The concept of superfetation: a critical review on a 'myth' in mammalian reproduction.

Superfetation is understood as another conception during an already ongoing pregnancy. This implies the existence of young of different developmental stages within the female reproductive tract during certain periods of pregnancy. Nevertheless, a clear definition of the term as well as distinct criteria to identify the occurrence of superfetation in a species is missing. The variable anatomy of mammalian reproductive tracts seems to make the occurrence of superfetation more or less likely but impedes the simple evaluation of whether it is present or not. Additionally, adequate determination methods are missing or are difficult to apply at the right time. Superfetation or rather superfetation-like pregnancies are reported for numerous species including humans, livestock and rodents. The usual criteria to assume a case of superfetation include the finding of discordantly developed young within the uterus during post mortem or parturition of young after a birth interval shorter than the assumed pregnancy length. Often the occurrence of superfetation is concluded because other explanations of reproductive artifacts are missing. Even severe reproductive pathologies are often confused with superfetation. True superfetation or superfetation as a reproductive strategy may exist in some mammals. In the American mink (Neovison (Mustela) vison) and the European badger (Meles meles) superfetation occurs in combination with embryonic diapause. In the European brown hare (Lepus europaeus), superfetation has long been assumed to exist but evidence is still controversial. Superfetation definitely occurs in certain species of poeciliid and zenarchopterid fish, some of which also exhibit viviparity and maternal care. In mammals, the evolution of such a reproductive mechanism poses many interesting evolutionary, endocrine, microbial and immunological questions that require further investigation. Here we review the scant and at times ancient literature on this poorly understood topic. The different manifestations of superfetation are defined and reliable criteria to detect superfetation are outlined. Also, the differentiation of superfetation into a reproductive strategy or as a disrupted, abnormal reproductive function is discussed. Due to the different discussed functional aspects of superfetation, it is appropriate to establish a more detailed scheme to classify the true natural superfetation cases into superfertilization, superconception and superfetation proper. To date, there is no mammal species known for which superfetation proper in terms of finding discordantly developed fetuses has been conclusively demonstrated to be not only a rare occurence but an evolved reproductive strategy.