Publications
2025
Burgin, C. J., Zijlstra, J. S., Becker, M. A., Handika, H., Widness, J., Liphardt, S., Huckaby, D. G., and Upham, N. S. 2025. How many mammal species are there now? Updates and trends in taxonomic, nomenclatural, and geographic knowledge. Journal of Mammalogy, gyaf047. https://doi.org/10.1093/jmammal/gyaf047
Giarla, T. C., Achmadi, A.S., Fabre, P. H., Handika, H., Chipps, A. S., Swanson, M.T., Nations, J. A., Morni, M. A., William-Dee, J., Inayah, N., Jayanti, E. D., Nurdin, M.R.T.J., Griffin, K. E., Khan, F. A. A., Heaney, L. R., Rowe, K. C., and Esselstyn, J. A. 2025. Systematics and historical biogeography of Crunomys and Maxomys (Muridae: Murinae), with the description of a new species from Sulawesi and new genus-level classification. Journal of Mammalogy, gyaf006. https://doi.org/10.1093/jmammal/gyaf006
Adams, A.L., Achmadi, A.S., Mursyid, A., Handika, H., Nurdin, M.R.T., Esselstyn, J.A., Perkins, S.L., Rowe, K.M., and Rowe, K.C. 2025. Wildlife disease surveillance from village to peak: Trypanosome infections of mammals on Sulawesi revealed higher prevalence in intact montane forests. THERYA, 16(1), 125-141.
2024
Handika, H. and Esselstyn, J. A. 2024. SEGUL: Ultrafast, memory-efficient and mobile-friendly software for manipulating and summarizing phylogenomic datasets. Molecular Ecology Resources, 24(7). https://doi.org/10.1111/1755-0998.13964
Nations, J.A., Kohli, B.A., Handika, H., Achmadi, A.S., Polito, M.J., Rowe, K.C., and Esselstyn, J. A. 2024. The roles of isolation and interspecific interaction in generating the functional diversity of an insular mammal radiation. Oikos, p.e10888. https://doi.org/10.1111/oik.10888
Nations, J. A., Handika, H., Mursyid, A., Butsa, R. D., Apandi, Achmadi, A. S., and Esselstyn, J. A. 2024. Three new species of shrew (Soricidae: Crocidura) from West Sumatra, Indonesia: elevational and morphological divergence in syntopic sister taxa. Journal of Mammalogy, 105 (2), 372-389. https://doi.org/10.1093/jmammal/gyad126
2023
Mursyid, A., Achmadi, A.S., Novarino, W., Handika, H., Nugroho, H.A., Anita, S., Adams, A.L., Rowe, K.M.C., and Rowe, K.C. 2023. Trypanosoma (Euglenozoa: Kinetoplastea) infections in rodents, bats, and shrews along an elevation and disturbance gradient in Central Sulawesi, Indonesia. Contributions to Mammalogy and Zooarchaeology of Wallacea, pp.663-671. http://dx.doi.org/10.3853/j.2201-4349.75.2023.1786
Achmadi, A. S., Fahri, F., Gazzard, A., Handika, H., Inayah, N., Kennerley, R. J., Lanusi, A. A., Nangoy, M., Nurdin, M. R., Rowe, K. C., and Sheherazade. 2023. Conservation status and priorities for Sulawesi's unique small mammal fauna. Journal of Mammalogy, 104 (3), 1-12. https://doi.org/10.1093/jmammal/gyad126
Swanson, M.T., Henson, M.W., Handika, H., Achmadi, A.S., Anita, S., Rowe, K.C., and Esselstyn, J.A. 2023. Mycoplasmataceae dominate microbial community differences between gut regions in mammals with a simple gut architecture. Journal of Mammalogy, 104(1), pp.146-158. https://doi.org/10.1093/jmammal/gyac098
2021
Esselstyn, J.A., Achmadi, A.S., Handika, H., Swanson, M. T., Giarla, T.C., and Rowe, K.C. 2021. Fourteen new, endemic species of shrew (genus Crocidura) from Sulawesi reveal a spectacular island radiation. Bulletin of the American Museum of Natural History, 454(1), pp.1-108. https://doi.org/10.1206/0003-0090.454.1.1
