From egg to adult: a developmental table of the ant Monomorium pharaonis
Ants are one of the most ecologically and evolutionarily successful groups of animals and exhibit a remarkable degree of phenotypic diversity. This success is largely attributed to the fact that all ants are eusocial and live in colonies with a reproductive division of labor between morphologically distinct queen and worker castes. Yet, despite over a century of studies on caste determination and evolution in ants, we lack a complete ontogenetic series from egg to adult for any ant species. We therefore present a developmental table for the Pharaoh ant Monomorium pharaonis, a species whose colonies simultaneously produce both reproductive queens and completely sterile workers. In total, M. pharaonis embryonic, larval, and pupal development lasts 45 days. During embryogenesis, the majority of developmental events are conserved between M. pharaonis and the fruit fly Drosophila melanogaster. However, we discovered two types of same-stage embryos prior to gastrulation: (1) embryos with internalized germ cells; and (2) embryos with germ cells outside of the blastoderm at the posterior pole. Moreover, we found two-types of embryos following germ band extension: (1) fertile embryos with primordial germ cells; and (2) sterile embryos with no germ cells. Together, our data shows that the queen (fertile) and worker (sterile) phenotypes are already determined and differentiated by early embryogenesis. During larval development, previous studies and our data find 3 larval instars reproductives and workers. However, there is considerable variation within each caste-specific instar, making it difficult to lineate instar boundaries. Here, we propose that developmental and anatomical markers can segregate larvae into gyne (unmatted queen), male and worker castes, including during the 1st larval instar. Overall, we hope that the ontogenetic series we present here will serve as a blueprint for the generation of future ant developmental tables.