†Camelosphecia fossor Boudinot et al., 2020 holotype female from Kachin amber.

Dr. Brendon E. Boudinot

Alexander von Humboldt Research Fellow
†Camelosphecia fossor Boudinot et al., 2020 holotype female from Kachin amber.
Image: Phil Barden
Dr. Brendon E. Boudinot
Dr. Brendon E. Boudinot
Image: Brendon E. Boudinot


Dr. Brendon E. Boudinot

Friedrich Schiller University Jena
Institute of Zoology and Evolutionary Research
Phyletisches Museum
Vor dem Neutor 1 (room 206)
07743 Jena

Tel.: +49 3641 9-49181

Research Interest

My overarching interest is the origin and evolution of complex phenotypic systems. The primary foci of my studies have been the malegenitalia of insects, the locomotory apparati of Hymenoptera, and the evolution and systematics of Formicidae and the Aculeata. My research integrates comparative anatomy, phylogenetic modeling, paleontology, and developmental biology to systematically resolve the patterns and pathways of morphological evolution. Through my work, I also provide identification resources for ants and other Hymenoptera, with special emphasis on the statistical resolution of character polarities for taxon diagnosis.

My work integrates fossils with molecular data to estimate phylogeny and the pathways of morphological evolution.
My work integrates fossils with molecular data to estimate phylogeny and the pathways of morphological evolution.
Graphic: Brendon Boudinot

Research Topics

  • Phylogenetics (morphological, molecular)
  • Arthropod anatomy (skeletomusculature)
  • Evolutionary morphology
  • Systematics of Formicidae and Hymenoptera
  • Paleontology

Research Methods

  • Sanger and UCE sequencing, analysis
  • Bayesian and maximum likelihood phylogenetics
  • Divergence dating, phylogeography, ancestral state estimation
  • µ-CT segmentation, 3D rendering
  • Stereomicroscopy, scanning electron microscopy
  • Linear and geometric morphometrics

Short CV


2014–2020      Ph.D., Entomology. University of California, Davis (advisor: Dr. Philip S. Ward).

2008–2012      B.Sc., Entomology. The Evergreen State College, Washington (mentor: Dr. John T. Longino).

Professional appointments

2020–        Alexander von Humboldt Research Fellow. Institut für Zoologie und Evolutionsforschung, Friedrich-Schiller-Universität Jena, Germany.

2012–13    Scientific Technician. The University of Utah.

2009–12    Fellow of the Natural History Collections. The Evergreen State College.

Honors & Awards

2022    Bernhard Rensch Prize. Society for Biological Systematics (GfBS, Germany).

2020    Robert Evans Snodgrass Memorial Research Award for Outstanding Ph. D. Dissertation. Systematics & Evolutionary Biology Section, Entomol. Soc. America.

2019    John Henry Comstock Graduate Student Award. Pacific Branch of the Entomol. Soc. America.

2019    Graduate Student Service Award. Department of Entomology & Nematology, UC Davis.

2018    Linnaean Games Regional & National Champion. Entomol. Soc. America.

2017    President’s Prize. Graduate level. Entomol. Soc. America: Systematics & Evolutionary Biology.

2016    Linnaean Games Regional & International Champion. International Congress of Entomol.

2015    Linnaean Games Regional & National Champion. Entomol. Soc. America.

2014    President’s Prize. Graduate level. Entomol. Soc. America: Systematics & Evolutionary Biology.

2012    President’s Prize. Undergraduate level. Entomol. Soc. America: Systematics & Evolutionary Biology.

Taxonomic patronym

2021    Syscia boudinoti, Longino & Branstetter, 2021. Insect Syst. Divers. 5, 1–51.

Scientific affiliations

American Association for the Advancement of Science (AAAS)
Coleopterists Society (CS)
Entomological Society of America (ESA)
International Society of Hymenopterists (ISH)
International Union for the Study of Social Insects (IUSSI)


Boudinot, B.E., Fikáček, M., Lieberman, Z.E., Kusy, D., Bocak, L., McKenna, D.D., Beutel, R.G., 2022. Systematic bias and the phylogeny of Coleoptera — a response to Cai et al. (2022) following the responses to Cai et al. (2020). Syst. Entomol., 2022, 1–10. https://doi.org/10.1111/syen.12570External link