Handika, H., Achmadi, A.S., Esselstyn, J.A., and Rowe, K.C. 2021. Molecular and morphological systematics of the Bunomys division (Rodentia: Muridae), an endemic radiation on Sulawesi. Zoologica Scripta, 50(2), pp. 141–154. https://doi.org/10.1111/zsc.12460
Nations, J.A., Mursyid, A., Darma Busta, R., Adrian, S.P., Handika, H., Apandi, Achmadi, A.S., and Esselstyn, J.A. 2021. The first report of albinism in a Sundaland endemic rodent. Mammalia, 85(2), pp.168-172. https://doi.org/10.1515/mammalia-2020-0047
2020
Harapan, T.S., Agung, A.P., Handika, H., Novarino, W., Tjong, D.H., and Tomlinson, K.W. 2020. New records and potential geographic distribution of Elongated Caecilian, Ichthyophis elongatus Taylor, 1965 (Amphibia, Gymnophiona, Ichthyophiidae), endemic to West Sumatra, Indonesia. Check List, , 16(6), pp. 1695-1701. https://doi.org/10.15560/16.6.1695
Winterhoff, M.L., Achmadi, A.S., Roycroft, E.J., Handika, H., Putra, R.T.J., Rowe, K.M.C., Perkins, S., and Rowe, K.C. 2020. Native and introduced trypanosome parasites in endemic and introduced murine rodents of Sulawesi. Journal of Parasitology, 106(5), pp.523-536. https://doi.org/10.1645/19-136
2019 and earlier
Esselstyn, J.A., Achmadi, A.S., Handika, H., Giarla, T.C., and Rowe, K.C. 2019. A new climbing shrew from Sulawesi highlights the tangled taxonomy of an endemic radiation. Journal of Mammalogy, 100(6), pp.1713-1725. https://doi.org/10.1093/jmammal/gyz077
Janra, M.N., Noske, R.A., Chornelia, A., Handika, H., Mursyid, A., Agung, A., and Saputra, A. 2018. A second breeding record of Chestnut-naped Forktails Enicurus ruficapillus in Sumatra, and observations suggesting females alone incubate. Kukila, 21, pp.13-16.
Demos, T.C., Achmadi, A.S., Handika, H., Rowe, K.C., and Esselstyn, J.A. 2017. A new species of shrew (Soricomorpha: Crocidura) from Java, Indonesia: possible character displacement despite interspecific gene flow. Journal of Mammalogy, 98(1), pp.183-193. https://doi.org/10.1093/jmammal/gyw183
Demos, T.C., Achmadi, A.S., Giarla, T.C., Handika, H., Rowe, K.C., and Esselstyn, J.A. 2016. Local endemism and within‐island diversification of shrews illustrate the importance of speciation in building Sundaland mammal diversity. Molecular ecology, 25(20), pp.5158-5173. https://doi.org/10.1111/mec.13820
Esselstyn, J.A., Achmadi, A.S., Handika, H., and Rowe, K.C. 2015. A hog-nosed shrew rat (Rodentia: Muridae) from Sulawesi Island, Indonesia. Journal of Mammalogy, 96(5), pp.895-907. https://doi.org/10.1093/jmammal/gyv093
Handika, H., Nurdin, J., and Rizaldi. 2013. Small Terrestrial Mammals Community on Mount Singgalang, West Sumatra. Jurnal Biologi Universitas Andalas, 2(2), pp.103-109. https://doi.org/10.25077/jbioua.2.2.%25p.2013
Manuscript in prep
Handika, H., Moncrieff, A. E., Del-Rio, G., Canton, R., Jaiswal, A., Gan, J. B., Henderson, C., Rutledge, S. L., Hernández-Canchola, G., Babb-Biernacki, S. J., Chipps, A. S., Pérez-Amaya, N., Chornelia, A., Morales-Martínez, D. M., McCallum, Q., Thom, G., and Esselstyn, J. A. In prep. NAHPU: A field cataloging application for efficient, repeatable, and inclusive collection-based evolutionary studies.
Handika, H., Del-Rio, G. D., and Esselstyn, J. A. In prep. ULLAR: An ULtrafast, scaLable, Accessible, and Reproducible phylogenomic pipeline.
