Higher Systematics of the Pentatomoidea 73
but their description and illustrations clearly indicate that they had a species of Palomena Mulsant and
Rey (Nezarini).
In spite of the small number of included taxa, this is a morphologically rather heterogenous group dif-
ficult to characterize. Specimens of this tribe tend to be medium to large in size; they are usually pale
yellowish-brown to dark brown (Figure 2.28F). The head is relatively large and usually distinctly trian-
gular in shape; the juga usually extend beyond the apex of the tylus and usually meet anterior to the tylus;
the lateral margins of the head are not reflexed. The rostrum reaches to and usually beyond the hind coxae.
The thoracic sterna are not distinctly sulcate, the prosternum is flat or shallowly concave, the mesoster-
num is distinctly carinate medially, and the metasternum is flat (note that the metasternum is relatively
small due to the nearly contiguous placement of the mid and hind coxae). The ostioles are relatively large;
the ostiolar rugae are quite long and slender, curving slightly cephalad, with the apices acuminate; the
evaporative area is relatively large. The anterolateral pronotal margins are often edged (obscurely so in
Halyabbas) and not reflexed; they are minutely toothed in Sennertus. The tarsi are three-segmented. The
abdominal venter is unarmed basally, although this area may appear somewhat swollen or hump-like.
Very little biological information is available for this tribe. Bolaca unicolor (Walker) has been
recorded from Pinus yunnanensis Franchet (Pinaceae) (Shi and Xu 2006) and Halyabbas unicolor has
been recorded from bamboo (Poaceae) (Hoffmann 1931, Zheng and Zou 1982, Zheng 1994, Wang et al.
2002) and sandalwood (Santalaceae) (Chatterjee 1934).
2.2.10.7.8 Antestiini Distant, 1902
Atkinson (1888) is actually the first worker to recognize this group at the family level, proposing two
names that currently belong here. His Hyllaria was based on the genus Hyllus Stål, 1868, which is preoc-
cupied by an arachnid genus of the same name described by Koch in 1846. A family-group name cannot
be based upon a preoccupied generic name, so the Hyllini is invalid. Bergroth (1891) recognized this
homonymy and proposed the generic name Anaca as a replacement name. The second family-level name
proposed by Atkinson was the Plautiaria, based on the genus Plautia Stål. As we discuss later in this
section, the tribal placement for Plautia is not certain, but if it does belong here, it could compete for
priority with Antestiaria Distant (1902). However, due to the uncertain placement of Plautia, we prefer
the continued usage of Antestiini for this tribe. Distant (1902) erected the Antestiaria to hold four genera,
two of which are the two discussed above (Anaca and Plautia); the other two were Apines Dallas and
Antestia Stål. The genus Apines is currently considered to be a member of the Menidini.
This is a relatively large tribe containing 29 genera and 171 species (Table 2.3). Linnavuori (1982)
divided the African genera into six subgroups. There is considerable variation in the color and structure
of members of this tribe. Usually they are small to medium in size, occasionally they are quite large (e.g.,
Porphyroptera); they are for the most part, robust, broadly ovate in shape. They tend to be somewhat
shiny dorsally; punctures are present, usually dark and may be coarse, dense or not. The coloration is
usually green or greenish (Figures 2.19E, 2.28G), but some species may have bright orange or reddish
markings on the dorsum (Figures 2.19D, 2.28H). The lateral margins of the head are margined, often
narrowly reflexed; the juga and tylus are usually subequal in length. The anterior pronotal margins and/
or the posterolateral pronotal margins are narrowly, but distinctly, reflexed. The scutellum is somewhat
broad at the base (basal width and medial length are subequal) but narrows to a narrowly rounded apex.
The mesosternum has a low but distinct medial carina. The ostiolar rugae are elongate, each reach-
ing beyond the middle of the metapleuron, and are usually acuminate apically; the associated evapora-
tive area is relatively extensive, distinct (except Bergrothina Schouteden). The base of the abdomen is
unarmed or at most slightly humped.
According to Gross (1976), the male parameres are C-shaped or T-shaped. The phallosoma of the
aedeagus may be sclerotized or not, and a narrow collar-like thecal shield may be present or absent;
the medial penial plates are simple and may be attached to the conjunctiva. The spermathecal bulb in
females is rounded with two long tubular processes (tubules are lacking in a few species of several dif-
ferent genera).
The placement of the genus Plautia Stål has been problematic and may bridge the gap between this
tribe and the Nezarini. Gross (1976) stated that “on the shape of the claspers and aedeagus I am con-
vinced that Plautia belongs, with a series of other genera from our region, to a later grouping here called