Summary: Hox and ParaHox genes are involved in patterning the anterior-posterior body axis in metazoans during embryo development. Body plan evolution and diversification are affected by variations in the number and sequence of Hox and ParaHox genes, as well as by their expression patterns. For this reason Hox and ParaHox gene investigation in the phylum Mollusca is of great interest, as this is one of the most important taxa of protostomes, characterized by a high morphological diversity. The comparison of the works reviewed here indicates that species of molluscs, belonging to different classes, share a similar composition of Hox and ParaHox genes. Therefore evidence suggests that the wide morphological diversity of this taxon could be ascribed to differences in Hox gene interactions and expressions and changes in the Hox downstream genes rather than to Hox cluster composition. Moreover the data available on Hox and ParaHox genes in molluscs compared with those of other Lophotrochozoa shed light on the complex and controversial evolutionary histories that these genes have undergone within protostomes. genesis 52:935-945, 2014. V C 2014Wiley Periodicals, Inc.Key words: Hox genes; ParaHox genes; molluscs; ProtoHox cluster
HOX AND PARAHOX GENES: EVOLUTIONARY HYPOTHESESBoth Hox and ParaHox genes belong to the ANTP homeobox gene class, which also includes the NK genes, the NK-related genes, the Dlx genes, and many more (Holland, 2013). It has been hypothesized that the ancestor genes of the current ANTP class were originally arranged in a single cluster called "Megacluster" (Garcıa-Fernandez, 2005a;Holland, 2013).The comparison of Hox and ParaHox gene number is interesting in order to reconstruct the evolutionary history of these gene families. In this regard the increase in sequenced genomes is providing significant data concerning the Hox and ParaHox gene complement in several organisms.Considering the early appearance of NK genes during evolution and their genomic proximity to the Hox genes in some species (Larroux et al., 2007), Holland (2013 proposed the ProtoHox cluster as originating from a ProtoNK cluster. Given the lack of Hox and ParaHox genes in basal metazoans, such as the demosponge Amphimedon queenslandica (Larroux et al., 2007) and the lobate ctenophore Mnemiopsis leidyi, it has been hypothesized (Ryan et al., 2010) that the ProtoHox gene/cluster could have arisen after the divergence of Porifera and Ctenophora from ParaHoxozoa (Placozoa, Cnidaria, and Bilateria). However, the phylogenetic relationships among basal metazoans are still