Higher Systematics of the Pentatomoidea 43
diagnoses and keys for identifying all genera and species. A checklist of Old World taxa was provided
by Lis (1990), a World catalog was provided by Rolston et al. (1996), and the Palearctic species also have
been catalogued (Lis 2006b). Information on the Neotropical species have been compiled by Schwertner
and Grazia (2015).
There are at least two fossil pentatomoids possibly belonging to this family: Dinidorites margiformis
described from Eocene deposits in Colorado (Cockerell 1921) and an unnamed fossil reported from
deposits in British Columbia (Archibald and Mathewes 2000), Canada, which was originally placed in
the subfamily Megymeninae.
Diagnostic characters of the family (Schuh and Slater 1995, Schwertner and Grazia 2015) include the
lateral margins of the head, which are usually carinate; and the bucculae, which are short and elevated,
almost flap-like. The antennae may be four or five-segmented, some segments may be flattened (Figure
2.2F), and the rostrum usually reaches to or beyond the middle coxae. The humeral angles of the pro-
notum are rounded, almost never developed, but the lateral margin usually bears an anteriorly directed
projection in the Megymeninae (Figure 2.16D). The scutellum is usually somewhat triangular in shape,
with the basal width subequal to the medial length, and it never covers the corium; the apex is usually
rounded. The hemelytral membrane usually has reticulate venation. The tarsi may be two- or three-
segmented; the tibiae lack distinct bristles or spines, and the coxae lack the coxal combs evident in the
Cydnidae and related families. The female spermatheca lacks a dilation and sclerotized rod. Eight of
twelve species that have been karyotyped have a diploid number of 12 + XY (Ueshima 1979, Kerzhner
et al. 2004, Rebagliati et al. 2005).
Typically, this family is divided into two subfamilies (i.e., Dinidorinae and Megymeninae) (Table
2.2), each with quite different facies. The dinidorines are somewhat rounded to oval, colored in browns,
blacks, and tans, occasionally with red or yellow, but are relatively smooth surfaced with typical pentato-
moid punctures. The megymenines are usually gray to dark grey or almost black, not quite so smoothly
ovoid in shape, and their dorsal surface is rather rough or granulated (Figure 2.16D). Eumenotes
Westwood (Figure 2.16C) and Afromenotes Kment and Kocorek, which superficially resemble members
of the Megymeninae, are of uncertain placement. It may be justified to place them in a third subfamily,
the Eumenotinae, or keep them as a tribe in either of the other two subfamilies.
The family Dinidoridae includes 17 genera and 109 species in two subfamilies and five tribes
(Table 2.2), distributed mostly in the Old World. The only representative in the New World is the
nominotypical genus, Dinidor, with six endemic species. The classification used in this chapter was
established by Durai (1987), followed by Rolston et al. (1996) in their World catalog of the group,
and includes updates from Kocorek and Lis (2000) and Lis et al. (2015). In the first work, Kocorek
and Lis (2000) proposed a new tribe (Byrsodepsini [Figure 2.16B]) within the Megymeninae and
placed the Eumenotini as a junior synonym of the Megymenini. The Eumenotini has had a complex
taxonomic history, at one time or another, having been classified within the Aradidae, Pentatomidae,
Tessaratomidae, Dinidoridae, or as a distinct family, the Eumenotidae (see Kment and Kocorek 2014).
The molecular analysis by Lis et al. (2012a) revealed that the Eumenotini is more closely related
to the Dinidorinae than to the Megymeninae, which resulted in its removal from the synonymy of
the Megymeninae and restoring it as a tribe without subfamily assignment. In the second work,
Lis et al. (2015) conducted a molecular investigation of the Madagascan genus Amberiana Distant.
Interestingly, the molecular data indicated that Amberiana was related to the cydnid subfamily
Sehirinae or the family Parastrachiidae, but the authors stated that the morphology was typical dini-
dorid; they ultimately decided to leave Amberiana in the Dinidoridae, but they erected a new tribe,
the Amberianini, for this single genus.
Gapud (1991) considered the Dinidoridae and Tessaratomidae as sister groups based on two syn-
apomorphies: the spiracles on abdominal segment II at least partially exposed (Figure 2.2B) and
laterotergites IX quite large in females. Grazia et al. (2008) found similar results, although they felt
that the Dinidoridae could be paraphyletic or monophyletic according to the analyses they performed
(morphological, molecular, or combined analyses). The sister-group relationship of Dinidoridae and
Tessaratomidae was supported morphologically (Kment and Vilímová 2010a) and by molecular data
(Lis et al. 2012a,c). The two families together certainly form a single monophyletic assemblage, but the
monophyly of each of them, particularly the Tessaratomidae, needs further support.