Enrique Lara

I am interested in ecology, diversity, biogeography and evolution of microbial eukaryotes in general, using from classical culture/morphological approaches to molecular phylogenetics /-omics and environmental DNA surveys in terrestrial, freshwater and marine systems. My pet group is the testate amoebae, a polyphyletic group of (very beautiful) amoeboid protists for which virtually nothing is known about their evolution, ecology and lifestyle. I am particularly interested in their micro-evolution to understand how their immense diversity is generated (how do protist speciate?). A related question is also which forces are driving the composition of their communities. I am basically using these questions to understand how far ecology and evolution follow the same rules in micro- and macro-organisms.




Blandenier, Q., Seppey, C. V.W., Singer, D., Vlimant, M., Simon, A., Duckert, C., Lara, E. (2016), Mycamoeba gemmipara nov. gen., nov. sp., the first cultured member of the environmental Dermamoebidae clade LKM74 and its unusual life cycle. Journal of Eukaryotic Microbiology doi:10.1111/jeu.12357

*Singer, D., *Lara, E., Steciow, M. M., Seppey, C. V. W., Paredes, N., Pillonel, A., Oszako, T. & Belbahri, L. 2016. High-throughput sequencing reveals diverse oomycete communities in oligotrophic peat bog micro-habitat. Fungal Ecology, 23:42-47.
* authors contributed equally to the work

Kosakyan, A., Lahr, D. J. G., Mulot, M., Meisterfeld, R., Mitchell, E. A. D., Lara, E. (2016), Phylogenetic reconstruction based on COI reshuffles the taxonomy of hyalosphenid shelled (testate) amoebae and reveals the convoluted evolution of shell plate shapes. Cladistics. doi:10.1111/cla.12167

Fernández, L. D., Fournier, B., Rivera, R., Lara, E., Mitchell, E. A. D. and Hernández, C. E. (2016), Water–energy balance, past ecological perturbations and evolutionary constraints shape the latitudinal diversity gradient of soil testate amoebae in south-western South America. Global Ecology and Biogeography. 25: 1216–1227

Qin, Y., Man, B., Kosakyan, A., Lara, E., Gu, Y., Wang, H., Mitchell, E. A. D. (2016), Nebela jiuhuensis nov. sp. (Amoebozoa; Arcellinida; Hyalospheniidae): A New Member of the Nebela saccifera - equicalceus - ansata Group Described from Sphagnum Peatlands in South-Central China. Journal of Eukaryotic Microbiology 63: 558–566.

Kosakyan, A., Lara, E., Lahr, D.J.G. (2016) Current and future perspectives on the systematics, taxonomy and nomenclature of testate amoebae. Accepted in European Journal of Protistology

Seppey, C.V.W., Fournier, B., Szelecz, I., Singer, D., Mitchell, E.A.D., Lara, E. (2015) Response of forest soil euglyphid testate amoebae (Rhizaria: Cercozoa) to pig cadavers assessed by high throughput sequencing. International Journal of Legal Medicine, 130: 551-562

Lara, E., Roussel-Delif, L., Fournier, B., Wilkinson, D.M., Mitchell, E.A.D (2016) Soil microorganisms behave like macroscopic organisms: patterns in the global distribution of soil euglyphid testate amoebae. In press, Journal of Biogeography, 43: 520-532


Jassey, V.E.J, Signarbieux, C., Hättenschwiler, S., Bragazza, L., Buttler, A., Delarue, F., Fournier, B., Gilbert, D., Laggoun-Défarge, F., Lara, E., Mills, R., Mitchell, E.A.D, Payne, R., Robroek, B. (2015) An unexpected role for mixotrophs in the response of peatland carbon cycling to climate warming. Scientific Reports 5:16931

Lahr, D.J.G., Bosak, T., Lara, E., Mitchell, E.A.D (2015) The Phanerozoic diversification of silica-cycling testate amoebae and its possible links to changes in terrestrial ecosystems. Peer Journal 5: e1234

