Painted Lady

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

Wander into our exotic Butterfly Pavilion, an environment bursting with beautiful butterflies, moths and glorious greenery. You will discover an astonishing landscape teeming with life and filled with color, growth, and transformation.

"Studies of model insects have greatly increased our understanding of animal development. Yet, they are limited in scope to this small pool of model species: a small number of representatives for a hyperdiverse group with highly varied developmental processes. One factor behind this narrow scope is the challenging nature of traditional methods of study, such as histology and dissection, which can preclude quantitative analysis and do not allow the development of a single individual to be followed. Here, we use high-resolution X-ray computed tomography (CT) to overcome these issues, and three-dimensionally image numerous lepidopteran pupae throughout their development..."

Cynthia cardui (Linnaeus), resurrected combination

Papilio cardui (including the synonym belladonna) Linnaeus, 1758, p. 475 [Article 11 (d), International Code of Zoological Nomenclature states that “a name first published as a synonym is not thereby made available.” Hence belladonna is not an available name.].—Fabricius, 1775, p. 499.—Esper, 1777, pp. 132–139, pl. 10: fig. 3.—Fabricius, 1787, p. 45; 1793, p. 103.—Godart, 1821, p. 102.

Vanessa cardui.—Hübner, [1819], p. 33.—Godart, 1819, pp. 323–324.—Boisduval, 1832, p. 119.—Boisduval and LeConte, 1834, pp. 178–179.—Boisduval, 1840, p. 21.—Herrich-Schaffer, 1843, pp. 38, 41.—Doubleday, 1844, p. 79.—Duponchel, 1844, p. 7.—Lucas, 1845, p. 57.—Noel, 1845, pl. 3.—Gray, 1846, p. 11.—Poey, 1847, p. 122.—Duponchel, 1849, pp. 107–109, pl. 12: figs. 42a-c.—Rambur, 1858, p. 15.—Berge, 1863, p. 51. figs. 1a-c.—Berce, 1867, p. 165, pl. 10: fig. 2.—Wagner, 1870, pp. 169–171.—Staudinger, 1871, p. 17.—Frey, 1880, p. 26.—De Waldeim and Eversmann, 1881, pp. 107–108, pl. 12: figs. 1, 2.—Kirby, 1882, p. 13, pl. 7: figs. 3a-d.—Kane, 1885, p. 62, pl. 9: fig. 1.—Jordan, 1886, pp. 84–85.—Pryer, 1888, p. 26; 1889, p. xii, pl. 7: fig. 2.—Scudder, 1888, pl. 2: fig. 1, pl. 67: fig. 7, pl. 83: figs. 60–62; 1889, pp. 470–487, pl. 12: fig. 7, pl. 21: fig. 1, pl. 33: fig. 31, pl. 61: figs. 35, 54, pl. 64: fig. 30, pl. 74: fig. 37, pl. 81: fig. 10, pl. 86: figs. 69–73.—Dale, 1890, pp. 137–148.—Barrett, 1893, pp. 149–155, pl. 21: figs. 1, a-e.—Fischer, 1895, pp. 18, 22–23, 24.—Schille, 1895, p. 218.—Standfuss, 1896, pp. 256–258, pl. 7: fig. 5.—Aurivillius, 1898, p. 130.—Favre, 1899, p. 99.—Meyrick, 1899, p. 194.—Moore, 1899–1900, pp. 105–107, pl. 320: figs. 2a-b.—Dyar, 1903, p. 24.—Comstock and Comstock, 1904, pp. 158–160, pl. 26: figs. 5, 6.—Bingham, 1905, pp. 365–366.—Kirkaldy, 1906, p. 139.—Barnes and McDunnough, 1917, p. 11.—Grinnell, 1918, p. 113.—Lhomme, 1923, pp. 51–52.—Antram, 1924, pp. 175–176.—Comstock, 1927, pp. 133–134, pl. 43: figs. 8, 9.—Hudson, 1828, p. 36, pl. 4: fig. 1.—Williams, 1930, pp. 168–212.—Holland, 1931, pp. 154–155, pl. 1: fig. 1, pl. 3: fig. 37, pl. 4: figs. 60–62.—Evans, 1932, p. 177.—Frohawk, 1934, pp. 16, 24, 25, 33, 140–147, pl. 11: figs. 1–6.—Bates, 1935, p. 166.—Peile, 1937, p. 139.—Davenport and Dethier, 1938, p. 164.—McDunnough, 1938, p. 21.—Hudson, 1939, p. 388.—Wild, 1939, p. 26, pl. 7: figs. 4, 5.—Field, 1940b, pp. 85–87, 274.—Hoffmann, 1940, p. 681.—Macy and Shepard, 1941, pp. 125–126, pl. 4: fig. [7].—Pierce and Beirne, 1941, p. 20, pl. 3: fig. 2.—Comstock, 1944, pp. 450–451.—Stokoe, 1944, pp. 94–96, pl. 9: fig. 3, pl. 10: fig. 3, pl. 15: fig. 1.—Ford, 1945, pp. 60, 76, 90, 154, pl. 2: fig. 2, pl. VIII: fig. 4, pl. 25.—Leighton, 1946, p. 59.—Richards, 1946, pp. 21–22.—Aubert, 1949, pp. 129, 133–135.—Paul and Robert, 1949, pl. 17.—Pinhey, 1949, p. 83, pl. 12: fig. 3.—Williams, 1949a, pp. 72–73; 1949b, p. 39.—Verity, 1950, pp. 329–334, pl. 51: figs. 26–30, pl. 52: figs. 1–6, pl. 18: fig. 2, pl. 19: fig. 14.—Woodhouse, 1950, pp. xiii, 49, pl. 15: fig. 1, pl. 37, fig. 4; pl. 42: fig. 4.—Clark and Clark, 1951, pp. 14, 44, 201, pl. 4: figs, d, e.—Klots, 1951, pp. 107–108, 113, pl. 6: fig. 14, pl. 14: fig. 10.—Agenjo, 1952, p. 283.—Peters, 1952, p. 76.—Ferguson, 1954, p. 195.—Brown, 1955, p. 102, 2 figs.—Forster and Wohlfahrt, 1955, p. 56, pl. 15: fig. 13.—Yokoyama, 1955, pp. 37, 96–97, pl. 29: fig. 89.—Lempke, 1956, pp. 185–186.—Wiltshire, 1957, p. 31.—Wynter-Blyth, 1957, pp. 210–212, pl. 38: fig. 8.—Zimmermann, 1959, pp. 453, 454, 455, 456, 457, 463, figs. 388, 389, 391, 396, 400.—Chalmers-Hunt, 1960–1961, pp. 60–62, 140.—Forbes, 1960, p. 159.—Dos Passos, 1964, pp. 77.—Niculescu, 1965, pp. 186–191, pl. 6: figs. 1, 2.—Gifford, 1965, p. 115.—Gaskin, 1966, pp. 88–89, pl. 1: fig. [5].—van Son, 1966, p. 66, fig. 1.—Dimock, 1968. p. 146.—Williams, 1970, pp. 157–175.

