 Thus, once termites have begun to build their monumental nests, it is the demands of living in those very nests to which future generations of termite must adapt.This inheritance may take many forms and operate on many scales, with a multiplicity of systems of inheritance complementing the genes.From position and maternal effects on gene expression to epigenetic inheritance  to the active construction and intergenerational transmission of enduring niches, development systems theory argues that not only inheritance but evolution as a whole can be understood only by taking into account a far wider range of ‘reproducers’ or ‘inheritance systems’ – genetic, epigenetic, behavioural and symbolic  – than neo-Darwinism’s ‘atomic’ genes and gene-like ‘replicators’.