Fernández, L. D., Lara, E., Mitchell, E.A.D (2015) Checklist, diversity and distribution of testate amoebae in Chile European Journal of Protistology, 51:409-425

Kosakyan, A., Mulot, M., Mitchell, E.A.D., Lara, E. (2015) Environmental DNA COI barcoding for quantitative analysis of protists communities at fine taxonomic level: a test using the Nebela collaris species complex (Amoebozoa; Arcellinida; Hyalospheniidae) European Journal of Protistology, 51: 311-320

Fournier, B., Lara, E., Jassey, V.E.J., Mitchell, E.A.D. (2015) Functional traits as a new approach for interpreting testate amoeba paleo-records in peatlands and assessing the causes and consequences of past changes in species composition. The Holocene, 25: 1375-1383

Tarnawski S. T., Lara, E. (2015) From environmental sequences to morphology: observation and characterisation of a paulinellid testate amoeba (Euglyphida, Paulinellidae) from soil using fluorescent in situ hybridization. Protist, 166: 264-270

Lara, E., Seppey, C.V.W., González Garraza, G., Singer, D., Quiroga, M.V., Mataloni, G. (2015) Molecular diversity of planktonic eukaryotes discriminate minerotrophic and ombrotrophic peatland pools in Tierra del Fuego (Argentina). Journal of Plankton Research 37:1-11.

De Vargas C, Audic, S., Henry, N., Decelle, J., Mahé, F., Logares, R., Lara, E., Berney, C., Le Bescot, N., Probert, I., Carmichael, M., Poulain, J., Romac, S., Colin, S., Aury, J.M., Bittner, L., Chaffron, S., Dunthorn, M., Engeler, S., Flegontova, O., Guidi, L., Horak, A., Jaillon, O., Mendez, G.L., Lukes, J., Malviya, S., Morard, R., Mulot, M., Scalco, E., Siano, R., Vincent, F., Zingone, A., Dimier, C., Picheral, M., Searson, S., Kandels-Lewis, S., Acinas, S., Bork, P., Bowler, C., Gaill, F., Gorsky, G., Grimsley, N., Hingamp, P., Iudicone, D., Not, F., Ogata, H., Pesant, S., Raes, J., Sieracki, M., Speich, S., Stemman, L., Sunagawa, S., Weissenbach, J., Wincker, P., Karsenti, E. (2015) Eukaryotic plankton diversity in the sunlit global ocean. Science, 348: 1261605.

Singer, D., Kosakyan, A., Pillonel, A., Mitchell E. A. D., Lara, E. (2015) Eight species in the Nebela collaris complex: Nebela gimlii (Arcellinida, Hyalospheniidae), a new species described from a Swiss raised bog. European Journal of Protistology 51: 79-85

Gomaa, F., Yang J., Mitchell, E. A. D., Zhang, W.-J., Zheng, Y., Todorov, M., Lara, E. (2015) Difflugia tuberspinifera and D. tuber n. sp. (Amoebozoa, Arcellinida): A case of fast morphological evolution in protists. Protist 166: 122-130


Steciow, M.M. *, Lara, E. *, Paul, C., Pillonel, A., Pelizza, S.A., Belbahri, L. (2014) Multiple barcode assessment within the Saprolegnia-Achlya clade (Saprolegniales, Oomycota, Stramenopiles) brings order in a neglected group of pathogens. IMA Fungus 5: 439-448
* authors contributed equally to the work

Gomaa, F., Kosakyan, A., Heger T. J., Corsaro, D., Mitchell E. A. D., Lara, E. (2014) One alga to rule them all: unrelated mixotrophic testate amoebae (Amoebozoa, Rhizaria and Stramenopiles) share the same symbiont (Trebouxiophyceae). Protist 165: 161-176


Steciow, M.M.*, Lara, E. *, Pillonel, A., Pelizza, S.A., Lestani, E.A., Rossi, G.C., Belbahri, L. (2013) Incipient loss of flagella in the genus Geolegnia: the emergence of a new clade within Leptolegnia? IMA Fungus 4:169-175
* authors contributed equally to the work

