The Gile Lab

Protist diversity, evolution, & symbiosis
Arizona State University - Tempe, AZ USA


What are Protists?

Microbial eukaryotes, as protists are sometimes called, can be found in all kinds of habitats on Earth. They can be primary producers (algae), grazers (eating bacteria), predators (eating other protists), or live on or within multicellular hosts as mutualists, commensals, or parasites. The very first eukaryotes were protists, and all of the multicellular lineages (e.g., animals, plants, fungi) have evolved from protist lineages. Many, or perhaps most, protists are associated with symbiotic bacteria that provide fixed nitrogen or fixed carbon or some other metabolic benefit. Symbiosis has played a pivotal role in eukaryotic evolution by providing us with mitochondria and plants with chloroplasts. The Gile lab studies protist diversity and evolution from both morphological and molecular and perspectives, including microbiomics, comparative genomics, and phylogenetics. We are currently working on protists from hot springs, marine plankton, and the hindguts of wood-feeding termites.


One of the most ancient and important symbioses of all is the establishment of plastids. Beginning over 1 billion years ago, when a host cell failed to digest its cyanobacterial food, an association between two single celled organisms gradually became a new, single organism. In the Gile lab we study the gene transfer and protein targeting that enabled the transition from symbiosis to new organism. We also study the loss of photosynthesis, which has happened many times in various lineages without loss of the plastid itself. Despite the importance of photosynthesis to life on earth, we do not fully understand how many times endosymbiosis has led to a new photosynthetic eukaryotic lineage, yet another topic of research in the Gile lab.


A major research focus in the Gile lab is the fascinating symbiosis occurs between termites and the protists that live in their hindguts. The termites rely on these protists to help them digest wood, and in return, the protists have a safe microcosm where they have flourished, evolving to become larger and morphologically more complex over about 150 million years of coevolution. Because most termite species have not yet had their hindgut communities investigated, the vast majority of symbiotic protist species are still unknown to science. We apply microscopy and molecular and computational techniques to study these protists, with particular interests in phylogeny, morphological convergence, genome evolution, and coevolution with their hosts.


Gillian Gile, PhD

  • Associate Professor i

Jonathon Hileman

  • PhD Candidate

Nicole Coots

  • PhD Candidate

Michael Breed

  • PhD Student

Kali Swichtenberg

  • PhD Student

Israa Jahan

  • Undergraduate Researcher

Daniel Jasso-Selles

  • Research Specialist

LeAnn Nguyen

  • 4+1 Master's Student

Noah Sharma

  • Undergraduate Researcher

Shaniya Deo

  • Undergraduate Researcher and LEAP Scholar

Serena Aguilar

  • Undergraduate Researcher

Lab Alumni

Francesca de Martini, PhD

  • Currently: Full Time Faculty in Biology at Mesa Community College

Gregory Gavelis, PhD

  • Postdoctoral Fellow i
    Currently: Bioinformatician, Bigelow Laboratory for Ocean Sciences

Stephen Taerum, PhD

  • Postdoctoral Fellow i
    Currently: Postdoctoral fellow, Triplett Lab, Connecticut Agricultural Experiment Station

Samantha Montoya

  • 4+1 Master’s Student

Viola Sanderlin

  • Master’s Student i

Bradley Bobbett

  • Undergraduate Researcher
    Currently: Clinical laboratory technician, ASU Biodesign Institute

Jordyn Shevat

  • Undergraduate Researcher

Xyonane Segovia

  • Undergraduate Researcher
    Currently: Biology PhD Program, Huijben Lab, Arizona State University

Tina Piarowski

  • Undergraduate Researcher
    Currently: Cell Biology, Stem Cells, and Development PhD program, University of Colorado at Denver

Keana Nguyen

  • Undergraduate Researcher
    Currently: Clinical research coordinator, ASU Biodesign Institute

