Vanessa cardui

Last updated

Vanessa cardui
0 Belle-dame (Vanessa cardui) - Echinacea purpurea - Havre (3).jpg
Upperside
Butterfly August 2008-3.jpg
Underside
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Nymphalidae
Genus: Vanessa
Species:
V. cardui
Binomial name
Vanessa cardui
Synonyms
  • Papilio carduiLinnaeus, 1758

Vanessa cardui is the most widespread of all butterfly species. It is commonly called the painted lady, [2] [3] or formerly in North America the cosmopolitan. [4] [5]

Contents

Description

Wing scales. Krylo babochki repeinitsy.jpg
Wing scales.

Male and female. Upperside. Ground-colour reddish-ochreous, basal areas olivescent-ochreous-brown; cilia black, alternated with white, Forewing with an outwardly-oblique black irregular-shaped broken band crossing from middle of the cell to the disc above the submedian vein; the apical area from end of cell and the exterior border also black; before the apex is a short white outwardly-oblique streak and a curved row of four round spots, the second and third being small; a marginal pale lunular line with its upper portion most defined and whitish. Hind-wing with a blackish patch from the costal vein across end of cell, a partly confluent recurved discal band, a submarginal row of lunules, and then a marginal row of somewhat scutiform spots; between the discal band and submarginal lunules is a row of five round black spots, which in some examples show a pale and dark outer ring. Underside. Forewing brighter reddish-ochreous, the apical area and outer margin much paler, the apex being olivescent ochreous-brown; discal irregular band as above, subapical white streak, row of spots and marginal lunules distinct; base of wing and interspace before end of cell white. Hindwing transversely-marbled with olivescent ochreous-brown and speckled with black scales; crossed by basal and discal sinuous whitish or pale fascia and intersected by white veins; an outer-discal row of five ocelli, the upper one smallest and usually imperfect, the second and fifth the largest, the fourth with black centre speckled with blue and ringed with yellow, and the second and fifth also with an outer black ring; submarginal lunules purpurescent-grey, bordered by a whitish fascia; outer margin ochreous. Body olivescent ochreous-brown, abdomen with ochreous bands; palpi blackish above, white beneath; body beneath and legs greyish-white; antennae black above, tip and beneath reddish.

Distribution

V. cardui is one of the most widespread of all butterflies, found on every continent except Antarctica and South America. [2] [3] [6] In Australia, V. cardui has a limited range around Bunbury, Fremantle, and Rottnest Island. However, its close relative, the Australian painted lady (V. kershawi, sometimes considered a subspecies) ranges over half the continent. Other closely related species are the American painted lady (V. virginiensis) and the West Coast lady (V. annabella).

Ecology and behavior

Food sources and host plants

Larvae feed on Asteraceae species, including Cirsium , Carduus , Centaurea , Arctium , Onopordum , Helianthus , and Artemisia . [7] [8]

The painted lady uses over 300 recorded host plants according to the HOSTS database. [9]

Adult butterflies feed on flower nectar and aphid honeydew. [10]

Migration

Vanessa cardui spring migration between North Africa and Europe Vanessa cardui migration in Europe-blank map.svg
Vanessa cardui spring migration between North Africa and Europe

V. 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 and Europe in May and June, [11] occasionally reaching Iceland, [12] and from the Red Sea basin, via Israel and Cyprus, to Turkey in March and April. [13] [14] The occasional autumn migration made by V. cardui is likely for the inspection of resource changes; it consists of a round trip from Europe to Africa. [15]

For decades, naturalists have debated whether the offspring of these immigrants ever make a southwards return migration. [16] Research suggests that British painted ladies do undertake an autumn migration, making 14,500 km (9,000 mi) round trip from tropical Africa to the Arctic Circle in a series of steps by up to six successive generations. [16] The Radar Entomology Unit at Rothamsted Research provided evidence that autumn migrations take place at high altitude, which explains why these migrations are seldom witnessed. [16] In recent years, thanks to the activity of The Worldwide Painted Lady Migration citizen science project, led by the Barcelona-based Institute of Evolutionary Biology (Catalan: Institut de Biologia Evolutiva), [17] the huge range of migration has begun to be revealed. For example, some butterflies migrated from Iceland to the Sahara desert, and even further south. [14]

