Dr Raymond Tobler

School of Biological Sciences

College of Sciences

Eligible to supervise Masters and PhD - email supervisor to discuss availability.


Ray is an ARC DECRA fellow whose central interest lies in understanding how evolutionary processes have shaped the remarkable biological diversity observed in humans and other species. In his PhD work, Ray used population genetic datasets to explore how fruit flies adapt to novel thermal environments. Active projects include working with Indigenous groups across Australia, New Guinea, and Island South East Asia to build a detailed understanding of the human population history of these regions – including the original settlement and subsequent movements and interactions. An emerging focus involves leveraging large genomic datasets to investigate how historical evolutionary processes, including adaptation, have shaped modern human disease, and exploring the potential of evolutionary genetic knowledge in the development of novel medical treatments (i.e. evolutionary medicine).

Date Position Institution name
2021 - ongoing Research Fellow Evolution of Cultural Diversity Initiative
2019 - ongoing ARC DECRA Fellow University of Adelaide
2016 - 2019 ARC DAATSIA Fellow University of Adelaide

Year Citation
2025 Souilmi, Y., Oliva, A., Davidson, R., Williams, M. P., Ravishankar, S., Roca-Rada, X., . . . Llamas, B. (2025). Lessons learned: Recommendations for reproducible paleogenomic data analyses.. Am J Hum Genet, 112(12), 2830-2841.
DOI
2025 Davidson, R., Roca-Rada, X., Ravishankar, S., Taufik, L., Haarkötter, C., Collen, E., . . . Llamas, B. (2025). Optimized in-solution enrichment of over a million ancient human SNPs. Genome Biology, 26(1), 190-1-190-20.
DOI Scopus5 WoS5 Europe PMC6
2025 Purnomo, G. A., Teixeira, J. C., Sudoyo, H., Llamas, B., & Tobler, R. (2025). Benchmarking Imputed Low Coverage Genomes in a Human Population Genetics Context. Molecular Ecology Resources, 25(8), e70007-1-e70007-12.
DOI
2025 Roca-Rada, X., Davidson, R., Williams, M. P., Villalba-Mouco, V., Carvalho, A. F., Ravishankar, S., . . . Teixeira, J. C. (2025). The genetic history of Portugal over the past 5,000 years.. Genome Biology, 26(1), 248-1-248-39.
DOI Scopus3 WoS4 Europe PMC1
2024 Purnomo, G. A., Kealy, S., O'Connor, S., Schapper, A., Shaw, B., Llamas, B., . . . Tobler, R. (2024). The genetic origins and impacts of historical Papuan migrations into Wallacea. Proceedings of the National Academy of Sciences of USA, 121(52), e2412355121.
DOI Scopus6 WoS4 Europe PMC2
2024 Souilmi, Y., Wasef, S., Williams, M. P., Conroy, G., Bar, I., Bover, P., . . . Mitchell, K. J. (2024). Ancient genomes reveal over two thousand years of dingo population structure. Proceedings of the National Academy of Sciences, 121(30), e2407584121-1-e2407584121-12.
DOI Scopus7 WoS7 Europe PMC6
2024 Johnson, O. L., Tobler, R., Schmidt, J. M., & Huber, C. D. (2024). Population genetic simulation: Benchmarking frameworks for non-standard models of natural selection. Molecular Ecology Resources, 24(3), e13930-1-e13930-13.
DOI Scopus3 WoS3 Europe PMC4
2023 Tobler, R., Souilmi, Y., Huber, C. D., Bean, N., Turney, C. S. M., Grey, S. T., & Cooper, A. (2023). The role of genetic selection and climatic factors in the dispersal of anatomically modern humans out of Africa.. Proc Natl Acad Sci U S A, 120(22), e2213061120.
DOI Scopus19 WoS15 Europe PMC14
2023 Davidson, R., Williams, M. P., Roca-Rada, X., Kassadjikova, K., Tobler, R., Fehren-Schmitz, L., & Llamas, B. (2023). Allelic bias when performing in-solution enrichment of ancient human DNA. Molecular Ecology Resources, 23(8), 1823-1840.
DOI Scopus16 WoS16 Europe PMC15
2023 Johnson, O. L., Tobler, R., Schmidt, J. M., & Huber, C. D. (2023). Fluctuating selection and the determinants of genetic variation. Trends in Genetics, 39(6), 491-504.
