Division of labor among workers is common in insect societies and is thought to be important in their ecological success. In most species, division of labor is based on age (temporal castes), but workers in some ants and termites show morphological specialization for particular tasks (physical castes). Large-headed soldier ants and termites are well-known examples of this specialization. However, until now there has been no equivalent example of physical worker subcastes in social bees or wasps. Here we provide evidence for a physical soldier subcaste in a bee. In the neotropical stingless bee Tetragonisca angustula, nest defense is performed by two groups of guards, one hovering near the nest entrance and the other standing on the wax entrance tube. We show that both types of guards are 30% heavier than foragers and of different shape; foragers have relatively larger heads, whereas guards have larger legs. Low variation within each subcaste results in negligible size overlap between guards and foragers, further indicating that they are distinct physical castes. In addition, workers that remove garbage from the nest are of intermediate size, suggesting that they might represent another unrecognized caste. Guards or soldiers are reared in low but sufficient numbers (1-2% of emerging workers), considering that <1% usually perform this task. When challenged by the obligate robber bee Lestrimelitta limao, an important natural enemy, larger workers were able to fight for longer before being defeated by the much larger robber. This discovery opens up opportunities for the comparative study of physical castes in social insects, including the question of why soldiers appear to be so much rarer in bees than in ants or termites.Meliponini | caste evolution D ivision of labor among workers is nearly universal in eusocial insects (1-5). In most species, this division is based on temporal subcastes, in which workers specialize in different tasks at different ages ("age polyethism") (2, 4, 5). Physical worker subcastes (i.e., groups of workers with morphological adaptations for particular tasks) are less common (4-6). Only ∼15% of all ant genera contain species with physical worker subcastes, and no species of bee or wasp is known to have a division of labor based on physical subcastes (4-6). Where physical worker subcastes occur, they appear to benefit the colony by increasing the efficiency of key tasks such as defense, but have the disadvantage of preventing a colony from rapidly adjusting caste ratios if necessary (4-6). In addition, extreme subcastes, such as large-bodied soldiers, are more costly to rear and have a limited task repertoire (4). It also has been argued that individual-level selection might prevent the evolution of worker polymorphism if workers of different subcastes have varying opportunities to reproduce (5, 7). Given the number of ant and termite species with physical worker subcastes, the lack of evidence for physical worker subcastes in bees and wasps is an intriguing and ongoing puzzle in the study ...