Higher Systematics of the Pentatomoidea 127
nymphal stage, which is unusual for European Pentatomoidea. Moreover, they may overwinter in any of
the first through fourth instars, depending on the weather and temperature during the preceding autumn.
All plataspids are phytophagous (mostly oligophagous or polyphagous). There seems to be a prefer-
ence of many plataspids for feeding on members of the plant family Fabaceae. The long and partially
coiled stylets (somewhat similar to some Aradidae) led China (1931) to suggest that plataspids might
feed on fungi, but more recent evidence involving a symbiotic relationship between plataspids and ants
in which the ants tended and collected honey dew from the plataspids (see below) seems to indicate that
at least these species of plataspids are phloem feeders, needing the long stylets to penetrate the bark of
their host plant.
Mutualistic relationship with ants has been reported in several plataspids, including Caternaultiella
rugosa Schouteden (tropical Africa), Tropidotylus servus Maschwitz, Fiala, and Dolling, T. minis-
ter Maschwitz, Fiala, and Dolling, Tetrisia vacca Webb, and Hemitrochostoma lambirense Bergroth
(Malaysia) (Maschwitz et al. 1987, Gibernau and Dejean 2001, Waldkircher et al. 2004, Tomokuni 2012).
The ants build pavilions to protect and shelter the egg batches, nymphs, and adults of the bug. In at least
C. rugosa, the adult female bugs are able to switch the maternal behavior as needed. That is, if there
are more egg batches outside of the pavilions, then the females will guard the eggs; if most of the eggs
are within the ant pavilions, then the adult females do not help with guarding of the eggs (Gibernau and
Dejean 2001).
Similarly to most other pentatomoids, plataspid females transmit symbionts to their offspring via the
eggs. However, they do it somewhat differently. In these bugs, the symbionts are contained in small,
dark capsules deposited between the individual eggs of the egg batch. This phenomenon, a potentially
unique apomorphy of the family, has been reported for the Palearctic species, Coptosoma scutella-
tum (Geoffroy) (Davidová-Vilímova 1987) and a number of East Asian species belonging to the gen-
era Coptosoma Laporte, Brachyplatys Boisduval, and Megacopta Hsiao and Jen (e.g., Hosokawa et al.
2006, 2008). The newly hatched nymphs feed on the capsules. The egg batches of at least two species
of Libyaspis Kirkaldy are nearly unique within the Heteroptera. The eggs are covered by an ootheca of
material originating from the midgut. A similar ootheca is known in some species of Reduviidae and
Urostylididae, but, in these examples, the oothecal material is a secretion produced by parts of the female
genital duct (Carayon 1949, 1952).
Some plataspids (e.g., Coptosoma lyncea Stål, from northern Australia) also are known to form large
aestivation aggregations (Monteith 1982). One such aggregation was described as covering every leaf of
a small tree, including lines of individuals on the petioles. When disturbed, the individuals would take
flight and disperse their defensive secretions but eventually would return to their previous roost.
At least one fossil species, Coptosoma eocenica Piton (1940), has been described.
The phylogeny and higher classification of the family is unresolved. Jessop (1983), based primarily on
somatic characters, first classified the plataspid genera into three suprageneric categories but refrained
from formally giving them tribal names (Brachyplatys group, Coptosoma group, Libyaspis group);
he also keyed the genera of the Libyaspis group. Ahmad and Moizuddin (1992) split the South Asian
Plataspidae into two subfamilies: Plataspinae (incorrectly as Brachyplatidinae) and Coptosomatinae
(incorrectly as Plataspidinae); the names were corrected by Davidová-Vilímova (2006) in the Palearctic
catalog. This classification cannot be adapted to the World fauna and it is difficult or impossible to place
many genera to any of the described subfamilies (Davidová-Vilímová 2006). As such, the key presented
below is based merely on the literature and will not allow placement of several described genera.
2.2.12.1 Key to the Subfamilies and Genus Groups of Plataspidae
(modified from Schuh and Slater 1995)
1 Ocelli placed laterally, near compound eye or anterior angles of pronotum (Figure 2.11A);
abdominal sterna usually clearly convex; head usually narrow, about 0.3-0.5 times width
of pronotum; base of scutellum (“pseudoscutellum”) usually elevated, set off from rest of
scutellum by an impressed line (Coptosomatinae) .....................................Coptosoma group