List of



Wilke F, Matthews H, Herrick N, Dopkins N, Claes P, Walsh S. A novel approach to craniofacial analysis using automated 3D landmarking of the skull. Scientific Reports. 2024.

Mohammed J, Arora N, Matthews H.S, Hansen K, Bader M, Walsh S, Shaffer J.R, Weinberg S.M, Swigut T, Claes P, Selleri L, Wysocka J. A common cis-regulatory variant impacts normal-range and disease-associated human facial shape through regulation of PKDCC during chondrogenesis.


Li Z, Melograna F, Hoskens H, Duroux D, Marazita M.L, Walsh S, Weinberg S.M, Shriver M.D, Müller-Myhsok B, Claes P, Van Steen K. netMUG: a novel network-guided multi-view clustering workflow for dissecting genetic and facial heterogeneity. Frontiers in Genetics. 2023

Goovaerts S, Hoskens H, Eller R.J, Herrick N , Musolf A.M, Justice C.M, Yuan M, Naqvi S, Lee M.K, Vandermeulen D, Szabo-Rogers H.L, Romitti P.A, Boyadjiev S.A, Marazita M.L, Shaffer J.R, Shriver M.D, Wysocka J, Walsh S , Weinberg S.M & Claes P Joint multi-ancestry and admixed GWAS reveals the complex genetics behind human cranial vault shape Nature Communications 2023

Herrick N,Walsh S ILIAD: A suite of automated Snakemake workflows for processing genomic data for downstream applications BMC Bioinformatics 2023

Riman S, Ghemrawi MG , Borsuk LA, Mahfouz R, Walsh S, Vallone PM. Sequence-based allelic variations and frequencies for 22 autosomal STR loci in the Lebanese population. FSI Genetics 2023

Peng F, Xiong Z, Zhu G, Hysi PG, Eller RJG , Wu S, Adhikari K, Chen Y, Li Y, Gonzalez-José R, Schüler-Faccini L, Bortolini MC, Acuña-Alonzo V, Canizales-Quinteros S, Gallo C, Poletti G, Bedoya G, Rothhammer F, Uitterlinden AG, Ikram MA, Nijsten T, Ruiz-Linares A, Wang S, Walsh S, Spector TD, Martin NG, Kayser M, Liu F Genome-wide association studies identify DNA variants influencing eyebrow thickness variation in Europeans and across continental populations. Investigative Dermatology 2023

Matthews H.S, Mahdi S, Penington A.J, Marazita M.L, Shaffer J.R, Walsh S , Shriver M.D, Claes P, Weinberg S.M. Using data-driven phenotyping to investigate the impact of sex on 3D human facial surface morphology. Journal of Anatomy 2023

Wilke F, Herrick N, Matthews H, Hoskens H, Singh S , Shaffer J.R, Weinberg S.M, Shriver M.D, Claes P, Walsh S. Exploring regional aspects of 3D facial variation within European individuals. Scientific Reports. 2023

Yuan M, Hoskens H, Goovaerts S, Herrick NG, Shriver MD, Walsh S, Claes P. Hybrid autoencoder with orthogonal latent space for robust population structure inference. Scientific Reports. 2023


Wang H, Fang S, Wilke F, Larsson M, Walsh S. Human Iris Image Analysis for the Classification of Fuchs’ Crypts and Peripupillary Rings Proceedings of SAI Intelligent Systems Conference. 2022

Naqvi S, Hoskens H, Wilke F, Weinberg S.M, Shaffer J.R, Walsh S, Shriver M.D, Wysocka J, Claes P. Decoding the Human Face: Challenges and Progress in Understanding the Genetics of Craniofacial Morphology. Annual Review of Genomics and Human Genetics. 2022


