Comparative fine structure of sperm of Heterodera schachtii and Punctodera chalcoensis, with phylogenetic implications for Heteroderinae (Nemata: Heteroderidae)

Abstract

The fine structure of sperm of Heterodera schachtii and Punctodera chalcoensis demonstrates a high degree of interspecific morphological and developmental diversity, particularly in comparison with previously examined Heteroderinae that develop cysts, including Globodera tabacum, as well as out-groups that lack cysts, including species of Verutus, Meloidodera, and Ekphymatodera. Sperm of P. chalcoensis are much smaller and have fewer filopodia than those of species of Heterodera and Globodera. However, the distribution of filopodia on only part of the body, the smooth surface of the filopodia, and the presence of cortical microtubules are traits shared by the three genera with cysts. Unique features shared by P. chalcoensis and Globodera species include the short persistence of fibrous bodies after spermiogenesis and the lack of sperm polarization in the female genital tract. These traits are absent in Heterodera species. Conversely, chromatin remains unchanged with respect to condensation during sperm development in P. chalcoensis and species of Heterodera but not in Globodera. Patterns of evolution of sperm may be useful for testing hypotheses of the phylogeny of Heteroderinae, but the diversity is so great that character coding will be required for a large number of representative species.