Cherise Stotts

  • Undergraduate Researcher

Erin Avilucea

  • Undergraduate Researcher

Shaurya Aggarwal

  • Undergraduate Researcher
    Currently: Intern at Applied Microarrays

Trevor Merrell

  • Undergraduate Researcher
    Currently: Attending Burrell College of Osteopathic Medicine, Las Cruces NM

Katalina Freeman

  • Undergraduate Researcher
    Currently: Evacuation Platoon Leader in the Regimental Support Squadron of the 2nd Cavalry Regiment, Germany

Mikaela Garcia

  • Undergraduate Researcher
    Currently: Master’s Student, Master of Public Health Program, University of Arizona Phoenix

Evan Mee

  • PhD Student i
    Currently: Bioinformatics specialist at TGen

Bailey Bressee

  • Post-baccalaureate researcher
    Currently: PhD student at Idaho State University

Emma Hammond

  • Undergraduate researcher

Maya Gaylor

  • Undergraduate Researcher
    Currently: MS student at Manchester University


The Gile lab has openings for graduate students and postdocs to work on protist diversity and evolution in termite hindguts or other environments. If you are interested in studying protists or symbiosis, please contact Dr. Gile


Macher J-N, Coots NL, Poh Y-P, Girard E, Langerak A, Muñoz-Gómez S, Sinha S, Jirsová D, Vos R, Wissels R, Gile GH, Renema W, Wideman J. 2023. Single-cell genomics reveals the divergent mitochondrial genomes of Retaria (Foraminifera and Radiolaria). mBio, in press.

Noda S, Kitade O, Radek R, pTaerum SJ, uJasso-Selles DE, Takayanagi M, Radek R, Lo N, Ohkuma M, Gile GH. 2023. Molecular phylogeny of Spirotrichonymphea (Parabasalia) with emphasis on Spironympha, Spirotrichonympha, and three new genera Pseudospironympha, Nanospironympha, and Brugerollina. Journal of Eukaryotic Microbiology, in press.

Maciejowski W, Gile GH, Jerlström-Hultqvist J, Dacks JB. 2023. Ancient and pervasive expansion of adaptin-related vesicle coat machinery across Parabasalia. International Journal for Parasitology, in press.

Kassaye A, Getu E, Wakgari M, Goftishu M, Seid A, gMontoya SJ, Gile GH. 2021. Survey and identification of termites (Insecta, Isoptera) using morphological and molecular methods from eastern, central, and western Ethiopia. SINET: Ethiopian Journal of Science 44:193-204.

Gile GH, pTaerum SJ, uJasso-Selles DE, Sillam-Dussès D, Ohkuma M, Kitade O, Noda S. 2021. Molecular phylogenetic position of Microjoenia (Spirotrichonymphea: Parabasalia) from Reticulitermes and Hodotermopsis hosts. Protist 172:125836.

Gavelis GS, Gile GH, Leander BS. 2021. Convergent evolution of animal-like organelles across the tree of eukaryotes. In: Hall B, Moody S editors. Origin and Evolution of Metazoan Cell Types. CRC Press, doi:10.1201/b21831-3.

Finn DR, Maldonado J, De Martini F, Yu J, Penton CR, Fontenele RS, Schmidlin K, Kraberger S, Varsani A, Gile GH, Barker B, Kollath DR, Muenich RL, Herckes P, Fraser M, Garcia-Pichel F. 2021. Agricultural practices drive biological loads, seasonal patterns, and potential pathogens in the aerobiome of a mixed-use dryland. Science of the Total Environment 798:149239.

De Martini F, Coots N, Jasso-Selles DE, Garcia MD, Shevat J, Ravenscraft A, Stiblík P, Šobotník J, Sillam-Dussès D, Scheffrahn RH, Carrijo T, Gile GH. 2021. Biogeography and independent diversification in the protist symbiont community of Heterotermes tenuis. Frontiers in Ecology and Evolution 9:640625.

