Research People Publications Links

My Google scholar profile (most up to date)


Pitchers, W.R., Klingenberg, C.P., Tregenza, T., Hunt., & Dworkin, I. 2014. The potential influence of morphology on the evolutionary divergence of an acoustic signal. Accepted. Journal of Evolutionary Biology.

Link to paper or pre-print on biorxiv Pitchers, W.R., Wolf, J.B.,Tregenza, T., Hunt, J. & Dworkin, I. 2014. Evolutionary rates for multivariate traits: the role of selection and genetic variation. Phil. Trans. Roy. Soc. 369(9). doi:10.1098/rstb.2013.0252 link to database of G matrices and scripts on github

Link to paper O ' Donnell, D.R., Fish, J.A., Parigi, A., Dworkin, I. & Wagner, A.P. 2014. The roles of standing genetic variation and evolutionary history in determining the evolvability of anti-predator strategies. PLoS One. 9(6): e100163. doi:10.1371/journal.pone.0100163

Link to paper Moghe, G.D., Hufnagel, D.E., Tang, H., Xiao, Y., Dworkin, I., Town, C.D., Conner, J.K. & Shiu, S. 2014.Consequences of Whole-Genome triplication as revealed by comparative genomic analyses of the Wild Radish Raphanus raphanistrium and three other Brassicaceae species. Plant Cell. doi:10.1105/tpc.114.124297

Link to paper Ledòn-Rettig, C., Pfennig, D., Chunco, C. & Dworkin, I. 2014. Cryptic Genetic variation in natural populations: A predictive framework. Integrative and Comparative Biology. In Press.

Link to paper Melicher, D.M., Torson, A.S., Dworkin, I. & Bowsher, J.H. 2014. A pipeline for the de novo assembly of the Themira biloba (Sepsidae: Diptera) transcriptome using a multiple k-mer length approach. BMC Genomics. 15(1):188 doi:10.1186/1471-2164-15-188

Link to paper Johnson, A., Strauss, E., Pickett, R., Adami, C., Dworkin, I., & Goldsby, H. More Bang For Your Buck: Quorum-Sensing Capabilities Improve the Efficacy of Suicidal Altruism. Artificial Life 14:121-128.

Link to paper or Pre-print on arXiV Chandler, C.H., Chari, S., Tack, D. & Dworkin, I. 2014. Causes and Consequences of Genetic Background Effects Illuminated by Integrative Genomic Analysis. Genetics. 196:1321-1336. doi:10.1534/genetics.113.159426

Link to paper Warren, I.A., Vera, J.C., Johns, A., Zinna, R., Marden, J.H., Emlen, D.J., Dworkin, I. & Lavine, L.C. 2014. Insights into the development and evolution of exaggerated traits using de novo transcriptomes of two species of horned scarab beetles. PLoS One. 9(2):e88364 doi:10.1371/journal.pone.0088364

Link to paper Ahmad, F., Roy, K., O ' Connor, B., Shelton, J., Dozier, G. & Dworkin, I. 2014. Fly Wing Biometrics using modified local binary pattern, SVMs and Random Forest. International Journal of Machine Learning and Computing. 4(3):279-285.doi:10.7763/IJMLC.2014.V4.425

Link to paper Lehmann, K.D.S., Goldman, B.W., Dworkin, I., Bryson, D.M. & Wagner, A.P. 2014. From Cues to Signals: Evolution of Interspecific Communication Via Aposematism and Mimicry in a Predator-Prey System. PLoS One. 9(3): e91783 doi:10.1371/journal.pone.0091783.

Link to paper Fritz, M.L., Yunker, A.J., Walker, E.D. & Dworkin, I. 2014. Daily blood feeding rhythms of laboratory-reared North American Culex pipiens. J. Circadian Rhythms. 12:1. doi:10.1186/1740-3391-12-1

Link to paper Chari, S & Dworkin, I. 2013. The conditional nature of genetic interactions: the consequences of wild-type backgrounds on mutational interactions in a genome-wide modifier screen. PLoS Genetics. 9(8):e1003661. doi/pgen.1003661.