Cynthia cardui.—Humphreys and Westwood, 1841, pp. 54, 56–57, pl. 15: figs. 7–10.—Stainton, 1857, pp. 36–37, fig.—Humphreys, [1859], pp. 21–22, pl. 11: figs. 1–4.—Morris, 1868, pp. 81–82, pl. 34.—Buckler, 1886, pp. 49–52, 174–176, pl. 8: figs. 1a-e.—Barnes and Benjamin, 1926, p. 15.—Gunder, 1928, p. 202, pl. 8: figs. 3b, 3c; 1929, p. 9.

Pyrameis cardui.—Doubleday, 1849, p. 205.—Horsfield and Moore, 1857, pp. 138–139, p. [3], pl. 5: figs. 3, 3a.—Lucas, 1857, pp. 543–544.—Morris, 1860, p. 8; 1862, pp. 59–60.—Behr, 1864, pp. 125–126.—Bremer, 1864, p. 17.—Trimen, 1862–1866, pp. 119–121.—Butler, 1870, pp. 77–78.—Kirby, 1871, pp. 185–186.—Newman, 1874, pp. 64–66, fig. 17.—Strecker, 1878, pp. 136–138, 190.—Moore, 1880–1881, pp. 50–51, pl. 27: figs. 1, 1a.—Godman and Salvin, 1882, pp. 217–218.—Saalmüller, 1884, p. 77.—Alphéraky, 1885, p. 598.—Staudinger, 1885. p. 97.—Distant, 1882–1886, pp. 48, 91.—Mabille, 1886, pp. 126–127.—Marshall and de Nicéville, 1886, pp. 227–228.—Shatz and Röber, 1886, pl. 16: fig. [9]; 1887, p. 125.—Trimen, 1887, pp. 200–203.—Edwards, 1889, p. 26.—Dixey, 1890, pp. 92, 93, 94, 96, 97, 98, 99, 101, 106, 109, 112, 115. 116, 117, 118, 122, 123, 126, 127, pl. 1: figs. 5, 8, pl. 2: fig. 24, pl. 3: fig. 38.—Maynard, 1891, p. 93, pl. 5: fig. 5.—Leech, 1892–1894, pp. xv, 351.—White, 1894, pp. 54–55, 63, pl. 2: fig. 5.—Hagen, 1896, p. 165.—Skinner, 1898, pp. 24–25.—Tutt, 1896, pp. 350–355, fig. 37.—Denton, 1898, pp. 266–267.—Fruhstorfer, 1898b, p. 151.—Holland, 1898, pp. 170–171, pl. 1: fig. 1, pl. 3: fig. 37, pl. 4: figs. 60–62.—Miyajima, 1899, pp. 3, [118], pl. 11: fig. 2.—Cannaviello, 1900, pp. 18–19.—Staudinger and Rebel, 1901, p. 24.—Wright, 1905, pp. 177–178, pl. 22: fig. 229.—Kirkaldy, 1906, p. 139.—Schultz, 1906, p. 108.—Stichel, 1908, pp. 199–200, pl. 62: fig. D 1.—Stichel, 1909, pp. 199–200, pl. 62: fig. D 1.—Rebel, 1910, p. 20.—Reuss, 1910a, pl. 1: figs. 1. 5. 9; 1910b, pp. 62, 63, 64, 65, 66, 67; 1910c, p. 90; 1910d. p. 303.—Fruhstorfer, 1912a, pp. 524–525; 1912b, p. 525.—Aurivillius, 1913a, p. 227; 1913b, p. 227.—Verity, 1913, p. 181.—Seitz, 1914a, p. 458; 1914b, p. 458.—Waterhouse and Lyell, 1914, p. 55.—Holland, 1915, p. 102, pl. 35.—Schultz, 1916, p. 27.—Verity, 1916b, p. 128.—Reuss, 1918, pp. 41–42.—Ragusa, 1919, p. 146.—Holland, 1920. pp. 143–144.—Stephan, 1923, p. 36.—Frohawk, [1924], pp. 151–157, pl. 26.—Ormiston, 1924, p. 32.—Stephan, 1924, p. 25.—Verity and Querci, 1924, p. 42.—Bang-Haas, 1926, p. 56; 1927. p. 55; 1928, p. 26.—Schrader, 1928, pp. 68–70, pl. 6.—Haas, 1929, p. 11.—Malsumura, 1929, pp. 9, 11, 19, pl. 5: fig. 5.—Schrader, 1929, p. 8.—Bang-Haas, 1930, p. 162.—Gaede, 1930a, p. 200; 1930b, p. 200; 1931, p. 343.—Clark, 1932, pp. 13, 14, 22, 88–91, pl. 8: figs. 1–3.—Bryk, 1940, p. 17.—South, 1941, pp. 78–81, fig. 24, pis: 44, 45.—Zerkowitz, 1946, p. 246.—Barrett and Bames, 1951, pp. 132–133.—Swanepoel, 1953, pp. 213–215.—Paulian, 1956, pp. 67–69.

Pyrameis cardui cardui.—Fruhstorfer, 1912a, p. 524; 1912b, p. 524.—Verity, 1919, p. 197.

