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In many yeast species, the three genes at the centre of the galactose catabolism pathway,  GAL1 ,  GAL10  and  GAL7 , are neighbours in the genome and form a metabolic gene cluster. We report here that some yeast strains in the genus  Torulaspora  have much larger  GAL  clusters that include genes for melibiase ( MEL1 ), galactose permease ( GAL2 ), glucose transporter ( HGT1 ), phosphoglucomutase ( PGM1 ) and the transcription factor  GAL4 , in addition to  GAL1 ,  GAL10 , and  GAL7 . Together, these eight genes encode almost all the steps in the pathway for catabolism of extracellular melibiose (a disaccharide of galactose and glucose). We show that a progenitor 5‐gene cluster containing  GAL 7‐1‐10‐4‐2  was likely present in the common ancestor of  Torulaspora  and  Zygotorulaspora . It added  PGM1  and  MEL1  in the ancestor of most  Torulaspora  species. It underwent further expansion in the  T. pretoriensis  clade, involving the fusion of three progenitor clusters in tandem and the gain of  HGT1 . These giant  GAL  clusters are highly polymorphic in structure, and subject to horizontal transfers, pseudogenization and gene losses. We identify recent horizontal transfers of complete  GAL  clusters from  T. franciscae  into one strain of  T. delbrueckii , and from a relative of  T. maleeae  into one strain of   T. globosa  . The variability and dynamic evolution of  GAL  clusters in  Torulaspora  indicates that there is strong natural selection on the  GAL  pathway in this genus.

Giant GAL gene clusters for the melibiose‐galactose pathway in Torulaspora