title: Geographic variation of melanisation patterns in a hornet species: genetic differences, climatic pressures or aposematic constraints?
abstract: Coloration of stinging insects is often based on contrasted patterns of light and black pigmentations as a warning signal to predators. However, in many social wasp species, geographic variation drastically modifies this signal through melanic polymorphism potentially driven by different selective pressures. To date, surprisingly little is known about the geographic variation of coloration of social wasps in relation to aposematism and melanism and to genetic and developmental constraints. The main objectives of this study are to improve the description of the colour variation within a social wasp species and to determine which factors are driving this variation. Therefore, we explored the evolutionary history of a polymorphic hornet, Vespa velutina Lepeletier, 1836, using mitochondrial and microsatellite markers, and we analysed its melanic variation using a colour space based on a description of body parts coloration. We found two main lineages within the species and confirmed the previous synonymy of V. auraria Smith, 1852, under V. velutina, differing only by the coloration. We also found that the melanic variation of most body parts was positively correlated, with some segments forming potential colour modules. Finally, we showed that the variation of coloration between populations was not related to their molecular, geographic or climatic differences. Our observations suggest that the coloration patterns of hornets and their geographic variations are determined by genes with an influence of developmental constraints. Our results also highlight that Vespa velutina populations have experienced several convergent evolutions of the coloration, more likely influenced by constraints on aposematism and Müllerian mimicry than by abiotic pressures on melanism.
authors.affiliation: UMR 7205 ISYEB, Muséum National d'Histoire Naturelle, Paris, France; Division of Invertebrate Zoology, American Museum of Natural History, New York, New York, United States of America.
authors.forename: Adrien
authors.initials: A
authors.lastname: Perrard
authors.affiliation: Unité de Recherche IRD 072-UPR9034 CNRS, Laboratoire Evolution Génomes et Spéciation, Gif-sur-Yvette, France; Université Paris-Sud 11, Orsay, France.
authors.forename: Mariangela
authors.initials: M
authors.lastname: Arca
authors.affiliation: UMR 7205 ISYEB, Muséum National d'Histoire Naturelle, Paris, France.
authors.forename: Quentin
authors.initials: Q
authors.lastname: Rome
authors.affiliation: UMR 7205 ISYEB, Muséum National d'Histoire Naturelle, Paris, France.
authors.forename: Franck
authors.initials: F
authors.lastname: Muller
authors.affiliation: School of Life Sciences, Northwest University, Xi'an, Shaanxi, China.
authors.forename: Jiangli
authors.initials: J
authors.lastname: Tan
authors.affiliation: Entomology Division, Nepal Agricultural Research Council (NARC), Lalitpur, Nepal.
authors.forename: Sanjaya
authors.initials: S
authors.lastname: Bista
authors.affiliation: Museum Zoologicum Bogoriense, Indonesian Institute of Sciences, Cibinong, Bogor, Indonesia.
authors.forename: Hari
authors.initials: H
authors.lastname: Nugroho
authors.affiliation: CBNBP, Muséum National d'Histoire Naturelle, Paris, France.
authors.forename: Raymond
authors.initials: R
authors.lastname: Baudoin
authors.affiliation: UMR 7205 ISYEB, Muséum National d'Histoire Naturelle, Paris, France.
authors.forename: Michel
authors.initials: M
authors.lastname: Baylac
authors.affiliation: Unité de Recherche IRD 072-UPR9034 CNRS, Laboratoire Evolution Génomes et Spéciation, Gif-sur-Yvette, France; Université Paris-Sud 11, Orsay, France.
authors.forename: Jean-François
authors.initials: JF
authors.lastname: Silvain
authors.affiliation: Division of Invertebrate Zoology, American Museum of Natural History, New York, New York, United States of America.
authors.forename: James M
authors.initials: JM
authors.lastname: Carpenter
authors.affiliation: UMR 7205 ISYEB, Muséum National d'Histoire Naturelle, Paris, France.