Chatelain, A., Meisterfeld, R., Roussel-Delif, L., Lara, E. (2013) Sphenoderiidae (fam. nov), a new clade of euglyphid testate amoebae characterized by small, round scales surrounding the aperture. Protist 164: 782-792

Qin, Y., Fournier, B., Lara, E., Gu, Y., Wang, H., Cui Y., Zhang, X., Mitchell E. A. D. (2013) Relationships between testate amoeba communities and water quality in Lake Donghu, a large alkaline lake in Wuhan, China. Frontiers of Earth Science, 7: 182-190

Qin, Y., Mitchell E. A. D., Lamentowicz, M., Payne, R. J., Lara, E., Gu, Y., Huang, X., Wang, H. (2013) Ecology of testate amoebae in peatlands of central China and development of a transfer function for paleohydrological reconstruction. Journal of Paleolimnology, 50: 319-330

Gomaa, F., Mitchell E. A. D., Lara, E. (2013) Amphitremida (Poche, 1913) is a new major, ubiquitous labyrinthulomycete clade. PLoS one,8: e53046


Guillou, L., Bachar D., Audic, S., Bass, D., Berney, C., Bittner, L., Boutte, C., Burgaud, G., de Vargas, C., Decelle J., del Campo, J., Dolan, J.R., Dunthorn M., Edvardsen B., Holzmann, M., Kooistra, W.H.C.F., Lara, E., Lebescot, N., Logares, R., Mahé, F., Massana, R., Montresor, M., Morard, R., Not F., Pawlowski, J., Probert, I., Sauvadet A.L., Siano, R., Stoeck, T., Vaulot, D., Zimmermann, P., Christen, R. (2012) The Protist Ribosomal Reference database (PR2): a catalog of unicellular eukaryote small subunit rRNA sequences with curated taxonomy. Nucleic Acids Research, 41: 597-604

Pawlowski, J., Audic, S., Adl, S.M., Bass, D., Belbahri, L., Berney, C., Bowser, S.S., Cepicka, I., Decelle, Dunthorn, M., Fiore-Donno, A.M., Gile, G.H., Holzmann, M., Jahn, R., Jirků, M., Keeling, P.J., Kotska, M., Kundryavstev, A., Lara, E., Lukeš, J., Mann, D.G., Mitchell, E.A.D., Nitsche, F., Romeralo, M., Saunders, G.W., Simpson, A.G.B., Smirnov, A.V., Spouge, J.L., Stern, R.F., 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.

Adl, S.M., Simpson, A.G.B., Lane, C. E., Lukeš, J., Bass, D., Bowser, S.S., Brown, M. W., Burki, F., Dunthorn, M., Hampl, V., Heiss, A., Hoppenrath, M., Lara, E., Le Gall, L., Lynn, D.H., McManus, H., Mitchell, E.A.D., Mozley-Stanridge, S.E., Parfrey, L. W., Pawlowski, J., Rueckert, S., Shadwick, L., Schoch, C.L., Smirnov, A.V., Spiegel, F.W. (2012). The Revised Classification of Eukaryotes. Journal of Eukaryotic Microbiology 59: 429-507

Kosakyan, A., Gomaa, F., Mitchell E. A. D., Heger T. J., Lara, E. (2012) Using DNA-barcoding for sorting out protist species complexes: A case study of the Nebela tincta-collaris-bohemica group (Amoebozoa; Arcellinida, Hyalospheniidae). European Journal of Protistology, 49: 222–237

Lahr, D.J.G., Lara, E., Mitchell E. A. D. (2012) Time to regulate microbial eukaryote nomenclature. Biological Journal of the Linnean Society 107: 469-476

Gomaa, F., Todorov, M., Heger T. J., Mitchell E. A. D., Lara, E. (2012) SSU rRNA phylogeny questions the monophyly of Arcellinida (Amoebozoa) and reveals that Difflugia, the largest Arcellinida genus, is not monophyletic. Protist 163: 389-399

Kosakyan, A., Heger, T. J., Leander, B.S., Todorov, M., Mitchell E. A. D., Lara, E. (2012) COI Barcoding of Nebelid Testate Amoebae (Amoebozoa: Arcellinida): Extensive Cryptic Diversity and Redefinition of the Hyalospheniidae Schultze. Protist 163: 415-434

Lara, E., Acosta-Mercado, D. (2012) A molecular perspective on ciliates as soil bioindicators. European Journal of Soil Biology 49: 107-111.

