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Stentor stipatus is a new unicellular species that demonstrates habituation and unique phototaxis
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  • Published: 20 February 2026

Stentor stipatus is a new unicellular species that demonstrates habituation and unique phototaxis

  • Deepa H. Rajan1,
  • Benjamin Lee2,
  • Ashley Albright1,
  • Eric Tang3,
  • Kristen Ressler4,
  • Arnie Maravillas2,
  • Carmen Vargas2,
  • Wallace F. Marshall1 &
  • …
  • Daniel B. Cortes4,5 

Scientific Reports , Article number:  (2026) Cite this article

We are providing an unedited version of this manuscript to give early access to its findings. Before final publication, the manuscript will undergo further editing. Please note there may be errors present which affect the content, and all legal disclaimers apply.

Subjects

  • Computational biology and bioinformatics
  • Ecology
  • Evolution
  • Microbiology

Abstract

Stentor, the genus of large trumpet-shaped ciliates, is well-known for its complex morphology and striking behaviors. Members of this genus are distributed throughout the world in a wide and diverse pool of freshwater ecosystems. Recently, the molecular phylogeny of Stentor has been explored through comparison of 18 S small subunit (SSU) ribosomal DNA (rDNA) sequences, clarifying several previously mischaracterized species and species complexes. However, despite their wide distribution, to-date, only about a dozen species of Stentor have been described and verified by phylogenetic means. Here, we introduce the discovery of a new species within genus Stentor: Stentor stipatus spec. nov., so named for their distinctive cytosolic dark pigmented granules which surround the macronucleus and are also present cortically alongside cortically-distributed green microalgae. We present morphological, phylogenetic, ecological, and behavioral characterizations of these cells. Phylogenetic analysis of S. stipatus spec. nov. by comparison of SSU rDNA sequence suggests it is a distinct species from its closest relative, S. amethystinus. We demonstrate that S. stipatus spec. nov. is capable of habituation in response to repeated mechanical stimulation. Further, S. stipatus spec. nov. exhibits strongly directed positive phototaxis, like its relative S. pyriformis, but with a distinct action spectrum from both S. coeruleus and S. pyriformis. Finally, S. stipatus phototaxis response strength varies in a consistent pattern throughout the day, providing evidence of potential circadian regulation. This work expands the current understanding of the ecological distribution of and behavioral features present within genus Stentor.

Data availability

Image and video files analyzed for phototaxis and habituation experiments are available upon request. 18 S SSU DNA sequences are included in the published work and are available on GenBank with the following accession numbers: Sequence 1 ( *S. coeruleus* ; wild strain) PX056129, Sequence 2 ( *S. muelleri* ; wild strain) PX056130, Sequence 3 ( *S. pyriformis* ; wild strain) PX056131, Sequence 4 ( *S. stipatus* ; spec. nov., wild strain) PX056132, Sequence 5 ( *S. stipatus* ; spec. nov., wild strain) PX056133, Sequence 6 ( *S. stipatus* ; spec. nov., wild strain) PX056134, Sequence 7 ( *S. stipatus* spec. nov. wild strain) PX393983. *S. stipatus* spec. nov. as described in this work, has been registered on ZooBank, LSID: urn: lsid: zoobank.org: act: D381C528-8486-424E-9C69-CBE1612C4295; with this work also registered under LSID: urn: lsid: zoobank.org: pub:0C20B702-EE37-47C9-8DDE-417C2B39CBC4.

References

  1. Lee, R. E. & Kugrens, P. Relationship between the flagellates and the ciliates. Microbiol. Rev. 56, 529–542 (1992).

    Google Scholar 

  2. Lischke, B. et al. Large biomass of small feeders: ciliates May dominate herbivory in eutrophic lakes. J. Plankton Res. 38, 2–15 (2016).

  3. Porter, K. G., Pace, M. L. & Battey, J. F. Ciliate protozoans as links in freshwater planktonic food chains. Nature. 277, 563–565 (1979).

  4. Liu, H. et al. Composition and distribution of planktonic ciliates in the Southern South China sea during late summer: comparison between surface and 75 m deep layer. J. Ocean Univ. China. 15, 171–176 (2016).

  5. Tartar, V. The Biology of Stentor. (1961).

  6. Slabodnick, M. M. et al. The kinase regulator mob1 acts as a patterning protein for stentor morphogenesis. PLoS Biol. 12, e1001861 (2014).

    Google Scholar 

  7. Homer Iliad. Book 1.

  8. Foissner, W. & Wölfl, S. Revision of the genus Stentor Oken (Protozoa, Ciliophora) and description of S. araucanus nov. spec, from South American lakes. J. Plankton Res. 16, 255–289 (1994).

    Google Scholar 

  9. Fernandes, N. M., Paiva, T. S., Silva-Neto, I. D., da, Schlegel, M. & Schrago, C. G. Expanded phylogenetic analyses of the class heterotrichea (Ciliophora, Postciliodesmatophora) using five molecular markers and morphological data. Mol. Phylogenet. Evol. 95, 229–246 (2016).

