Structural aspects of leg-to-gonopod metamorphosis in male helminthomorph millipedes (Diplopoda)

Background: In the adult males of helminthomorph millipedes, one or two pairs of legs in the anterior part of the trunk are strongly modified into sexual appendages (gonopods) used for sperm transfer during the copula. Gonopods differentiate in an advanced phase of post-embryonic development, in most cases as replacement for the walking legs of the seventh trunk ring, as these first regress to tiny primordia, to eventually develop into gonopods at a subsequent stadium. These extremely localized but dramatic changes have been described as a non-systemic metamorphosis. In the present study we describe morphological and anatomical changes of trunk ring VII associated with non-systemic metamorphosis in four helminthomorph species. Results: As documented here for the first time by means of traditional histology methods and new techniques based on confocal laser scanning microscopy, the external odifications caused by non-systemic metamorphosis are associated to a huge rearrangement of internal anatomy, mostly due to the development of gonopod apodemes and extrinsic muscles. Conclusions: Internal changes in the seventh trunk ring, locally leading to the dorsal displacement of the ventral nerve cord and the digestive tract, are modulated in a taxon-specific manner, and are very conspicuous in the blaniulids Nopoiulus kochii and Blaniulus guttulatus, with likely major functional consequences.