Identification Keys and Illustrated Monographs

A collection of useful keys and illustrated monographs for testate amoeba identification and taxonomy.

Feel free to add what you have!

Please note that testate amoeba taxonomy is very far from being reliable. Ongoing work combining molecular and morphological approaches is revealing huge unsuspected diversity. As a result most species will need to be redescribed with high resolution light and scanning electron microscopy documentation, biometry data and molecular data (e.g. ideally SSU and COX). This is a huge task but a very rewarding one!

Nevertheless we provide some keys for common species as even with imperfect taxonomy testate amoeba analysis is a very powerful and useful tool in palaeoecology and other applications such as biomonitoring. The species listed in these keys should mostly be considered as morpho-species and even species complexes. Any suggestion for improving these keys is of course welcome!


A list of references is provided, please add any relevant information here!

An Atlas of Sphagnum-Dwelling Testate Amoebae in Bulgaria While few of us may have a chance to visit Bulgaria on a regular basis, this book may entice us to visit this beautiful country! As many Sphagnum peatland testate amoebae have broad geographical distributions (ignoring cryptic species and within taxa contrasted distribution ranges), this excellent monograph will certainly be a very useful reference for teaching and research. Congratulations and thanks a lot to Milcho and Nokola for this work! []. Milcho Todorov, Nikola Bankov (2019) An Atlas of Sphagnum-Dwelling Testate Amoebae in Bulgaria

Heads up everyone!!! Recently released, but quite likely not seen by most of the community, Andrey Tsyganov and colleagues published an extremely useful guide to testate amoebae with keys to genera Tsyganov, A.N., Babeshko, K.V. & Mazei, Y.A. (2016) A Guide to Testate Amoebae with the Keys to Genera - Monograph. Publishing house of Penza State University, Penza, Russia. This is a simplified and updated version of the larger monograph published by Yuri Mazei and Andrey Tsyganov in Russian in 2006 [Mazei, Y. & Tsyganov, A. (2006) Freshwater Testate Amoebae. KMK Sci Press, Moscow.] that has proven very useful to many of us - even if most of us could only read the latin names, look at images and understand the information on the size, this was already hugely helpful! Thanks a lot for this very nice work!

Key for Nebela and similar taxa
Key for Centropyxis and similar taxa
Key for Heleopera

Key for Trinema
Key for Euglypha
Key for Euglypha Decloitre 1962

General identification guides
Grospietsch 1958 monograph
Harnisch 1959 monograph
Mazei-Tsyganov 2006 Freshwater Testate Amoebae (in Russian)

Historical monographs
Leidy J. 1879 Freswhater Rhizpopds of North America
Penard E. 1902 La faune Rhizopodique du Bassin du Léman


Phylogeny & Taxonomy papers:

