In exploring the evolution and adaptive significance of epididymal function, we have studied the male excurrent duct and spermatozoa of a monotreme mammal--the echidna. Sperm maturation in the echidna excurrent duct appears simpler than that in most therians examined. Furthermore, neither the duct nor the spermatozoa of the echidna display specific therian characteristics; they bear a much closer resemblance to those of non-passerine birds. The echidna spermatozoon is filiform, the sperm tail has no distinctive features, and the anterior seventh of the undulating nucleus is covered by a modest acrosome. Immediately behind this a restricted apposition between plasma membrane and nuclear envelope constitutes a post-acrosomal ring. This is evident also in some reptiles and marsupials, whereas in Eutheria such a membrane association appears as the posterior ring at the base of the sperm nucleus. Maturation of spermatozoa in the Wolffian duct of the echidna appears to be expressed only in a changing capacity for motility and in loss of the cytoplasmic droplet. Neither surface, structural nor acrosomal changes that characterize sperm maturation in therian mammals have been detected in maturing echidna spermatozoa. The echidna duct displays little of the regional complexity of the epithelium that typifies this duct in the Theria. Of five regions distinguishable on the basis of epithelial morphology, the first two appear to be counterparts of efferent ducts by virtue of a low columnar, partially ciliated epithelium. The tall pseudo-stratified Golgi-rich epithelium of the major portion of the duct broadly resembles that of the therian epididymis, but it displays only two structurally distinguishable regions, the more distal being the site of a dense luminal secretion. The foamy epithelial cells of the fifth and terminal region, characterized by a mass of supra-nuclear vesicles and rough ER, suggest a secretory function that may in some way contribute significantly to the ejaculate, for accessory glands are poorly developed in monotremes. The possibility is considered that the relative complexity of epididymal function and sperm structure in therian mammals could have been determined by evolutionary change in the milieu of the female tract, and/or in the character of the egg vestments that the fertilizing spermatozoon must penetrate.