authors.forename: Claire
authors.initials: C
authors.lastname: Villemant
mesh_terms: D000818:Animals; D002980:Climate; D014644:Genetic Variation; D005843:Geography; D006239:Haplotypes; D008543:Melanins / Q000378:metabolism*; D018895:Microsatellite Repeats; D013045:Species Specificity; D014863:Wasps / Q000033:anatomy & histology / Q000235:genetics* / Q000378:metabolism
chemical_list: D008543:Melanins
references.citation: Semin Cell Dev Biol. 2009 Feb;20(1):65-71
references.pmid: 18977308
references.citation: Naturwissenschaften. 2008 Aug;95(8):681-95
references.pmid: 18542902
references.citation: Nature. 2005 Feb 3;433(7025):481-7
references.pmid: 15690032
references.citation: J Evol Biol. 2011 Jun;24(6):1160-8
references.pmid: 21457173
references.citation: Evol Bioinform Online. 2007 Feb 23;1:47-50
references.pmid: 19325852
references.citation: Genetics. 2008 Dec;180(4):2135-49
references.pmid: 18245845
references.citation: Dev Biol. 2002 Feb 1;242(1):15-30
references.pmid: 11795937
references.citation: Evolution. 2009 Feb;63(2):469-78
references.pmid: 19154362
references.citation: Bioinformatics. 2004 Jan 22;20(2):289-90
references.pmid: 14734327
references.citation: Proc Biol Sci. 2005 Feb 22;272(1561):431-8
references.pmid: 15734698
references.citation: Development. 2002 Apr;129(8):1849-58
references.pmid: 11934851
references.citation: Nat Rev Genet. 2013 Mar;14(3):179-90
references.pmid: 23381120
references.citation: Philos Trans R Soc Lond B Biol Sci. 2005 Oct 29;360(1462):1959-67
references.pmid: 16214753
references.citation: Front Zool. 2012 Aug 20;9(1):20
references.pmid: 22901602
references.citation: Mol Biol Evol. 1999 Jan;16(1):37-48
references.pmid: 10331250
references.citation: Development. 1999 Aug;126(16):3495-507
references.pmid: 10409497
references.citation: J Insect Physiol. 2009 Apr;55(4):358-68
references.pmid: 19200435
references.citation: Oecologia. 2003 Sep;137(1):42-50
references.pmid: 12838404
references.citation: Am Nat. 2010 May;175(5):495-503
references.pmid: 20302424
references.citation: Mol Ecol. 2005 Jul;14(8):2611-20
references.pmid: 15969739
references.citation: Nat Rev Genet. 2007 Mar;8(3):206-16
references.pmid: 17304246
references.citation: Dev Genes Evol. 2003 Jan;212(12):571-84
references.pmid: 12536321
references.citation: Mol Ecol. 2009 Nov;18(21):4463-76
references.pmid: 19796331
references.citation: Dev Genes Evol. 2004 Dec;214(12):628-34
references.pmid: 15449055
references.citation: Heredity (Edinb). 2006 Sep;97(3):157-67
references.pmid: 16835591
references.citation: Mol Biol Evol. 1987 Jul;4(4):406-25
references.pmid: 3447015
references.citation: Cell. 2006 Jun 30;125(7):1387-99
references.pmid: 16814723
references.citation: Zoolog Sci. 2006 Jan;23(1):95-104
references.pmid: 16547411
references.citation: Evolution. 1980 Jul;34(4):622-637
references.pmid: 28563986
references.citation: Evolution. 2008 Nov;62(11):2742-59
references.pmid: 18764916
references.citation: Proc Biol Sci. 2012 Feb 7;279(1728):417-26
references.pmid: 22113031
references.citation: Mol Ecol. 2006 May;15(6):1419-39
references.pmid: 16629801
references.citation: Dev Genes Evol. 2000 Nov;210(11):536-44
references.pmid: 11180804
references.citation: Dev Biol. 1995 Mar;168(1):112-23
references.pmid: 7883067
references.citation: J Evol Biol. 2013 Jul;26(7):1406-16
references.pmid: 23675858
references.citation: Pigment Cell Res. 1993 Jun;6(3):117-26
references.pmid: 8234196
references.citation: Nature. 2011 Aug 14;477(7363):203-6
references.pmid: 21841803
references.citation: Bioinformatics. 2005 Dec 15;21(24):4338-47
references.pmid: 16234323
references.citation: Evolution. 2010 Jul;64(7):1944-54
references.pmid: 20148951
references.citation: EXS. 1993;67:153-75
references.pmid: 8400687
references.citation: Proc Natl Acad Sci U S A. 2003 Feb 18;100(4):1808-13
references.pmid: 12574518
references.citation: Evol Dev. 2003 Jan-Feb;5(1):9-18
references.pmid: 12492404
references.citation: Genetics. 2000 Jun;155(2):945-59
references.pmid: 10835412
references.citation: Mol Mar Biol Biotechnol. 1994 Oct;3(5):294-9
references.pmid: 7881515
references.citation: Nature. 2000 Nov 30;408(6812):553-9
references.pmid: 11117736
references.citation: Dev Biol. 1991 Sep;147(1):32-45
references.pmid: 1879614