Higher Systematics of the Pentatomoidea 129
Schaefer and Ashlock (1970) gave a summary of the similarities between the Urostylididae and the
Saileriolidae. They stated that the two groups “have in common the placement of the antennae (dorsal
to the midline of the eye), the annulate antennifers, the proximity of the ocelli, the sutures of the vertex,
and the general structure of the genital capsule.” China and Slater (1956) had noted some of these same
similarities, and they placed “great importance” on the structure of the antennifer and the position of
its insertion onto the head. Hsiao (1964) described the genus and species Bannacoris arboreus Hsiao,
placing it in the Saileriolinae, but Schaefer and Ashlock (1970) were apparently unaware of its descrip-
tion as they made no mention of it. The phylogenetic study by Grazia et al. (2008) indicated that the
Urostylididae in the traditional sense was not a monophyletic group, which was solved by moving the
Saileriolinae to its own family. Furthermore, they concluded that the Saileriolidae is the sister group to
all of the non-urostylidid Pentatomoidea.
The only published biological information available comes from the type series of Ruckesona vitrella
Schaefer and Ashlock, which Schaefer and Ashlock (1970) reported had been collected on “palm at
water margin.” The authors also noted that the green gut contents of specimens contained what looked
like fragmented chloroplasts that led them to propose that specimens might not feed exclusively on plant
sap. Specimens of Bannacoris arboreus (Figure 2.25I) were collected from leaves of banana species
(D. Rédei, unpublished data).
2.2.15 Scutelleridae Leach, 1815
The Scutelleridae was proposed by Leach (1815) as a family group, and included it together with the
Pentatomidae in a taxon currently equivalent to the Pentatomoidea. Amyot and Serville (1843) proposed
the tribe Orbiscutes in the family Longiscutes to include all genera with the scutellum enlarged. In their
classification, scutellerids were divided into five different groups (Scutellérides, Pachycorides, Tétyrides,
Eurygastrides, and Odontoscélides), but they also included within these groups a number of genera now
placed in other pentatomoid families. Dallas (1851) recognized many of the same taxa, but now at the
family level (Eurygastridae, Pachycoridae, and Odontoscelidae) within the group Scutelleroidea, but
the first and last of these also included members now placed in the Podopinae and the Thyreocoridae,
respectively. Fieber (1861) recognized only one family-group, the Tetyridae, but this still contained sev-
eral non-scutelleroid members. Stål (1865) was the first to treat this group in a fashion similar to our
present-day classification, although he considered the scutelleroids to form a subfamily of a more inclu-
sive Pentatomidae. He refined his classification in a later work (Stål 1873), which became the classifica-
tion followed by most subsequent workers (e.g., Lethierry and Severin 1893, Kirkaldy 1909, Oshanin
1912, Leston 1952a, China and Miller 1955, Lattin 1964, Linnavuori 1982). The only major change is
that Reuter (1912), without comment, elevated the scutelleroids to the family level (Tsai et al. 2011). The
works of Singh-Pruthi (1925) and Pendergrast (1957) supported the recognition of this group as a valid
family, which has been followed by most modern workers (e.g., Leston 1958, McDonald 1966, Gross
1975b, Rolston and McDonald 1979, Froeschner 1988d, Schuh and Slater 1995, Cassis and Gross 2002,
Cassis and Vanags 2006, Göllner-Scheiding 2006b, Tsai et al. 2014, Barcellos et al. 2015).
Although limited in scope and not the direct focus of the studies, explicit phylogenetic studies have
been conducted that include the Scutelleridae as a terminal taxon (e.g., Gapud 1991, Fischer 2001, Grazia
et al. 2008). In all of those analyses, whether based on morphology, molecules, or a combination of the
two, the monophyly of the Scutelleridae and its familial status were corroborated, but the phylogenetic
relationship within the Pentatomoidea remains inconclusive. Tsai et al. (2011) argue that the monophyly
of the Scutelleridae is currently weakly supported, considering no unique morphological apomorphies
are recognized, and the molecular data are too fragmentary. These same arguments can be applied to
most families within the Pentatomoidea.
Members of this family (Figures 2.26A-L) are medium to relatively large in size (5 to 20 mm in
length), ovoid to elongate ovoid in shape, and the body is moderately to strongly convex. The color var-
ies greatly among scutellerids, with some of the most brightly colored species in the Heteroptera. Many
species are polymorphic. Whether the colorations represent aposematism or mimicry has been discussed
(Tsai et al. 2011, Eger et al. 2015a). The juga and tylus are usually subequal in length. The antennae are
usually five-segmented, but sometimes (exceptionally) only four-segmented (Augocoris Burmeister) or