DOI Scopus37 WoS33 Europe PMC44
2023 Taufik, L., Teixeira, J. C., Llamas, B., Sudoyo, H., Tobler, R., & Purnomo, G. A. (2023). Human Genetic Research in Wallacea and Sahul: Recent Findings and Future Prospects. Genes, 13(12), 1-11.
DOI Scopus6 WoS4 Europe PMC3
2022 Lan, D., Purnomo, G., Tobler, R., Souilmi, Y., & Llamas, B. (2022). Genozip Dual-Coordinate VCF format enables efficient genomic analyses and alleviates liftover limitations.
DOI
2022 Souilmi, Y., Tobler, R., Johar, A., Williams, M., Grey, S. T., Schmidt, J., . . . Huber, C. D. (2022). Admixture has obscured signals of historical hard sweeps in humans. Nature Ecology & Evolution, 6(12), 2003-2015.
DOI Scopus20 WoS19 Europe PMC31
2021 Cooper, A., Turney, C. S. M., Palmer, J., Hogg, A., McGlone, M., Wilmshurst, J., . . . Zech, R. (2021). A global environmental crisis 42,000 years ago. Science in China Series C: Life Sciences, 371(6531), 811-818.
DOI Scopus94 WoS82 Europe PMC23
2021 Lan, D., Tobler, R., Souilmi, Y., & Llamas, B. (2021). Genozip - a universal extensible genomic data compressor. Bioinformatics, 37(16), 2225-2230.
DOI Scopus36 WoS31 Europe PMC26
2021 Aguiar Macedo Teixeira, J. C., Jacobs, G. S., Stringer, C., Tuke, S., Hudjashov, G., Purnomo, G. A., . . . Helgen, K. M. (2021). Widespread Denisovan ancestry in Island Southeast Asia but no evidence of substantial super-archaic hominin admixture. Nature Ecology & Evolution, 5(5), 616-624.
DOI Scopus45 WoS33 Europe PMC20
2021 Oliva, A., Tobler, R., Cooper, A., Llamas, B., & Souilmi, Y. (2021). Systematic benchmark of ancient DNA read mapping. Briefings in bioinformatics, 22(5), 1-12.
DOI Scopus33 WoS30 Europe PMC33
2021 Souilmi, Y., Lauterbur, M. E., Tobler, R., Huber, C. D., Johar, A. S., Moradi, S. V., . . . Enard, D. (2021). An ancient viral epidemic involving host coronavirus interacting genes more than 20,000 years ago in East Asia. Current Biology, 31(16), 3504-3514.
DOI Scopus65 WoS61 Europe PMC58
2021 Purnomo, G. A., Mitchell, K. J., O'Connor, S., Kealy, S., Taufik, L., Schiller, S., . . . Tobler, R. (2021). Mitogenomes reveal two major influxes of Papuan ancestry across Wallacea following the last glacial maximum and Austronesian contact. Genes, 12(7), 1-21.
DOI Scopus26 WoS24 Europe PMC16
2021 Oliva, A., Tobler, R., Llamas, B., & Souilmi, Y. (2021). BWA-mem is not the best aligner for ancient DNA short reads.
DOI Europe PMC2
2021 Herrando-Pérez, S., Tobler, R., & Huber, C. D. (2021). smartsnp, an r package for fast multivariate analyses of big genomic data. Methods in Ecology and Evolution, 12(11), 2084-2093.
DOI Scopus24 WoS23
2021 Souilmi, Y., Lauterbur, M. E., Tobler, R., Huber, C. D., Johar, A. S., Moradi, S. V., . . . Enard, D. (2021). An ancient viral epidemic involving host coronavirus interacting genes more than 20,000 years ago in East Asia.. Curr Biol, 31(16), 3704.
DOI Scopus4 WoS2 Europe PMC11
2021 Alpaslan-Roodenberg, S., Anthony, D., Babiker, H., Bánffy, E., Booth, T., Capone, P., . . . Zahir, M. (2021). Ethics of DNA research on human remains: five globally applicable guidelines. Nature, 599(7883), 41-46.
DOI Scopus95 WoS77 Europe PMC55
2021 Cooper, A., Turney, C. S. M., Palmer, J., Hogg, A., McGlone, M., Wilmshurst, J., . . . Zech, R. (2021). Response to Comment on "A global environmental crisis 42,000 years ago". Science New York N Y, 374(6570), eabi9756.
DOI Scopus2 WoS2 Europe PMC2
2021 Cooper, A., Turney, C. S. M., Palmer, J., Hogg, A., McGlone, M., Wilmshurst, J., . . . Zech, R. (2021). Response to comment on “A global environmental crisis 42,000 years ago”. Science, 374(6570), 3 pages.
DOI Scopus2
2021 Oliva, A., Tobler, R., Llamas, B., & Souilmi, Y. (2021). Additional evaluations show that specific
BWA‐aln
settings still outperform
BWA‐mem
for ancient DNA data alignment. Ecology and Evolution, 11(24), 18743-18748.