  10. Thamm, M., Schmidt, S. L. & Bernhard, D. Insights into the phylogeny of the genus Stentor (Heterotrichea, Ciliophora) with Special emphasis on the evolution of the macronucleus based on SSU rDNA data. 2010, 149–157 (2010).

  11. Foissner, W. & Berger, H. A user-friendly guide to the ciliates (Protozoa, Ciliophora) commonly used by hydrobiologists as bioindicators in rivers, lakes, and waste waters, with notes on their ecology. Freshw. Biol. 35, 375–482 (1996).

    Google Scholar 

  12. Hoshina, R., Tsukii, Y., Harumoto, T. & Suzaki, T. Characterization of a green Stentor with symbiotic algae growing in an extremely oligotrophic environment and storing large amounts of starch granules in its cytoplasm. Sci. Rep. 11, 2865 (2021).

    Google Scholar 

  13. Boudreau, V. et al. The cell biology and genome of Stentor pyriformis, a giant cell that embeds symbiotic algae in a microtubule meshwork. Mol. Biol. Cell. 36, ar44 (2025).

    Google Scholar 

  14. Kim, I. H., Prusti, R. K., Song, P. S., Häder, D. P. & Häder, M. Phototaxis and photophobic responses in Stentor coeruleus action spectrum and role of Ca2+ fluxes. Biochim. Biophys. Acta (BBA) - Gen. Subj. 799, 298–304 (1984).

    Google Scholar 

  15. Wood, D. C. Action spectrum and electrophysiological responses correlated with the photophobic response of Stentor coeruleus. Photochem. Photobiol. 24, 261–266 (1976).

    Google Scholar 

  16. Dexter, J. P., Prabakaran, S. & Gunawardena, J. A. Complex hierarchy of avoidance behaviors in a single-cell eukaryote. Curr. Biol. 29, 4323–4329e2 (2019).

    Google Scholar 

  17. Rajan, D. et al. Single-cell analysis of habituation in Stentor coeruleus. Curr. Biol. 33, 241–251e4 (2023).

    Google Scholar 

  18. Morgan, T. H. Regeneration of proportionate structures in Stentor. Biol. Bull. 2, 311 (1901).

    Google Scholar 

  19. Marshall, W. F. Regeneration in Stentor coeruleus. Front. Cell. Dev. Biol. 9, 753625 (2021).

    Google Scholar 

  20. Taher, M. A., Kabir, A. S., Shazib, S. U. A., Kim, M. S. & Shin, M. K. Morphological redescriptions and molecular phylogeny of three Stentor species (Ciliophora: Heterotrichea: Stentoridae) from Korea. Zootaxa 4732, zootaxa473236 (2020).

    Google Scholar 

  21. Lin, A., Makushok, T., Diaz, U. & Marshall, W. F. Methods for the study of regeneration in Stentor. J. Vis. Exp. https://doi.org/10.3791/57759 (2018).

    Google Scholar 

  22. Guindon, S. et al. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst. Biol. 59, 307–321 (2010).

  23. Anisimova, M., Gil, M., Dufayard, J. F., Dessimoz, C. & Gascuel, O. Survey of branch support methods demonstrates accuracy, power, and robustness of fast Likelihood-based approximation schemes. Syst. Biol. 60, 685–699 (2011).

  24. Rajan, D., Chudinov, P. & Marshall, W. Studying habituation in Stentor coeruleus. J. Vis. Exp. https://doi.org/10.3791/64692 (2023).

    Google Scholar 

  25. Rajan, D. H. & Marshall, W. F. A receptor-inactivation model for single-celled habituation in Stentor coeruleus. Curr. Biol. 35, 3327–3340e4 (2025).

    Google Scholar 

  26. Motzkin, G. Atlantic White Cedar Wetlands of Massachusetts (Massachusetts Agricultural Experiment Station, College of Food and Natural Resources, University of Massachusetts, 1991).

  27. Höfle, G., Reinecke, S., Laude, U. & Spitzner, D. Amethystin, the coloring principle of Stentor amethystinus. J. Nat. Prod. 77, 1383–1389 (2014).

    Google Scholar 

  28. Wood, D. Habituation in stentor: a response-dependent process. J. Neurosci. 8, 2248–2253 (1988).

    Google Scholar 

  29. Chen, H., Ma, Y., Zhang, W. F., Ma, T. & Wu, H. X. Molecular phylogeny of Colletotrichum (Sordariomycetes: Glomerellaceae) inferred from multiple gene sequences. Genet. Mol. Res. 14, 13649–13662 (2015).

