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Overview

Brief Summary

Biology

In Britain, one or more generations may be produced during a single year, depending on the climate (2). The female lays eggs singly on the upperside of leaves of the foodplants (3). The eggs hatch after around a week, and the caterpillars create a tent-like shelter of leaves spun together with silk, within which feeding takes place. They pupate inside these shelters, and the adults emerge after around two weeks (3). In 1997 it was first shown that adults are able to overwinter in Britain, although it is not known if adults are capable of breeding after overwintering (2). There is currently no evidence to suggest that this species migrates south at the end of the season (2). In some years, the migration of this species involves enormous numbers of individuals; in 1996, many millions of painted ladies arrived in Britain, and the event made the headlines at the time (2).
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North American Ecology (US and Canada)

Vanessa cardui, the painted lady, is resident to southern California, Arizona, New Mexico and Texas, south to Venezuela, the Bahamas and Antilles. It is the most widely distributed butterfly in the world, occurring also in Eruasia, Africa, Australia, New Zealand and elsewhere (Scott 1986). Hightly migratory, it migrates throughout the North America. Habitats are everywhere, mostly in open or disturbed areas. Host plants are mostly herbaceous, but rarely shrubs or trees, and include species from many families, especially preferring Compositae, and sometimes is a pest on Iowa soybeans (Leguminosae). Eggs are laid on the host plant singly. Individuals overwinter as adults, but only in mild winters or in the south. There are multiple flights all year in south Texas, Florida and California, with migrations many thousands of kilometers between spring to fall (Scott 1986).
  • Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
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Introduction

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Description

The beautiful painted lady butterfly is a long-distance migratory species, and is easily recognised by its orange and black patterns and white spots (1). The caterpillar reaches 2.8 cm in length, has a black body with white dots, yellowish-black spines, and a yellow line along each side (3).
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Comprehensive Description

General Description

There are two other similar Vanessa; the first, V. virginiensis, occurs only very rarely as a migrant in Alberta, and the Painted Lady can immediately be distinguished from virginiensis by the row of three to four smaller eyespots on the hindwing underside; virginiensis has only two, much larger spots. Compared to the West Coast Lady (V. annabella), cardui has a large white spot two-thirds up the leading edge of the forewing, which is orange in annabella; cardui is also larger.
Royal Alberta Museum page
 There are no named subspecies.
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Description

Size: 54-58 mm. Unmistakable.The forewings are brown. On the wing leading edge there is a big white spot. The wing is black and there is a chain of little white spots. There is a chain of big, black spots on the wing. The underside is a copy from upside, but there are some differences. The wing is grey. Nearby the body there is a pink area. The hind wings are brown. At the margin there are three chains of black spots. In the middle of the wing there is a black spot. The underside is brown and marmorate. At the margin there is a chain of little eyes. The body is brown. Sex differences: None. The caterpillar reaches 2.8 cm in length, has a black body with white dots, yellowish-black spines, and a yellow line along each side .

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Distribution

This butterfly is more widespread tha nany other species in the world; although it does not tolerate hard winter forsts, migrants have been found on every continent save Antarctica; recorded as far north as northern Greenland (Scott 1986).
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Geographic Range

This is one of the most common butterfly species in the world. The only places it doesn't live are on Antarctica and some remote islands. It even migrates to Hawaii and Iceland!

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native ); oceanic islands (Native )

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Geographic Range

The Painted Lady is found everywhere in the world except South America, the Arctic, and Australia.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native ); oceanic islands (Native )

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Records

102 records. Latest in 2007 (St Katherine)

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Distribution in Egypt

Widespread

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Global Distribution

Widespread (Holarctic). everywhere in the world except South America, the Arctic, and Australia.

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occurs (regularly, as a native taxon) in multiple nations

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National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Breeding

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Holarctic: in North America, Panama north to the southern United States. Migrates north to sub-Arctic North America, but dies off in winter. Established virtually worldwide; the most widespread species of butterfly in the world.

