Bivalents are cooriented but only a small fraction separate into half-bivalents in the beginning of anaphase in mutant testes. Where anaphase occurs, whole bivalents rather than dyads are transported polewards, resulting in a wide variation in nuclear size and some cysts containing more than 32 nuclei. Multipolar spindles are seen, some of them located outside the nuclear region. Primary spermatocytes show irregularities of nuclear shape. The nucleus is larger than usual and has various evaginations and protrusions. Chromosomes sometimes leave the nucleus and enter the evagination zone, where they condense for meiosis. Centriolar bodies are underdeveloped and difficult to observe. If seen, the tiny elongated bodies are found attached or detached but close to the nuclei. The sperm heads are deformed and are found spread along the length of the tail bundle, never attached to the basal body end of the tails. Males show shaking upon etherisation.