V. cardui is known for its distinct migratory behaviour. In California, they are usually seen flying from north to north-west. These migrations appear to be partially initiated by heavy winter rains in the desert where rainfall controls the growth of larval food plants. [18] In March 2019, after heavy rain produced an abundance of vegetation in the deserts, Southern California saw these butterflies migrating by the millions across the state. [19]

Similarly, heavier than usual rain during the 2018-2019 winter seems to have been the cause of the extraordinarily large migration observed in Israel at the end of March, estimated at a billion individual butterflies. [20] Painted lady migration patterns are highly erratic and they do not migrate every year. [21] Some evidence suggests that global climatic events, such as el Niño, may affect the migratory behaviour of the painted lady butterflies, causing large-scale migrations. [22] The first noticeable wave of migration in eastern Ukraine was noted in the 20s of April 2019. From May 15, numbers began to grow and it was possible to observe hundreds of this species in the Kharkiv region of Ukraine, including in the city streets of Kharkiv. [23]

Based on experimental data, the painted lady's migration pattern in northern Europe apparently does not follow a strict north-west heading. The range of headings suggests that migrating butterflies may adjust their migration patterns in response to local topographical features and weather, such as strong wind patterns. Laboratory-raised autumn-generation painted lady butterflies were able to distinguish a southern orientation for a return migration path. According to the same laboratory-based study, when butterflies were isolated from the sun, they were unable to orient themselves in a specific direction, opposed to those that did have access to the sun. This suggests that V. cardui requires a direct view of the sky, implying the use of a solar compass to orient its migratory direction and maintain a straight flight path. [24]

A 2024 Nature Communications article provided the first evidence that the painted lady, or any insect, had traveled across an ocean. Specimens were captured on a beach in French Guiana, outside the painted lady's natural habitat. Pollen grains from the butterflies' bodies matched species of West African shrubs that flowered at the same time of year. The researchers also analyzed the butterflies' genomes and used isotope tracing to confirm that they were born in Europe or Africa. Finally, the study found that the trade wind conditions from Africa to South America were "exceptionally favorable" at that time, which would have allowed the butterflies to be propelled over 2,600 miles (4,200 km)—one of the longest journeys of an insect ever recorded. [25]

Behavior

Groups of two to eight painted lady butterflies have been observed to fly in circles around each other for about one to five seconds before separating, symbolizing courtship. Groups of butterflies usually will not fly more than 4.5 m (15 ft) away from the starting point. [26] To establish and defend their territories, adult males perch in the late afternoon in areas where females are most likely to appear. [27] Once the male spots a female of the same species, he begins pursuit of her. If the foreign butterfly is a male, the original male will give chase, flying vertically for a few feet before returning to his perch. [28]

V. cardui establishes territories within areas sheltered by hedgerows. [29] Vanessa cardui tend to inhabit sunny, brightly lit, open environments and are often attracted to open areas of flowers and clovers. [27] Adults spend time in small depressions in the ground on overcast days. [29]

Mating

V. cardui displays a unique system of continuous mating, throughout all seasons, including the winter. This may be attributed to its migratory patterns, thus significantly affecting its mating behaviour. During European migrations, the butterflies immediately begin to mate and lay eggs upon arrival in the Mediterranean in the spring, starting in late May. [30] In the United States, painted lady butterflies migrating towards the north experience poor mating conditions, and many butterflies have limited breeding capabilities. [31] The "local adult generation" develops during this time, roughly from the middle of May through early June in conjunction with the butterfly progression throughout their flight. [30]

During its migratory process, these painted lady butterflies start breeding, and reproduce entirely throughout their migration. [32] Scientists have not been able to find evidence of their overwintering; this may be because they migrate to warmer locations to survive and reproduce. [31] Female painted lady butterflies may suspend their flight temporarily when they are "ready to oviposit"; [33] this allows them the opportunity to continually reproduce throughout their migrations. Because these butterflies are constantly migrating, male butterflies are thought to lack consistent territory. Instead of requiring territory to mate with females and developing evolutionary behaviour to defend this territory, the mating butterflies appear to establish a particular "time and place" in certain locations that they find to be suitable for reproduction. More specifically, they locate certain perches, hilltops, forest-meadow edges, or other landmarks where they will stay until, presumably, a female arrives to mate.