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Range

Every year this species migrates northwards from 'source populations' in North Africa, central Asia and the Middle East to Europe, including Britain and Ireland, where it breeds. Numbers peak in late summer. This species has a worldwide distribution, but is absent from South America. Some authorities claim that the race found in Australia is a separate species (2).
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Physical Description

Morphology

Physical Description

Eggs are pale green, placed one by one on the uppersides of leaves. The caterpillars vary in color, they are grayish-brown or darker, and darker at each end than in the middle. They have a yellow stripe down the back and along each side, and many spines on their back and sides. The pupa can also be different colors, metallic-greenish, or bluish-white, or brown. Adult Painted Lady Butterflies have speckled wings with brown, black, red and white markings. The upper side has more red and sometimes pink or orange, the underside is more brown and black. The forewings have a white bar, and the hindwing has a row of 5 tiny black dots. When the wings are folded, they appear camouflaged.

Other Physical Features: ectothermic ; bilateral symmetry

Sexual Dimorphism: sexes alike

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Physical Description

The Painted Lady has a pointed forewing which bears a distinct white bar. The hindwing has a submarginal row of 5 tiny black dots. The upperside of the freshly emerged butterfly is orange with rose-like overtones. The underside is a mottled gray, brown, and black.

Other Physical Features: ectothermic ; bilateral symmetry

Sexual Dimorphism: sexes alike

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Ecology

Habitat

A migrant that can be found in almost any habitat in years that it is common.
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Painted Lady Butterflies are found almost anywhere, but they prefer brightly lit and open environments like clover fields, flowery meadows and hilly country.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: tundra ; taiga ; desert or dune ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp ; bog

Other Habitat Features: suburban ; agricultural

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The Painted Lady is found almost anywhere, but they tend to inhabit brightly lighted and open environments. They prefer clover fields, flowery meadows and hilly country. Marshes, dunes, and thorn scrubs also attract the Painted Lady.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: tundra ; taiga ; desert or dune ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp ; bog

Other Habitat Features: suburban ; agricultural

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Occurs in any area with flowers.

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Comments: A migratory species that potentially might find and use any patch of foodplants in good years. Normal habitats when not migrating include deserts, any kind of dry open fields with thistles, pastures etc. Overwintering areas seem to be frost-free portions of southwestern deserts and Mexico. Habitats checked off do not include casual use areas like downtown cities and alpine tundra.

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Found in a huge variety of habitats. Adults seem to prefer open areas with good populations of thistles (Cirsium and Carduus spp.), which are used as foodplants for the caterpillars, although a very wide range of foodplants may be used, including common nettle (Urtica dioica) and viper's bugloss (Echium vulgare) (2).
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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Trophic Strategy

Many composites (Asteraceae) have been recorded as larval hosts, but thistles (Cirsium spp.) seem to be favoured in western Canada (Guppy & Shepard 2001).
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Food Habits

The caterpillars of this species prefer the leaves of plants in the daisy family (Compositaceae) especially thistles, but can eat many different kinds of plants. Adult painted ladies sip nectar from flowers, and sometimes take "honeydew" from aphids (See Aphids).

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Food Habits

The Painted lady consumes more than 100 different plants, some include thistles, Burdock, and Groundsel. The Larval foodplants are thistles and members of the families Asteraceae and Malvaceae.

Primary Diet: herbivore

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Host-plants: a wide range, especially Malva.

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Comments: Larvae most often on thistles but many others reported.