Vanessa cardui cardui.—Field, 1940b, p. 85.—Corbet and Pendlebury, 1956, p. 207, pl. 6: fig. 69, pl. 40: fig. 100.—Okano and Ohkura, 1959, p. 42, pl. 41: fig. 124.

Papilio carduelis Seba, 1765, p. 6, pl. 1.—Cramer, 1775. pp. 40–41, pl. 26: figs, E, F; 1776, p. 152.—Doubleday, 1844, p. 79 [synonym of cardui]; 1849, p. 205.—Horsfield and Moore, 1857, p. 138.—Lucas, 1857, p. 544.—Saalmuller, 1884, p. 77.—Mabille, 1886, p. 126.—Marshall and de Nicéville. 1886, p. 227.—Leech, 1892–1894, p. 351.—Aurivillius. 1898. p. 130.—Moore, 1899–1900, p. 105.

Vanessa carduelis.—Dyar, 1903, p. 24 [synonym of cardui].—McDunnough, 1928, p. 21.—Comstock, 1944, p. 450.—Dos Passos, 1964, p. 77.

Cynthia carduelis.—Barnes and Benjamin, 1926, p. 15 [synonym of cardui].

Pyrameis cardui universa form carduelis.—Verity, 1919, p. 198.

Pyrameis cardui variety carduelis.—Verity, 1920, p. 61.

Pyrameis cardui race carduelis.—Verity and Querci, 1924, p. 42.

Vanessa cardui form carduelis: Verity, 1936, p. 87; 1950, p. 334.

Vanessa cardui carduelis.—Field, 1940a, p. 342; 1940b, pp. 85, 86, 274.—Klots, 1951, p. 108.

Vanessa cardui aberration carduelis.—Verity, 1950, p. 330.

Papilio belladonna Godart, 1821, pp. 102–104, pl. 14: fig. 2.

Vanessa elymi Rambur, 1829, pp. 256–257, pl. 5: figs. 1, 2.—Dyar, 1903, p. 24 [synonym of cardui].

Vanessa cardui aberration elymi.—Boisduval, 1840, p. 21.—Duponchel, 1844, p. 7.—Staudinger, 1871, p. 17.—Jordan, 1886, pp. 84–85.—Scudder, 1889, p. 473.—Fischer, 1896, pp. 17–18, 57, 66, pl. 2: fig. 5b.—Skinner, 1898, p. 25.—Fischer, 1903, p. 224; 1907, pp. 170, 201, figs. 5, 7.—Barnes and McDunnough, 1917, p. 11.—Lhomme, 1923, p. 51.—Comstock, 1927, p. 134.—Gunder, 1925, p. 198.—McDunnough, 1938, p. 21.—Field, 1940a, p. 87.—Leighton, 1946, p. 59.—Verity, 1950, p. 332.—Chalmers-Hunt, 1960–1961, p. 61.—Dos Passos, 1964, p. 77.

Pyrameis cardui variety elymi.—Kirby, 1871, p. 186.

Pyrameis cardui aberration elymi.—Strecker, 1878, p. 137.—Tutt, 1896, p. 352.—Staudinger and Rebel, 1901, p. 24.—Schultz, 1906, p. 108.—Stichel, 1908, p. 200; 1909, p. 200.—Rebel, 1910, p. 20.—Reuss, 1910a, pp. 63, 65, 66.—Fischer, 1900, p. 5.

Vanessa cardui variety elymi.—Dale, 1890, p. 138.

Pyrameis cardui elymi.—Fruhstorfer, 1912a, p. 525.

Vanessa cardui elymi.—Fox, 1921, p. 46, pl. 2: fig. 3.

Cynthia cardui aberration elymi.—Barnes and Benjamin, 1926, p. 15.

Cynthia cardui transition form elymi.—Gunder, 1927, p. 133, pl. 2: fig. 2a; 1927c, p. 270, pl. 9: figs. 2–4; 1929, p. 9.

Vanessa cardui form elymi.—Dimock, 1968, p. 146, fig. 1.

Pyrameis cardui variety leachiana Doubleday [nomen nudum], 1849, p. 205.

Pyrameis cardui aberration ate Strecker [excluded name, type 2], 1878, pp. 137–138.—Skinner, 1898, p. 25.

Vanessa cardui aberration ate.—Scudder, 1889, pp. 474–475.—Barnes and McDunnough, 1917, p. 11.—McDunnough, 1938, p. 21.—Field, 1940b, p. 86.—Dos Passos, 1954, p. 77.

Cynthia cardui aberration ate.—Barnes and Benjamin, 1926, p. 14.

Cynthia cardui transition form ate.—Gunder [excluded name, type 2], 1927c, p. 270, pl. 9: fig. [6].

Vanessa cardui aberration pallens Noel [excluded name, type 2], 1881, p. 102.—Field, 1940b, p. 86.—Verity, 1950, p. 331.—Niculescu, 1965, p. 190.

Pyrameis cardui aberration pallens.—Stichel, 1908, p. 199; 1909, p. 199.

Pyrameis cardui form pallens.—Seitz [excluded name, type 3], 1914a, p. 458; 1914b, p. 458.

Pyrameis cardui transition form pallens.—Gunder [excluded name, type 2], 1927, p. 53.

Vanessa cardui variety pallida Schøyen, 1881, pp. 77–79.

Pyrameis cardui aberration pallida.—Tutt, 1896, p. 352.—Rebel, 1910, p. 20.—Pionneau, 1926, p. 4.

Pyrameis cardui variety pallida.—Staudinger and Rebel, 1901, p. 24.—Schultz, 1906, p. 108.

Pyrameis cardui form pallida.—Stichel, 1908, p. 200; 1909, p. 200.

Pyrameis cardui cardui form pallida.—Verity, 1919, p. 197.

Vanessa cardui aberration pallida.—Lhomme, 1923, p. 51.—Verity, 1950, p. 330.—Niculescu, 1965, p. 190.

Pyrameis cardui race pallida.—Gaede, 1930a, p. 200; 1930b, p. 200.

Vanessa cardui form pallida.—Field, 1940b, p. 84.—Lempke, 1956, p. 187.

