Higher Systematics of the Pentatomoidea 75
margins of the propleura are not swollen; the mesosternum is distinctly carinate medially; and the meta-
sternum is swollen and produced posteriorly, meeting with the basal abdominal spine. The ostiolar rugae
are short and spout-like.
In general, axiagastine species have some superficial resemblance to several genera placed in the
Menidini or Eurysaspini, but they lack the apically free, basal spine on the abdomen. In fact, Distant
(1902) treated Acesines in the Menidaria and Axiagastus in the Hoplistodaria. And as mentioned previ-
ously, Gross (1976) indicated that Axiagastus might be related to the Antestiini.
Axiagastus cambelli Distant (Figure 2.28I) is considered to be an occasional pest on coconut palm in
the Bismarck Archipelago (Smee 1965, Lever 1969, Baloch 1973).
2.2.10.7.10 Bathycoeliini Atkinson, 1888
Stål (1876), without formally naming it, first recognized this group at a higher level when he referred to
it as ”Bathycoelia et affinia.” Atkinson (1888), following Stål, established the Bathycoeliaria for three
genera: Abeona Stål, Bathycoelia Amyot and Serville (Figures 2.19F, 2.28J), and Jurtina Stål. Bergroth
(1913) placed Jurtina as a junior synonym of Bathycoelia. Abeona is a junior homonym of Abeona
Girard, 1855, a genus in the Pisces; Bergroth (1891) recognized this homonymy and proposed Amblycara
as a replacement name. Amblycara currently is considered to be a member of the Pentatomini, but Tsai
and Rédei (2014) speculated that Amblycara and Bathycoelia are closely related (see discussion later in
this section) and should probably be placed together in the same tribe.
This tribe at present contains a single genus, Bathycoelia, and 32 species (Table 2.3). Most spe-
cies occur in Africa and adjacent islands (Madagascar, Mascarenes, Seychelles, Socotra), but other
species occur in India, China, Southeast Asia, and many islands of Malesia and Oceania. The ability
of species of this genus to colonize remote islands and evolve endemic species is quite remarkable. Two
species were described from South America (Jensen-Haarup 1931a, 1937), but these are probably mis-
labeled; Rolston et al. (1980) examined the types for both species and confirmed that they belong in the
genus Bathycoelia.
Most species are greenish with black or red spots or lines on the connexiva and along the margins of
the head and anterolateral margins of the pronotum (Figure 2.19F); the basal angles of the scutellum
are often black, foveate, or sometimes have a pale yellowish callus (Figure 2.28J). The head is usually
triangular to subtriangular in shape, flattish, with the juga and tylus subequal in length. The anterolat-
eral pronotal margins are usually narrowly to broadly reflexed, and the anterior margin also is beaded,
forming a collar posterior to the vertex that disappears laterally behind the eyes. The humeral angles
are usually rounded to angulate, never spinose. The prosternum is shallowly concave, the mesosternum
is distinctly carinate medially, and the metasternum is usually elevated, somewhat tectiform. The ostio-
lar rugae are quite elongate, curving cephalad, extending over three-quarters of the way to the lateral
metapleural margin, becoming sharply acuminate apically; the associated evaporative area is extensive
(Figure 2.9C). The connexiva are often distinctively patterned. The abdominal venter is usually dis-
tinctly furrowed longitudinally along the median with the elongate rostrum fitting into the abdominal
groove (Figure 2.8A). The posterior margin of the male pygophore is usually emarginate medially, usu-
ally with variously shaped or produced lobes on each side of emargination; dorsal plates are often pres-
ent, sometimes well-developed. The parameres are somewhat T-shaped. In the females, the spermathecal
bulb is usually broader than long and provided with two diverticula of unequal lengths.
Linnavuori (1982) indicated a possible relationship with the antestiines with which they share the
greenish coloration and the elongate ostiolar rugae. Bathycoelia horvathi Schouteden (western Africa)
seems to represent a separate lineage, being smaller and more robust; its coloration is a deeper green, and
the abdominal sulcation is less developed. It may not be congeneric with other Bathycoelia species. More
recently, Tsai and Rédei (2014) discussed the relationship between the genera Amblycara and Bathycoelia.
Amblycara typically has been considered to be a member of the Pentatomini, but they proposed that
Amblycara might be closely related to Bathycoelia based on the genitalia and the fact that both genera
have a ventral sulcus on the abdomen. They stated “Bathycoelia is not of such an isolated position which
would justify a placement in a tribe of its own, or even with Amblycara,” but they chose to not formally
synonymize the Bathycoeliini with the Pentatomini until a more conclusive study could be conducted.