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Associations

Flowering Plants Visited by Vanessa cardui in Illinois

Vanessa cardui Linnaeus: Nymphalidae, Lepidoptera
(observations are from Robertson, Graenicher, Hilty, Reed, Macior, Broyles & Wyatt, and Wist; this butterfly is the Painted Lady)

Apiaceae: Eryngium yuccifolium sn (H); Asclepiadaceae: Asclepias exaltata [plup sn] (BW), Asclepias verticillata [plup sn] (H); Asteraceae: Aster lanceolatus sn (Rb), Aster novae-angliae sn (Rb), Aster pilosus sn (Rb), Aster salicifolius sn (Rb), Bidens aristosa sn fq (Rb), Boltonia asterioides sn (Rb), Chrysopsis villosa sn (Re), Cirsium arvense sn (Re), Cirsium discolor sn (Rb, H), Echinacea angustifolia sn (Ws), Echinacea purpurea sn (Rb, H), Eupatorium serotinum sn (Rb), Euthamia graminifolia sn (Gr), Helianthus grosseserratus sn fq (Rb), Heliopsis helianthoides sn (Re), Liatris aspera sn (H, Re), Liatris pycnostachya sn (H), Silphium integrifolium sn (H), Silphium perfoliatum sn (Rb), Solidago canadensis sn (Rb), Solidago speciosa sn (Re); Campanulaceae: Campanulastrum americanum sn (Rb); Fabaceae: Trifolium pratense sn (Rb, Re); Lamiaceae: Agastache foeniculum sn (Re), Blephilia hirsuta sn (Rb), Monarda fistulosa sn (Rb), Pycnanthemum tenuifolium sn (Rb); Lythraceae: Lythrum alatum sn (Rb); Ranunculaceae: Delphinium tricorne sn np (Mc); Rubiaceae: Cephalanthus occidentalis sn (Rb); Scrophulariaceae: Linaria vulgaris sn np (Rb); Verbenaceae: Verbena stricta sn (Re)

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Animal / parasitoid / endoparasitoid
larva of Phryxe vulgaris is endoparasitoid of larva of Cynthia cardui

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Predation

Adult painted ladies' main defenses are flight and camouflage. The caterpillars hide in small silk nests on top of leaves, and may have chemical defenses, but this is uncertain.

Known Predators:

  • wasps (Hymenoptera)
  • spiders (Araneae)
  • ants (Formicidae)
  • birds (Aves)

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Predation

Adult painted ladies' main defenses are flight and camouflage. The caterpillars hide in small silk nests on top of leaves, and may have chemical defenses, but this is uncertain.

Known Predators:

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Known predators

Vanessa cardui is prey of:
Aves
Araneae
Hymenoptera
Formicidae

This list may not be complete but is based on published studies.
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

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Global Abundance

10,000 to >1,000,000 individuals

Comments: Population fluctuates greatly.

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Life History and Behavior

Behavior

Adults feed flower nectar and sometimes aphid honeydew (Scott, 1986).
  • Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
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Cyclicity

Can be found throughout the season, from late April into October.
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Life Cycle

The light green cylindrical eggs have vertical ribs. Mature larvae are grey-brown with fine yellow and black markings and yellow branched spines (Guppy & Shepard 2001). Larvae live in silken leaf-nests on the hostplant (Layberry et al. 1998). This species is rare or absent in Alberta in most years, but in years that populations build up in the southern United States, worn and often tattered migrants appear in May to lay their eggs, having flown thousands of miles. The offspring of these migrants emerge from July onwards. Since it is rare or absent in most years in Alberta, it is thought that Painted Ladies cannot overwinter here, and re-colonize in good migrant years. It is not known if summer brood individuals attempt a southward return migration in the fall here; they may in the southwestern US (Scott 1986), but there is no evidence they do so in British Columbia (Guppy & Shepard 2001).  
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Development

Females lay eggs on the plants their babies will eat. The caterpillars that hatch out feed continuously and molt several times. After a few weeks they transform into a pupa, go through a complete metamorphosis, and emerge as an adult butterfly. The timing of this depends on the climate, the warmer it is the faster they grow.

Development - Life Cycle: metamorphosis

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The eggs are laid singly and hatched after about a week. The larvae create a tent-like shelter of leaves spun together with silk, within which feeding takes place, and pupate inside these shelters. The adults emerge after about two weeks and it has several generations per year.