Pyrameis kershawi M'Coy, Godman and Salvin not M'Coy [a misidentification], 1882, p. 218.

Vanessa cardui aberration inornata Bramson [excluded name, type 2], 1886, p. 284.—Scudder, 1889, pp. 473, 474.—Frohawk, 1934, p. 147; 1938, pp. 84, 88, pl. 21: figs. 1, 2.—Field, 1940b, p. 86.—Verity, 1950, p. 331.—Chalmers-Hunt, 1960–1961, p. 62.

Pyrameis cardui aberration inornata.—Tutt, 1896, p. 352.—Staudinger and Rebel, 1901, p. 24.—Stichel, 1908, pp. 199–200; 1909, pp. 199–200.—Rebel, 1910, p. 20.—Pionneau, 1926, p. 4.

Pyrameis cardui form inornata.—Seitz [excluded name, type 3], 1914a, p. 458; 1914b, p. 458.—Van Mellaerts, 1928, p. 105.

Pyrameis cardui variety inornata.—Stephan [excluded name, type 3], 1924, p. 25.

Vanessa cardui form inornata.—Kuijken, 1967, pp. 224, 225.

Vanessa cardui aberration minor Failla-Tedaldi [excluded name, type 2], 1887, pp. 70–71.—Verity, 1950, p. 330.

Pyrameis cardui aberration minor.—Ragusa, 1919, p. 146.

Pyrameis cardui universa form minor.—Verity [excluded name, type 1 and type 2], 1919, p. 198.

Vanessa cardui carduelis form minor.—Field, 1940b, pp. 86, 274.

Vanessa cardui form minor.—Lempke, 1956, p. 187.

Vanessa cardui semisuffusa Cockerell [excluded name, type 2, as an individual variation], 1889, p. 54.

Vanessa cardui form semisuffusa.—Lempke [excluded name, type 3], 1956, p. 187.

Vanessa cardui aberration semisuffusa.—Chalmers-Hunt [excluded name, type 2], 1960–1961, p. 62.

Vanessa cardui aberration wiskotti Standfuss [excluded name, type 2], 1895, pp. 90–91; 1896, 392, pl. 7: fig. 6.—Verity, 1950, p. 332.

Pyrameis cardui variety wiskotti.—Fischer [excluded name, type 3], 1900, p. 5.

Pyrameis cardui aberration wiskotti.—Fischer, 1901, p. 326.—Schultz, 1906, p. 108.—Stichel, 1908, p. 200; 1909, p. 200.—Reuss, 1910a, fig. 19; 1910b, p. 64; 1910c, p. 90.

Vanessa cardui warm variety wiskotti.—Fischer [excluded name, type 3], 1903, p. 224.

Vanessa cardui cold variety wiskotti.—Fischer [excluded name, type 3], p. 170.

Pyrameis cardui form minor Cockerell [excluded name, type 3 and homonym of minor Failla-Tedaldi], 1890, p. 57.

Pyrameis cardui aberration minor.—Skinner [excluded name, type 2], 1898, p. 25.

Vanessa cardui aberration minor.—Barnes and McDunnough, 1917, p. 11.—McDunnough, 1938, p. 21.

Cynthia cardui aberration minor.—Barnes and Benjamin, 1926, p. 15.—Gunder, 1928, p. 202, pl. 8: fig. 3a.

Pyrameis cardui aberration minor Cannaviello [excluded name, type 2 and homonym of minor Failla-Tedaldi], 1900, pp. 19–20.—Stichel, 1908, p. 199; 1909, p. 199.—Rebel, 1910, p. 20.—Stephan, 1923, p. 36; 1924, p. 25.

Pyrameis cardui variety minor.—Schultz [excluded name, type 3], 1905, p. 67; 1906, p. 108.

Pyrameis cardui form minor.—Seitz [excluded name, type 3], 1914a, p. 458; 1914b, p. 458.—Gaede, 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration minor.—Lhomme, 1923, p. 51.—Verity, 1950, p. 330.

Pyrameis cardui aberration carduelis Schultz [excluded name, type 2 and homoym of carduelis Seba], 1906, p. 108.—Stichel, 1908, p. 199; 1909, p. 199.—Rebel, 1910, p. 20.

Pyrameis cardui variety carduelis.—Stephan [excluded name, type 3], 1924, p. 25.

Vanessa cardui aberration carduelis.—Verity, 1950, p. 331.

Pyrameis cardui aberration priameis Schultz [excluded name, type 2], 1906, p. 108.—Stichel, 1908, p. 199; 1909, p. 199.—Rebel, 1910, p. 20.—Van Mellaerts, 1928, p. 105.—Zerkowitz, 1946, p. 246.

Pyrameis cardui variety priameis.—Stephan [excluded name, type 3], 1924, p. 25.

Vanessa cardui aberration priameis.—Verity, 1950, p. 332.—Chalmers-Hunt, 1960–1961, p. 62.

Pyrameis cardui aberration carduelina Alphéraky [excluded name, type 2], 1908, pp. 573–574.—Bang-Haas, 1926, p. 56.

Pyrameis cardui form carduelina.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration carduelina.—Field, 1940b, p. 86.—Verity, 1950, p. 330.

Pyrameis cardui japonica Stichel [new synonymy], 1908, p. 200, pl: 62, fig. D 2; 1909, p. 200, pl. 62: fig. D 2.—Fruhstorfer, 1912a, p. 525; 1912b, p. 525.—Matsumura, 1929, p. 19.—Cho, 1934, pl. 20: fig. 2.—Doi, 1934, p. 29.—Mori, 1934, pp. 72–73.—Bryk, 1946, p. 39.

Pyrameis cardui universa form japonica.—Verity, 1919, p. 197.

Vanessa cardui japonica.—Field, 1940b, p. 85.

Vanessa cardui aberration japonica.—Verity, 1950, p. 331.

Vanessa cardui form japonica.—Lempke, 1956, p. 188.

Pyrameis cardui aberration ocellata Rebel [excluded name, type 2], 1910, p. 20.—Reuss, 1910b, pp. 63, 64.—Bang-Haas, 1926, p. 56.

