b Onthophagus binodis produced many brood balls from a dung pad in soil of 4% moisture, but few at 2% or 1% (wilting point 3.5%). Euoniticellus intermedius produced consistently high numbers of brood balls at all 3 soil moistures. At the 2 drier levels O. binodis placed brood balls only in the soil moistened by the dung, but E. intermedius always placed brood balls throughout the available soil. E. intermedius in dry soil produced clumps averaging 8.7 brood balls and lined the walls of the tunnels with dung, thus preventing their collapse. In moist soil both species produced brood balls singly in tunnels throughout the available soil.
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Barkhouse et al. (1986) studied this question.
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