Liu C, Lee MK, Naqvi S, Hoskens H, Liu D, White JD, Indencleef K, Matthews H, Eller RJ, Li J, Mohammed J, Swigut T, Richmond D, Manyama M, Hallgrímsson B, Spritz RA, Feingold E, Marazita ML, Wysocka J, Walsh S, Shriver MD, Claes P, Weinberg SM, Shaffer JR. Genome scans of facial features in East Africans and cross-population comparisons reveal novel associations. PLoS Genetics. 2021

Matthews HS, Palmer RL, Baynam GS, Quarrell OW, Klein OD, Spritz RA, Hennekam RC, Walsh S, Shriver MD, Weinberg SM, BenHallgrimsson B, Hammond P, Penington AJ, Peeters H, Claes P. Large-scale open-source three- dimensional growth curves for clinical facial assessment and objective description of facial dysmorphism. Scientific Reports. 2021

Mahdil SS, Nauwelaers N, Joris P, Bouritsas G, Gong S, Bokhnyak S, Walsh S, Shriver MD, Bronstein M, Claes P. 3D Facial Matching by Spiral Convolutional Metric Learning and a Biometric Fusion-Net of Demographic Properties. IEEE Transactions on Biometrics, Behavior, and Identity Science. 2021

Hoskens H, Liu L, Naqvi S, Lee MK, Eller RJ, Indencleef K, White JD, Li J, Larmuseau MHD, Hens G, Wysocka J, Walsh S, Richmond S, Shriver MD, Shaffer JR, Peeters H, Weinberg SM, Claes P. 3D facial phenotyping by biometric sibling matching used in contemporary genomic methodologies. PLoS Genetics 2021

Naqvi S, Sleyp Y, Hoskens H, Indencleef K, Spence J.P., Bruffaerts R, Radwan A, Eller R.J. , Richmond S, Shriver M.D., Shaffer J.R., Weinberg S.M, Walsh S, Thompson J, Pritchard J.K, Sunaert S, Peeters H, Wysocka J & Claes P. Shared heritability of human face and brain shape. Nature Genetics 2021

Katsara M, Branicki W, Walsh S, Kayser M,Nothnagel M, VISAGE Consortium Evaluation of supervised machine-learning methods for predicting appearance traits from DNA. Forensic Science International: Genetics 2021

Ekrami O, Claes P, Shriver M.D, Weinberg S.M, Marazita M.L, Walsh S, Van Dongen S. Effects of Male Facial Masculinity on Perceived Attractiveness. Adaptive Human Behavior and Physiology 2021

Indencleef K, Hoskens H, Lee M, White J.D, Liu C, Eller R.J , Naqvi S, Wehby G.L, Moreno Uribe L.M, Hecht J.T, Long R.E, Christensen K, Deleyiannis F.W., Walsh S , Shriver M.D., Richmond S, Wysocka J, Peeters H, Shaffer J.R., Marazita M.L., Hens G, Weinberg S.M, Claes P. The Intersection of the Genetic Architectures of Orofacial Clefts and Normal Facial Variation Frontiers in Genetics 2021

White J.D, Indencleef K, Naqvi S, Eller R.J , Hoskens H, Roosenboom J, Lee M.K, Li J, Mohammed J, Richmond S, Quillen E.E, Norton H.L, Feingold E, Swigut T, Marazita M.L, Peeters H, Hens G, Shaffer J.R, Wysocka J, Walsh S, Weinberg S.M, Shriver M.D & Claes P. Insights into the genetic architecture of the human face. Nature Genetics 2021

Ekrami O, Claes P, Van Assche E, Shriver M.D, Weinberg S.M, Marazita M.L, Walsh S, Van Dongen S. Fluctuating Asymmetry and Sexual Dimorphism in Human Facial Morphology: A Multi-Variate Study Symmetry 2021


Katsara M.A, Branicki W, Pośpiech E, Hysi P, Walsh S, Kayser M, Nothnagel M, VISAGE Consortium. Testing the impact of trait prevalence priors in Bayesian-based genetic prediction modeling of human appearance traits. Forensic Science International:Genetics 2020

