Higher Systematics of the Pentatomoidea 55
convergent adaptation for avoidance of parasitoids that are known to key on the scent to find their hosts.
Moreover, at least one genus of asopines, Tylospilus Stål, has species that are at least partially, and pre-
sumably secondarily, herbivorous (Stoner et al. 1974), and this latter genus has the least crassate rostrum
in the subfamily.
Although predation, and its accompanying adaptations, could result from convergence, important syn-
apomorphies of the genitalia strongly support asopine monophyly. Although neither feature, the thecal
shield (apical part of theca sensu Gapon and Konstantinov 2006) or the genital plates (parandria sensu
Tuxen 1970) is unique to the asopines, these bugs are the only pentatomids that have both in combination
(McDonald 1966).
Members of the Asopinae are found in both the Old and New World and also in the Pacific islands.
The Old World fauna consists of 40 genera (Thomas 1994b) and the New World 26 (Thomas 1992b),
with four genera found in both. Gapon (2010a) added a new New World genus bringing the total to 63
genera, with 303 species worldwide (Tables 2.2, 2.3). The exact number of species is in doubt because
many of the larger Old World genera have not been revised recently. Nomenclatural notes and a check-
list of the Chinese species was provided by Rider and Zheng (2002). When making identifications of
asopine specimens, the two monographs by Thomas should be consulted. His 1992b publication pro-
vides keys to the genera and species of the New World taxa; his 1994 work provides keys to the Old
World genera and checklists of the known species. Also, the Euro-Mediterranean taxa were treated by
Péricart (2010). Two older papers that still provide very useful information for identifying taxa are those
by Schouteden (1905a, 1907). The Chinese species of the genera Amyotea Ellenrieder and Picromerus
Amyot and Serville have been reviewed by Zhao et al. (2011, 2013) respectively. Other genera that have
been reviewed, either in part or in whole, include Anasida Karsch (Ahmad and Rana 1991), Arma Hahn
(Ahmad and Önder 1990a, Zheng 1981), Blachia Walker (Ahmad and Rana 1994), Canthecona Amyot
and Serville (Ahmad and Rana 1988, Khuong and Lam 2001), Oechalia Stål (Usinger 1941, 1942),
Perillus Stål (Knight 1952), Picromerus (Ahmad and Önder 1990b), and Rhacognathus Fieber (Josifov
and Kerzhner 1978).
The tribal classification is unresolved. Various suprageneric names have been proposed but not in
the context of a formal classification. The oldest name applied to the group, Spissirostres, by Amyot
and Serville (1843) was not based on a genus name and, therefore, has no significance in tribal clas-
sification. Kirkaldy (1909) used the name “Cimicinae” for the group based on the genus Cimex, but a
subsequent ICZN decision restricted this name to the bed bugs and, thus, cannot apply to Asopinae.
The name Asopinae, based on the genus Asopus Burmeister, was proposed by Amyot and Serville
(1843) as “Asopides.” Tribal names that have been proposed, and their type genera, are provided in
Table 2.4.
When Amyot and Serville (1843) divided their “Spissirostres” into two subordinate taxa (i.e., Asopides
and Stirétrides), they were following Burmeister’s (1835) Handbuch that recognized three genera:
Stiretrus Laporte, Discocera Laporte, and Asopus. Their “Asopides” included only Asopus, and their
“Stirétrides” included Stiretrus and Discocera. Thus, Kirkaldy’s (1909) selection of Asopus gibbus
Burmeister, a junior synonym of Discocera cayennensis Laporte, as the type-species of Asopus, was
unfortunate as it rendered Asopini, Stiretrini, and Discocerini as synonymous. Kirkaldy was a strict
priorist and his selection serves as an excellent example for not always adhering to strict priority.
TABLE 2.4
Proposed Asopine Tribal Names, Authority and Nominate Genus
Asopini Amyot & Serville (1843) based on Asopus Burmeister
Stiretrini Amyot & Serville (1843) based on Stiretrus Laporte
Armini Bergroth (1904) based on Arma Hahn
Discocerini Schouteden (1907) based on Discocera Laporte
Amyotini Schouteden (1907) based on Amyotea Ellenreider
Jallini Dupuis (1949) based on Jalla Hahn
Stilbotini Gapud (2015) based on Stilbotes Stål