing on wild lupine can be substantial. Wasps are

11700-484: The same area that were occupied by Karner blue butterflies were also surrounded by a relatively low percentage of unoccupied patches. In addition, wild lupine patches in the Allegan State Game Area that were occupied by Karner blue butterflies were closer to other occupied patches than wild lupine patches without Karner blue butterflies. This led to the recommendation that habitat patches be within 230 feet (70 m) of each other. Isolation of habitat patches has been suggested as

11817-520: The second. Timing and size of both flights can exhibit substantial variation, depending on local weather conditions among other factors. A wide range of values related to Karner blue butterfly recruitment have been reported. Adult Karner blue sex ratios vary from 1 male to 1.44 females to 2 males to 1 female. Wild caught Karner blue butterfly females have been observed to lay from 7.7 to 83 eggs on average. Reported percentages of eggs that reach adulthood under controlled conditions vary from 21.4% to 75.2%. In

11934-408: The shortest observed larval durations and gained the most weight for the amount of lupine eaten. Prior to the heat and drought of 2012 that extirpated several populations in the southern parts of the species range, habitat loss through direct conversion to other land uses and through succession were considered the major causes of the decline of the Karner blue butterfly. Notably, two reviews summarized

12051-496: The status of Karner blue butterfly across its range. Recovery criteria were addressed, while Christenson and Lentz discussed lessons learned while developing a statewide Habitat Conservation Plan for Wisconsin. In 2023, the National Park Service published a 154-page report, using studies and recommendations that were motivated by population extirpations attributed to the 2012 drought and warmth that were regarded as

12168-509: The subfamilies Formicinae , Myrmicinae , and Dolichoderinae , which are the most common in the area. The species of ant is likely to influence the degree of benefit gained by Karner blue butterfly larvae. At sites in Wisconsin and Minnesota, ant tending rates increased significantly with increasing larval age. The Karner blue larvae also has fine hairs on it to protect itself. In a study of the effect of different experimental feeding treatments, Karner blue butterfly larvae tended by ants had one of

12285-710: The summer. Common cinquefoil ( Potentilla simplex ) is used as a nectar species for both broods. Karner blue butterflies' preferred nectar species may include butterfly weed in New York and Michigan and lyrate rockcress ( Arabis lyrata ), lanceleaf tickseed ( Coreopsis lanceolata ), white sweetclover, and northern dewberry ( Rubus flagellaris ) at Indiana Dunes National Park. Differences in nectar species used between male and female Karner blue butterflies and across locations have been reported. Other often mentioned nectar sources include New Jersey tea ( Ceanothus americanus ), wild lupine, goldenrods ( Solidago spp. ), and spotted knapweed ( Centaurea maculosa ). Spiders and many insects are

12402-441: The sun. The same trend was observed in 2nd-flight ovipositions. In cool temperatures 65% (n=11) of ovipositions occurred in the sun, while in hot temperatures only 40% (n=37) occurred in the sun. Temperature also influences Karner blue butterfly phenology and brood success. Weather had strong influence on Karner blue butterfly phenology at Fort McCoy in west-central Wisconsin. In a cool year the 2nd-brood flight began 6 June, while in

12519-511: The varied requirements of different Karner blue butterfly broods, sexes, and life stages over a wide range of environmental conditions. Research at Indiana Dunes National Park led to recommendations for canopy openings for adult males and nectaring of both sexes, as well as areas with 30-60% cover for ovipositing females. In the Allegan State Game Area, wild lupine patches occupied by Karner blue butterflies had higher density of edge habitats that unoccupied wild lupine patches. In west-central Wisconsin

12636-506: The vast majority of their eggs on wild lupine. These eggs develop into the adults of the second Karner blue butterfly flight, which generally occurs in July and August. Although always near a wild lupine plant, second brood females lay more eggs on grasses, other plants, and litter than 1st brood females. The 2nd flight is typically two to four times the size of the first flight. However, the first flight of Karner blue butterflies can be larger than

