28In mealybugs, transcriptional inactivation of the entire paternal genome in males, due to 29 genomic imprinting, is closely correlated with sex determination. The sequencing, de-novo 30 assembly and annotation of the mealybug, Maconellicoccus hirsutus genome and its 31 comparison with Planococcus citri genome strengthened our gene identification. The expanded 32 gene classes, in both genomes relate to the high pesticide and radiation resistance; the 33 phenotypes correlating with increased gene copy number rather than the acquisition of novel 34 genes. The complete repertoire of genes for epigenetic regulation and multiple copies of genes 35 for the core members of polycomb and trithorax complexes and the canonical chromatin 36 remodelling complexes are present in both the genomes. Phylogenetic analysis with 37 Drosophila shows high conservation of most genes, while a few have diverged outside the 38 functional domain. The proteins involved in mammalian X-chromosome inactivation are 39 identified in mealybugs, thus demonstrating the evolutionary conservation of factors for 40 facultative heterochromatization. The transcriptome analysis of adult male and female 41 M.hirsutus indicates the expression of the epigenetic regulators and the differential expression 42 of metabolic pathway genes and the genes for sexual dimorphism. The depletion of 43 endosymbionts in males during development is reflected in the significantly lower expression 44 of endosymbiont genes in them.45 46 47 48 49 3 50Author summary 51 The mealybug system offers a unique model for genomic imprinting and differential regulation 52 of homologous chromosomes that pre-dates the discovery of dosage compensation of X 53 chromosomes in female mammals. In the absence of robust genetics for mealybugs, we 54 generated and analysed the genome and transcriptome profile as primary resources for effective 55 exploration. The expanded gene classes in the mealybugs relate to their unique biology; the 56 expansion of pesticide genes, trehalose transporter, SETMAR and retrotransposons correlate 57 with pesticide, desiccation and radiation resistance, respectively. The similarity in the genomic 58 profile of two species of mealybugs strengthens our gene prediction. All the known epigenetic 59 modifiers and proteins of the primary complexes like the PRC1,2 and the trithorax are 60 conserved in mealybugs, so also the homologues of mammalian proteins involved in X 61 chromosome inactivation. The high copy number of genes for many partners in these 62 complexes could facilitate the inactivation of a large part of the genome and raise the possibility 63 of formation of additional non-canonical complexes for sex specific chromosome inactivation.
64In adult males and females, the status of epigenetic regulation is likely to be in a maintenance 65 state; therefore, it is of interest to analyze the expression of epigenetic regulators during 66 development. 67 68 69 70 71 72 4 73 Abbreviations 74 Mhir-Maconellicoccus hirsutus, Pcit-Planococcus citri, Dmel-Drosoph...