The gut of termites (Isoptera) comparative anatomy, systematics, phylogeny. II. - Higher termites (Termitidae)

The gut of the Termitidae is much more diverse than that of the previously studied "lower" termites (NOIROT, 1995). Nevertheless the monophyly of this family is confirmed. The characters of the mesentero-proctodeal junction (presence and morphology of the mixed segment) and of the Malpighian tubules (attachment, fusion, closure, presence of a "Malpighian knot") seem especially significant. The gut allows the definition of twelve groups of genera. The Macrotermes group is equivalent with the subfamily Macrotermitinae. The Apicotermes, Anoplotermes and Speculitermes groups make the Apicotermitinae, the monophyly of which is also confirmed. Among the Nasutitermitinae the genera with mandibulate soldiers (Syntermes group) are very distinct from the genera with nasute soldiers (Nasutitermes and Subulitermes groups), which suggests a paraphyly of the Nasutitermitinae. The genera now included in the subfamily Termitinae can be distributed into five groups. The Cubitermes group is very homogeneous. The genera having soldiers with "snapping" mandibles could make two distinct groups: Termes and Pericapritermes. The Amitermes group seems more heterogenous. The gut of the Foraminitermes group is idiosyncratic (new subfamily?). Thus the Termitinae don't make a clade. At least the Foraminitermes group must be removed. Finally a phylogeny of the Termitidae is proposed. As it is based essentially on the gut characters it must not be regarded as more than a working hypothesis.