Nezara viridula ( L .) 355
(Slovenia, France, Greece, Italy, Japan, Guadeloupe, Galápagos, California, Brazil, and Botswana) and
supported the ‘out-of-Africa’ hypothesis for the origin of N. viridula followed by dispersal to Asia and a
more recent expansion to Europe, the Americas, and Oceania. They also suggested that the African and
non-African gene pools have been separated for 3.7–4.0 million years according to the standard insect
molecular clock of 2.3% pairwise divergence per million years. However, to better understand the origin
of N. viridula and its related species, a more comprehensive study, including integrative data sources
(e.g., molecular, pheromones, reproduction, bioacoustics) of all species of the Oriental clade (including
N. antennata and N. yunnana) and some of the African species is required (e.g., Aldrich et al. 1987, Čokl
A BCEFGHDFIGURE 7.1 (See color insert.) Nezara viridula eggs, nymphs, and adults. A, egg mass at ≈2 days old (from laboratory
culture); B, fifth instars and adults feeding on fruit of Cucurbita foetidissima; C, mating adults on Rapistrum rugosum
(B–C in Central Texas); D, adults expressing varying degree of overwintering russet coloration in Kyoto, Japan; E, (l. to
r.) – representative adult color morphs – form smaragdula (common all green form, form G-type), form torquata (O -type),
form viridula (R-type), and form “OR-type” resulting from cross of O-type and R-type; F, form “OY-type” resulting from
cross of O-type female and either a G-type or O-type male; G–H, several other color morphs include blue, orange, and
black forms. (Images A–C, Courtesy of Jesus F. Esquivel, USDA, Agricultural Research Service, College Station, TX;
D, Courtesy of Dmitry L. Musolin, Saint Petersburg, Russia; E–F, Adapted from K. Ohno and Md. Z. Alam, Applied
Entomology and Zoology 27: 133–139, 1992, with permission; G, Blue form and orange form aurantiaca, courtesy of
Antônio R. Panizzi, Passo Fundo, RS, Brazil; H, Black form, modified from J. F. Esquivel, V. A. Brown, R. B. Harvey, and
R. E. Droleskey, Southwestern Entomologist 40: 649–652, 2015, with permission.)