DOI Scopus17 WoS15 Europe PMC19
2020 Tobler, R., Johar, A., Huber, C., & Souilmi, Y. (2020). PolyLinkR: A linkage-sensitive gene set enrichment R package.
2020 Lan, D., Tobler, R., Souilmi, Y., & Llamas, B. (2020). genozip: a fast and efficient compression tool for VCF files. Bioinformatics, 36(13), 4091-4092.
DOI Scopus12 WoS10 Europe PMC12
2020 Teixeira, J., Jacobs, G., Stringer, C., Tuke, J., Hudjashov, G., Purnomo, G., . . . Helgen, K. (2020). Introgression, hominin dispersal and megafaunal survival in Late Pleistocene Island Southeast Asia.
DOI
2020 Souilmi, Y., Tobler, R., Johar, A., Williams, M., Grey, S., Schmidt, J., . . . Cooper, A. (2020). Ancient human genomes reveal a hidden history of strong selection in Eurasia.
DOI Europe PMC1
2020 Souilmi, Y., Lauterbur, E., Tobler, R., Huber, C., Johar, A., & Enard, D. (2020). An ancient viral epidemic involving host coronavirus interacting genes more than 20,000 years ago in East Asia.
DOI Europe PMC3
2019 Richards, S. M., Mitchell, K. J., Tobler, R., & Cooper, A. (2019). Recombinase Polymerase Amplification (RPA) versus PCR for ancient DNA library amplification.
DOI
2019 Richards, S., Mitchell, K., Tobler, R., & Cooper, A. (2019). Recombinase Polymerase Amplification (RPA) versus PCR for ancient DNA library amplification.
DOI Europe PMC1
2019 Barghi, N., Tobler, R., Nolte, V., Jakšić, A. M., Mallard, F., Otte, K. A., . . . Schlötterer, C. (2019). Genetic redundancy fuels polygenic adaptation in Drosophila. PLoS Biology, 17(2), 31 pages.
DOI Scopus185 WoS178 Europe PMC229
2018 Kofler, R., Senti, K. -A., Nolte, V., Tobler, R., & Schlötterer, C. (2018). Molecular dissection of a natural transposable element invasion.. Genome research, 28(6), 824-835.
DOI Scopus54 WoS51 Europe PMC69
2018 Mallard, F., Nolte, V., Tobler, R., Kapun, M., & Schlötterer, C. (2018). A simple genetic basis of adaptation to a novel thermal environment results in complex metabolic rewiring in Drosophila. Genome Biology, 19(1), 119-1-119-15.
DOI Scopus73 WoS74 Europe PMC77
2018 Rohrlach, A. B., Bean, N., Glonek, G., Holland, B., Tobler, R., Tuke, J., & Cooper, A. (2018). Unsupervized Quantification of Demographic Structure for Single-copy Alignments.
DOI
2018 Barghi, N., Tobler, R., Nolte, V., Jaksic, A. M., Mallard, F., Otte, K. A., . . . Schlöetterer, C. (2018). Polygenic adaptation fuels genetic redundancy in<i>Drosophila</i>.
DOI
2017 Mallard, F., Nolte, V., Tobler, R., Kapun, M., & Schlötterer, C. (2017). A simple genetic basis of adaptation to a novel thermal environment results in complex metabolic rewiring in Drosophila.
DOI Europe PMC1
2017 Tobler, R., Rohrlach, A., Soubrier, J., Bover Arbos, P., Llamas, B., Tuke, J., . . . Cooper, A. (2017). Aboriginal mitogenomes reveal 50,000 years of regionalism in Australia. Nature, 544(7649), 180-184.
DOI Scopus169 WoS156 Europe PMC69
2017 Barghi, N., Tobler, R., Nolte, V., & Schlötterer, C. (2017). Drosophila simulans: a species with improved resolution in evolve and resequence studies. G3: Genes, Genomes, Genetics, 7(7), 2337-2343.
DOI Scopus22 WoS21 Europe PMC20
2017 Tobler, R., Nolte, V., & Schlötterer, C. (2017). High rate of translocation-based gene birth on the Drosophila Y chromosome. Proceedings of the National Academy of Sciences of the United States of America, 114(44), 11721-11726.
DOI Scopus34 WoS34 Europe PMC34
2016 Schlötterer, C., Kofler, R., Versace, E., Tobler, R., & Franssen, S. U. (2016). Erratum: Combining experimental evolution with next-generation sequencing: A powerful tool to study adaptation from standing genetic variation (Heredity (2015) 114 (431-440)). Heredity, 116(2), 248.
DOI Scopus6 WoS7 Europe PMC11
2016 Bergland, A. O., Tobler, R., González, J., Schmidt, P., & Petrov, D. (2016). Secondary contact and local adaptation contribute to genome-wide patterns of clinal variation in Drosophila melanogaster. Molecular Ecology, 25(5), 1157-1174.
DOI Scopus102 WoS103 Europe PMC128
2016 Nouhaud, P., Tobler, R., Nolte, V., & Schlötterer, C. (2016). Ancestral population reconstitution from isofemale lines as a tool for experimental evolution. Ecology and Evolution, 6(20), 7169-7175.
DOI Scopus20 WoS20 Europe PMC24
2015 Franssen, S. U., Nolte, V., Tobler, R., & Schlötterer, C. (2015). Patterns of linkage disequilibrium and long range hitchhiking in evolving experimental Drosophila melanogaster populations.. Molecular biology and evolution, 32(2), 495-509.
DOI Scopus65 WoS67 Europe PMC76
2015 Tobler, R., Hermisson, J., & Schlötterer, C. (2015). Parallel trait adaptation across opposing thermal environments in experimental Drosophila melanogaster populations.. Evolution; international journal of organic evolution, 69(7), 1745-1759.
DOI Scopus32 WoS32 Europe PMC28
2015 Schlötterer, C., Kofler, R., Versace, E., Tobler, R., & Franssen, S. (2015). Combining experimental evolution with next-generation sequencing: a powerful tool to study adaptation from standing genetic variation. Heredity, 114(5), 431-440.
DOI Scopus198 WoS199 Europe PMC199
2014 Schlötterer, C., Tobler, R., Kofler, R., & Nolte, V. (2014). Sequencing pools of individuals - mining genome-wide polymorphism data without big funding.. Nature reviews. Genetics, 15(11), 749-763.
DOI Scopus558 WoS544 Europe PMC527
2014 Tobler, R., Franssen, S. U., Kofler, R., Orozco-Terwengel, P., Nolte, V., Hermisson, J., & Schlötterer, C. (2014). Massive habitat-specific genomic response in D. melanogaster populations during experimental evolution in hot and cold environments.. Molecular biology and evolution, 31(2), 364-375.
DOI Scopus111 WoS110 Europe PMC116
2014 Versace, E., Nolte, V., Pandey, R. V., Tobler, R., & Schlötterer, C. (2014). Experimental evolution reveals habitat-specific fitness dynamics among Wolbachia clades in Drosophila melanogaster.. Molecular ecology, 23(4), 802-814.
DOI Scopus34 WoS33 Europe PMC32
2014 Bergland, A., Tobler, R., Gonzalez, J., Schmidt, P., & Petrov, D. (2014). Secondary contact and local adaptation contribute to genome-wide patterns of clinal variation in Drosophila melanogaster.
DOI Europe PMC3
2013 Bastide, H., Betancourt, A., Nolte, V., Tobler, R., Stöbe, P., Futschik, A., & Schlötterer, C. (2013). A Genome-Wide, Fine-Scale Map of Natural Pigmentation Variation in Drosophila melanogaster. Plos Genetics, 9(6), 8 pages.
DOI Scopus122 WoS115 Europe PMC127
2010 Tobler, R., Pledger, S., & Linklater, W. (2010). No evidence for ovarian synchrony or asynchrony in hamadryas baboons. Animal Behaviour, 80(5), 829-837.
DOI Scopus9 WoS8