Pyrameis cardui variety ocellata.—Wize [excluded name, type 3], 1917, p. 5.—Stephan, 1924, p. 25.

Pyrameis cardui form ocellata.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration ocellata.—Field, 1940b, p. 86.—Verity, 1950, p. 331.

Vanessa cardui form ocellata.—Lempke, 1956, p. 187.

Pyrameis cardui aberration carnea Fritsch [excluded name, type 2], 1912, p. 136.—Bang-Haas, 1926, p. 56.

Pyrameis cardui form carnea.—Reuss [excluded name, type 3], 1918, p. 41.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui variety carnea.—Stephan [excluded name, type 3], 1924, p. 25.

Vanessa cardui aberration carnea.—Field, 1940b, p. 86.—Verity, 1950, p. 330.

Vanessa cardui form carnea.—Lempke, 1956, p. 187.

Pyrameis cardui aberration melanosa Cabeau [excluded name, type 2], 1913, p. 43.—Bang-Haas, 1926, p. 56.

Vanessa cardui aberration melanosa Lhomme, 1923, p. 51.—Verity, 1950, p. 332.

Pyrameis cardui form melanosa.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui form melanosa.—Lempke, 1956, p. 187.

Pyrameis cardui form brunneaalbimaculata Reuss [excluded name, type 3], 1916, pp. 130–131.—Bang-Haas, 1926, p. 56.—Gaede, 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration brunneaalbimaculata.—Field [excluded name, type 2], 1940b, p. 86.—Verity, 1950, p. 331.

Pyrameis cardui form rosacea Reuss [excluded name, type 3], 1916, p. 131; 1918, p. 41.—Bang-Haas, 1926, p. 56.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui variety roseacea.—Stephan [excluded name, type 3], 1924, p. 25.

Vanessa cardui aberration rosacea.—Field [excluded name, type 2], 1940b, p. 86.—Verity, 1950, p. 330.

Pyrameis cardui form hunteri Reuss [excluded name, type 3], 1918, p. 41.

Vanessa cardui form hunteri.—Lempke, 1956, p. 187.

Pyrameis cardui form ochracea Reuss [excluded name, type 3], 1918, p. 41.

Vanessa cardui form ochracea.—Lempke, 1956, p. 187.

Pyrameis cardui form conjuncta Verity [excluded name, type 1 and type 3], 1919, p. 197.

Pyrameis cardui form conjuncta.—Bang-Haas [excluded name, type 3], 1927, p. 55.—Gaede, 1930a, p. 200; 1930b. p. 200.

Vanessa cardui cardui aberration conjuncta.—Field [excluded name, type 1 and type 2], 1940b, p. 85.

Vanessa cardui aberration conjuncta.—Verity [excluded name. type 2], 1950, pp. 331, 336.—Niculescu, 1965, p. 190.

Pyrameis cardui aberration emielymi Verity [excluded name. type 2], 1919, p. 198.—Bang-Haas, 1927, p. 55.

Pyrameis cardui form emielymi.—Gaede [excluded name. type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration emielymi.—Field, 1940b, p. 86.—Verity, 1950, p. 331.—Chalmers-Hunt, 1960–1961, p. 62.

Vanessa cardui form emielymi.—Lempke, 1956, p. 189.

Pyrameis cardui variety infrabrunnea Verity [excluded name, type 3], 1919, p. 198.

Pyrameis cardui inops form infrabrunnea.—Bang-Haas [excluded name, type 1 and type 3], 1927, p. 56.

Pyrameis cardui form infrabrunnea.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration infrabrunnea.—Verity [excluded name, type 2], 1950, p. 331.

Pyrameis cardui form infraflava Verity [excluded name, type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infraflava.—Bang-Haas, 1927, p. 56.

Vanessa cardui aberration infraflava.—Verity [excluded name, type 2], 1950, p. 331.

Pyrameis cardui form infragrisea Verity [excluded name, type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infragrisea.—Bang-Haas, 1927, p. 56.

Vanessa cardui aberration infragrisea.—Verity [excluded name, type 2], 1950, p. 331.

Pyrameis cardui form infranigrans Verity [excluded name, type 3], 1919. p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infranigrans.—Bang-Haas, 1927, p. 55.

Vanessa cardui aberration infranigrans.—Verity [excluded name, type 2], 1950, p. 331.

Pyrameis cardui form infraochracea Verity [excluded name, type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui inops form infraochracea.—Bang-Haas, 1927, p. 56.

Vanessa cardui aberration infraochracea.—Verity [excluded name, type 2], 1950, pp. 331, 334.

Pyrameis cardui universa form inops Verity [excluded name, type 1], 1919, p. 198.

Pyrameis cardui race inops Verity and Querci, [new synonymy], 1924, p. 42.—Bang-Haas, 1927, p. 55.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui form inops.—Verity, 1936, p. 87.

Vanessa cardui inops.—Field, 1940b, p. 85.

Vanessa cardui aberration inops.—Verity, 1950, pp. 331, 334.

Vanessa cardui form inops.—Lempke, 1956, pp. 186–187.

Pyrameis cardui form sexiespupillata Verity [excluded name, type 3], 1919, p. 198.—Gaede, 1930a, p. 200; 1930b, p. 200.

Pyrameis cardui aberration sexiespupillata.—Bang-Haas [excluded name, type 2], 1927, p. 55.—Van Mellaerts, 1928, pp. 104–105.

Vanessa cardui aberration sexiespupillata.—Field, 1940b, p. 86.—Verity, 1950, p. 330.—Chalmers-Hunt, 1960–1961, p. 62.

Vanessa cardui carduelis aberration sexiespupillata.—Field [excluded name, type 1], 1940b, p. 274.

Vanessa cardui form sexiespupillata.—Lempke, 1956, p. 188.

Pyrameis cardui form septiespupillata Verity [excluded name, type, 3], 1919, p. 198.—Gaede, 1950a, p. 200; 1950b, p. 200.

Pyrameis cardui aberration septiespupillata.—Bang-Haas [excluded name, type 2], 1927, p. 55.—Van Mellaerts, 1928, p. 104.