Handika, H., Rowe, K.C., Achmadi, A.S., Zulkurnia Irsaf, Swanson, M. T., and Esselstyn, J.A. In prep. An island of islands: complex gene tree discordance and local endemism on a recent, rapid rodent radiation on Sulawesi.
Handika, H., Rowe, K.C., Achmadi, A.S., Swanson, M. T., and Esselstyn, J.A. In prep. A new species of Frateromys from the Northern Peninsula of Sulawesi, Indonesia.
Teaching, Mentoring, & Workshops
Courses Taught
Lousiana State University, USA
BIOL 2153 Principles of Genetics
Teaching assistant · Fall 2024
This course covers several topics, including Mendelian inheritance, gene mapping, population genetics, and genomics. I taught recitation sections and assisted with homework, exam preparation, and grading. My class size was about 60 students divided into two sections.
BIOL 4141 Mammalogy
Teaching assistant · Fall 2021
This course covers the evolution and systematics of mammals. I taught lab section using specimens from the museum collection. My class size was 16 students.
BIOL 1209 Introduction to Biology
Teaching assistant · Spring 2021, 2025
This course is an introduction to the principles of biology that covers topics on ecology, evolution, and diversity. I taught lab section and assisted with grading. My class size was about 50 students.
Andalas University, Indonesia
PAB125 Computer Application for Biology
Instructor · 2nd semester 2012
This course introduces students to computer applications in biology that covers topics on basic computational statistics, geographic information system (GIS), and scientific photography. I developed the syllabus, taught lectures, and graded assignments. My class size was 70 students.
Workshops Taught

Molecular Analyses of Indonesian Mammals
Instructor · 18, 19, 25, and 26 May 2023
Organized by Generasi Biologi Indonesia
This workshop covers an introduction to mammalian phylogenetics, multi-locus DNA sequencing analyses, and genomic analyses using current methods and software. The workshop was tailored with data from Indonesian mammals, but the methodologies are applicable to studies utilizing both Sanger and short-read sequencing data.
Coding for Conservation: Integrating GitHub to Your Workflow
Instructor · 8-9 April 2023
Organized by Tambora Muda, RLadies Jakarta, and RLadies Bogor
This workshop was designed to introduce students to version control and collaborative coding using GitHub. I taught the basics of Git and GitHub, including creating a repository, staging, making commits, and creating pull requests. I also introduced students to advanced approach to R dependency management and writing cross-platform code.
Introduction to Phylogenetics
Instructor · 15 January 2020
Organized by Graduate Student Association at Andalas University, Indonesia
This workshop covers the fundamental concepts of phylogenetics, including phylogenetic tree construction and interpretation. It also introduces students to multi-locus DNA sequencing analyses using IQ-TREE.
Student Mentees
Fieldwork, survey design, scientific, and/or grant writing
- Fatimah (BS student in Biology, Andalas University, Indonesia).
2014–2017.
Received $5,300 from the Mohammed Bin Zayed Conservation Fund to study the Mentawai Langurs.
- Ahmad Mursyid (BS student in Biology, Andalas University, Indonesia). 2012–2018.
- Zulkurnia Irsaf (BS student in Biology, Tadulako University, Indonesia). 2018–2020.
- Tri S. Harapan (BS student in Biology, Andalas University, Indonesia). 2020.
- Fajar Alfitrian (BS student in Biology, Soedirman University, Indonesia). 2023–2025.
- Alfath Fanidya (MS student in Biology, Bogor Agricultural
University, Indonesia). 2023–now.
Received $13,000 from the Fonseca Species Conservation Fund to study the Javan bamboo rats.
- Lutfi S. Maulidiyah (BS student in Biology, University of Lampung, Indonesia). 2023–2025.
- Akbar F. Hilmi (BS student in Biology, University of Lampung, Indonesia). 2023–2025.
Specimen curation
- Ericka Boyden (BS student in Biology, Louisiana State University). Fall 2019–Spring 2020.
- Brittany E. Thames (BS student in Biology, Louisiana State University). Spring 2019.
- Emily A. Bowers (BS student in Biology, Louisiana State University). Fall 2020–Spring 2021.
Genomic Analyses
- Murky Mutchler (BS student in Biology, Louisiana State
University / PhD, the University of Chicago). 2021.