Lahr, D.J.G., Wegener Parfrey, L., Mitchell, E. A. D., Katz, L. A., Lara, E. (2011) The chastity of amoebae: re-evaluating evidence for sex in amoeboid organisms. Proceedings of the Royal Society B: Biological Sciences 278: 2081–2090


Lara, E., Belbahri, L. (2011). SSU rRNA reveals major trends in oomycete evolution. Fungal Diversity. DOI 10.1007/s13225-011-0098-9

Lara, E., Heger, T.J., Scheihing, R., Mitchell, E.A.D. (2011). COI gene and ecological data suggestsize-dependent high dispersal and lowintra-specific diversity in free-livingterrestrial protists (Euglyphida: Assulina). J .Biogeogr. 38: 640-650.

Deschamps, P., Lara, E., Marande, W., López García, P., Ekelund, F., Moreira, D.  (2011) Phylogenomic analysis of kinetoplastids supports that trypanosomatids arose from within bodonids. Mol. Biol. Evol. 28: 53-58

Lara, E., Mitchell, E.A.D., Moreira, D., López García, P. (2011) Highly diverse and seasonally dynamic protist community in a pristine peat bog. Protist 162:14-32

Lara, E., Heger, T.J., Scheihing, R., Mitchell, E.A.D. (2011) Cytochrome c oxidase subunit I gene and ecological data suggest size-dependent high dispersal and low intra-specific diversity in free-living terrestrial protists (Euglyphida; Assulina). J. Biogeography (in press, early view: DOI: 10.1111/j.1365-2699.2010.02426.x)

Heger, T.J., Pawlowski, J., Lara, E., Leander, B.S., Todorov, M., Golemansky, V., Mitchell, E.A.D. (2011) Comparing COI and SSU rDNA barcodes for assessing the diversity and phylogenetic relationships of cyphoderiid testate amoebae (Rhizaria: Euglyphida), Protist 162:131-141


Heger, T.J., Mitchell, E.A.D, Todorov, M., Golemansky, V., Lara, E., Leander, B.S., Pawlowski, J. (2010) Molecular phylogeny of euglyphid testate amoebae (Cercozoa: Euglyphida) suggests transitions between marine supralittoral and freshwater/terrestrial environments are infrequent. Mol. Phyl. Evol. 55: 113-122

Jousset, A., Lara, E., Nikolausz, M., Harms, H., Chatzinotas, A. (2010) Application of the denaturing gradient gel electrophoresis (DGGE) technique as an efficient diagnostic tool for ciliate communities in soil. Sci. Tot. Environm. 408: 1221-1225

Lara, E., Moreira, D., López García, P. (2010) The environmental clade LKM11 and Rozella form the deepest branching clade of Fungi. Protist 161: 116-121


Lara, E., Moreira, D., Vereshchaka, A., López García, P. (2009) Pan-oceanic distribution of new highly diverse clades of deep-sea diplonemids. Environm. Microbiol. 11: 47-55


Simpson, A.G.B., Perley, T. A., Lara, E. (2008) Lateral transfer of the gene for a widely used marker, alpha tubulin, indicated by multi-protein study of the phylogenetic position of Andalucia (Excavata). Mol. Phyl. Evol. 47: 366-377

Lara, E., Heger T. J., Lamentowicz, M., Ekelund, F. and Mitchell, E. A. D. (2008) Ribosomal RNA genes challenge the monophyly of Hyalospheniidae (Amoebozoa: Arcellinida). Protist 159: 165-176

Smith, H., Bobrov, A., Lara, E. (2008) Diversity and biogeography of testate amoebae. Biodiv. Conserv. 17: 329-343