Vanessa cardui aberration septiespupillata.—Field, 1940b, p. 86.—Verity, 1950, p. 331.

Vanessa cardui form septiespupillata.—Lempke, 1956, p. 188.

Pyrameis cardui race universa Verity, [new synonymy], 1919, p. 197; 1920, p. 61.—Verity and Querci, 1924, p. 42.—Bang-Haas, 1926, p. 56.—Gaede, 1930a, p. 200; 1930b, p. 200.

Vanessa cardui form universa.—Verity, 1950, p. 334.—Lempke, 1956, p. 186.

Vanessa cardui race universa.—Verity, 1936, p. 87; 1937, p. 22.—Field, 1940b, p. 85.

Pyrameis cardui aberration maria Stephan [excluded name, type 2], 1924, p. 25.

Pyrameis cardui aberration marthamaria Stephen [excluded name, type 2], 1924, p. 25.—Bang-Haas, 1927, p. 55.

Vanessa cardui aberration marthamaria.—Field, 1940b, p. 86.—Verity, 1950, p. 331.

Pyrameis cardui aberration schoenfellneri Hoffman [excluded name, type 2], 1924, p. 25.—Bang-Haas, 1927, p. 55.

Vanessa cardui aberration marthamaria.—Field, 1940b, p. 86.—Verity, 1950, p. 331.

Pyrameis cardui aberration schoenfellneri Hoffman [excluded name, type 2], 1925, pp. 29–30.

Pyrameis cardui litoralis aberration schoenfellneri.—Bang-Haas [excluded name, type 1], 1927, p. 56.

Pyrameis cardui form schoenfellneri.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration schoenfellneri.—Field, 1940b, p. 86.—Verity, 1950, p. 332.

Pyrameis cardui aberration subfracta Stack [excluded name, type 2], 1925, p. 114, fig. 3.—Bang-Haas, 1928, p. 26.

Pyrameis cardui litoralis aberration subfracta.—Bang-Haas [excluded name, types 1 and 2], 1927, p. 56.

Pyrameis cardui form subfracta.—Gaede [excluded name, type 3], 1930a, p. 200; p. 1930b, p. 200.

Vanessa cardui aberration subfracta.—Field, 1940b, p. 86.—Verity, 1950, p. 330.—Niculescu, 1965, p. 190.

Vanessa cardui form subfracta.—Lempke, 1956, p. 188.

Pyrameis cardui aberration takesakiana Kato [excluded name, type 2], 1925, pp. 145–146, fig.

Vanessa cardui aberration takesakiana.—Field, 1940b, p. 86.

Pyrameis cardui litoralis de Souza [new synonymy], 1926, p. 3.—Bang-Haas, 1927, p. 56.

Pyrameis cardui aberration rosea Pionneau [excluded name, type 2], 1926, p. 4.—Bang-Haas, 1927, p. 55.

Pyrameis cardui form rosea.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration rosea.—Field, 1940b, p. 86.

Cynthia cardui jacksoni Clark [as an individual variant, excluded name, type 2], 1927, p. 127.

Pyrameis cardui form jacksoni.—Clark [excluded name, type 3], 1932, p. 89, pl. 8: figs. 2, 3.

Vanessa cardui carduelis aberration jacksoni.—Field [excluded name, type 1], 1940b, pp. 86, 87, 274.

Vanessa cardui form jacksoni.—Lempke, 1956, p. 187.—Dos Passos, 1964, p. 77.

Pyrameis cardui aberration browni Meilhan [excluded name, type 2], 1928, pp. 102–103.

Pyrameis cardui form browni.—Gaede [excluded name, type 3], 1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration browni.—Verity, 1950, p. 331.

Pyrameis cardui aberration flava Bandermann [excluded name, type 2], 1928, pp. 236–237.—Bang-Haas, 1929, p. 11.

Pyrameis cardui form flava.—Gaede [excluded name, type 3], 1930a, p. 200; 1930b, p. 200.

Vanessa cardui aberration flava.—Field, 1940b, p. 86.

Pyrameis cardui aberration rogeri Meilhan [excluded name, type 2], 1928, pp. 100–103.—Bang-Haas, 1930, p. 162.

Pyrameis cardui subspecies rogeri.—Gaede [new synonymy], 1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration rogeri.—Field, 1940b, p. 86.—Verity, 1950, p. 331.—Chalmers-Hunt, 1960–1961, p. 62.

Pyrameis cardui aberration varini Meilhan [excluded name, type 2], 1928, p. 103.—Bang-Haas, 1930, p. 162.

Pyrameis cardui form varini.—Gaede [excluded name, type 3], 1931a, p. 343; 1931b, p. 343.

Vanessa cardui aberration varini.—Field, 1940b, p. 86.—Verity, 1950, p. 331.—Chalmers-Hunt, 1960–1961, p. 62.

Vanessa cardui form varini.—Lempke, 1956, p. 189.

Pyrameis cardui form hemielymi Gaede [excluded name, type 3], 1930a, pl. 13: fig. a 1; 1930b, pl. 13: fig. a 1.

Pyrameis cardui aberration johni Fischer [excluded name, type 2], 1932, p. 158, fig.

Vanessa cardui aberration johni.—Field, 1940b, p. 86.

Vanessa cardui aberration pallida Frohawk [excluded name, type 2 and homonym of pallida Schøyen]. 1938, p. 84.

Vanessa cardui ushuwaia Bryk [new synonymy], 1944, pp. 10–11, pl. 1: fig. 6; 1946, p. 39; 1953, p. 93.

Vanessa cardui japonica form koreana Bryk [excluded name, types 1, 2, 3], 1946, p. 39.

Vanessa cardui aberration albicans Verity [excluded name, type 2], 1950, p. 331.

Vanessa cardui aberration infralutea Verity [excluded name, type 2], 1950, p. 334.

Vanessa cardui form albipuncta Lempke [excluded name, type 3], 1956, p. 188.

Vanessa cardui form nigripuncta Lempke [excluded name, type 3], 1956, pp. 188–189.