Velenovsky JF, Gile GH, Su N-Y, Chouvenc T. 2021. Dynamic protozoan abundance of Coptotermes kings and queens during the transition from biparental to alloparental care. Insectes Sociaux 68:33-40.

Taerum SJ, Jasso-Selles DE, Hileman JT, De Martini F, Mizumoto N, Gile GH. 2020. Spirotrichonymphea (Parabasalia) symbionts of the termite Paraneotermes simplicicornis. European Journal of Protistology 76:125742.

Jasso-Selles DE, De Martini F, Velenovsky JF, Mee ED, Hileman JT, Montoya SJ, Garcia MD, Su N-Y, Chouvenc T, Gile GH. 2020. The protist symbiont communities of Coptotermes formosanus and Coptotermes gestroi: Morphological and molecular characterization of five new species. Journal of Eukaryotic Microbiology 67:626-641.

Mizumoto N, Gile GH, Pratt SC. 2021. Behavioral rules for soil excavation by colony founders and workers in termites. Annals of the Entomological Society of America 114:654-661.

Mee ED, Gaylor MG, Jasso-Selles DE, Mizumoto N, Gile GH. 2020. Molecular phylogenetic position of Hoplonympha natator (Trichonymphea, Parabasalia): Horizontal symbiont transfer or differential loss? Journal of Eukaryotic Microbiology 67:268-272.

Crowther CV, Sanderlin V, Hayes MA, Gile GH. 2019. Effects of surface treatments on trapping with DC insulator-based dielectrophoresis. Analyst 144:7478-7488.

Piarowski CM, De Martini F, Gile GH. Distribution and relative abundance of three protist genera within the Zootermopsis nevadensis nuttingi hindgut. 2019. Symbiosis 79: 231-238.

Taerum SJ, Jasso-Selles DE, Wilson M, Ware JL, Sillam-Dussès D, Šobotník J, Gile GH. Molecular phylogenetic position of Holomastigotes (Spirotrichonymphea, Parabasalia) with descriptions of Holomastigotes flavipes n. sp. and Holomastigotes tibialis n. sp. 2019. Journal of Eukaryotic Microbiology 66:882-891.

Schmidlin K, Kraberger S, Fontenele RS, De Martini F, Chouvenc T, Gile GH, Varsani A. 2019. Genome sequences of microviruses associated with Coptotermes formosanus. Microbiology Resource Announcements 8:e00185-19.

Dorrell RG, Azuma T, Nomura M, Auren de Kerdrel G, Paoli L, Bowler C, Ishii K, Miyashita H, Gile GH, Kamikawa R. 2019. Principles of plastid reductive evolution illuminated by non-photosynthetic chrysophytes. PNAS 116:6914-6923.

Gavelis GS, Herranz M, Wakeman KC, Ripken C, Mitarai S, Gile GH, Keeling PJ, Leander BS. 2019. Dinoflagellate nucleus contains an extensive endomembrane network, the nuclear net. Scientific Reports 9:839.

Gavelis GS, Gile GH. 2018. How did cyanobacteria first embark on the path to becoming plastids?: Lessons from protist symbioses. FEMS Microbiology Letters 367:fny209.

Taerum SJ, De Martini F, Liebig JR, Gile GH. Incomplete co-cladogenesis between Zootermopsis termites and their associated protists. 2018. Environmental Entomology 47:184-195.

Čepička I, Dolan MF, Gile GH. 2017. Parabasalia. In: Handbook of Protists, Archibald JM, Simpson AGB, Slamovits CH eds. Springer, doi:10.1007/978-3-319-32669-6_9-1.

Jasso-Selles DE, De Martini F, Freeman KD, Garcia MD, Merrell TL, Scheffrahn RH, Gile GH. 2017. The parabasalid symbiont community of Heterotermes aureus: Molecular and morphological characterization and description of four new species and reestablishment of the genus Cononympha. European Journal of Protistology 61:48-63.