Tozetto, L., Chaul, J.C.M., Boudinot, B.E., Lattke, J.E., 2022. Review of the Leptogenys unistimulosa species group (Hymenoptera: Formicidae) with the description of a new Amazonian species. Rev. Bras. Entomol., 66(3), e20220045. https://doi.org/10.1590/1806-9665-RBENT-2022-0045External link

Richter, A., Boudinot, B.E., Yamamoto, S., Katzke, J., Beutel, R.G., 2022. The first reconstruction of the head anatomy of a Cretaceous insect, †Gerontoformica gracilis (Hymenoptera, Formicidae), and the early evolution of the ants. Insect Syst. Divers., 6(5), 4, 1–80. https://doi.org/10.1093/isd/ixac013External link

Boudinot, B.E., Richter, A.K., Hammel, J., Szwedo, J., Bojarski, B., Perrichot, V., 2022. Genomic-phenomic reciprocal illumination: †Desyopone gen. nov., an exceptional aneuretine-like fossil ant from Ethiopian amber (Hymenoptera: Formicidae: Ponerinae). Insects, 13, 796. https://doi.org/10.3390/insects13090796External link

Boudinot, B.E., Yan, E.V. Prokop, J., Luo, X.-Z., Beutel, R.G., 2022. Permian parallelisms: Reanalysis of †Tshekardocoleidae sheds light on the earliest evolution of the Coleoptera. Syst. Entomol., XX, 1–28. https://doi.org/10.1111/syen.12562External link

Perrichot, V., Boudinot, B.E., Engel, M.S., Xu, C., Bojarski, B., Szwedo, J., 2022. Ants (Hymenoptera: Formicidae) in Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota. Zool. J. Linn. Soc., 1–17. https://doi.org/10.1093/zoolinnean/zlac053External link

Lieberman, Z., Billen, J., van de Kamp, T., Boudinot, B.E., 2022. The abdominal anatomy of the worker of Amblyopone australis ants (Formicidae: Amblyoponinae), with special reference to skeletomusculature. J. Morphol. https://doi.org/10.1002/jmor.21471External link

Aibekova, L., Boudinot, B.E., Beutel, R.G., Richter, A., Keller, R.A., Hita-Garcia, F., Economo, E.P., 2022. The skeletomuscular system of the mesosoma of Formica rufa workers (Hymenoptera: Formicidae). Insect. Syst. Divers. 6, 1–26. https://doi.org/10.1093/isd/ixac002External link

Boudinot, B.E., Borowiec, M.L., Prebus, M.M. 2022. Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini, and the subfamily Formicinae (Hymenoptera: Formicidae). Syst. Entomol. 47, 113–151. https://doi.org/10.1111/syen.12522External link

Boudinot, B.E., Richter, A., Beutel, R.G., Katzke, J., Keller, R.A., Economo, E.P., Yamamoto, E., 2022. Evidence for the evolution of eusociality in stem ants and a systematic revision of †Gerontoformica (Hymenoptera, Formicidae). Zool. J. Linn. Soc. https://doi.org/10.1093/zoolinnean/zlab097External link

Mapalo, M.A., Robin, N., Boudinot, B.E., Ortega-Hernández, J. and Barden, P. 2021. A tardigrade in Dominican amber. Proc. Roy. Soc. B 288, 20211760. Doi: 10.1098/rspb.2021.1760.

Boudinot, B.E., Moosdorf, O.T.D., Beutel, R.G. and Richter, A. 2021. Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): Patterns of sex- and caste-limited traits in the sausagefly and the driver ant. J. Morphol. 282, 1616–1658. Doi: 10.1002/jmor.21410.

Ward, P.S. and Boudinot, B.E. 2021. Grappling with homoplasy: taxonomic refinements and reassignments in the ant genera Camponotus and Colobopsis (Hymenoptera: Formicidae). Arthropod Syst. Phylog. 79, 37–56. Doi: 10.3897/asp.79.e66978.

Richter, A., Keller, R.A., Hita Garcia, F., Billen, J., Katzke, J., Boudinot, B.E., Economo, E.P. and Beutel, R.G. 2021. Head anatomy of Protanilla lini (Leptanillinae, Formicidae, Hymenoptera) and a hypothesis of their mandibular movement. Myrmecol. News 31, 107–136. Doi: 10.25849/myrmecol.news_031:107.

