Darwin's special difficulty: the evolution of neuter insects and current theory

In the Origin of Species, Darwin discussed several challenges that worker insects presented to his theory of natural selection. Complex instincts such as building of combs of hexagonal cells were one problem and were explained by showing plausible intermediate stages. A more serious challenge was posed by the multiple worker castes seen in many ants. How could sterile individuals continue to evolve? A careful reading of the Origin suggests that Darwin was not primarily concerned by the evolution of worker sterility itself, which he considered a minor difficulty. Some modern commentaries on Darwin and insect workers seem to be cases of present interests interfering with the interpretation of the past. From a modern perspective, the evolution of a worker caste, and its corollary altruism, are evolutionary puzzles inasmuch as natural selection normally favors greater, not lesser, individual reproduction. These puzzles were resolved by Hamilton's theory of inclusive fitness. We now have a good functional understanding of how natural selection can cause both the origin of workers and their elaboration into greater levels of sterility and multiple morphological castes. Mechanistic understanding of morphological castes is also increasing via research into alternative developmental pathways. When the Origin was written, genetics did not exist and it would have been virtually impossible for Darwin to elaborate such ideas. However, the Origin probably addressed the main questions in the minds of Victorian readers in relation to insect workers. Darwin was prescient in having insights with close relationships to modern-day interests and the key principles involved, including kinship and benefits to the colony, even if these are not exact precursors to modern thinking.