Lara, E., Berney, C. Harms, H. and Chatzinotas, A. (2007) Cultivation -independent analysis reveals a shift in ciliate 18S rRNA gene diversity in a polycyclic aromatic hydrocarbon polluted soil. FEMS Microbiol. Ecol. 62: 365-373

Lara, E., Heger T. J., Mitchell, E. A. D. and Ekelund, F. (2007) SSU rRNA reveals a sequential increase in shell complexity among the Euglyphid testate amoebae Protist 158: 229-237

Lara, E., Berney, C., Ekelund, F., Harms H., and Chatzinotas A. (2007) Molecular comparison of cultivable Protozoa from a pristine and a Polycyclic Aromatic Hydrocarbons polluted site. Soil Biol. Biochem. 39: 139-148


Jousset, A., Lara, E., Wall, L. G., and Valverde, C. (2006) Secondary metabolites help biocontrol strain Pseudomonas fluorescens CHA0 to escape protozoan grazing. Appl. Environm. Microbiol. 72: 7083-7090

Lara, E., Chatzinotas, A. and Simpson, A.G.B. (2006) Andalucia (gen. nov,): a new taxon for the deepest branch within Jakobids (Jakobida; Excavata), based on morphological and molecular study of a new flagellate from soil. J. Euk. Microbiol., 52: 112-120


Heger, T.J., Lara, E., Mitchell, E.A.D. (2010). Arcellinida testate amoebae (Arcellinida: Amoebozoa): model of organisms for assessing microbial biogeography. In: Fontaneto D. (Ed.) The importance of being small: does size matter in biogeography? Cambridge University Press. (in press)



Lara, E., Heger, T.J., Mitchell, E.A.D. (2009) Pseudocryptic speciation and long range migration in the euglyphid testate amoeba genus Assulina. International Symposium on Testate Amoebae, Montbéliard (France). Oral Presentation.

Lara, E. Environmental diversity of eukaryotes: what clone libraries can tell us? 2009, Geneva (Switzerland). Oral Presentation.


Lara, E., Moreira, D., López García, P. Pan-oceanic distribution of new highly diverse clades of deep-sea diplonemids. 1st Spanish-French Congress on Protistology, 2008, Sevilla (Spain). Oral Presentation.


Lara, E., Heger, T. J., Ekelund, F., Lamentowicz, and Mitchell, E. A. D. Ribosomal RNA genes challenge the monophyly of Nebela and Hyalosphenia (Arcellinida: Hyalospheniidae). Meeting of the International Society for Evolutionary Protistology. 2006, Wroclaw (Poland). Poster.

Lara, E., Heger, T. J., Ekelund, F., Lamentowicz, M., Pawlowski, J., Mitchell, E. A. D. Evolution of testate amoebae: insights from molecular biology. International Symposium on Testate Amoebae, 2006, Antwerpen (Belgium). Oral Presentation.

Lara, E., Heger T. J., Ekelund, F., Mitchell, E. A. D. Testate amoebae under molecular light: Phylogeny of family Hyalosphaeniidae revisited. Meeting of the German Society of Protozoology, 2006, Liebenwalde bei Berlin, Germany. Oral presentation.

Lara, E., Heger, T. J., Meisterfeld, R:, Mitchell, E. A. D., Ekelund, F. A contribution to the understanding of evolution within Euglyphidae as shown by SSU-based molecular phylogeny). Meeting of the International Society for Evolutionary Protistology, 2006, Wroclaw, Poland. Poster.


Lara, E., Jousset, A., Harms, H. and Chatzinotas Analysis of ciliate diversity in soils using 18S rRNA gene clone libraries and denaturing gradient gel electrophoresis (DGGE). Meeting of the German Society of Protozoology, 2004, Innsbruck, Austria. Poster.


Lara, E., Harms, H. and Chatzinotas, A. Drastic differences of ciliate diversity between PAH polluted and pristine soils with 18S rDNA based molecular tools. Meeting of the German Society of Environmental Microbiology, 2003, Marburg, Germany. Poster.