Csősz, S., Seifert, B., Mikó, I., Boudinot, B.E., Borowiec, M.L., Fisher, B.L., Prebus, M., Puniamoorthy, J., Rakotonirina, J.-C., Rasoamanana, N., Schultz, R., Trietsch, C., Ulmer, J.M. and Elek, Z. 2021. Insect morphometry is reproducible under average investigation standards. Ecol. Evol. 11, 547–559. Doi: 10.1002/ece.3.7075.

Boudinot, B.E., Beutel, R.G., Gorb, S.N. and Polilov, A.A. 2021a. Functional diversity of attachment and grooming leg structures is retained in all but the smallest insects. J. Zool. (Lond.) 313, 99–11. Doi: 10.1111/jzo.12840.

Sharaf, M., Mohamed, A.A., Boudinot, B.E., Wetterer, J.K., Aldawood, A.S., al Dhafer, H.M. and Hita Garcia, F. 2021.Monomorium (Hymenoptera: Formicidae) of the Arabian Peninsula with description of two new species, M. heggyi sp. n. and M. khalidi sp. n. PeerJ 9, e10726. Doi: 10.7717/peerj.10726.

Pohl, H., Wipfler, B., Boudinot, B.E. and Beutel, R.G. 2021. On the value of Burmese amber for understanding insect evolution: Insights from †Heterobthmilla – an exceptional stem-group genus of Strepsiptera (Insecta). Cladistics 37, 211–229. Doi: 10.1111/cla.12433.

Boudinot, B.E., Perrichot, V. and Chaul, J.C.M. 2020. †Camelosphecia gen. nov., lost ant-wasp intermediates from the mid Cretaceous (Hymenoptera, Formicidae). ZooKeys 1005, 21–55. Doi: 10.3897/zookeys.1005.57629.

Cao*, H., Boudinot*, B. E., Wang, Z., Miao, X., Shih, C., Ren, D. and Gao., T. 2020b. Two new iron-maiden ants from Burmese amber (Hymenoptera: Formicidae: †Zigrasimeciini). Myrmecol. News 30, 161–173. *Authors contributed equally.

Cao*, H., Boudinot*, B.E., Shih, C., Ren, D. and Gao, T. 2020a. Cretaceous ants shed new light on the origins of worker polymorphism. Sci. China: Life Sci.. Doi: 10.1007/s11427-019-1617-4. *Authors contributed equally.

Boudinot, B.E. and Dungey, D.R. 2020. †Eorhopalosoma lohrmanni, a new species of Rhopalosomatidae from mid-Cretaceous amber of northern Myanmar (Hymenoptera, Aculeata: Vespoidea). Cretac. Res. 108, 104339.

Wipfler, B., Kočárek, P., Richter, A., Boudinot, B.E., Bai, M. and Beutel, R.G. 2019. Structural features and life habits of †Alienoptera (Polyneoptera, Dictyoptera, Insecta). Palaeoentomol. 2, 465–473.

Boudinot, B.E. 2018. A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences, with emphasis on the Endopterygota. Arthropod Struct. Devel. 47, 563–613.

Barden, P., Boudinot, B.E. and Lucky, A. 2017. Where fossils dare and males matter: combined morphological and molecular analysis untangles the evolutionary history of the spider ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Invert. Syst. 31, 765–780.

Boudinot, B.E., Probst, R.S., Brandão, C.R.F., Feitosa, R.M. and Ward, P.S. 2016a. Out of the Neotropics: newly discovered relictual species sheds light on the biogeographic history of spider ants (Leptomyrmex, Dolichoderinae, Formicidae). Syst. Entomol. 41, 658–671.

Boudinot, B.E., Wachkoo, A.A. and Bharti, H. 2016b. The first ergatoid male of Platythyrea (Hymenoptera: Formicidae: Ponerinae), with contribution to colony labor suggested by observation and comparative morphology. Myrmecol. News 22, 59–64.

Probst*, R.S., Guénard, B. and Boudinot*, B.E. 2015. Towards understanding the predatory ponerine ant Myopias(Hymenoptera: Formicidae), including a key to global species, male-based generic diagnosis, and new species description. Sociobiol. 62, 192–212. *Authors contributed equally.

Boudinot, B.E. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): A new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. Euro. J. Taxon. 102, 1–62.