This species, probably one of the two or three best known and most described and illustrated species of butterflies in the world, hardly needs redescription except to compare it to the other species of the genus that are not nearly as well known. C. cardui, as mentioned earlier in the group description, differs from all other species of Cynthia, except C. kershawi, in having the fuscous color in interspace Cu2 on upper surface of forewing not extending to vein Cu1. On the upper surface of the hindwing it differs from C. kershawi in having five, not four, submarginal ocular spots, these located between veins Rs and Cu2, and all lacking blue centers except for a rare individual variant that sometimes occurs with a blue center in the spot located in interspace Cu1.

MALE (Figures 89–96).—Forewing above with subapical white bar just beyond cell, and subapical and submarginal white spots very much as in C. kershawi and quite similar to those of the other Cynthia species except that the white spot found in interspace Cu1 in the species of the virginiensis group is lacking in C. cardui and C. kershawi; with fuscous bar in interspace Cu1 connecting apical and subapical fuscous areas to central fuscous spot in interspace Cu2 lacking; with the central fuscous spot in interspace Cu2 just mentioned usually not so strongly connected to the fuscous spot at base and above base of Cu2 as it is in C. kershawi and with the fuscous spots in middle of cell distinctly separated, not nearly confluent as is true of C. kershawi.

Hindwing on the upper surface as mentioned before with five, not four, submarginal ocular spots between vein Rg and Cu2, none with blue centers except very rarely the one in interspace Cu1; with an upwardly projected hooklike fuscous marking below and beyond cell, all that remains of the fuscous and usually sinuous band that crosses the wing from costa to just above anal angle in the species of the virginiensis group, this hooklike fuscous spot nearly or sometimes entirely joining a downward projected fuscous streak from the fuscous area of the costal margin and base of wing.

Under surfaces of forewing with fuscous of central area of interspace Cu2 extending upward nearly, or at least halfway, through interspace Cu1 and with two fuscous spots in middle of cell just inward from the white area distinctly separated from each other.

Hindwing on the under surface with a lighter overall appearance than in C. kershawi, with the submarginal ocular markings distinctly larger than in that species, and with the subtriangular white spot at the end of the cell in interspace M2 also much larger.

Length of forewing, 22–39.5 mm (average 29.3 mm).

Male genitalia as illustrated by Figure 8 (drawn from my preparation no. 6104), with uncus tapered distally to a point; gnathos longer, with uncus with distal part after “heel” nearly at right angles to the base; valva with costal margin broadly projected upward, forming a broad obtuse angle with a small upward projecting spine at its summit just below uncus; distal margin of valva also broadly projected and terminating in one or two spines; lower margin of valva broadly convex and continuous with distal margin, lacking completely the sickle-shaped projection found in C. kershawi; clasper directed toward outer margin and curved inward at its extremity; cuiller smooth with free part curved in a shallow figure toward distal margin; aedeagus very similar to that of C. myrinna, with base slightly bent downward.

FEMALE (Figures 97–104).—This sex hardly differs from the male except that most females have a slightly more rounded outer margin on the hindwing and have the outer margin of forewing near vein Cu2 slightly more produced.

Length of forewing, 24–37 mm (average 30.9 mm).

Female genitalia as illustrated by Figure 24 (drawn from my preparation no. 3675), with anterior margin of eighth and posterior margin of seventh sternite greatly undulated and with these sternites forming two large posteriorly directed lobes; ostium bursae swollen and located in eighth sternite between lobes of seventh sternite; with a pair of laterally placed pockets between seventh and eighth sternites; ribbonlike signa extending nearly full length of bursa copulatrix.

INDIVIDUAL AND ABERRATIONAL VARIATION.—Individual variation in this species commonly consists of differences in the ground color of the upper surfaces, which varies from a pale tan to a salmon red. The fuscous markings vary in extent and sometimes there are blue centers in the lowermost submarginal round spots of the upper surfaces of the hindwings. Aberrational variation has been studied by numerous persons who unfortunately have given some forty-nine formal names to individuals displaying this “more rare” type of individual variation. Specimens having an extra white spot, or lacking one, or displaying any slight variation from the “normal” have received names. Names have usually always been given also to any remarkably different looking specimen. All of these names are synonymized above or are excluded from nomenclature. Gaede (1930a, 1930b, p. 200), Verity (1950, pp. 330–332), and Lempke (1956, pp. 186–189) treat many of these names and should be consulted by those interested.

SEASONAL VARIATION.—Variation in habitus between the three broods (where there are three broods) does not seem to occur in this species. There are “wet” and “dry” forms in material collected in microenvironments that would be thought of (at least temporarily) as wet and dry areas. Where there is sufficient rain, specimens occur that are larger, brighter, and more pinkish above and darker on the under surfaces.

GEOGRAPHICAL VARIATION.—Verity (1919, p. 197; 1920, p. 21; 1950, p. 334), and Verity and Querci (1924, p. 42) have maintained that there is a great deal of geographical variation and have recognized several subspecies as Vanessa cardui universa, V. cardui inops, V. cardui carduelis, and V. cardui cardui. Stichel (1908, p. 200) describes what he considers to be a subspecies as Pyrameis cardui japonica. None of these hold up as subspecies. Of the more than two hundred and fifty specimens examined, the characters supposedly differentiating these subspecies occur without relation to geography, so all of these proposed subspecies are synonymized above.

METHOD OF IDENTIFICATION.—The original description of Linnaeus (1758, p. 475) and his references to previous descriptions and illustrations clearly identify his name with the species here treated as Cynthia cardui.

Since Linnaeus in his original description refers to his 1746 work “Fauna Svecica,” I consider Sweden to be the type locality of C. cardui. Verity (1913, p. 181), in his study of the Linnean types, states of cardui: “There is nothing noteworthy about the one typical specimen.” This specimen is in the Linnean collection of the Linnean Society, London.

SYNONYMY.—See synonymy and literature citations above. In addition to the names proposed for so-called subspecies, which are synonymized above, Papilio belladonna Linnaeus, a name used earlier by Linnaeus (and therefore not available), is listed by him as a synonym (1758, p. 485). Vanessa elymi Rambur (1829, pp. 256–257), described as a species, was based upon an aberrant specimen of C. cardui and is a synonym.