Link to paper Swanson, E.M., McElhinny, T.L., Dworkin, I., Weldele, M.L., Glickman, S.E. & Holekamp, K.E. 2013. Ontogeny of sexual size dimorphism in the spotted hyena (Crocuta crocuta). J. Mammalogy. 94(6):1298-1310

PDF Warren, I.A., Gotoh, H., Dworkin, I.M., Emlen, D.J. & Lavine, L.C. 2013. A general mechanism for conditional expression of exaggerated sexually-selected traits. BioEssays. 35(10):889-99. doi:10.1002/bies.201300031

PDF or Pre-print on ArXiv Chandler, C.H., Chari, S & Dworkin, I. 2013. Does your gene need a background check? How genetic background impacts the analysis of mutations, genes, and evolution. Trends in Genetics. 29(6):358-366. DOI:10.1016/j.tig.2013.01.009

PDF Pitchers, W.R., Brooks, R., Jennions, M.D., Tregenza, T., Dworkin, I. & Hunt, J. 2013. Limited plasticity in the phenotypic variance-covariance matrix for male advertisement calls in the black field cricket, Teleogryllus commodus. J. Evolutionary Biology. 26(5):1060-1078. DOI:10.1111/jeb.12120

PDF Chandler, C.H. Ofria, C. & Dworkin, I. 2013. Runaway Sexual Selection leads to Good Genes. Evolution 67(1):110-119. DOI:10.1111/j.1558-5646.2012.01750.x

PDF Pitchers, W.R. Pool, J & Dworkin, I. 2013. Altitudinal Clinal Variation in Wing Form in African Drosophila melanogaster: One Cline or Many? Evolution 67(2):438-452. DOI:10.1111/j.1558-5646.2012.01774.x

PDF Ingleby, F.C., Hosken, D.J., Flowerts, K., Hawkes, M.F., Lane, S.M., Rapkin, J, Dworkin, I. & Hunt, J. 2013. Genotype-by-environment interactions for cuticular hydrocarbon expression in Drosophila simulans. J. Evolutionary Biology. 26(1):94-107. DOI:10.1111/jeb.12030

PDF Corley-Lavine, L., Hanhn, L.L., Garczynski, S.F., Warren, I.A., Dworkin, I. & Emlen, D.J. 2013. Cloning and characterization of a mRNA encoding an insulin receptor from the horned scarab beetle Onthophagus nigriventris (Coleoptera: Scarabaeidae). Archives of Insect Biochemistry & Physiology. 82(1):43-57. DOI: 10.1002/arch.21072

Payne, M., Dozier, G., Shelton, J., Turner, J., Adams, J., Carter, J., Williams, H., Hansen, C. & Dworkin, I. 2013. Fly wing biometrics using genetic and evolutionary feature extraction. 2013 IEEE Symposium Series on Computational Intelligence and Biometrics. Pp. 42-46. doi: 10.1109/CIBIM.2013.6607912

Colbry, D., Dyer, F., Dworkin, I., Wang, Y and Wang, L. 2013. Speeding up Scientific Imaging Workflows: Design of Automated Image Annotation Tools. User-Center Computer Vision (UCCV). In Press.

PDF Emlen, D.J., Warren, I., Johns, A., Dworkin, I. & Corley Lavine, L. 2012. A mechanism of extreme growth and reliable signaling in sexually selected ornaments and weapons. Science. 337:860-864. DOI: 10.1126/science.1224286

PDF Malek, T., Boughman, J., Dworkin, I., & Peichel, K. 2012. Admixture mapping of male nuptial color and body shape in a recently formed hybrid population of threespine stickleback. In Press. Molecular Ecology. DOI: 10.1111/j.1365-294X.2012.05660.x

PDF Christopher H. Chandler, Genna E. Chadderdon, Patrick C. Phillips, Ian Dworkin, and Fredric J. Janzen. 2012. Experimental evolution of the Caenorhabditis elegans sex determination pathway. Evolution. 66(1):82-93. DOI: 10.1111/j.1558-5646.2011.01420.x

R. Craig Stillwell, Ian M. Dworkin, Alexander W. Shingleton,  W. Anthony Frankino. 2011. Experimental manipulation of body size to estimate morphological scaling relationships in Drosophila. J. Vis. Exp. (56), e3162, DOI : 10.3791/316.

PDF Moczek,A.P., Sultan, S., Foster, S., Ledon-Rettig, C., Dworkin, I., Nijhout, H.F., Abouheif, E., & David W. Pfennig, D.W. 2011.The role of developmental plasticity in evolutionary innovation. Proc. R. Soc. B. 278:2705-13.

PDF Dworkin, I., Anderson, J.A, Idaghour, Y., Kennerly, E.M., Stone, E.A., Gibson, G. 2011. The effects of weak genetic perturbations on the transcriptome of the wing imaginal disc, and its association with wing shape in Drosophila melanogaster. GENETICS. 187:1171-1184. PMID 21288875. DOI: 10.1534/genetics.110.125922.

