Anamorphosis in millipedes (Diplopoda)-the present state of knowledge with some developmental and phylogenetic considerations

Abstract

Abstract A review of the postembryonic development of millipedes (Diplopoda) is given, based mainly on published information. Original observations are, however, also included. Millipedes hatching from the pupoid usually have three pairs of legs; during their postembryonic growth they acquire more segments and more legs. This process is known as anamorphosis. Three types of anamorphosis are recognized. In euanamorphosis , every moult is accompanied by addition of new segments, even after the attainment of sexual maturity. In hemianamorphosis , the addition of new segments goes on until a certain stadium, and further moults take place without addition of segments. In teloanamorphosis , the addition of segments stops at a certain stadium (the adult, and ultimate, stadium) after which no further moults occur. Available information on anamorphosis in each of the millipede orders is reviewed. General patterns are emphasized, but variations are also considered in detail. It is shown that the so-called 'law of anamorphosis' is valid only for the 'ring-forming' millipedes (Merocheta and Juliformia) in which tergites, pleurites, and sternites of each diplosegment are firmly fused into a complete 'ring', and for some other forms (Polyzoniida, Chordeumatida), where there is a constant relationship between rings and legs. The chapter on the order Julida is particularly detailed and includes discussions of patterns in the variation and a section on periodomorphosis. The general chapter on developmental patterns includes inter alia an interpretation of the variations in millipede anamorphosis in terms of the 'biometabolic modi' of Remane. The hypothetical ancestral millipede is shown to have developed by hemianamorphosis. Euanamorphosis was acquired by the ancestral species of Helminthomorpha. Within this clade, Chordeumatida and Merocheta have secondarily become teloanamorphic, whereas some Juliformia seem to have returned to hemianamorphosis. The contrasting principles of elongation and contraction, subject of much debate among diplopodologists, are shown both to have played a role in the course of millipede evolution.