76 Invasive Stink Bugs and Related Species (Pentatomoidea)
Bathycoelia thalassina (Herrich-Schäffer) is considered to be a major pests of cacao in tropical Africa
(Gerald 1965; Lodos 1967; Owusu-Manu 1971, 1976, 1977, 1990), and B. natalicola Distant (= B. dis-
tincta Distant) has been recorded as a pest on both avocado (Erichsen and Schoeman 1992; van den Berg
et al. 1999b, 2000) and macadamia nut in southern Africa (van den Berg et al. 1999a, Schoeman 2013).
2.2.10.7.11 Cappaeini Atkinson, 1888
This group was recognized first, but not formally named, by Stål (1876) who referred to it as “Cappaea et
affinia”; he originally included three genera Cappaea Ellenrieder (Figure 2.28K), Halyomorpha Mayr
(Figure 2.28L), and Tolu m nia Stål. In 1888, Atkinson formally erected two tribes, the Tropicorypharia
(based on Tropicorypha Mayr, currently included in this tribe), and the Cappaearia (based on Cappaea).
Distant (1902) treated both tribes as synonyms of the Carpocorini, but most recent workers have recog-
nized the Cappaeini as a valid tribe (the exception being Zaidi and Shaukat, 1993, who also treated both
of the above tribes as synonyms under the Carpocorini). Members of the Cappaeini tend to be brown in
color (several species of Caura Stål and Halyomorpha are dark bluish), lack a spine or tubercle on the
base of the abdomen, and have ostiolar rugae that are elongate, tapering to a point apically. This tribe
contains 24 genera and 151 species (Table 2.3), all of which are restricted to the Old World with the
exception of Halyomorpha halys (Stål) (Figure 2.28L), which has been introduced accidentally into
and is spreading throughout North America, and also has become established in Chile, South America
(Faúndez and Rider 2017). Linnavuori (1982) dealt with the African genera included here in the Veterna
group and the Halyomorpha group.
The head is typically shaped, usually subtriangular (elongate, spatulate in Tropicorypha), with the juga
and tylus subequal in length (juga longer than and meeting in front of tylus in Benia Schouteden and
Tripanda Berg). The body shape is often broadly rounded, somewhat robust, convex below. The antero-
lateral margins of the pronotum may be narrowly reflexed or not, and the anterior margin is usually flat
behind the head (a row of punctures may form a transverse ridge behind the vertex in some species of
Tolu m nia). The humeral angles are usually rounded (Figures 2.28K, L) but may be weakly spined (some
species of Caura) to strongly spinose (some species of Veterna Stål [Figure 2.19H]). The prosternum is
shallowly to distinctly sulcate medially, the mesosternum is distinctly carinate medially, and the meta-
sternum is generally flat. The ostiolar rugae tend to be quite elongate, usually curving cephalad, and
sharply acuminate apically (more rounded apically in Benia, Tripanda, and Tropicorypha, and relatively
short in Veterna). The abdominal venter is unarmed.
Linnavuori (1982) described the male pygophore as having excavations and lobe-like processes, and
the parameres are rather short, incrassate, and variously modified. The female spermatheca is ball-
shaped without diverticula (Benia, Mabusana Distant, Tripanda subgenus Te n er va Cachan), or with two
diverticula (Boerias Kirkaldy, Paralerida Linnavuori, Tripanda sensu stricto, Veterna) or three diver-
ticula (Caura, Lerida Karsch); or not ball-like but rather with an irregular bulb (Lokaia Linnavuori).
There seems to be some overlap in characters with members of the Carpocorini.
Several cappaeinine genera have received attention in recent years. For example, Cappaea was stud-
ied by Zaidi (1993), Caura was revised by Leston and Dutton (1957), a second species was added to
Massocephalus Dallas (Rider 2008), Prytanicoris Gross was recently described and reviewed (Gross
1978), the Indian species of Tolu m nia have been treated (Zaidi 1995), and the genus Tripanda recently
has been revised (Kment and Jindra 2009).
One species in particular has been the focus of much study recently - Halyomorpha halys (Figure
2.28L). This species, native to eastern Asia, has been introduced accidentally into Europe, North
America, and South America and is spreading quickly through all three continents. It attacks several
fruit trees and vegetable crops and is considered to be a nuisance because it congregates around and
enters homes in the fall for overwintering (see Chapter 4).
2.2.10.7.12 Carpocorini Mulsant and Rey, 1866
Mulsant and Rey (1866) proposed three family-level names that we now treat under the tribe Carpocorini.
The three were the Rubiconiaires, based on the genus Rubiconia Dohrn; Aulacetraires, based on
the genus Aulacetrus Mulsant and Rey; and Carpocorates, based on the genus Carpocoris Kolenati.
Rubiconia was transferred to the Eysarcorini (Puton 1869), but most current workers treat it as a member