Chen Y, Branicki W, Walsh S, Nothnagel M, Kayser M, Liu F, VISAGE Consortium. The impact of correlations between pigmentation phenotypes and underlying genotypes on genetic prediction of pigmentation traits. Forensic Science International:Genetics 2020

Li J, González Zarzar T, White J.D, Indencleef K, Hoskens H, Matthews H, Nauwelaers N, Zaidi A, Eller R.J, Herrick N, Günther T, Svensson E.M, Jakobsson M, Walsh S, Van Steen K, Shriver M.D, Claes P. Robust genome-wide ancestry inference for heterogeneous datasets: illustrated using the 1,000 genome project with 3D facial images. Scientific Reports 2020

Ekrami O, Claes P, White J.D, Weinberg S.M, Marazita M.L, Walsh S , Shriver M.D, Van Dongen S. A Multivariate Approach to Determine the Dimensionality of Human Facial Asymmetry. Symmetry 2020


Wang J, Fang S, Fang M, Wilson J, Herrick N, Walsh S. Automatic Landmark Placement for Large 3D Facial Image Dataset. In Proceedings of the IEEE International Conference on Big Data, Los Angeles, CA, USA, 9-13 December 2019; pp. 5088-5093.

K Breslin , B Wills , A Ralf, M Ventayol Garcia, M Kukla-Bartosze, E Pośpiech, A Freire-Aradas, C Xavier, S Ingold, M de La Puente, K.J van der Gaag, N Herrick , C Haas, W Parson, C Phillips, T Sijen, W Branicki, S Walsh, M Kayser. HIrisPlex-S system for eye, hair, and skin color prediction from DNA: massively parallel sequencing solutions for two common forensically used platforms. Forensic Sci. Int. Genet. 2019

Daniels-Higginbotham J, Gorden E.M, Farmer S.K , Spatola B, Damann F, Bellantoni N, Gagnon K.S, de la Puente M, Xavier C, Walsh S , Parson W, McMahon T.P, Marshall C DNA testing reveals the putative identity of JB55, a 19th century vampire buried in Griswold, Connecticut. Genes 2019

R.J Eller, S.C Janga and S Walsh Odyssey: a semi-automated pipeline for phasing, imputation, and analysis of genome-wide genetic data. BMC Bioinformatics 2019

S Brace, Y Diekmann, T Booth, L van Dorp, Z Faltyskova, N Rohland, S Mallick, I Olalde, M Ferry, M 2019 Michel, J Oppenheimer, N Broomandkhoshbacht, K Stewardson, R Martiniano, S Walsh, M Kayser, S Charlton, G Hellenthal, I Armit, R Schulting, O Craig, A Sheridan, M Parker Pearson, C Stringer, D Reich, M Thomas, and I Barnes. Ancient Genomes Indicate Population Replacement in Early Neolithic Britain. Nature Ecology & Evolution. 2019

Peng F, Zhu G, Hysi P.G, Eller R.J, Chen Y, Li Y, Hamer M.A, Zeng C, Hopkins R.L, Jacobus C.L, Wallace P.L, Uitterlinden A.G, Ikram M.A, Nijsten T, Duffy D.L, Medland S.E, Spector T.D, Walsh S, Martin N.G, Liu F, Kayser M. Genome-wide association studies identify multiple genetic loci influencing eyebrow color variation. Investigative Dermatology. 2019


Pośpiech E, Chen Y, Kukla-Bartoszek M, Breslin K, Aliferi A, Andersen J.D, Ballard D, Chaitanya L, Freire-Aradas A, van der Gaag K.J, Girón-Santamaría L, Gross T.E, Gysi M, Huber G, Mosquera- Miguel A, Muralidharan C, Skowron M, Carracedo Á, Haas C, Morling N, Parson W, Phillips C, Schneider P.M, Sijen T, Syndercombe-Court D, Vennemann M, Wu S, Xu S, Jin L, Wang S, Zhu G, Martin N.G, Medland S.E, Branicki W, Walsh S, Liu F, Kayser M, EUROFORGEN-NoE Consortium. Towards broadening Forensic DNA Phenotyping beyond pigmentation: Improving the prediction of head hair shape from DNA. Forensic Sci. Int. Genet. 2018

Kukla-Bartoszek M, Pośpiech E, Spólnicka M, Karlowska-Pik J, Strapagiel D, Zadzińska E, Rosset I, Sobalska M, Słomka M, Walsh S, Kayser M, Sitek A, Branicki W. Investigating the impact of age-depended hair colour darkening during childhood on DNA-based hair colour prediction with the HIrisPlex system. Forensic Sci. Int. Genet. 2018

Chaitanya L, Breslin K, Zuñiga S, Wirken L, Pospiech E, Kukla-Bartoszek M, Sijen T, de Knijff P, Liu F, Branicki W, Kayser M, Walsh S. The HIrisPlex-S system for eye, hair and skin colour prediction from DNA: Introduction and forensic developmental validation. Forensic Sci. Int. Genet. 2018

Liu F, Chen Y, Zhu G, Hysi P.G, Wu S, Adhikari K, Breslin K, Pospiech E, Hamer M.A, Peng F, Muralidharan C, Acuna-Alonzo V, Canizales-Quinteros S, Bedoya G, Gallo C, Poletti G, Rothhammer F, Bortolini M.C, Gonzalez-Jose R, Zeng C, Xu S, Jin L, Uitterlinden A.G, Ikram M.A, van Duijn C.M, Nijsten T, Walsh S , Branicki W, Wang S, Ruiz-Linares A, Spector T.D, Martin N.G, Medland S.E, Kayser M. Meta-analysis of genome-wide association studies identifies 8 novel loci involved in shape variation of human head hair. Human Molecular Genetics 2018


Walsh S , Chaitanya L, Breslin K, Muralidharan C, Bronikowska A, Pospiech E, Koller J, Kovatsi L, Wollstein A, Branicki W, Liu F, Kayser M. Global skin colour prediction from DNA. Human Genetics. 2017

Caliebe A, Walsh S , Liu F, Kayser M, Krawczak M. Likelihood ratio and posterior odds in forensic genetics: Two sides of the same coin. Forensic Science International: Genetics. 2017.

Wollstein A, Walsh S , Liu F, Chakravarthy U, Rahu M, Seland JH, Soubrane G, Tomazzoli L, Topouzis F, Vingerling JR, Vioque J Böhringer S, Fletcher AE, Kayser M. Novel quantitative pigmentation phenotyping enhances genetic association, epistasis, and prediction of human eye colour. Scientific Reports. 2017 Feb 27;7:43359.


Haeusler M, Haas C, Seiler R, Lösch S, Moghaddam N, Walsh S, Kayser M, Rühli F, Janosa M, Papageorgopoulou C. Multidisciplinary identification of the controversial freedom fighter Jörg Jenatsch, assassinated 1639 in Chur, Switzerland. Plos One. 2016.

Chaitanya L, Pajnic IZ, Walsh S, Balažic J, Zupanc T, Kayser M. Bringing colour back after 70 years: Predicting eye and hair colour from skeletal remains of World War II victims using the HIrisPlex system. Forensic Sci. Int. Genetics 2016.

Walsh S , Pospiech E, Branicki W. Commentary: Hot on the Trail of Genes that Shape Our Fingerprints. Invest. Dermat. 2016.


Liu F, Visser M, Duffy D.L, Hysi P.G, Jacobs L.C, Lao O, Zhong K, Walsh S , Chaitanya L et al. Genetics of skin color variation in Europeans: genome-wide association studies with functional follow-up. Human Genetics 2015.


King T.E, Fortes G.G, Balaresque P, Thomas M.G, Balding D, Delser P.M, Neumann R, Parson W, Knapp M, Walsh S, Tonasso L, Holt J, Kayser M, Appleby J, Forster P, Ekserdjian D, Hofreiter M, Schu¨rer K. Identification of the remains of King Richard III. Nat Commun. 2014.