Equally important for the reproduction of the painted lady butterflies is the males' exhibition of polygynous mating behaviour, in which they often mate with more than one female. [34] This is important for painted lady butterflies because the benefits may supersede the costs of polygyny [35] since no permanent breeding ground is used. Upon mating, which typically occurs in the afternoon, female painted lady butterflies lay eggs one by one in their desired breeding locations. The variety of eclosion locations ultimately dictates the male painted lady behaviour. [36]

Female painted lady butterflies have been observed to have a relatively "high biotic potential", meaning they each produce large numbers of offspring. This perpetual influx of reproduction may be a reason why these painted lady butterflies have propagated so successfully. One interesting aspect that scientists have observed is that these butterflies like to fly towards rain. Further studies have suggested that the large amounts of rainfall may somehow "activate more eggs or induce better larval development". [37] Inhabited locations begin to observe a large influx of new generations of painted lady butterflies in the fall, particularly in September and October. Their reproductive success declines relatively throughout the winter, primarily through November. [30] However, they still continue to reproduce—an aspect of butterfly behaviour that is quite unique. Scientists hypothesize that these extensive migratory patterns help the painted lady butterflies find suitable conditions for breeding, thus offering a possible reason as to why these butterflies mate continuously.

Oviposition

Females oviposit on plants with nectar immediately available for the adults even if it leads to high mortality of the larvae. This lack of discrimination indicates they do not take into account volatile chemicals released from potential host plants when searching for oviposition choices. [38]

The availability of adult resources dictates a preference for specific areas of flowers. Flowers with more available nectar result in a larger number of eggs deposited on the plants. This reinforces the idea that the painted lady butterfly does not discriminate host plants and chooses mainly on the availability of adult food sources even if it increases the mortality rate of the offspring. The data also suggest that the painted lady butterfly favors quantity of offspring over quality. [39]

Defence mechanisms

The main defence mechanisms of painted lady butterflies include flight and camouflage. The caterpillars hide in small silk nests on top of leaves from their main predators that include wasps, spiders, ants, and birds. [27]

Vision

Painted lady butterflies have a visual system that resembles that of a honey bee. Adult V. cardui eyes contain ultraviolet, blue, and green opsins. [40] Unlike other butterflies, such as the monarch or red postman butterflies, painted ladies lack red receptors, which means that they are not sensitive to red light. [41] Behavioral studies on the related species, Vanessa atalanta , have demonstrated that V. atalanta cannot distinguish yellow light from orange light or orange light from red light. [42]

Human interaction

Vanessa cardui and other painted lady species are bred in schools for educational purposes and used for butterfly releases at hospices, memorial events, and weddings. [43]

Related Research Articles

<span class="mw-page-title-main">Butterfly</span> Group of insects in the order Lepidoptera

Butterflies are winged insects from the lepidopteran suborder Rhopalocera, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The group comprises the superfamilies Hedyloidea and Papilionoidea. The oldest butterfly fossils have been dated to the Paleocene, about 56 million years ago, though they likely originated in the Late Cretaceous, about 101 million years ago.

<span class="mw-page-title-main">Lepidoptera</span> Order of insects including moths and butterflies

Lepidoptera or lepidopterans is an order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organisms, making it the second largest insect order with 126 families and 46 superfamilies, and one of the most widespread and widely recognizable insect orders in the world.

<i>Cynthia</i> (butterfly) Subgenus of insects

Cynthia is a group of colourful butterfly species that used to be considered a subgenus of the genus Vanessa, in the family Nymphalidae. Before that, it was first described as a genus. Nowadays, this group is not considered a valid taxon anymore, because it is paraphyletic. The name CynthiaFabricius, 1807 is now a junior subjective synonym of VanessaFabricius, 1807.

