title: Morphological outcomes of gynandromorphism in Lycaeides butterflies (Lepidoptera: Lycaenidae).
abstract: The genitalia of male insects have been widely used in taxonomic identification and systematics and are potentially involved in maintaining reproductive isolation between species. Although sexual selection has been invoked to explain patterns of morphological variation in genitalia among populations and species, developmental plasticity in genitalia likely contributes to observed variation but has been rarely examined, particularly in wild populations. Bilateral gynandromorphs are individuals that are genetically male on one side of the midline and genetically female on the other, while mosaic gynandromorphs have only a portion of their body developing as the opposite sex. Gynandromorphs might offer unique insights into developmental plasticity because individuals experience abnormal cellular interactions at the genitalic midline. In this study, we compare the genitalia and wing patterns of gynandromorphic Anna and Melissa blue butterflies, Lycaeides anna (Edwards) (formerly L. idas anna) and L. melissa (Edwards) (Lepidoptera: Lycaenidae), to the morphology of normal individuals from the same populations. Gynandromorph wing markings all fell within the range of variation of normal butterflies; however, a number of genitalic measurements were outliers when compared with normal individuals. From these results, we conclude that the gynandromorphs' genitalia, but not wing patterns, can be abnormal when compared with normal individuals and that the gynandromorphic genitalia do not deviate developmentally in a consistent pattern across individuals. Finally, genetic mechanisms are considered for the development of gynandromorphism in Lycaeides butterflies.
authors.affiliation: Program in Ecology, Evolution, and Conservation Biology, Department of Biology, University of Nevada, Reno, NV 89557, USA jpjahner@gmail.com.
authors.forename: Joshua P
authors.initials: JP
authors.lastname: Jahner
authors.affiliation: Department of Biology, Texas State University, San Marcos, TX 78666, USA.
authors.forename: Lauren K
authors.initials: LK
authors.lastname: Lucas
authors.affiliation: Department of Biology, Utah State University, Tooele, UT 84074, USA.
authors.forename: Joseph S
authors.initials: JS
authors.lastname: Wilson
authors.affiliation: Program in Ecology, Evolution, and Conservation Biology, Department of Biology, University of Nevada, Reno, NV 89557, USA.
authors.forename: Matthew L
authors.initials: ML
authors.lastname: Forister
mesh_terms: D000818:Animals; D002080:Butterflies / Q000033:anatomy & histology* / Q000254:growth & development; D005260:Female; D005835:Genitalia / Q000033:anatomy & histology / Q000254:growth & development; D008297:Male; D014481:United States; D014921:Wings, Animal / Q000033:anatomy & histology* / Q000254:growth & development
keywords: Lycaeides anna; Lycaeides melissa; developmental plasticity; genitalia; gynandromorph,; local outlier factor
references.citation: J Exp Zool B Mol Dev Evol. 2011 Jul 15;316(5):313-8
references.pmid: 21404423
references.citation: Trends Ecol Evol. 2004 Feb;19(2):87-93
references.pmid: 16701234
references.citation: Mol Phylogenet Evol. 2008 Aug;48(2):481-90
references.pmid: 18539051
references.citation: Nat Rev Genet. 2002 Jun;3(6):442-52
references.pmid: 12042771
references.citation: Nature. 1996 Nov 21;384(6606):236-42
references.pmid: 12809139
references.citation: Proc Natl Acad Sci U S A. 2003 Apr 15;100(8):4873-8
references.pmid: 12672961
references.citation: Evolution. 2013 Apr;67(4):1055-68
references.pmid: 23550755
references.citation: Bioessays. 2013 Aug;35(8):725-32
references.pmid: 23804281
references.citation: Trends Ecol Evol. 2011 Jan;26(1):17-21
references.pmid: 21093955
references.citation: Chromosome Res. 2012 Jan;20(1):177-90
references.pmid: 22124858
references.citation: Mol Ecol. 2010 Aug;19(15):3171-92
references.pmid: 20618903
references.citation: Anat Rec. 1956 Feb;124(2):223-43
references.pmid: 13302819
references.citation: Mol Ecol. 2005 May;14(6):1741-54
references.pmid: 15836646
references.citation: Evolution. 2012 Jul;66(7):2167-81
references.pmid: 22759293
references.citation: Science. 2006 Dec 22;314(5807):1923-5
references.pmid: 17138866
references.citation: Mol Ecol. 2014 Sep;23(18):4555-73
references.pmid: 24866941
references.citation: Oecologia. 2009 Jun;160(3):551-61
references.pmid: 19271241
references.citation: BMC Evol Biol. 2013 Aug 12;13:168
references.pmid: 23937355
references.citation: Oecologia. 2013 May;172(1):177-88
references.pmid: 23053234
references.citation: Genetica. 2010 Jan;138(1):5-18
references.pmid: 19308664
references.citation: J Exp Zool. 1969 Jan;170(1):61-75
references.pmid: 5780530
references.citation: Wilhelm Roux Arch Entwickl Mech Org. 1931 Sep;123(3-4):719-746
references.pmid: 28354041
references.citation: J Environ Monit. 2012 May;14(5):1317-24
references.pmid: 22456673
references.citation: Nature. 2010 Mar 11;464(7286):237-242
references.pmid: 20220842