Eukaryota, Organisms with nucleated cellsPatrick Keeling, Brian S. Leander, and Alastair Simpson
This tree diagram shows the relationships between several groups of organisms.
The root of the current tree connects the organisms featured in this tree to their containing group and the rest of the Tree of Life. The basal branching point in the tree represents the ancestor of the other groups in the tree. This ancestor diversified over time into several descendent subgroups, which are represented as internal nodes and terminal taxa to the right.
You can click on the root to travel down the Tree of Life all the way to the root of all Life, and you can click on the names of descendent subgroups to travel up the Tree of Life all the way to individual species.close box
In the last decade, advances in a variety of fields ranging from molecular biology and genomics, to statistics and phylogenetic reconstruction methods, to environmental surveys and the description of new species, have in combination led to a significant improvement of our understanding of protist diversity and evolution. To recognize these advances, and to establish a flexible resource to concentrate and distribute our knowledge about the biology and diversity of microbial eukaryotes in the context of the phylogenetic relationships among protists, a massive update of the “protist” sections of the Tree of Life Web Project is currently underway.
In July 2008, on the eve of the combined meeting of the International Society for Evolutionary Protistology and the International Society of Protistologists, members of the protist research community gathered in Halifax to present an overview of the basic biology, structure, molecular biology, and phylogenetics of different protist groups and to report on their plans for developing/updating relevant branches of the Tree of Life Web Project (see Tree of Life Web Project Protist Diversity Workshop July 2008). The first products of this effort are the revised phylogeny above and several newly published protist pages, e.g.: Dinoflagellates, Fornicata, Arcellinida.
This page will soon present an updated discussion of eukaryote diversity, characteristics, and relationships. For the time being, please refer to the archived version of the previous version of the Tree of Life Eukaryotes page by David Patterson and Mitch Sogin.
Adl, S., Simpson, A. G. B., Farmer, M. A., Adersen, R. A., Anderson, O. R., Barta, J. R., Bowser, S. S., Brugerolle, G., Fensome, R. A., Fredericq, S., James, T. Y., Karpov, S., Kugrens, P., Krug, J., Lane, C. E., Lewis, L. A., Lodge, J., Lynn, D. H., Mann, D. G., McCourt, R. M., Mendoza, L., Moestrup, ?., Mozley-Standridge, S. E., Nerad, T. A., Shearer, C. A., Smirnov, A. V., Spiegel, F. W., and Taylor, M. F. J. R. 2005. The new higher level classification of eukaryotes with emphasis on the taxonomy of protists. J. Eukaryot. Microbiol. 52:399?451.
Arisue, N., M. Hasegawa, and T. Hashimoto. 2005. Root of the Eukaryota tree as inferred from combined maximum likelihood analyses of multiple molecular sequence data. Molecular Biology and Evolution 22(3):409-420.
Baldauf, S. L. 1999. A search for the origins of animals and fungi: Comparing and combining molecular data. American Naturalist 154(suppl.):S178-S188.
Baldauf, S. L. and W. F. Doolittle. 1997. Origin and evolution of the slime molds (Mycetozoa). Proceedings of the National Academy of Sciences (USA) 94:12007-12012.
Baldauf, S. L. and J. D. Palmer. 1993. Animals and fungi are each other's closest relatives: Congruent evidence from multiple proteins. Proceedings of the National Academy of Sciences (USA) 90:11558-11562.
Berney, C. and J. Pawlowski. 2006. A molecular time-scale for eukaryote evolution recalibrated with the continuous microfossil record. Proceedings of the Royal Society Series B 273(1596):1867-1872.
Borchiellini C., N. Boury-Esnault, J. Vacelet, and Y. Le Parco. 1998. Phylogenetic analysis of the Hsp70 sequences reveals the monophyly of metazoa and specific phylogenetic relationships between animals and fungi. Molecular Biology and Evolution 15:647-655.
Budin, K. and H. Philippe. 1998. New insights into the phylogeny of eukaryotes based on Ciliate Hsp70 sequences. Molecular Biology and Evolution 15:943-956.
Burki, F. and J. Pawlowski. 2006. Monophyly of Rhizaria and multigene phylogeny of unicellular bikonts. Molecular Biology and Evolution 23(10):1922-1930.
Canning, E. U. 1998. Evolutionary relationships of Microsporidia. Pages 77-90 in Evolutionary Relationships among Protozoa (G. H. Coombs, K. Vickerman, M .A. Sleigh, and A. Warren, eds.) Chapman & Hall, London.
Cavalier-Smith, T. 1993. Kingdom Protozoa and its 18 phyla. Microbiol. Rev. 57:953-94.
Cavalier-Smith, T. and Chao, E. E. 1995. The opalozoan Apusomonas is related to the common ancestor of animals, fungi and choanoflagellates. Proceedings of the Royal Society of London Series B 261:1-6.
Clark C. G. and A. J. Roger. 1995. Direct evidence for secondary loss of mitochondria in Entamoeba histolytica. Proceedings of the National Academy of Sciences (USA) 92:6518-6521.
Copeland, H. F. 1956. The Classification of Lower Organisms. Pacific Books, Palo Alto, California.
Delwiche, C. F. 1999. Tracing the thread of plastid diversity through the tapistry of life. American Naturalist 154 (suppl.):S164-S177.
Douzery, E. J. P., E. A. Snell, E. Bapteste, F. Delsuc, and H. Philippe. 2004. The timing of eukaryotic evolution: Does a relaxed molecular clock reconcile proteins and fossils? Proceedings of the National Academy of Sciences (USA) 101(43):15386-15391.
Edlind, T. D. 1998. