Higher Systematics of the Pentatomoidea 137
and biological information available for the Neotropical species was compiled by Matesco and Grazia
(2015). There are currently 30 genera and 223 species in this family (Table 2.2). The systematics of this
family has been studied as a thesis project (Matesco 2014); a recent review of the genus Alkindus Distant
also has been provided (Matesco and Grazia 2013).
All species of this family are phytophagous, with many different host plants reported for various spe-
cies. According to Štys and Davidová (1979), the most preferred host plant for the European Thyreocoris
species is Viola tricolor L. (Violaceae). Feeding, especially in Corimelaena spp., occurs mainly in
the reproductive parts of the hosts, such as flowers and developing fruits (Schaefer 1988). McPherson
(1971, 1972) observed preference of Corimelaena lateralis (F.) for the mature parts of Daucus carota
L. (Apiaceae) rather than the inflorescence; however, such a preference was not observed in C. pulicaria
(Germar) (McPherson 1972). Polyphagy seems to be the rule with some species recorded from host plant
associations from more than ten plant families; yet host specialization may occur in some species.
Thyreocorids may be uni-, bi-, or multivoltine (McPherson 1972; Lung and Goeden 1982; Bundy
and McPherson 1997, 2009). Adults overwinter under litter, soil, or stones. Precopulatory behavior was
described by Bundy and McPherson (1997), and copulation may take several hours. Thyreocorids lay
eggs singly, glued laterally to the substrate, often the reproductive parts of the host plant (less frequently
on other parts of the host plant). Egg development varies from eight to eleven days, depending on the spe-
cies and the temperature and humidity. First instars do not show gregarious behavior. Nymphal develop-
mental time varies from 30 to 45 days, influenced by diet, temperature, and humidity. Several species of
tachinids (Diptera) and parasitic wasps (formerly Scelionidae, now Platygastridae in the Hymenoptera)
have been recorded as attacking thyreocorids. Occasionally, thyreocorids can occur in rather large
numbers in their natural habitats (McPherson 1974, Mendonça et al. 2009). In field studies in southern
Brazil, thyreocorids were especially abundant, and they were one of the most speciose groups collected
(Schmidt and Barcellos 2007, Mendonça et al. 2009).
2.2.18.1 Key to the Subfamilies of Thyreocoridae (modified from Schuh
and Slater 1995)
1 Hind wing with jugal lobe entire (Figure 2.11E); Eastern Hemisphere (Figure 2.25K) ...........
..............................................................................................................................Thyreocorinae
1’ Hind wing with an oval perforation in jugal lobe (Figures 2.11F, G); Western Hemisphere .....
.............................................................................................................................Corimelaeninae
2.2.19 Urostylididae Dallas, 1851
Urostylidids are an Old World group occurring from India through the Oriental region and into Japan
and Southeast Asia. The family currently contains eight genera and 172 species (Table 2.2). They are
variously colored, frequently green to yellowish green (Figure 2.25L), but many species are contrasting
brown and yellow, others are aposematically red and black, resembling pyrrhocorids, and still others
have a complex color pattern. They are medium in size (8 to 15 mm), and, in general, they look more
like a coreoid or pyrrhocoroid than a pentatomoid. They share several unique characters with the closely
related family Saileriolidae such as more dorsal antennal insertions and ocelli that are situated close
to each other. They differ, however, in several important characters. For example, in urostylidids, the
abdominal spiracles are all ventral (they are situated close to the lateral margins of the abdomen in saile-
riolids), the scent gland rugae are distinct and spoutlike (external scent efferent structures are strongly
reduced or absent in saileriolids), and a hamus is present in each hind wing (not present in saileriolids);
in addition, saileriolids are much smaller (less than 5 mm in length). Most of the early work on this group
was under the family name Urostylidae, which was homonymous with Urostylidae in the Ciliophora.
Berger et al. (2001) emended the name of the heteropteran family to Urostylididae to remove this hom-
onymy. The Palearctic species recently have been catalogued (Rider 2006c). At least six fossil species
have been described, all in the genus Urochela (Zhang 1989).
Members of this family can be characterized further by a rather short, declivent head with rounded
margins (not edged or reflexed). The antennae are five-segmented with segment I relatively long, reaching