Developmental abnormalities in Glomeris marginata (Villers 1789) (Myriapoda: Diplopoda): implications for body axis determination in a myriapod.

Abnormally developing embryos (ADEs) of the common pill millipede Glomeris marginata have been investigated by means of nuclear staining and mRNA in situ hybridization. It showed that all ADEs represent cases of Duplicitas posterior, which means that the posterior body pole is duplicated. The severity of the duplication ranges from duplicated posterior trunk segments in one specimen to an almost completely duplicated specimen that only shares the very anterior head region. Remarkably, none of the encountered ADEs represents a case of Duplicitas anterior (duplicated anterior pole) or a case of Duplicitas cruciata (cruciate duplication with two anterior and two posterior poles). This observation is discussed in the light of earlier reports on G. marginata ADEs that claim to have found these abnormalities. The lack of any other axial abnormality aside from D. posterior implies that early axis determination in G. marginata, and possibly myriapods in general, underlies the developmental mechanisms that prevent the formation of any other type of axial duplication. It is proposed that the formation of D. posterior-type embryos could be caused by the formation of two instead of only one posterior cumulus early during development.