HRM analysis of the 134 wild-caught individuals afforded genotyping data consistent with those obtained by direct sequencing in all but one sample ( Table 1). The sensitivities of the delta T m values and shape discriminations were 40% and 20%, respectively. memnon showed that the H and h alleles exhibited unique, non-overlapping melting curves, enabling differentiation of the three dsx genotypes ( Fig. HRM analysis using primers targeting the dsx sequences of P.
polytes revealed that the divergence of H and h occurred independently within each species ( Fig. The reconstructed phylogenetic relationship among the four dsx allelic sequences from P. polytes h sequence contained a 14-bp insertion in the 5′-UTR (untranslated region) that was absent in P. None of the non-synonymous substitutions between the H and h sequences were shared between the two species. polytes by 33 nucleotide substitutions (5.8%), of which 5 were non-synonymous. memnon h sequence differed from that of P. polytes by 57 nucleotide substitutions (10.1%), of which 13 were non-synonymous. memnon H sequence differed from that of P. The 567-bp sequence contained 18 nucleotide substitutions (3.2% of all bp) between H and h, of which 4 were non-synonymous ( Fig. Next, we discriminated the H and h (hh) sequences by subtracting the h sequence from the Hh sequence ( Fig. These sequences were comprised of only two genotypes, probably corresponding to Hh and hh. Upon sequencing of a 567-bp portion of contig c1658_g1_i1, we obtained three heterozygous and three homozygous sequences from 6 female cDNA samples and six heterozygous and five homozygous sequences from 11 male cDNA samples. The two female contigs were identified as portions of the dsx sequence separated by a 59-bp gap. polytes, but no male contig yielded a significant match ( Table S2, Supplementary Information).
In a BLAST search of the dsx sequences, two of the female contigs (c1658_g1_i1, c17110_g5_i1) yielded significant matches with the F1 isoform of the dsx H and h alleles of P. memnon transcriptomes (DDBJ DRA accession nos.: SAMD00052594-SAMD00052595) yielded 45,887 and 39,035 contigs for one female and one male, respectively ( Table S1, Supplementary Information).
Our de novo assembly of 101-bp paired-end reads from P. memnon would also have doublesex alleles corresponding to mimetic and non-mimetic forms. 2), we predicted that the genetic basis of female-limited Batesian mimicry in P. Based on the phylogenetic proximity of P. These phenotypic traits are tightly linked with a putative supergene 9, 11. 1), which allow them to mimic unpalatable species in the genera Pachliopta and Atrophaneura 9, 11. Mimetic females have tails in their hindwings and aposematic colours in their hindwings and abdomens ( Fig. memnon males exhibit only the non-mimetic form ( Fig. memnon females can exist as either the mimetic or non-mimetic form, whereas P. polytes and exhibits similar female-limited Batesian mimicry polymorphism 15, 16. The Great Mormon butterfly, Papilio memnon, is closely related to P. These findings would facilitate the comparative genomic study of female-limited mimicry polymorphisms in Papilio butterflies to understand the evolutionary diversification of genes underlying the complex adaptive traits involved in mimicry. polytes, a single ~130-kb autosomal region including the sex-determinant gene doublesex (dsx), with mimetic ( H-type) and non-mimetic ( h-type) allelic sequences, controls polymorphism, and these two alleles are protected against recombination by an inversion that covers the entire region of the dsx gene 12, 14. dardanus, and the allelic differences associated with mimicry polymorphism have been identified 12, 13, 14. Recently, the genetic basis of Batesian mimicry was revealed in two Papilio species, P. Some Papilio butterflies are textbook examples of female-limited Batesian mimicry polymorphism, which is controlled by a supergene locus 7, 8, 9, 10, 11. One of the most intriguing types of Batesian mimicry in butterflies is the female-limited polymorphism in which females display both mimetic and non-mimetic forms 4, 5, 6. Batesian mimicry is the phenomenon in which palatable mimics avoid predation by resembling unpalatable species 1, 2, 3.
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