Boudinot, B.E. 2014. Rediscovery of Tricytarus Donisthorpe, 1947, a new genus-level synonym of Meranoplus F. Smith 1853 (Hymenoptera: Formicidae: Myrmicinae). Myrmecol. News 20, 95–100.

MacGown, J.A., Boudinot, B.E., Deyrup, M. & Sorger, D.M. 2014. A review of the Nearctic Odontomachus(Hymenoptera: Formicidae: Ponerinae) with a treatment of the males. Zootaxa 3802, 515–552.

Boudinot, B.E., Sumnicht T.P. and Adams, R.M.M. 2013. Central American ants of the genus Megalomyrmex Forel (Hymenoptera: Formicidae): six new species and keys to workers and males. Zootaxa 3732, 1–82.

Longino, J.T. and Boudinot B.E. 2013. New species of Central American Rhopalothrix Mayr 1870 (Hymenoptera, Formicidae). Zootaxa 3616 301–324.

Boudinot, B.E. and Fisher, B.L. 2013. A taxonomic revision of the Meranoplus F. Smith of Madagascar (Hymenoptera: Formicidae: Myrmicinae) with descriptions of queens and males. Zootaxa 3635, 301–339.

Boudinot, B.E. 2013. The male genitalia of ants: musculature, homology, and functional morphology (Hymenoptera: Aculeata: Formicidae). J. Hym. Res. 30, 29–49.

Book chapters

Delsinne, T., Boudinot, B.E., Leponce, M. and Serna, F. 2019. Capítulo 13: Glosario de morfología. In: Fernandez, F., Guerrero, R. & Delsinne, T. (eds.) Hormigas de Colombia. Bogotá: Universidad Nacional de Colombia, Facultad de Ciencias, pp. 387–457.

Boudinot, B.E. 2019. Capítulo 15: Clave para las subfamilias y géneros basado en los machos. In: Fernandez, F., Guerrero, R. & Delsinne, T. (eds.) Hormigas de Colombia. Bogotá: Universidad Nacional de Colombia, Facultad de Ciencias, pp. 487–499.


Boudinot, B.E. 2020. Systematic and Evolutionary Morphology: Case Studies on Formicidae, Mesozoic Aculeata, and Hexapodan Genitalia. Department of Entomology & Nematology, University of California, Davis. 573 pp.


Boudinot, B.E. 2019. Toward phylomics in entomology: Current systematic, evolutionary, and phylogenetic morphology. Insect Systematics and Diversity, 3(6:4): 1–4.

Boudinot, B.E. 2017. Letter to the Editor: “21st Century Morphology”. Hamuli, the Newsletter for the International Society of Hymenopterists.

Media Coverage

2022 February 22.  Division of labour in ants goes back over 100 million years.External link idw scientific information service.

2022 January 27. Ant Research: The story behind the Lasius work by three alumni of the Phil Ward lab. External link Entomology & Nematology News.

2021 October 5. “Once-in-a generation” tardigrade fossil discovery reveals new species in 16-million-year-old amberExternal link.EurekAlert!

2021 July. UC Davis Alumnus Receives Robert Snodgrass AwardExternal link. California Ag Today.

2021 July 2. The Art of the AntExternal link. University of California Agriculture and Natural Resources.

2020 December 22. Beast ant – a lost ant-wasp intermediateExternal link. Myrmecological News.

2020 August 7. Brendon Boudinot Receives ESA's Robert E. Snodgrass Memorial Research AwardExternal link. UC ANR Entomology & Nematology News.

2020 July 22. Exploring two new iron maiden ants of the CretaceousExternal link. Myrmecological News.

2020 July 14. 'Ant Man' Brendon Boudinot Off to Do Research in GermanyExternal link. University of California Agriculture and Natural Resources.

2019 November 12. New collection showcases cutting-edge techniques in insect morphology and systematicsExternal link. EurekAlert!

2015 January 23. Male ants “don’t get no respect” until nowExternal link. The Daily Democrat.

Research Talks


2022          “Origin and Evolution of Ants and the Aculeata” (lead author). International Conference on Insect Systematics and Evolutionary Biology. Indian Society of Evolutionary Biologists (ISEB) and Punjabi University Patiala. *** Expected 16–18 Feb.

2021          “Origin and Morphological Evolution of the Ants” (sole author). Rockefeller University.