    Google Scholar 

  30. Adl, S. M. et al. Diversity, Nomenclature, and taxonomy of protists. Syst. Biol. 56, 684–689 (2007).

  31. Doerder, F. P. Barcodes reveal 48 new species of tetrahymena, dexiostoma, and glaucoma: Phylogeny, ecology, and biogeography of new and established species. J. Eukaryot. Microbiol. 66, 182–208 (2019).

    Google Scholar 

  32. Heep, T., Rohozinski, J., Simpson, A. G. B. & Patterson, D. J. Stentor amethystinus (Protista: Ciliophora: Heterotrichida), a common protozoan member of fresh-water plankton in Australia. Records Australian Museum. 50, 211–216 (1998).

    Google Scholar 

  33. Cragg, P. R. An explanation for the limited contractility of Stentor amethystinus Leidy, 1880. J. Protozoology. 18, 672–676 (1971).

    Google Scholar 

  34. Yin, X. X., Zhang, Y. Y., Yang, J. & Zhu, Y. G. Rapid biotransformation of arsenic by a model protozoan tetrahymena thermophila. Environ Pollut . 159, 837–840 (2011).

  35. Xiong, W. et al. Construction of tetrahymena strains with highly active arsenic methyltransferase genes for arsenic detoxification in aquatic environments. Ecotoxicol. Environ. Saf. 275, 116258 (2024).

    Google Scholar 

  36. Alharbi, R. M., Sholkamy, E. N., Alsamhary, K. I., Abdel-Raouf, N. & Ibraheem, I. B. M. Optimization study of the capacity of Chlorella vulgaris as a potential bio-remediator for the bio-adsorption of arsenic (III) from aquatic environments. Toxics. 11, (2023).

  37. Wille, J. J. J. & Ehret, C. F. Circadian rhythm of pattern formation in populations of a free-swimming organism, Tetrahymena. J. Protozool. 15, 789–792 (1968).

    Google Scholar 

  38. Wood, D. C. Stimulus specific habituation in a protozoan. Physiol. Behav. 11, 349–354 (1973).

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Acknowledgements

Initial discovery and collection of this novel Stentor species was made possible through funding by the Grass Foundation in 2023. Initial culturing and most of the phototaxis experiments were performed at the Marine Biological Laboratory (MBL) in the summers of 2023 and 2024. We also thank Mark Slabodnick for guidance on phylogenetic analysis.

Funding

Initial discovery of S. stipatus spec. nov. was supported by Grass Fellowship funding to DC from the Grass Foundation. Molecular analysis was performed in the CCC Summer Course supported by NSF grant DBI-1548297 and NIH 5K12GM081266. Experiments on S. stipatus spec. nov. habituation were supported by NIH grant R35GM130327 and work on the phylogeny of wild Stentor species was supported by a grant from the Moore Foundation.

Author information

Authors and Affiliations

  1. UCSF Genentech Hall N376, 600 16th Street, San Francisco, CA, 94158, USA

    Deepa H. Rajan, Ashley Albright & Wallace F. Marshall

  2. Hensill Hall 534, 1600 Holloway Ave, San Francisco, CA, 94132, USA

    Benjamin Lee, Arnie Maravillas & Carmen Vargas

  3. UCSF Cardiovascular Research Institute, 555 Mission Bay Boulevard South, San Francisco, CA, 94158, USA

    Eric Tang

  4. Steger 162B, 1015 Life Science Circle, Blacksburg, VA, 24061, USA

    Kristen Ressler & Daniel B. Cortes

  5. Marine Biological Laboratory, 7 MBL Street, Woods Hole, MA, 02543, USA

    Daniel B. Cortes

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Contributions

DHR performed the habituation experiments as well as the phylogenetic analysis and drafted the manuscript. AA, ET, AM, CV and BL all contributed to single cell 18 S SSU sequencing. BL also contributed to phylogenetic tree assembly. WFM supervised DHR and provided support and guidance for experimental design as well as feedback and edits on the manuscript. KR performed phototaxis and action spectrum experiments. DBC performed phototaxis and action spectrum experiments, did the imaging for morphological analysis, supervised KR, and drafted and edited the manuscript. All authors read and approved of the final manuscript.

Corresponding author

Correspondence to Daniel B. Cortes.

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Supplementary Information

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Supplementary Material 11

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Cite this article

Rajan, D.H., Lee, B., Albright, A. et al. Stentor stipatus is a new unicellular species that demonstrates habituation and unique phototaxis. Sci Rep (2026). https://doi.org/10.1038/s41598-026-40277-0

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  • Received: 14 July 2025

  • Accepted: 11 February 2026

  • Published: 20 February 2026

  • DOI: https://doi.org/10.1038/s41598-026-40277-0

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Keywords

  • Stentor
  • Heterotrich
  • Morphology
  • Phylogeny
  • SSU rDNA
  • Protist
  • Cell behavior
  • Cell learning
Supplementary Material 1Supplementary Material 2Supplementary Material 3Supplementary Material 4Supplementary Material 5Supplementary Material 6Supplementary Material 11
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