Penetrant deletions for microcephaly outlined two SROs (Fig. 2 a); SRO1 (p: 0.25, 2/8) including the BCL11A (cp: 0.23) and PAPOLG (cp: 0.13) genes, and SRO2 (p: 0.55, 6/11) overlapping the USP34 (cp: 0.77) and XPO1 (cp: 0.18) genes. Here, USP34 is linked to microcephaly.