2021          “Origin and Evolution of the Ants” (sole author). University of Copenhagen, Ecology & Evolution Section.

2020          Snodgrass Memorial Lecture (sole author). Entomol. Soc. America, National Meeting.

2020          “Morphological Insights into the Evolution of Ant Eusociality” (sole author). Entomol. Soc. America, National Meeting.

2020          “The Aculeate Ancestry and Evolution of the Ants (Hymenoptera)” (lead author). Entomol. Soc. America, Pacific Branch, Spokane, WA. *Cancelled due to COVID-19.

2020          “Insects, Species, and Extinction” (sole author). UC Davis Law School Symposium on Insect Decline, Davis, CA.

2019          “Altered States: Aculeate Anatomy and the Early Evolution of the Formicoidea” (lead author). XIV Simpósio de Mirmecologia, Belo Horizonte, Minas Gerais, Brazil.

2019          “A General Theory of Genital Homologies for the Hexapoda (Pancrustacea)” (sole author). 9th Dresden Meeting on Insect Phylogeny, Dresden, Germany.

2019          “Cambrian to the Quaternary: Insect Phylogeny and Evolutionary History” (sole author). Entomol. Soc. America, Pac. Branch, San Diego, CA. *John Henry Comstock Award Presentation.

2018          “Bayesian Phylogenetics: Morphological Modeling” (sole author). Entomol. Soc. America, Pac. Branch, Reno NV.

2017          “Stem or Crown: The Cretaceous Fauna and the Ant Tree of Life” (sole author). Barry Bolton Tribute Symposium. XXIII Simpósio de Mirmecologia, Curitiba, Paraná, Brazil.

2016          “Male Ants: Past, Present, and Prospects” (sole author). Advances in Ant Systematics Symposium. International Congress of Entomology, Orlando, FL.

2015          “Prospects for the Study of Male Ants” (sole author). XXII Simpósio de Mirmecologia, Ilhéus, Bahia, Brazil.

2015          “Updating Our Understanding of Ants” (sole author). California Department of Food and Agriculture Seminar Series, Sacramento, CA.

2015          “Relict Spider Ants: A Tale of Discovery and Intrigue” (first author). UC Berkeley Graduate Entomology Seminar Series, Berkeley, CA.


2021          “Head Anatomy and Evolution of the Sausagefly and the Driver Ant (Formicidae: Dorylinae)” (lead author). Entomol. Soc. America, Denver, CO. November 2021.

2020          “Abdomens and Ants: Evolutionary and Phylogenetic Morphology” (sole author). Ph.D. dissertation defense seminar, University of California, Davis, 4 Mar 2020.

2018          “Evolutionary and Phylogenetic Morphology” (sole author). Entomol. Soc. America, Vancouver, Canada.

2017          “The Protopodal Theory of Genitalic Evolution in the Hexapoda” (sole author). Entomological Society of America, Denver, CO. *Awarded ESA President’s Prize.

2016          “Hunting the Martian Ant, Martialis heureka” (sole author). International Union for the Study of Social Insects Meeting, Orlando, FL.

2016          “African Fossiliferous Amber: A Review” (second author). 7th International Conference on Fossil Insects, Arthropods, and Amber, Edinburgh, U.K.

2014          “Male Ants of the New World” (sole author). Entomological Society of America, Portland, OR. *Awarded ESA President’s Prize.

Conference Abstracts/Poster


Perrichot, V., Boudinot, B.E., Chény-Bergeronne, C., Cole, J., Jeanneau, L., Schmidt, A.R., Szwedo, J. & Wang, B. (2018). The age and paleobiota of Ethiopian amber revisited. 5th International Palaeontological Congress, Paris, France.


2015          “The Evolution of Leptomyrmex (Formicidae: Dolichoderinae): Insights from the Discovery of an Extant Neotropical Relict Species” (first author). Entomol. Soc. America, Minneapolis, MN.

2014          “Phylogenetic Placement of the Ant Lasius atopus, with Implications for the Evolution and Subgeneric Classification of the Genus” (second author). Entomol. Soc. America, Portland, OR.

2012          “Male Ant Genitalia: Musculature, Functional Morphology, and Homology” (sole author). Entomol. Soc. America, Knoxville, TN. *Awarded ESA President’s Prize.