<i>Vanessa atalanta</i> Species of butterfly

Vanessa atalanta, the red admiral or, previously, the red admirable, is a well-characterized, medium-sized butterfly with black wings, red bands, and white spots. It has a wingspan of about 2 inches (5 cm). It was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. The red admiral is widely distributed across temperate regions of North Africa, the Americas, Europe, Asia, and the Caribbean. It resides in warmer areas, but migrates north in spring and sometimes again in autumn. Typically found in moist woodlands, the red admiral caterpillar's primary host plant is the stinging nettle ; it can also be found on the false nettle. The adult butterfly drinks from flowering plants like Buddleia and overripe fruit. Red admirals are territorial; females will only mate with males that hold territory. Males with superior flight abilities are more likely to successfully court females. It is known as an unusually calm butterfly, often allowing observation at a very close distance before flying away, also landing on and using humans as perches.

<span class="mw-page-title-main">Small tortoiseshell</span> Species of butterfly in the family Nymphalidae

The small tortoiseshell is a colourful Eurasian butterfly in the family Nymphalidae. Adults feed on nectar and may hibernate over winter; in warmer climates they may have two broods in a season. While the dorsal surface of the wings is vividly marked, the ventral surface is drab, providing camouflage. Eggs are laid on the common nettle, on which the larvae feed.

<i>Pieris brassicae</i> Species of butterfly

Pieris brassicae, the large white, also called cabbage butterfly, cabbage white, cabbage moth (erroneously), or in India the large cabbage white, is a butterfly in the family Pieridae. It is a close relative of the small white, Pieris rapae.

<span class="mw-page-title-main">West Coast lady</span> Species of butterfly

The West Coast lady is one of three North American species of brush-footed butterflies known colloquially as the "painted ladies". V. annabella occurs throughout much of the western US and southwestern Canada. The other two species are the cosmopolitan Vanessa cardui and the eastern Vanessa virginiensis. This species has also been considered a subspecies of the South American Vanessa carye and is frequently misspelled as "anabella".

<i>Heliconius charithonia</i> Species of butterfly

Heliconius charithonia, the zebra longwing or zebra heliconian, is a species of butterfly belonging to the subfamily Heliconiinae of the family Nymphalidae. It was first described by Carl Linnaeus in his 1767 12th edition of Systema Naturae. The boldly striped black and white wing pattern is aposematic, warning off predators. It is the state butterfly of Florida.

<span class="mw-page-title-main">Small pearl-bordered fritillary</span> Species of butterfly

Boloria selene, known in Europe as the small pearl-bordered fritillary and in North America as the silver-bordered fritillary, is a species of butterfly of the family Nymphalidae. It is found across Europe, Asia and North America, and feeds exclusively on violets in its larval stages. This species prefers wet grassland habitats, where its larval food source, violets, are found. It overwinters in its larval stage, and eggs hatch in the late summer to early autumn. Members of this species are prey for multiple types of birds and other insects.

<span class="mw-page-title-main">Cabbage moth</span> Species of moth

The cabbage moth is primarily known as a pest that is responsible for severe crop damage of a wide variety of plant species. The common name, cabbage moth, is a misnomer as the species feeds on many fruits, vegetables, and crops in the genus Brassica. Other notable host plants include tobacco, sunflower, and tomato, making this pest species particularly economically damaging.

<i>Vanessa indica</i> Species of butterfly

Vanessa indica, called the Indian red admiral or the Asian admiral in the United States, is a butterfly found in the higher altitude regions of India, primarily the Himalayas and the Nilgiri Hills. It is also found in Sri Lanka and Myanmar, China, Korea, SE Russia, Japan. It is a close relative of the painted lady.

<i>Vanessa</i> (butterfly) Genus of brush-footed butterflies in the family Nymphalidae with a near-global distribution

Vanessa is a genus of brush-footed butterflies in the tribe Nymphalini. It has a near-global distribution and includes conspicuous species such as the red admirals, the Kamehameha, and the painted ladies of the Cynthia group : Painted lady, American painted lady, West Coast lady, Australian painted lady, etc. For African admirals, see genus Antanartia. Recently, several members traditionally considered to be in the genus Antanartia have been determined to belong within the genus Vanessa.

<span class="mw-page-title-main">American lady</span> Species of butterfly

The American painted lady or American lady is a butterfly found throughout North America.