Gile GH, James ER, Tai V, Harper JT, Merrell TL, Boscaro V, Husník F, Scheffrahn RH, Keeling PJ. 2017. New species of Spirotrichonympha from Reticulitermes and the relationships among genera in Spirotrichonymphea (Parabasalia). Journal of Eukaryotic Microbiology, 65:159-169.

Dorrell RG, Gile GH, Méheust R, McCallum G, Klinger CM, Brillet-Guéguen L, Freeman KD, Richter DJ, Bapteste EP, Bowler C. 2017. Reconstruction of an ancestral plastid proteome reveals the origin and fate of complex plastid lineages. eLife 6:e23717.

Sierra R, Cañas-Duarte SJ, Burki F, Schwelm A, Fogelqvist J, Dixelius C, González-García LN, Gile GH, Slamovits CH, Klopp C, Restrepo S, Arzul I, and Pawlowski J. 2016. Evolutionary origins of rhizarian parasites. Molecular Biology and Evolution 33:980-983.

Gile GH, Moog D, Slamovits CH, Maier U-G, Archibald JM. 2015. Dual organellar targeting of aminoacyl-tRNA synthetases in diatoms and cryptophytes. Genome Biology and Evolution 7:1728-1742.

Gile GH, James ER, Okamoto N, Carpenter KJ, Scheffrahn RH, Keeling PJ. 2015. Molecular evidence for the polyphyly of Macrotrichomonas (Parabasalia: Cristamonadea) and a proposal for Macrotrichomonoides n. gen. Journal of Eukaryotic Microbiology 62:494-504.

Tai V, Gile GH, Pan J, James ER, Carpenter KJ, Scheffrahn RH, Keeling PJ. 2015. The phylogenetic position of Kofoidia loriculata (Parabasalia) and its implications for the evolution of the Cristamonadea. Journal of Eukaryotic Microbiology 62:255-259.

Gile GH and Slamovits CH. 2014. Transcriptomic analysis reveals evidence for a cryptic plastid in the colpodellid Voromonas pontica, a close relative of chromerids and apicomplexan parasites. PLoS ONE 9:e96258.

Gile GH†, Carpenter KJ†, James ER, Scheffrahn RH, and Keeling PJ. 2013. Morphology and molecular phylogeny of Staurojoenina mulleri sp. nov. (Trichonymphida, Parabasalia) from the hindgut of the kalotermitid Neotermes jouteli. Journal of Eukaryotic Microbiology 60:203-213. †Contributed equally.

Curtis BA, Tanifuji G, Burki F, Gruber A, Irimia M, Maruyama S, Arias MC, Ball SG, Gile GH, Hirakawa Y, Hopkins JF, Kuo A, Rensing SA, Schmutz J, Symeonidi A, Elias M, Eveleigh RJM, Herman EK, Klute MJ, Nakayama T, Oborník M, Reyes-Prieto A, Armbrust EV, Aves SJ, Beiko RG, Coutinho P, Dacks JB, Durnford DG, Fast NM, Green BR, Grisdale CJ, Hempel F, Henrissat B, Höppner MP, Ishida K-I, Kim E, Kořený L, Kroth PG, Liu Y, Malik S-B, Maier U-G, McRose D, Mock T, Neilson JAD, Onodera NT, Poole AM, Pritham EJ, Richards TA, Rocap G, Roy SW, Sarai C, Schaack S, Shirato S, Slamovits CH, Spencer DF, Suzuki S, Worden AZ, Zauner S, Barry K, Bell C, Bharti AK, Crow JA, Grimwood J, Kramer R, Lindquist E, Lucas S, Salamov S, McFadden GI, Lane CE, Keeling PJ, Gray MW, Grigoriev IV, Archibald JM. 2012. Algal genomes reveal evolutionary mosaicism and the fate of nucleomorphs. Nature 492:59-65.

