SEX RATIOS AND CONDITIONS REQUIRED FOR ENVIRONMENTAL SEX DETERMINATION IN ANIMALS

Environmental sex determination (ESD) is a system of sexual determination that is influenced by a variable environment. Once sex is determined it is then fixed for life. The model of Charnov & Bull (1977) proposes that ESD is favoured by natural selection when an individual's fitness as a male or female is strongly influenced by environmental conditions and when the individual has little control over which environment it will experience. Adaptive sex ratio variation is considerably easier for organisms with ESD, and this feature is the ultimate cause for the evolution and maintenance of ESD. ESD is taxonomically widely expressed, and more cases are likely to be discovered. Both environmental and genotypic sex determination mechanisms are found in closely related species. Evidence of geographical variation in the degree and in the critical environmental values of ESD within the same species has also been discovered, e.g. in the fish Menidia menidia and in the crustacean Gammarus duebeni. The factors causing sex determination in invertebrates include temperature, daylength, nutrition, density, humidity, ionic composition of the environment, pH, carbon dioxide, UV light, metabolic products, parasites, exposure to the opposite sex of the same species, and in parasitoids also host size, age and type. In vertebrates temperature is the dominant factor causing sex determination, though in fish also pH, salinity, light, water quality and nutrition, and in turtles water potential of the substrate have some effect on the sex expression. Most of these factors influence growth through resource availability or developmental speed. In most cases of ESD in invertebrates and fish, the environmental factor has a gradual effect on the sex expression, in contrast to the typical steep threshold mode found in reptiles. These differences might be due to the fact that invertebrates exhibiting ESD are commonly parasitic or confined to aquatic environments, where less spatial microhabitat differentiation exists. Sex ratio data available from nature for animals with ESD are quite limited, except for reptiles. In the laboratory sex ratios can be varied more widely than what is observed in nature. There are a number of characteristic features some of which are found in each species exhibiting ESD: (1) Patchy environments, (2) variable sex ratios, (3) parthenogenesis in addition to bisexuality, (4) parasitism, (5) aquatic habitats, (6) sexual dimorphism, (7) females larger than males, and (8) local mate competition.