Year Citation
2025 Bowern, C., Llamas, B., Miceli, L., Tobler, R., & Veth, P. (2025). AUSTRALIAN ARCHAEOLINGUISTICS. In Oxford Handbook of Archaeology and Language (pp. 518-539). Oxford University Press.
DOI Scopus1

Year Citation
2021 Souilmi, Y., Lauterbur, E., Tobler, R., Huber, C., Johar, A. S., & Enard, D. (2021). An ancient coronavirus-like epidemic 25,000 years in ancestral populations from East Asia. Poster session presented at the meeting of AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY. WILEY.

Year Citation
- Souilmi, Y., & Tobler, R. (n.d.). Frequency data and SweepFinder2 output.
DOI
- Tobler, R., Souilmi, Y., & Huber, C. (n.d.). Supplemental Dataset S57.
DOI

Year Citation
- Tobler, R., Huber, C., & Souilmi, Y. (n.d.). Supplementary data 58-114.
DOI
- Tobler, R., Huber, C., & Souilmi, Y. (n.d.). Supplementary data 1-57.
DOI
- Tobler, R., Souilmi, Y., & Huber, C. (n.d.). Supplemental Datasets S1-S56.
DOI

Year Citation
2024 Purnomo, G., Kealy, S., O’Connor, S., Schapper, A., Shaw, B., Llamas, B., . . . Tobler, R. (2024). The genetic origins and impacts of historical Papuan migrations into Wallacea.
DOI
2024 Purnomo, G., Teixeira, J., Sudoyo, H., Llamas, B., & Tobler, R. (2024). Benchmarking Imputed Low Coverage Genomes in a Human Population Genetics Context.
DOI
2024 Roca-Rada, X., Davidson, R., Williams, M., Ravishankar, S., Collen, E., Haarkötter, C., . . . Teixeira, J. (2024). The genetic history of Portugal over the past 5,000 years.
DOI
2024 Williams, A., Tobler, R., Griffiths, B., Ulm, S., Nitschke, M., Bird, M., . . . Bradshaw, C. (2024). Large size of the Australian Indigenous population prior to its massive decline following European invasion.
DOI

Date Role Research Topic Program Degree Type Student Load Student Name
2024 Co-Supervisor Genetic Continuity and Population Dynamics in Eastern Indonesia Doctor of Philosophy Doctorate Full Time Mr Leonard .
2024 Co-Supervisor Genetic Continuity and Population Dynamics in Eastern Indonesia - Doctorate Full Time Mr Leonard .

Date Role Research Topic Program Degree Type Student Load Student Name
2021 - 2022 Co-Supervisor Advances in Genomic Data Compression Doctor of Philosophy Doctorate Full Time Mr Divon Mordechai Lan
2021 - 2024 Co-Supervisor Fluctuating Selection and its Effects on Neutral Genetic Variation Doctor of Philosophy Doctorate Full Time Miss Olivia Lenore Johnson
2018 - 2022 Co-Supervisor Quantifying and Reducing Biases in Paleogenomic Research Doctor of Philosophy Doctorate Full Time Mr Adrien Oliva
2018 - 2022 Co-Supervisor Reconstructing the Population History of Wallacea and Sahul Doctor of Philosophy Doctorate Full Time Dr Gludhug Ariyo Purnomo
2017 - 2022 Co-Supervisor Exploring the Genetic History of the Ancient Near East through the Bronze
and Iron Ages
Doctor of Philosophy Doctorate Full Time Mr Matthew Peter Williams

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