Abstract
In many social insects, including bumblebees, the division of labor between workers relates to body size, but little is known about the factors influencing larval development and final size. We confirmed and extend the evidence that in the bumblebee Bombus terrestris the adult bee body size is positively correlated with colony age. We next performed cross-fostering experiments in which eggs were switched between incipient (before worker emergence) and later stage colonies with workers. The introduced eggs developed into adults similar in size to their unrelated nestmates and not to their same-age full sisters developing in their mother colony. Detailed observations revealed that brood tending by the queen decreases, but does not cease, in young colonies with workers. We next showed that both worker number and the queen presence influenced the final size of the developing brood, but only the queen influence was mediated by shortening developmental time. In colonies separated by a queen excluder, brood developmental time was shorter in the queenright compartment. These findings suggest that differences in body size are regulated by the brood interactions with the queen and workers, and not by factors inside the eggs that could vary along with colony development. Finally, we developed a model showing that the typical increase in worker number and the decrease in brood contact with the queen can account for the typical increase in body size. Similar self-organized social regulation of brood development may contribute to the optimization of growth and reproduction in additional social insects.






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Acknowledgments
This work was supported by research grants from the US–Israel Binational Science Foundation (BSF #2007465), and the US–Israel Binational Agricultural Research and Development Fund (BARD #IS-4418-11R) to GB and the “Hoffman Leadership and Responsibility” fellowship to HS. We thank two anonymous reviewers for providing helpful comments on a previous version of this manuscript.
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Supplementary Fig. 1
Flow chart for experiment 4. We placed a queen at the colony founding stage and 10 workers in a small cage. After the queen laid a batch of eggs, we transferred her together with two workers to a second cage. The remaining 10 workers stayed to care for the developing brood (treatment 1). After the queen laid in the second cage, we transferred her together with a worker to a third small cage and left the other worker to care alone for the batch of eggs (treatment 2). After the queen laid in the third cage, we removed the worker and left the queen to care alone for the batch of eggs (treatment 3). When 10 workers had emerged from the brood in the third cage, we transferred the entire colony to a larger cage, and the queen and the 10+ workers cared for a new batch of eggs that was laid in the large cage (treatment 4). Q queen, w worker. Arrows indicate transfer of bees from one cage to another. Transferred bees are highlighted in a gray box. (PDF 80 kb)
Supplementary Fig. 2
Brood developing in colonies at the founding stage is smaller than in later stages of development. 1st—the first batch of brood in the colony; 2nd—the batch of brood developing from eggs laid when the colony contained a queen and approximately 10 workers. Values are mean ± SE; sample size is given inside the bars; p values were obtained from a two-tailed Student’s t test. The results were obtained from repetitions with four colonies in experiment 3. (PDF 64 kb)
Supplementary Fig. 3
The body size of emerging bees is positively correlated with colony age. Similar to the analysis presented in Fig. 1, but for bees emerging in a different colony. Each filled circle represents a worker (n = 56); each open triangle represents a drone (n = 46). Day 1 is the day of first worker emergence. There is a positive correlation between adult body size and colony age: workers—R 2 = 0.52, p < 0.001; drones—R 2 = 0.69, p < 0.001. (PDF 44 kb)
Supplementary Fig. 4
Signals from the queen shortens larval developmental time. Colonies were split into two similar parts using plastic meshes. The colonies in the upper row were separated by a mesh that enabled workers but not the queen to move between the two compartments (“Queen excluder colonies”). The colonies in the middle row were separated by a mesh that enabled both workers and the queen to move freely between the two compartments (“Queenright colonies”). The colonies in the bottom row have a mesh allowing free worker movement but did not have a queen (“Queenless colonies”). Gray bars—the side of the hive in which sugar syrup was provided (“S”); open bars—the side of the hive in which pollen was provided (“P”). In the queen excluder colonies, gray is also the queenless side (QL) and the open bars are the queenright side (QL). Values are mean ± SE; sample size is given inside the bars. Body size (left column) and developmental time (right column) were compared between the two sides in each colony; p values were obtained using Student’s t tests (N.S. not significant). (PDF 153 kb)
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Shpigler, H., Tamarkin, M., Gruber, Y. et al. Social influences on body size and developmental time in the bumblebee Bombus terrestris . Behav Ecol Sociobiol 67, 1601–1612 (2013). https://doi.org/10.1007/s00265-013-1571-0
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DOI: https://doi.org/10.1007/s00265-013-1571-0