<span class="mw-page-title-main">Insect migration</span> Seasonal movement of insects

Insect migration is the seasonal movement of insects, particularly those by species of dragonflies, beetles, butterflies and moths. The distance can vary with species and in most cases, these movements involve large numbers of individuals. In some cases, the individuals that migrate in one direction may not return and the next generation may instead migrate in the opposite direction. This is a significant difference from bird migration.

<span class="mw-page-title-main">Australian painted lady</span> Species of butterfly

The Australian painted lady is a species of butterfly mostly confined to Australia, although westerly winds have dispersed it to islands east of Australia, including New Zealand. Debate surrounds the taxonomy of this species. Some believe that the Australian painted lady should be a subspecies of the painted lady due to the similarity in lifestyle and behaviour. Furthermore, the painted lady is found around the globe, but Australia is the only location in which it varies enough to be considered a separate species.

<i>Boloria eunomia</i> Species of butterfly

Boloria eunomia, the bog fritillary or ocellate bog fritillary is a butterfly of the family Nymphalidae.

<span class="mw-page-title-main">Lepidoptera migration</span>

Many populations of Lepidoptera migrate, sometimes long distances, to and from areas which are only suitable for part of the year. Lepidopterans migrate on all continents except Antarctica, including from or within subtropical and tropical areas. By migrating, these species can avoid unfavorable circumstances, including weather, food shortage, or over-population. In some lepidopteran species, all individuals migrate; in others, only some migrate.

<i>Catopsilia florella</i> Species of butterfly

Catopsilia florella, the African migrant, African emigrant, or common vagrant, is a butterfly of the family Pieridae. It is found in Africa, Arabia and the Canary Islands. Like Catopsilia pomona, this species also has a habit of migration.Many early authors mentioned the presence of this species in Asia; but those were probably due to confusion arises as Catopsilia pyranthe females exhibit a lot of seasonal variations. Catopsilia florella is not included as a species in India in any recent checklists.

<i>Metacrias erichrysa</i> Species of moth

Metacrias erichrysa is a species of moth of the family Erebidae. It is endemic to New Zealand. This species can be found in the lower half of the North Island and western alpine areas of the South Island. The adult female of the species is flightless and buff coloured whereas the male is brightly coloured and a rapid flier during daylight hours. The male of the species is on the wing from mid-November to early January. The species inhabits open herb and tussock fields in mountainous terrain at altitudes of between 900 and 1200 m. Larvae feed on Brachyglottis bellidioides,Festuca novae-zealandiae and indigenous species from the genera Acaena, Muehlenbeckia, Wahlenbergia and Raoulia.

<span class="mw-page-title-main">Monarch butterfly migration</span> Migrations, mainly across North America

Monarch butterfly migration is the phenomenon, mainly across North America, where the subspecies Danaus plexippus plexippus migrates each autumn to overwintering sites on the West Coast of California or mountainous sites in Central Mexico. Other populations from around the world perform minor migrations or none at all. This massive movement of butterflies has been recognized as "one of the most spectacular natural phenomena in the world".

