Danaus, Kluk, 1802

Smith, David A. S., Lushai, Gugs & Allen, John A., 2005, A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA, Zoological Journal of the Linnean Society 144 (2), pp. 191-212 : 209

CHARACTERS FOR DANAUS TAXA AND TWO TIRUMALA

SPECIES (OUTGROUPS)

Bold Roman numerals indicate datasets (I-VI); character numbers (1–67) are column headings in the matrix (Appendix 1B))

I Allozymes 1: 1 Acid phosphatase bands A/B, bands C/D, band F; (other); 2 Aldolase, band A (band C); 3 Hexokinase-1, bands F/G, band B; (other); 4 Esterase- D, polymorphic for bands B/C/D, C unique; polymorphic for bands A/B, A unique; (other); 5 Malic enzyme-2, band B; band D; band E; (other); 6 Superoxide dismutase-1, bands A/B; band G; (other); 7 Aspartate aminotransferase-1, bands F1/G; band C; (other); 8 Malate dehydrogenase, monomorphic for band S; polymorphic for bands F/S; polymorphic for bands B/S; 9 Phosphohexose isomerase, functional allele; (silent allele); 10 6-phosphogluconate dehydrogenase, band F; (other).

II Dihydropyrrolizines 2: Hairpencil secretion contains 11 hydroxy-danaidol, 12 danaidal, 13 danaidone. III Eggs 3: 14 In polar view, ribs raised and narrow, cells concave (longitudinal ribs wide and flat, cells generally convex).

IV Larvae 3: 15 Cuticular ornamentation on body surface pointed (blunt); 16 Filaments present on segment 5; 17 Subprimary labial seta lateral to seta M 1 (mesial); 18 Epipharyngeal spines between seta 1 and 2 present distally; 19 Number of campaniform sensillae on ventral surface of galea, two (one); 20 Submentum, basally, at least as far as insertion of primary setae, sclerotized (unsclerotized); 21 Submental cuticular ornamentation not extending laterally of primary setae at the level of their insertion (extending almost to edge of stipes); 22 Major hypopharyngeal spines in one regular row (in one regular row and a slightly more distal irregular row); 23 Depressed area on hypopharynx, distal to major spines, smooth (covered in small spines).

V Pupae 3: 24 Grooving on stalk deep (shallow); 25 Abdominal segments 4–7 smooth; slightly rugose; distinctly rugose; 26 Median dorsal spot on abdominal segment 9 absent (present); 27 Angle between abdominal segments 1–3 and 3–8 obtuse (approximately 90∞); 28 Transverse yellow band on abdominal third segment of exuviae; 29 ‘Monarch gold spot array’ only (additional median spots on abdominal segments 1 and 2, a lateral pair on segment 2 and a pair at the base of the hindwing); 30 Mesothorax continuous in outline with abdomen (slightly swollen dorsally).

VI Imagines 2,4-7: 31 Male with a section of vein 1 A + 2 A swollen; 32 Deep pouch containing particle- producing hairs in male hindwing cell Cu 1b; 33 Welldeveloped, lateral, ventrally directed projections from anteroventral lip of female 8th sternite; 34 Clasper with an annularly corrugated process; 35 Male alar organ formed as a pocket (in which the anterolateral ‘roof ’ or flap originates as a dorsal outgrowth); 36 Male terminalia with pseudovalves; 37 Forewing with four (three) pale pattern elements in forewing cell M 1; 38 Aedeagus long and narrow; 39 Ductus long and narrow; 40 Saccus long, somewhat swollen in dorsal aspect; 41 Hairpencils long; short; vestigial; 42 Fifth tarsal segment of mid- and hind-legs long and armed with long spines; 43 Juxta narrow; 44 Aedeagus incised dorsally or dorso-laterally (ventrally); 45 Aedeagus with well-developed lateral spines mounted on processes; 46 Aedeagus with fewer, smaller, lateral spines 4 not mounted on processes; 47 Hindwing vein Sc + R 1 curved in an even, shallow arc; 48 Androconia distributed in cell Cu 1b about the entrance of the pouched alar organ; 49 Hindwing cell elongate, c. 2 / 3 length of wing (c. 1 / 2 length of wing); 50 Forewing cell R 4 with single, large, marginal, white spot; single smaller spot; two small spots; 51 Clasper deeper than long, with one (two) curved (corrugated) processes; 52 Clasper very short, lacking postero-dorsal bulge; 53 Clasper extremely short, lacking postero-dorsal bulge; 54 Intracellular, postdiscal, pale markings in cells Cu 1, M 3, M 2, M 1 and Rs of hindwing underside; 55 Hindwing crossvein m 1 - m 2 sharply angled; 56 Course of hindwing long veins beneath black and bordered with black (marked with pale, whitish scales); 57 Series of three black spots on m-cu crossveins of hindwing; 58 4 Marginal white spots on hindwing underside in a double row (single row); 59 4 Black margin of hindwing broad (narrow); 60 hindwing marginal area pale posteriorly; 61 Wings with blue (phycobilin) pigment; 62 6 Wings with extensive areas of yellow pigment; tawny orange pigment (bb genotype in D. chrysippus ); nutbrown pigment (B- genotype in D. chrysippus ); 63 5 The apical third (or more) of the forewing upperside black with white spotting (black pigment predominantly confined to wing margins; 64 6 Central, small white patch on hindwing underside; 65 5 Hindwing predominantly white (aa genotype in D. chrysippus ); 66 5 Subapical pale spots in forewing cells M 3, M 2, M 1 & R 5 (cc genotype in D. chrysippus ); (spots absent, C- genotype in D. chrysippus ); 67 7 Subapical pale spots in forewing cells M 3, M 2, M 1 & R 5 (if present) large and fused. (L- genotype in D. chrysippus ).

References: 1 Kitching (1986); 2 Ackery & Vane-Wright (1984); 3 Kitching (1985); 4 Smith et al. (2002); 5 Smith (1998); Lushai et al. (2003a-b, 2005a); 6 Common & Waterhouse (1972); 7 Smith & Owen (1997) .