Pawlowski J, Audic S, Adl S, Bass D, Belbahri L, Berney C, Bowser SS, Čepička I, Decelle J, Dunthorn M, Fiore-Donno A-M, Gile GH, Holzmann M, Jahn R, Jirků M, Keeling PJ, Kostka M, Kudryavtsev A, Lara E, Lukeš J, Mann DG, Mitchell EAD, Nitsche F, Romeralo M, Saunders GW, Simpson AGB, Smirnov AV, Spouge JL, Stern RF, Stoeck T, Zimmermann J, Schindel D, de Vargas C. 2012. CBOL Protist Working Group: Barcoding eukaryotic richness beyond the animal, plant and fungal kingdoms. PLoS Biology 10:e1001419.

Gile GH and Slamovits CH. 2012. Phylogenetic position of Lophomonas striata Bütschli (Parabasalia) from the hindgut of the cockroach Periplaneta americana. Protist 163:274-283.

Saldarriaga J†, Gile GH†, James ER, Horák A, Scheffrahn RH, and Keeling PJ. 2011. Morphology and molecular phylogeny of Pseudotrichonympha hertwigi and Pseudotrichonympha paulistana (Trichonymphea, Parabasalia) from neotropical rhinotermitids. Journal of Eukaryotic Microbiology 58:487-496. †Contributed equally.

Gile GH, James ER, Scheffrahn RH, Carpenter KJ, and Keeling PJ. 2011. Molecular and morphological analysis of the Calonymphidae with a description of Calonympha chia sp. nov., Snyderella kirbyi sp. nov., Snyderella swezyae sp. nov., and Snyderella yamini sp. nov. International Journal of Systematic and Evolutionary Microbiology 61:2547-2558.

Cocquyt E, Gile GH, Leliaert F, Verbruggen H, Keeling PJ, and De Clerck O. 2010. Complex phylogenetic distribution of a non-canonical code in green algae. BMC Evolutionary Biology 10:e327.

Hirakawa Y, Gile GH, Ota S, Keeling PJ, and Ishida K. 2010. Characterization of periplastid compartment targeting signals in chlorarachniophytes. Molecular Biology and Evolution 27:1538-1545.

Gile GH†, Stern RF†, James ER, and Keeling PJ. 2010. DNA barcoding of chlorarachniophytes using nucleomorph ITS sequences. Journal of Phycology 46:743-750. †Contributed equally.

Gile GH, Novis PM, Cragg DS, Zuccarello GC, and Keeling PJ. 2009. The distribution of EF-1α, EFL, and a non-canonical genetic code in the Ulvophyceae: Discrete genetic characters support a consistent phylogenetic framework. Journal of Eukaryotic Microbiology 56:367-372.

Harper JT, Gile GH, James ER, Carpenter KJ, and Keeling PJ. 2009. The inadequacy of morphological concepts for species and genus delineation in microbial eukaryotes: An example from the parabasalian termite symbiont, Coronympha. PLoS ONE 4:e6577.

Gile GH, Faktorová D, Castlejohn CA, Burger G, Lang BF, Farmer MA, Lukeš J, and Keeling PJ. 2009. Distribution and phylogeny of EF-1α and EFL in Euglenozoa suggest an ancient introduction of EFL followed by differential loss. PLoS ONE 4:e51162.

Gile GH and Keeling PJ. 2008. Nucleus-encoded periplastid-targeted EFL in chlorarachniophytes. Molecular Biology and Evolution 25:1967-1977.

Gile GH, Patron NJ, and Keeling PJ. 2006. EFL GTPase in cryptomonads and the distribution of EFL and EF-1α in chromalveolates. Protist 157:435-444.

Gould SB, Sommer MS, Kroth PG, Gile GH, Keeling PJ, and Maier UG. 2006. Nucleus-to-nucleus gene transfer and protein retargeting into a remnant cytoplasm of cryptophytes and diatoms. Molecular Biology and Evolution 23:2413-2422.




Phone: 480-727-4761
Room: LSE 607


Phone: 480-727-4295
Room: LSE 604