Won a poster award at the 2021 Wilson Ornithological Society Meeting.
- Ryan Zucker (BS student in Biology, Cornell University). 2024.
Software Development
- Filipe Delgado (MS student in Computer Science, Georgia Institute of Technology). Fall 2025.
- Cody Henderson (MS student in Computer Science, Georgia Institute of Technology). Fall 2025–now.
- Lawrence Bohan Gan (MS student in Computer Science, Georgia Institute of Technology). Fall 2025–now.
- Aakanksha Jaiswal (MS student in Computer Science, Georgia Institute of Technology). Fall 2025–now.
- Kira L. Nichtawitz (BS student in Computer Science, University of Florida). Fall 2025–now.
- Julian D. Perez (BS student in Computer Engineering, University of Florida). Fall 2025–now.
- Ethan E. Wolfe (BS student in Data Science, University of Florida). Fall 2025–now.
- Zahara M. Shuemaker (BS student in Computer Science, University of Florida). Fall 2025–now.
Outreach & Service
Outreach Events

KCA-LH Rafflesia FMIPA Unand Conservation Club, Indonesia: "Manusia Hebat Tidak Hanya Terlahir dari Bakat (Great People Are Not Only Born from Talents)", 18 October 2020.
Tambora Muda Indonesia: "Bincang Hari Keanekaragaman Hayati (Biodiversity Day Talk)", 22 May 2020.
LSU Museum of Natural History, USA: "Night at the Museum", 14 November 2019.
LSU Museum of Natural History, USA: "Night at the Museum", 11 August 2018.
Museums Victoria, Australia: "Meet the Scientists", 22 August 2015.
KCA-LH Rafflesia FMIPA Unand Conservation Club, Indonesia: "Peringatan Hari Bumi (Earth Day Celebration)", 2009-2012.
Interviews

Science Next: "What Could Recovery Look Like for Blazing Australia?", 27 January 2020.
The Indonesian Ministry of Finance Magazine, Media Keuangan: "Berniat Terus Meneliti Tikus (Intending to Keep Researching Rats)", Volume XII / NO. 118 / July 2017.
Kompas.Com-ABC Australia Plus: "Kisah Mahasiswa Indonesia Terlibat Penemuan Tikus Hidung Babi (The Story of Indonesian Students Involved in the Discovery of the Hog-Nosed Rat)", 26 November 2015.
Popular Articles
Republika-ABC AustraliaPlus: "Hal yang Perlu Dilakukan agar Kuliah S2 di Luar Negeri (What You Need to Do to Study Abroad for a Master's Degree)", 16 February 2017.
DetikNews-ABC AustraliaPlus: "Langkah-Langkah Kuliah ke Luar Negeri: Dari A-Z (Steps to Study Abroad: From A-Z)", 7 June 2016.
Republika-ABC AustraliaPlus: "Menentukan Topik Penelitian: Dilema Kecepatan Tamat dan Kualitas Penelitian (Determining Research Topics: The Dilemma of Graduating Quickly and Research Quality)", 10 May 2016.
TribunNews-ABC AustraliaPlus: "Pengalaman Heru Handika Menyikapi Kerja Sama dengan Peneliti Asing (Heru Handika's Experience in Dealing with Cooperation with Foreign Researchers)", 8 May 2016.
DetikNews-ABC AustraliaPlus: "Jurus Kuliah ke Luar Negeri Berkat Jaringan Pertemanan (Study Abroad Tips, Thanks to Friendship and Networks)", 15 April 2016.
Republika-ABC AustraliaPlus: "Kiat Meningkatkan TOEFL Melebihi 550 (Tips to Improve TOEFL Scores Beyond 550)", 7 March 2016.
DetikNews-ABC AustraliaPlus: "Mengenal LoA dan Cara Mendapatkan 'Surat Sakti' S2/S3 di Luar Negeri (Getting to Know LoA and How to Get a 'Magic Letter' for Master's/PhD Abroad)", 3 February 2016.
Republika-ABC AustraliaPlus: "Bagaimana Menulis Esai Untuk Mendapatkan Beasiswa LPDP (How to Write an Essay to Get an LPDP Scholarship)", 11 January 2016.