References

  1. "Vanessa cardui". IUCN Red List of Threatened Species .
  2. 1 2 Varshney, R.K.; Smetacek, Peter (2015). A Synoptic Catalogue of the Butterflies of India. New Delhi: Butterfly Research Centre, Bhimtal & Indinov Publishing. p. 219. doi:10.13140/RG.2.1.3966.2164. ISBN   978-81-929826-4-9.
  3. 1 2 Savela, Markku. "Vanessa cardui (Linnaeus, 1758)". Lepidoptera and Some Other Life Forms. Retrieved July 3, 2018.
  4. PD-icon.svg One or more of the preceding sentences incorporates text from this source, which is in the public domain : Bingham, Charles Thomas (1905). Fauna of British India. Butterflies Vol. 1. pp. 365–366.
  5. 1 2 PD-icon.svg One or more of the preceding sentences incorporates text from this source, which is in the public domain : Moore, Frederic (1899–1900). Lepidoptera Indica. Vol. IV. London: Lovell Reeve and Co. pp. 105–107.
  6. "Painted Lady". A-Z of Butterflies. Butterfly Conservation. Archived from the original on October 22, 2012. Retrieved October 22, 2012.
  7. "Painted Lady". A-Z of Butterflies. Butterfly Conservation. Archived from the original on October 22, 2012. Retrieved October 22, 2012.
  8. Vanessa cardui, Butterflies of Canada
  9. Robinson, Gaden S.; Ackery, Phillip R.; Kitching, Ian; Beccaloni, George W.; Hernández, Luis M. (2023). "Vanessa cardui". HOSTS - A Database of the World's Lepidopteran Hostplants. The Natural History Museum. doi:10.5519/havt50xw . Retrieved October 22, 2012.
  10. Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
  11. Kuczkowski, Sławek (May 27, 2009). "Afrykańska Rusałka osetnik (Vanessa cardui) zalewa Europę". Motyle.info - Portal Lepidopterologiczny (in Polish). Retrieved April 24, 2019.
  12. Nature Spot
  13. ארליכמן, ארז (April 2, 2014). "הרימו ראש: מיליוני פרפרים נודדים מעל ישראל". ynet . Retrieved April 24, 2019.
  14. 1 2 רינת, צפריר (March 21, 2019). "הפרפרים נעים בכמויות: מסע הנדידה של מיליוני נמפיות חורשף הגיע לישראל". Haaretz . Retrieved April 24, 2019.
  15. Puig‐Montserrat, X., Izquierdo, R., Ubach, A., & Arrizabalaga, A. (2017, July 20). Back to Africa: autumn migration of the painted lady butterfly Vanessa cardui is timed to coincide with an increase in resource availability. Retrieved March 6, 2020, from https://onlinelibrary.wiley.com/doi/full/10.1111/een.12442
  16. 1 2 3 "Butterfly Conservation: Secrets of Painted Lady migration unveiled". BirdGuides Ltd. October 22, 2012. Archived from the original on October 31, 2012. Retrieved November 6, 2024.
  17. "The worldwide painted lady migration project" . Retrieved April 24, 2019.
  18. Orsak, L.J. (1977). The Butterflies of Orange County, California. Center for Pathobiology Miscellaneous Publication #3. University of California Press, New York. 349 pp.
  19. Netburn, Deborah (March 12, 2019). "In the middle of a butterfly crisis, California sees a burst of painted ladies". Los Angeles Times . Retrieved March 20, 2019.
  20. "בוא אליי פרפר נחמד: גל חריג של פרפרים ברחבי הארץ". mako . March 21, 2019. Retrieved April 24, 2019.
  21. Larsen, T.B. 1984. Butterflies of Saudi Arabia and Its Neighbours. Stacey International, London, 160 pp.
  22. Tilden, J.W. (1962). General characteristics of the movements of Vanessa cardui (L.). Journal of Research on the Lepidoptera 1: 43-49.
  23. Annual report of the Department of Science and nature monitoring, National nature park "Gomil`shans`ky Lisy" (Ukraine, Kharkiv region). 2018 and 2019 years.
  24. Nesbit, R.L., J.K. Hill, I.P. Woiwod, D. Sivell, K.J. Bensusan, and J.W. Chapman (2009). "Seasonally Adaptive Migratory Headings Mediated By A Sun Compass In The Painted Lady Butterfly, Vanessa cardui. Animal Behaviour 78: 1119-1125. Print.
  25. Multiple sources:
  26. Hardesty, Richard. "Roosting Behavior in Adult Vanessa cardui" Journal of the Lepidopterists' Society 41.2 (1987): 116-117
