title: Modularity of a leaf moth-wing pattern and a versatile characteristic of the wing-pattern ground plan.
abstract: BACKGROUND
  One of the most intriguing questions in evolutionary developmental biology is how an insect acquires a mimicry pattern within its body parts. A striking example of pattern mimicry is found in the pattern diversity of moth and butterfly wings, which is thought to evolve from preexisting elements illustrated by the nymphalid ground plan (NGP). Previous studies demonstrated that individuality of the NGP facilitates the decoupling of associated common elements, leading to divergence. In contrast, recent studies on the concept of modularity have argued the importance of a combination of coupling and decoupling of the constituent elements. Here, we examine the modularity of a mimicry wing pattern in a moth and explore an evolvable characteristic of the NGP.
  
  
  RESULTS
  This study examined the wings of the noctuid moth Oraesia excavata, which closely resemble leaves with a leaf venation pattern. Based on a comparative morphological procedure, we found that this leaf pattern was formed by the NGP common elements. Using geometric morphometrics combined with network analysis, we found that each of the modules in the leaf pattern integrates the constituent components of the leaf venation pattern (i.e., the main and lateral veins). Moreover, the detected modules were established by coupling different common elements and decoupling even a single element into different modules. The modules of the O. excavata wing pattern were associated with leaf mimicry, not with the individuality of the NGP common elements. For comparison, we also investigated the modularity of a nonmimetic pattern in the noctuid moth Thyas juno. Quantitative analysis demonstrated that the modules of the T. juno wing pattern regularly corresponded to the individuality of the NGP common elements, unlike those in the O. excavata wing pattern.
  
  
  CONCLUSIONS
  This study provides the first evidence for modularity in a leaf mimicry pattern. The results suggest that the evolution of this pattern involves coupling and decoupling processes to originate these modules, free from the individuality of the NGP system. We propose that this evolution has been facilitated by a versatile characteristic of the NGP, allowing the association of freely modifiable subordinate common elements to make modules.
authors.affiliation: Laboratory for Evolutionary Morphology, Center for Developmental Biology, RIKEN, Chuo-ku Kobe, Japan. homaresuzuki@gmail.com
authors.forename: Takao K
authors.initials: TK
authors.lastname: Suzuki
mesh_terms: D000818:Animals; D005075:Biological Evolution; D005260:Female; D008297:Male; D009036:Moths / Q000033:anatomy & histology / Q000737:chemistry*; D018515:Plant Leaves / Q000033:anatomy & histology / Q000737:chemistry*; D014921:Wings, Animal / Q000033:anatomy & histology / Q000737:chemistry*
references.citation: Evolution. 1998 Oct;52(5):1363-1375
references.pmid: 28565401
references.citation: Trends Ecol Evol. 2006 Sep;21(9):488-92
references.pmid: 16806575
references.citation: Brief Bioinform. 2006 Jun;7(2):151-8
references.pmid: 16772265
references.citation: Evolution. 1996 Jun;50(3):967-976
references.pmid: 28565291
references.citation: Bioinformatics. 2004 Sep 22;20(14):2242-50
references.pmid: 15130938
references.citation: Evolution. 2013 Jan;67(1):241-59
references.pmid: 23289575
references.citation: Evolution. 2008 Oct;62(10):2688-708
references.pmid: 18691262
references.citation: PLoS One. 2012;7(9):e45035
references.pmid: 23028749
references.citation: Evolution. 2001 Sep;55(9):1734-45
references.pmid: 11681729
references.citation: Evolution. 2012 May;66(5):1525-42
references.pmid: 22519788
references.citation: Cell. 1996 Jun 28;85(7):951-61
references.pmid: 8674123
references.citation: Proc Natl Acad Sci U S A. 1973 Jul;70(7):1936-8
references.pmid: 4198660
references.citation: Nat Rev Genet. 2010 Sep;11(9):623-35
references.pmid: 20697423
references.citation: Science. 1983 May 13;220(4598):671-80
references.pmid: 17813860
references.citation: Evol Dev. 2009 Jul-Aug;11(4):405-21
references.pmid: 19601974
references.citation: Evolution. 2008 Apr;62(4):943-58
references.pmid: 18194472
references.citation: Evolution. 2006 Jan;60(1):169-83
references.pmid: 16568641
references.citation: Insect Biochem Mol Biol. 2012 Feb;42(2):148-54
references.pmid: 22178129
references.citation: Evolution. 1993 Apr;47(2):593-618
references.pmid: 28568736
references.citation: Am Nat. 2012 Jan;179(1):64-78
references.pmid: 22173461
references.citation: Evol Dev. 2009 May-Jun;11(3):302-11
references.pmid: 19469857
references.citation: Evolution. 1982 May;36(3):499-516
references.pmid: 28568050
references.citation: Evolution. 1987 May;41(3):491-503
references.pmid: 28563806
references.citation: Nat Biotechnol. 2000 Jan;18(1):81-4
references.pmid: 10625397
references.citation: Phys Rev Lett. 2005 Jan 14;94(1):018102
references.pmid: 15698136
references.citation: Evolution. 1979 Mar;33(1Part2):402-416
references.pmid: 28568194
references.citation: Genetics. 2003 Nov;165(3):1329-40
references.pmid: 14668386
references.citation: Syst Biol. 2007 Oct;56(5):818-36
references.pmid: 17934997
references.citation: Nat Rev Genet. 2002 Jun;3(6):442-52
references.pmid: 12042771
references.citation: Am Nat. 2010 Mar;175(3):289-301
references.pmid: 20095825
references.citation: Science. 2010 Jan 1;327(5961):51
references.pmid: 20044568
references.citation: Nat Rev Genet. 2007 Dec;8(12):921-31
references.pmid: 18007649
references.citation: Evolution. 2010 Oct;64(10):2935-51
references.pmid: 20482613
references.citation: Proc Natl Acad Sci U S A. 2012 Jul 31;109(31):12632-7
references.pmid: 22802635
references.citation: Mol Biol Evol. 2010 Dec;27(12):2864-78
references.pmid: 20624848
references.citation: Biol Lett. 2010 Oct 23;6(5):597-9
references.pmid: 20410028
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2006 Jul;74(1 Pt 2):016110
references.pmid: 16907154
references.citation: Insect Biochem Mol Biol. 2008 Dec;38(12):1165-73
references.pmid: 18992341
references.citation: PLoS Genet. 2012;8(3):e1002486
references.pmid: 22412378
references.citation: Bioinformatics. 2003 May 22;19(8):1019-26
references.pmid: 12761066
references.citation: Science. 1994 Jul 1;265(5168):109-14
references.pmid: 7912449
references.citation: Curr Biol. 1999 Feb 11;9(3):109-15
references.pmid: 10021383
references.citation: Nature. 2002 Feb 7;415(6872):609-13
references.pmid: 11832937
references.citation: Proc Natl Acad Sci U S A. 2000 Oct 24;97(22):12182-6
references.pmid: 11027309
references.citation: J Insect Physiol. 2011 Feb;57(2):231-45
references.pmid: 21078327
references.citation: J Exp Zool. 2001 Oct 15;291(3):213-25
references.pmid: 11598911
references.citation: Am Nat. 2006 Aug;168(2):270-80
references.pmid: 16874636
references.citation: Evolution. 1996 Aug;50(4):1585-1597
references.pmid: 28565721