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A new paper using analytical and simulation models to explore the relationships between particle-level heritability and collective-level heritability during major transitions has been published in BMC Biology:

Herron et al. 2018 Figure 3
Figure 3 from Herron et al. 2018.  Relative heritability of four collective-­level traits to cell-­level heritability for size.

From the Abstract:

Rather than being an impediment to major transitions, we show that, under a wide range of conditions, the heritability of collective-level traits is actually higher than that of the corresponding particle-level traits. High levels of collective-level trait heritability thus arise “for free,” with important implications not only for major transitions but for multilevel selection in general.

Herron MD, Zamani-Dahaj SA, & Ratcliff WC. 2018. Trait heritability in major transitions. BMC Biology 16:145.