  • Bhattacharya, D., Helmchen, T. & Melkonian, M. (1995) Molecular evolutionary analyses of nuclear-encoded small subunit ribosomal RNA identify an independent rhizopod lineage containing the Euglyphina and the Chlorarachniophyta. Journal of Eukaryotic Microbiology, 42, 65-69.
  • Heger TJ, Mitchell EAD, Todorov M, Golemansky V, Lara E, Pawlowski J. 2010. Molecular phylogeny of euglyphid testate amoebae (Cercozoa: Euglyphida) suggest transitions between marine supralittoral and freshwater/terrestrial environments are infrequent. Molecular Phylogenetics and Evolution 55: 113-122. doi: 10.1016/j.ympev.2009.11.023
  • Heger TJ. Pawlowski J, Golemanski V, Todorov M, Lara E, Mitchell EAD. 2010. Comparing potential COI and SSU rDNA barcodes for assessing the diversity and phylogenetic relationships of cyphoderiid testate amoebae (Rhizaria: Euglyphida). Protist. doi: 10.1016/j.protis.2010.05.002
  • Lahr, D.J.G. & Lopes, S.G.B.C. (2006) Morphology, biometry, ecology and biogeography of five species of Difflugia Leclerc, 1815 (Arcellinida : Difflugiidae), from Tiete River, Brazil. Acta Protozoologica, 45, 77.
  • Lahr, D.J.G. & Lopez, S.G.B.C. (2007) Ultra-structure and Biometry of Three Lobose Testate Amoebae of the Family Lesquereusiidae (Tubulinea: Arcellinida) Based on Specimens from Sao Paulo, Brazil Acta Protozoologica, 46, 339–348.
  • Lahr, D.J.G., Bergmann, P.J. & Lopes, S. (2008) Taxonomic Identity in Microbial Eukaryotes: A Practical Approach Using the Testate Amoeba Centropyxis to Resolve Conflicts Between Old and New Taxonomic Descriptions. Journal of Eukaryotic Microbiology, 55, 409-416.
  • Lara E, Heger TJ, Mitchell EAD, Meisterfeld R, Ekelund F. 2007. SSU rRNA reveals a sequential increase in shell complexity among the Euglyphid testate amoebae (Rhizaria: Euglyphida). Protist 158: 229-237. doi:10.1016/j.protis.2006.11.006
  • Lara E, Heger TJ, Ekelund F, Lamentowicz M, Mitchell EAD. 2008. Ribosomal RNA Genes Challenge the Monophyly of the Hyalospheniidae (Amoebozoa: Arcellinida). Protist 159(2): 165-176. doi: 10.1016/j.protis.2007.09.003
  • Lara E, Heger TJ, Mitchell EAD. COI gene reveals size-dependent high dispersal and low intra-specific diversity in free-living terrestrial protists (Euglyphida; Assulina). Journal of Biogeography. doi:10.1111/j.1365-2699.2010.02426.x
  • Nikolaev SI, Mitchell EAD, Petrov NB, Berney C, Fahrni J, Pawlowski J, 2005. The testate lobose amoebae (order Arcellinida Kent, 1880) finally find their home within Amoebozoa. Protist, 156(2): 191-202.
  • Tekle, Y.I., Grant, J., Anderson, O.R., Nerad, T.A., Cole, J.C., Patterson, D.J. & Katz, L.A. (2008) Phylogenetic placement of diverse amoebae inferred from multigene analyses and assessment of clade stability within 'Amoebozoa' upon removal of varying rate classes of SSU-rDNA. Molecular Phylogenetics and Evolution, 47, 339-352.
  • Todorov M, Golemanski V, Mitchell EAD, Heger TJ. 2009. Morphology, Biometry, and Taxonomy of Freshwater and Marine Interstitial Cyphoderia (Cercozoa: Euglyphida). Journal of Eukaryotic Microbiology 56(3): 279-289. doi: 10.1111/j.1550-7408.2009.00394.x
  • Wylezich, C., Meisterfeld, R., Meisterfeld, S. & Schlegel, M. (2002) Phylogenetic analyses of small subunit ribosomal RNA coding regions reveal a monophyletic lineage of euglyphid testate amoebae (order Euglyphida). Journal of Eukaryotic Microbiology, 49, 108-118.

Papers discussing the problems of testate amoeba taxonomy (these illustrate some "hot" debates")

  • Finlay, B.J., Esteban, G.F. & Fenchel, T. (2004) Protist diversity is different? Protist, 155, 15-22.
  • Heger TJ, Mitchell EAD, Ledeganck P, Vincke S, Van De Vijver B, Beyens L. 2009. The curse of taxonomic uncertainty in biogeographical studies of free-living terrestrial protists: A case study of testate amoebae from Amsterdam Island Journal of Biogeography 36: 1551-1560. doi:10.1111/j.1365-2699.2009.02094.x
  • Lahr, D.J.G., Bergmann, P.J. & Lopes, S. (2008) Taxonomic Identity in Microbial Eukaryotes: A Practical Approach Using the Testate Amoeba Centropyxis to Resolve Conflicts Between Old and New Taxonomic Descriptions. Journal of Eukaryotic Microbiology, 55, 409-416.
  • Medioli, F.S., Scott, D.B. & Abbott, B.H. (1987) A case study of protozoan intraclonal variability: Taxonomic implications. Journal of Foraminiferal Research, 17, 28-47.
  • Mitchell EAD, Meisterfeld R. 2005. Taxonomic confusion blurs the debate on cosmopolitanism versus local endemism of free-living protists. Protist 156(3): 263-267.
  • Schlegel, M. & Meisterfeld, R. (2003) The species problem in protozoa revisited. European Journal of Protistology, 39, 349-355.
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