Chaitanya L*, Walsh S*, et al. Collaborative EDNAP exercise on the IrisPlex system for DNA-based prediction of human eye colour. Forensic Sci. Int. Genet. 2014.

Pospiech E, Wojas-Pelc A, Walsh S, Liu F, Maeda H, Ishikawa T, Skowron M, Kayser M, Branicki W. The common occurrence of epistasis in the determination of human pigmentation and its impact on DNA-based pigmentation phenotype prediction. Forensic Sci. Int. Genet. 2014.

Walsh S, Chaitanya L, Clarisse L, Wirken L, Draus-Barini J, Kovatsi L, Maeda H, Ishikawa T, Sijen T, de Knijff P, Branicki W, Liu F, Kayser M. Developmental Validation of the HIrisPlex system: DNA-based eye and hair colour prediction for forensic and anthropological usage. Forensic Sci. Int. Genet. 2014.


Liu F, Walsh S, Kayser M. Of sex and IrisPlex eye colour prediction: A reply to Martinez-Cadenas et al. Forensic Sci. Int. Genet. 2013.

Keating B, Bansal A.T, Walsh S, Millman J, Newman J, Kidd K, Budowle B, Eisenberg A, Donfack J, Gasparini P, Budimlija Z, Henders A.K, Chandrupatla H, Duffy D.L, Gordon S.D, Hysi P, Liu F, Medland S.E, Rubin L, Martin N.G, Spector T.D, Kayser M, on behalf of the International Visible Trait Genetics (VisiGen) Consortium. First All-in-One Tool for DNA Forensics: Parallel Genome-wide Inference of Bio-Geographic Ancestry, Appearance, Relatedness and Gender With Identitas Forensic Chip Int. J. Legal Med. 2013.

Draus-Barini J, Walsh S, Pośpiech E, Kupiec T, Głąb H, Branicki W, Kayser M. Bona fide colour: DNA prediction of human eye and hair colour from ancient and contemporary skeletal remains Investigative Genetics 2013.


Walsh S, Liu F, Wollstein A, Kovatsi L, Ralf A, Kosiniak-Kamysz A, Branicki W, Kayser M. The HIrisPlex system for simultaneous prediction of hair and eye colour from DNA Forensic Sci. Int. Genet 2012.


Walsh S, Wollstein A, Liu F, Chakravarthy U, RahuM, Seland J.H., Soubrane G, Tomazzoli L, Topouzis F, Vingerling J.R., Vioque J, Fletcher A.E., Ballantyne K.N., Kayser M. DNA-based eye colour prediction across Europe with the IrisPlex system. Forensic Sci. Int. Genet 2011.

Branicki W, Liu F, van Duijn K, Draus-Barini J, Pospiech E, Walsh S, Kupiec T, Wojas-Pelc A, Kayser M Model-based prediction of human hair color using DNA variants. Hum. Genet. 2011.


Walsh S, Lindenbergh A, Zuniga S.B, Sijen T, de Knijff P, Kayser M, Ballantyne K.N. Developmental validation of the IrisPlex system: Determination of blue and brown iris colour for forensic intelligence. Forensic Sci. Int. Genet 2010.

Liu F, Wollstein A, Hysi P.G, Ankra-Badu G.A, Spector T.D, Park D, Zhu G, Larsson M, Duffy D.L, Montgomery G.W, Mackey D.A, Walsh S, Lao O, Hofman A, Rivadeneira F, Vingerling J.R, Uitterlinden A.G, Martin N.G, Hammond C.J, Kayser M. Digital quantification of human eye color highlights genetic association of three new loci. PLoS Genet. 2010.

Walsh S, Liu F, Ballantyne K.N, van Oven M, Lao O, Kayser M. IrisPlex: A sensitive DNA tool for accurate prediction of blue and brown eye colour in the absence of ancestry information. Forensic Sci. Int. Genet 2010.