  27. 1 2 3 Harris, M. 1999. "Vanessa cardui" (On-line), Animal Diversity Web. Accessed September 16, 2013 at http://animaldiversity.ummz.umich.edu/accounts/Vanessa_cardui/
  28. "Behaviors to Watch." University of Minnesota Extension. Regents of the University of Minnesota, n.d. Web. 16 Sept. 2013. < http://www.extension.umn.edu/distribution/horticulture/components/DG6711d.html>
  29. 1 2 Hoskins, Adrian. "Butterflies of Europe - Vanessa cardui". learnbutterflies.com. N.p., n.d. Web. 16 September 2013. <https://learnbutterflies.com/painted-lady/>.
  30. 1 2 3 Stefanescu, C., Páramo, F., Åkesson, S., Alarcón, M., Ávila, A., Brereton, T., Carnicer, J., Cassar, L. F., Fox, R., Heliölä, J., Hill, J. K., Hirneisen, N., Kjellén, N., Kühn, E., Kuussaari, M., Leskinen, M., Liechti, F., Musche, M., Regan, E. C., Reynolds, D. R., Roy, D. B., Ryrholm, N., Schmaljohann, H., Settele, J., Thomas, C. D., van Swaay, C. and Chapman, J. W. (2012), Multi-generational long-distance migration of insects: studying the painted lady butterfly in the Western Palaearctic. Ecography. doi: 10.1111/j.1600-0587.2012.07738.x
  31. 1 2 Shapiro, Art. "Vanessa cardui". Art Shapiro's Butterfly Site. Information Center for the Environment (ICE). Archived from the original on September 27, 2013. Retrieved September 25, 2013.
  32. Vandenbosch, R. (2003), Fluctuations of Vanessa cardui butterfly abundance with El Niño and Pacific Decadal Oscillation climatic variables. Global Change Biology, 9: 785–790. doi: 10.1046/j.1365-2486.2003.00621.x
  33. Tilden, J. W. (1962). "General Characteristics of the Movements of Vanessa cardui (L.)" Journal of Research on the Lepidoptera 43-49
  34. Harris, Marie. "Vanessa cardui". Animal Diversity Web. University of Michigan Museum of Zoology. Retrieved September 25, 2013.
  35. Davies, N., Krebs, J., & West, S. (2012). An introduction to behavioural ecology. (4th ed.). West Sussex, UK: Wiley-Blackwell.
  36. Rutowski, Ronald L. (1991). "The Evolution of Male Mate-Locating Behavior in Butterflies". The American NaturalistVol. 138, No. 5: 1121-1139.
  37. Abbott, Charles H. (1951). "A Quantitative Study of the Migration of the Painted Lady Butterfly, Vanessa cardui L." EcologyVol. 32, No. 2: 155-171.
  38. O'Neill, Bridget F., Arthur R. Zangerl, Evan H. Delucia, and May R. Berenbaum. "Olfactory Preferences Of, AndforGrown Under Elevated CO." Environmental Entomology 39.4 (2010): 1291-1301. Print.
  39. Janz, Niklas. "The Relationship Between Habitat Selection And Preference For Adult And Larval Food Resources In The Polyphagous Butterfly Vanessa Cardui (Lepidoptera: Nymphalidae)". Journal of Insect Behavior 18.6 (2005): 767-780. Print.
  40. Briscoe, Adriana D.; Bernard, Gary D.; Szeto, Allan S.; Nagy, Lisa M.; White, Richard H. (2003). "Not all butterfly eyes are created equal: Rhodopsin absorption spectra, molecular identification, and localization of ultraviolet-, blue-, and green-sensitive rhodopsin-encoding mRNAs in the retina of Vanessa cardui". Journal of Comparative Neurology. 458 (4): 334–349. doi:10.1002/cne.10582. ISSN   1096-9861. PMID   12619069. S2CID   33070553.
  41. Bernard, Gary D.; Briscoe, Adriana D. (February 15, 2005). "Eyeshine and spectral tuning of long wavelength-sensitive rhodopsins: no evidence for red-sensitive photoreceptors among five Nymphalini butterfly species". Journal of Experimental Biology. 208 (4): 687–696. doi: 10.1242/jeb.01453 . ISSN   0022-0949. PMID   15695761.
  42. Briscoe, Adriana D.; Sison-Mangus, Marilou P.; Kelber, Almut; Zaccardi, Guillermo (May 15, 2006). "Color discrimination in the red range with only one long-wavelength sensitive opsin". Journal of Experimental Biology. 209 (10): 1944–1955. doi: 10.1242/jeb.02207 . ISSN   0022-0949. PMID   16651559.
  43. "Live butterfly release for special events". Cloverlawn Butterflies Farm. Retrieved January 15, 2016.

Further reading