12753-420: The wide range of nectar species used by Karner blue butterfly (see section "Food habits"), planting several often used or preferred nectar species is favored over selecting 1 or 2 specific species. Species selection should attempt to provide nectar sources in both open and partially shaded conditions throughout the growing season despite yearly variation in environmental conditions. Planting nonnative nectar sources

12870-479: The wild, loss of eggs may be substantial. Karner blue butterflies do not typically move very far, with males usually moving further than females, with most studies showing average distances moved by individual butterflies of well under 1,000 feet (300 m). Wild lupine is the sole larval food source of the Karner blue butterfly. Adult Karner blue butterflies obtain nectar from several native and nonnative species. Karner blue butterflies have been reported feeding on

12987-526: Was grown in shade and was in seed. In addition, on sites in Wisconsin and Minnesota, the number of 1st flight oviposition sites on nonreproductive and reproductive wild lupine was similar. Many other factors may influence the quality of wild lupine ( Lupinus perennis ) as a Karner blue butterfly food resource. At Indiana Dunes National Park, wild lupine with Karner blue butterfly feeding damage had significantly larger leaves and longer and thicker stems than plants without feeding damage. In addition, larval length

13104-408: Was highest in the wild lily-of-the-valley-starflower type. In this habitat, mosses ( Bryophyta , 6.9%), wild lily-of-the-valley (4.4%), grasses (Poaceae, 4.4%), and starflower (2.1%) had the highest cover. Coverage in the sweetfern-whorled yellow loosestrife type was dominated by grasses (40.9%), sweetfern (12.1%), mosses (9.4%), and whorled yellow loosestrife (5.2%). In the blackberry-sheep sorrel type,

13221-401: Was negatively associated with the abundance of reproductive lupine and positively related with the frequency of immature wild lupine. However, a feeding trial found that Karner blue butterfly fed 1st year wild lupine had one of the lowest survival rates observed and significantly longer larval durations than larvae fed older wild lupine that did not flower, was flowering, had recently flowered, or

13338-424: Was positively associated with wild lupine height in central Wisconsin. Higher nitrogen concentrations in wild lupine leaves resulted in significantly shorter larval durations in a feeding trial. In 1995, in west-central Wisconsin, significantly more Karner blue butterfly larvae were observed in oak-pine barren plots where mildew infection was delayed compared to areas where wild lupine were infected earlier. However, in

13455-739: Was positively related to trails, and the percent of lupine stems with larval feeding damage was significantly (p<0.05) greater in areas disturbed by military vehicle traffic. Karner blue butterflies are directly affected by temperature. A laboratory investigation of temperature on Karner blue butterfly found that flight typically begins at 76 °F (24.6 °C) for females and 80 °F (26.4 °C) for males (p=0.25). Signs of heat stress started at 96 °F (35.6 °C) for females and 98 °F (36.8 °C) for males (p=0.25). In pine-oak barrens in Wisconsin, observation rates of Karner blue butterflies increased significantly (p=0.000) with increasing temperature. The effect of temperature can influence

13572-418: Was significantly (p=0.0001) influenced by temperature, with butterflies, especially females, increasing in partial (15-75%) and shaded (>75%) habitats with increasing temperature. In addition, 80% (n=45) of 1st-flight ovipositions when temperatures were cool (68-79 °F, 20-26 °C) were in sun, while only 40% (n=17) of 1st-flight ovipositions in hot temperatures (86-97 °F, 30-36 °C) occurred in

13689-493: Was very weak. At Indiana Dunes National Park, cover at late summer oviposition sites was significantly higher than at late spring oviposition sites. In Wisconsin and Minnesota, late spring oviposition sites occurred in partial and closed habitats significantly more than expected based on the number of nectar flowers and the cover and number of wild lupine stems, while summer oviposition sites did not differ significantly from expected. The different brood responses to shade may be due to

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