LIFE HISTORY.—The food plants of the larvae are principally members of the Compositae and most especially the various thistles. Among its recorded hosts are: Anaphalis margaritaceae, Arctium leppa, Artemisia vulgaris, Blumea, Carduus nutans, C. crispus, C. acanthoides, Centaurea benedicta, Cirsium, Cnicus lanceatatus, C. acaulis, Filago arvenis, Gnaphalium indicum, Lappa officinalis, Senecio cineraria, Silybum marianum, and Xanthium, all members of the Compositae (order Asterales). Occasional hosts are: Althaea officinalia, Malva sylvestris, and M. rotundifolia of the Malvaceae (order Malvales); Anchusa officinales, Borago officinalis, and Echium vulgare of the Boraginaceae (order Polemoniales), and Nicotiana glauca (Solanaceae, order Polemoniales). It has even been reported upon Zornia (Leguminosae, order Rosales) and upon Urtica dioica (Urticaceae)! I believe these last two records are open to question.

Fabricius (1775, p. 499) gave brief descriptions of the larva and pupa. Life-history notes and descriptions of the immature stages are numerous. Edwards (1889, p. 26) and Davenport and Dethier (1938, p. 164) listed many of the references. Some other references are: Esper (1777, p. 136), Latreille (1824, p. 325), Boisduval and LeConte (1834, p. 179), Duponchel (1849, pp. 107–109, pl. 12: figs. 42 a-c), Morris (1862, p. 60), Trimen (1862–1866, pp. 120–121), Newman (1874, p. 64), Buckley (1886, pp. 49–52, 174–176, pl. 8: figs. 1, 1a-e), Marshall and de’ Nicéville (1886, pp. 227–228), Tutt (1886, pp. 352–354), Trimen (1887, p. 202), Dale (1890, pp. 138–140), Barrett (1893, pp. 151–154), Moore (1899–1900, p. 106), Stichel (1909, p. 200), Frohawk (1934, pp. 143–146, pl. 11: figs. 1–4), Stokoe (1944, pp. 94–96, pl. 9: fig. 3, pl. 10: fig. 3, pl. 15: fig. 1), Woodhouse (1950, p. 49), and Wynter-Blyth (1957, pp. 210–211).

DISTRIBUTION.—This butterfly is the most cosmopolitan of all butterflies being found in most parts of the world except Australia, New Zealand, and only rarely in South America south of Venezuela. It is absent in the Antartic and the Artie regions. Its migrations have attracted attention for nearly two hundred years and much of this information has been summarized by Williams (1930, pp. 168–212).

MATERIAL STUDIED.—One hundred and thirty-eight males and one hundred and twenty-one females were studied from the following localities.

NORTH AMERICA: BRITISH COLUMBIA: Duncans; Vancouver (August); Victoria (August). ALBERTA: Calgary (June). MANITOBA: Manchester (July). ONTARIO: Cedarhurst (September); Hymers. MAINE: Mount Desert Island (July). MASSACHUSETTS: Essex (July). NEW YORK: Allegany State Park; Staten Island (September). MARYLAND: College Park (August); Prince Frederick County October). DISTRICT OF COLUMBIA: (June, August). VIRGINIA: Bald Mountain (August); Limeton (October); Salem (July, August, September, October). NORTH CAROLINA: Asheville (July). SOUTH CAROLINA: Sullivans Island. NORTH DAKOTA: Tower City (August). MINNESOTA: No specific locality. MICHIGAN: Chelsey (October); Detroit (July); Saint Ignace (July). ILLINOIS: Decatur (September). MISSOURI: Webster Groves (September). NEBRASKA: Lincoln (September). KANSAS: Concordia (June); Eureka (April, June, November); Lawrence (June); Scandia (June). KENTUCKY: Frankfort (September). LOUISIANA: Baton Rouge (December). TEXAS: Brownsville; San Benito (May). WASHINGTON: Brewster (June, July); Cooney Lake (July); Godman Springs, Blue Mountains (July); Yakima (October). UTAH: Stockton (June). WYOMING: Fremont County; Yellowstone Park. COLORADO: Golden (June). NEW MEXICO: Mount Tasco (August). ARIZONA: Huachuca Mountains (September); Palmerlee, Cochise County; Readington. CALIFORNIA: LOS Angeles; San Diego (June. September); Taft (May). HAWAII: Oahu (March). MEXICO: Aguascalientes (August); Jalapa; Mazatlan, Sinaloa; Mexico City (June); Orizaba (Novvember). GUATEMALA: San Sebastian (June, 3,000 ft.). HONDURAS: NO specific locality. COSTA RICA: San Jose (January, 4,000 ft.). COSTA RICA: San Jose (January, 4,000 ft). PANAMA: NO specific locality. CUBA: Loma del Gato, Sierre del Cobre, Oriente (September, 2,600 ft); Vinales (June). HAITI: Beaumont (October).

EUROPE: FRANCE: Royan (October). SWITZERLAND: Zurich. GERMANY: Miesbach. CZECHOSLOVAKIA: Carpathian Mountains; Friedland. HUN-GARY: Budapest (July); Galgamacsa (August). ALBANIA: NO specific locality. GREECE: Athens (May); Spetsai Island (July, August).

ASIA: TURKEY: NO specific locality. SYRIA: Beyrout. JORDAN: Jordan Valley (March). INDIA: Kasmir; Khasia Hills; Madras; Sikkim (May). BHUTAN: Thimbu (September, 7,700 ft). CHINA: Foochow; Ishang; near Mupin (14,000 ft); Mount Omei, Szechuan (August); Ts'ao P'o, Szechuan (August, 6,300 ft); Tschili Province (September). JAPAN: Kotanijochi (July); Musaki (October); Oeda (September); Oita (September); Sifu; Tokosanchi (June); Tosari (July). JAVA: Buitenzorg. SUMATRA: Deli.

AFRICA: UGANDA: NO specific locality. KENYA: Kibwezi (December); Nairobi (May); Thomsons Falls (January). RHODESIA: Sinoia. NYASALAND: NO specific locality. MADAGASCAR: NO specific locality.