4 Invasive Stink Bugs and Related Species (Pentatomoidea)
or jewel bugs), Thyreocoridae (black bugs or ebony bugs), and, recently, Plataspidae (plataspids). Most
species in these families are phytophagous, the major exception being the asopine pentatomids, which
are predaceous. Within each of these families are species that cause economic injury to crops in the New
World, Old World, and worldwide (Schuh and Slater 1995, Schaefer and Panizzi 2000, Eger et al. 2010,
Ruberson et al. 2013). The Pentatomidae, largest of the six pentatomoid families, contains the highest
number of economically important species (Schuh and Slater 1995, Schaefer and Panizzi 2000).
1.2 Classification Overview
The Pentatomoidea are members of the order Hemiptera and suborder Heteroptera (true bugs). The
Heteroptera are recognized by a segmented beak that arises from the front of the head; and wings
that, when present and well developed, have the first pair leathery basally and membranous distally
(Figure 1.1A), the second pair membranous, with both pairs lying flat on the abdomen. The other sub-
orders (i.e., Auchenorrhyncha, Sternorrhyncha, Coleorrhyncha) have the segmented beak appearing to
arise ventrally from the rear of the head or between the front coxae; and wings, that when present and
well developed, usually are held rooflike over the abdomen with both pairs of uniform texture throughout.
As with most other insect groups, the higher classification of the Pentatomoidea has changed con-
siderably over time. Various families and subfamilies have been raised, lowered, then raised again
between those two categories on a number of occasions. With few exceptions, the family and subfam-
ily classification now seems relatively stable (see Grazia et al. 2008). However, the tribal classification
remains in a state of chaos and requires a thorough phylogenetic analysis. The higher classification of the
Pentatomoidea is discussed in Chapter 2. A list of the families, subfamilies, and tribes currently in use
is presented along with notes concerning how these taxa are defined. The generic groups of Gross (1975,
1976) and Linnavuori (1982) also are discussed. Keys are provided to the families of the Pentatomoidea
and subfamilies and tribes of the Pentatomidae. Some preliminary speculation is given on the validity of
these taxa and their phylogenetic relationships.
1.3 Biology
As noted above, most pentatomoids are phytophagous. Phytophagous pentatomoids feed on a wide vari-
ety of fruit, vegetable, nut, and grain crops as well as wild hosts. Generally, the bugs feed on roots,
stems, and leaves but most often are associated with developing seeds, fruits, or growing shoots. Adults
and nymphs obtain nutrients by piercing the plant tissues with their mandibular and maxillary stylets.
ABCDFIGURE 1.1 Adult and eggs of a typical stink bug, Chlorochroa ligata. A, Adult; B, Egg mass; C, Individual egg;
D, Empty egg showing displaced pseudoperculum. (Modified from Morrill 1910).