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The BEP-linked genes in *O. nivara*, which co-existed with the *NIPK*-linked gene in b-*Oa*-*NIPK* (Fig. [5b](#Fig5){ref-type=”fig”}), are likely to derive from a non-C~4~ lineage. The clade of O*nipk*-associated genes (*OaCORE, OaGLK, OaHAD*, and *OaPROG*) are distantly related to corresponding genes in b-*NIPK* (Fig. [6d](#Fig6){ref-type=”fig”}), but gene similarity indicates a common ancestry with the non-C~4~ ancestor of C~4~ monocots \[[@CR12]\]. The close proximity of *NIPK* and *CBSP5* to *NIPK* in *O. nivara*, however, suggests their ancestry is C~4~. A closer look at the phylogeny also reveals a close relationship between *NIPK* and *OaCBSP5* in *P. contorta* (Fig. [6e](#Fig6){ref-type=”fig”}). Future experimentation with these accessions of *P. contorta* and *O. nivara* is needed to determine if the monophyly of these genes is ancestral to C~4~ monocots or they are derived from another non-C~4~ lineage that existed prior to the divergence of C~4~ grasses.
Conclusions {#Sec7}
===========
Here we demonstrate that CBP5 was likely derived from the *NIPK* gene in monocots. Although CBP5 is not a direct orthologue of *NIPK*, which is a gene necessary for the de novo synthesis of C~4~-associated NAD(P)^+
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