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Development

Females lay eggs on the plants their babies will eat. The caterpillars that hatch out feed continuously and molt several times. After a few weeks they transform into a pupa, go through a complete metamorphosis, and emerge as an adult butterfly. The timing of this depends on the climate, the warmer it is the faster they grow.

Development - Life Cycle: metamorphosis

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The Flight Period

February-November

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Life Expectancy

Lifespan/Longevity

Lifespan depends on the climate, but is probably never more than one winter. Only adults survive through winter, and even then only in mild climates.

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Lifespan/Longevity

Lifespan depends on the climate, but is probably never more than one winter. Only adults survive through winter, and even then only in mild climates.

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Reproduction

Mating System: polygynous

In the temperate zone, reproductive behavior stops in the fall, but it may go on year-round in warmer climates.

Breeding season: All year in the tropics, spring and summer in cooler climates.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; sexual ; fertilization (Internal ); oviparous

Parental Investment: no parental involvement

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Mating System: polygynous

A green, barrel-shaped egg is laid singly on a host plant. The color of the larva varies from chartreuse with black marbling to a purple with a yellow hue.

Breeding season: All year in the tropics, spring and summer in cooler climates.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; sexual ; fertilization (Internal ); oviparous

Parental Investment: no parental involvement

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Vanessa cardui

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 18 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

ACATTATATTTTATTTTCGGAATTTGAGCAGGAATAGTAGGAACTTCACTT---AGTTTATTAATTCGAACTGAATTAGGTAATCCAGGATCTTTAATTGGAGAT---GATCAAATTTATAATACAATTGTTACAGCTCATGCTTTTATTATAATTTTTTTCATAGTTATACCAATTATAATTGGAGGATTTGGTAATTGATTAGTTCCTTTAATA---TTAGGAGCTCCTGATATAGCCTTTCCACGTATAAATAATATAAGATTTTGACTTTTACCCCCGTCACTAATATTATTAATTTCTAGTAGAATTGTCGAAAACGGAGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCACACAGAGGATCATCTGTAGATTTA---GCAATTTTTTCCCTTCATTTAGCTGGTATTTCATCAATTCTAGGAGCAATTAACTTTATTACAACTATTATTAATATACGGGTTAATAGTATATCCTTTGATCAAATACCTTTATTTGTTTGAGCTGTGGGTATTACAGCATTACTTTTATTACTTTCTTTACCTGTTTTAGCTGGG---GCTATTACTATACTTTTAACAGATCGAAATATTAATACATCATTTTTCGATCCAGCAGGAGGAGGAGATCCAATTCTTTATCAACATTTATTTNNNNNNNNNNNNNNNNNNNNNNNNNNNATTTTAATTTTACCAGGTTTCGGAATAATTTCACACATTATTTCTCAAGAAAGAGGAAAAAAG---GAAACCTTTGGATGTTTAGGAATAATTTATGCTATAATAGCAATTGGATTATTAGGATTTATTGTATGAGCACATCATATATTTACAGTAGGTATAGATATTGATACTCGAGCTTATTTTACTTCAGCAACTATAATTATTGCTGTACCCACAGGAATTAAAATTTTTAGTTGATTA---GCAACATTACATGGAACT---CAAATTAATTATAGACCTTCTATATTATGAAGATTAGGATTTATTTTTTTATTTACAGTAGGAGGATTAACAGGAGTAATTTTAGCTAATTCATCTATTGATATTACTCTTCATGATACTTATTATGTTGTAGCTCATTTCCATTATGTA---TTATCTATAGGAGCTGTATTTGCTATTTTAGGTGGATTTATTCATTGATATCCTTTATTTACAGGATTAATAATAAATAACTATTTATTAAAAATTCAATTTATTTCAATATTTATTGGAGTTAATTTAACATTTTTCCCTCAACATTTTTTAGGATTAGCTGGTATACCTCGA---CGATACTCAGATTATCCAGATAGATTTATA---TCATGAAATATTATTTCATCTTTTGGATCTTATATTTCCTTATTATCTATAATAATAATTATTATTATTATTTGAGAATCTATAATTAATCAACGAATTATT---TTATTTTCATTAAATATACCTTCTTCT
-- end --

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Statistics of barcoding coverage: Vanessa cardui

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 19
Specimens with Barcodes: 117
Species With Barcodes: 1
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Conservation

Conservation Status

Not of concern.
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These are such common butterflies that they need no special conservation efforts.