PDF Swanson, E.M, Dworkin, I., and K. Holekamp. 2011. Lifetime selection on a hypoallometric size trait in the spotted hyena. Proc. R. Soc. B. 278:3277-3285.
See commentary on this paper by: Stephanie Pappas at

PDF Debat, V. Debelle, A. Dworkin, I. 2009. Plasticity, canalization and developmental stability of the Drosophila wing: joint effects of mutations and of developmental temperature. Evolution, 63:2864-2876.

PDF Shingleton, A.W., Estep, C.M., Driscoll, M.V., Dworkin, I. 2009. Many ways to be small: Different environmental regulators of size generate different scaling relationships in Drosophila melanogaster. Proceedings of the Royal Society, London. Series B. 276:2625-2633.

PDF Dworkin, I. Kennerly, E. Tack, D. Hutchinson, J. Brown, J. Mahaffey, J and Gibson, G. 2009. Genomic consequences of background effects on scalloped mutant expressivity in the wing of Drosophila melanogaster. Genetics 181: 1065-1076.

PDF Dworkin, I. and Jones, C.D. 2009. Genetic changes accompanying the evolution of host specialization in Drosophila sechellia. Genetics 181:721-736.


PDF Supplement Hall, C.M. Dworkin, I. Ungerer, M.V, Purugannan, M. 2007. Genetics of canalization in Arabidopsis thaliana. PNAS USA. 104(34): 13717-13722.

PDF Dworkin, I. Lee, W. McCloskey, F and Larsen, E. 2007. Complex genetic interactions govern the temporal effects of Antennapedia on antenna-to-leg transformations in Drosophila melanogaster.


PDF Supplement Dworkin, I. and Gibson, G. 2006. EGF-R and TGF-β signalling contributes to variation for wing shape in Drosophila melanogaster. Genetics. 173:1417-1431.

PDF Emlen, D.J. Szafran, S. Corley, L. Dworkin, I. 2006. Candidate genes for the development and evolution of beetle horns. Heredity. 97:179-91.


PDF Dworkin, I. 2005. Towards a genetic architecture of cryptic genetic variation and genetic assimilation: The contribution of K.G. Bateman. Journal of Genetics. 84(3):223-226.

PDF Dworkin, I. 2005. A Study of Canalization and Developmental Stability in the Sternopleural Bristle System of Drosophila melanogaster. Evolution. 59(7):1500-1509.

PDF Dworkin, I. Palsson, A. Gibson, G. 2005. Replication of an Egfr-wing shape association in a wild-caught cohort of Drosophila melanogaster. Genetics 169:2115-2125.

PDF Palsson, A. Dodgson, J. Dworkin, I. Gibson, G. 2005. Tests for the replication of an association between Egfr and natural variation in Drosophila melanogaster wing morphology. BMC Genetics.6:44.

PDF Dworkin, I. 2005. Canalization, Cryptic Variation and Developmental Buffering: A critical examination and analytical perspective. Chapter 8 in: VARIATION, pp.131-158. B. Hallgrimsson and B.K. Hall editors. Academic Press.
Link to Book.

PDF Dworkin, I. 2005. Evidence for Canalization of Distal-less function in the leg of Drosophila melanogaster. Evolution and Development.7(2): 89-100.


PDF Gibson, G. and Dworkin, I. 2004. Uncovering Cryptic Genetic Variation. Nature Reviews Genetics. 5:681-690.

PDF Palsson, A. Rouse, A. Riley-Berger,R. Dworkin, I. Gibson, G. 2004. Nucleotide Variation in the Egfr Locus of Drosophila melanogaster. Genetics. 167:1999-1212.

PDF Atallah, J*. Dworkin, I.* Cheung, U.* Greene, A. Ing, B. Larsen, E*. 2004. The environmental and genetic regulation of obake expressivity: morphogenetic fields as evolvable systems. Evolution and Development 6(2):114-22 *(co-first authors).


PDF Supplement Dworkin, I. Palsson, A. Birdsall, K. Gibson, G. 2003. Evidence that Egfr contributes to cryptic genetic variation for photoreceptor determination in natural populations of Drosophila melanogaster. Current Biology. 13:1888-1893.

See commentary on this paper by: T. Casci in Nature Review Genetics html, G.Wagner in Current Biology PDF, and in The Economist html PDF.

PDF Dworkin, I. Tanda, S. Larsen, E. 2001. Are entrenched characters developmentally constrained? Creating biramous limbs in an insect. Evolution and Development. 3(6):424-31.