IUCN Red List of Threatened Species: no special status

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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These are such common butterflies that they need no special conservation efforts.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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Status in Egypt

Migrant

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IUCN

Not Assessed (not resident in Egypt)

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Abundance

Abundant

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National NatureServe Conservation Status

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N5 - Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: Abundant migratory. Occurs virtually worldwide.

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Status

Not listed under any conservation designations or legislation (2).
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Threats

Degree of Threat: D : Unthreatened throughout its range, communities may be threatened in minor portions of the range or degree of variation falls within natural variation

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The appearance of this species in Britain, and indeed its survival as a whole, relies on the persistence of suitable habitat in its source populations, outside of Europe (2).
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Management

Global Protection: Many to very many (13 to >40) occurrences appropriately protected and managed

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Conservation

No conservation measures are in place for this species.
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Wikipedia

Vanessa cardui

Vanessa cardui is a well-known colourful butterfly, known as the Painted Lady, or in North America as the Cosmopolitan. This butterfly has a strange pattern of flying in a sort of screw shape.

Contents

Distribution

V. cardui is one of the most widespread of all butterflies, found on every continent except Antarctica and South America.[1] In Australia, V. cardui has a limited range around Bunbury, Fremantle, and Rottnest Island. However, its close relative, the Australian Painted Lady (Vanessa kershawi, sometimes considered a subspecies) ranges over half the continent. Other closely related species are the American Painted Lady (Vanessa virginiensis) and the West Coast Lady (Vanessa annabella).

Migration

Vanessa cardui occurs in any temperate zone, including mountains in the tropics. The species is resident only in warmer areas, but migrates in spring, and sometimes again in autumn. It migrates from North Africa and the Mediterranean to Britain in May and June, but for decades natualists have debated whether the offspring of these immigrants ever make a southwards return migration.[2] Recent research suggests that British painted ladies do undertake an autumn migration.[2] Using an entomological radar, scientists at Rothamsted Research provided evidence that autumn migrations take place at high altitude, which could be why these migrations are seldom witnessed.[2]

Relationship with humans

Vanessa cardui butterflies are raised in many preschool and elementary classrooms to demonstrate the life cycle of a butterfly. Naturally, this is one reason they are so popular amongst children. They are also often found in science fair projects.

Life cycle with notes for rearing in classrooms

As these animals are cold blooded and their life cycle does not depend on a certain number of day/night cycles, temperature can greatly effect the times presented here.

At 90 °F (32 °C) the entire life cycles will take roughly 16 days. At 65 °F (18 °C) the life cycles will take months. At such extreme temperatures one can expect some deaths. At room temperatures the egg takes three to five days to hatch. The eggs are tiny, as tiny as a sugar crystal. They are green and ribbed and can be observed best with a magnifying glass. It is possible to view the cap at the top of the egg where the caterpillar will emerge.

The embryo can be viewed growing inside the egg using a hand lens or dissecting scope. A high powered dissecting scope allows for watching hatching quite clearly. If eggs turn deep green, or become dented and wrinkled, the eggs do not contain living embryos. Just before hatching the embryos fill the whole egg and make the eggs look black or brown. As protection against disease, newly laid eggs may be knocked off the leaf, or left attached to the leaf, and dipped in dilute household bleach solution (1 part household bleach to 200 parts water) for 1–2 minutes and swished about. Afterwards, the eggs are left on a paper towel to dry. This will kill disease on the surface of the eggs and increase caterpillar survival.

The caterpillars will emerge as small and black and will begin to eat immediately. As they grow they will shed their skins three times, called instars. At each instar the caterpillar will need much more food as it has expanded in size. It will also become more spiky. These spikes do not contain poison and are not sharp. The moulted skin appears as a black speck, what looks like dirt, near the caterpillar. Many people believe this to be the excretion of the caterpillar. Occasionally the moult will look like an entire, dead, caterpillar, as snake's skin does. If under stress they will sometimes shed into a fifth instar, which is a very large caterpillar. A fifth instar is a sign that care is incorrect in some way, typically due to diet.

The four instars take 7–11 days to turn into a chrysalis. The caterpillar will spin a patch of silk and attach its hind end to the silk. At this point it begins changing internally, forming a "j" shape. Once the caterpillar forms a J, it should not be disturbed as it can no longer reattach itself to the silk pad. A fallen "J" caterpillar can be laid on its side on a flat piece of cotton and may shed successfully. The chrysalis is very soft at first and will dent if resting on a hard surface. After hardening, the chrysalis will crack if dropped or struck. The chrysalis can be dark or light colored depending on conditions during development of the caterpillar. It takes 7–11 days for the chrysalis to turn into a butterfly.

When emerging from the chrysalis the butterfly pumps its wings with fluid to expand them. This happens within a few minutes of emerging or cannot happen at all. Once the wings are expanded they are still soft for up to a day. Initially the butterfly prefers not to move as its wings harden, but after the wings harden for a few hours the painted lady will become incredibly sensitive to movement and will damage its still soft wings when frightened. It is best to wait a full day after emergence from the chrysalis to handle a painted lady. Its wing span is 2 inches (50 mm).

Host plants

Larvae feed on Asteraceae spp., including Cirsium, Carduus,Centaurea, Arctium, Helianthus, and Artemisia spp.[1][3]

The painted lady uses over 300 recorded host plants according to the HOSTS database.[4] For raising in the classroom one need only sprout a bed of black oil sunflower seeds, as are used for bird seed. The caterpillars will eat the true adult leave (not the sprouts) and in this way one may inexpensively produce many host plants. Soak the seeds for eight hours in 10% diluted household bleach (1 pt bleach, 9 pts water) to ensure a disease free, even sprouting. Place the seeds upon the surface of the soil and keep moist until they grow roots and can be watered normally.

See also

References

  1. ^ a b "Painted Lady". A-Z of Butterflies. Butterfly Conservation. Retrieved 22 October 2012. 
  2. ^ a b c "Butterfly Conservation: Secrets of Painted Lady migration unveiled". BirdGuides Ltd. Retrieved 22 October 2012. 
  3. ^ Vanessa cardui, Butterflies of Canada
  4. ^ "Vanessa cardui". HOSTS - a Database of the World's Lepidopteran Hostplants. The Natural History Museum. Retrieved 22 October 2012. 

Further reading

  • Opler, Paul A.; Wright, Amy Bartlett (1999). A Field Guide to Western Butterflies. Peterson Field Guides. Boston: Holt McDougal. ISBN 978-0-547-35114-8. 
  • Chapman, Jason W.; Nesbit, Rebecca L.; Burgin, Laura E.; Reynolds, Don R.; Smith, Alan D.; Middleton, Douglas R.; Hill, Jane K. (2010). "Flight Orientation Behaviors Promote Optimal Migration Trajectories in High-Flying Insects". Science 327 (5966): 682–5. doi:10.1126/science.1182990. PMID 20133570. 
  • Nesbit, R.L.; Hill, J.K.; Woiwod, I.P.; Sivell, D.; Bensusan, K.J.; Chapman, J.W. (2009). "Seasonally adaptive migratory headings mediated by a sun compass in the painted lady butterfly, Vanessa cardui". Animal Behaviour 78 (5): 1119–25. doi:10.1016/j.anbehav.2009.07.039. 
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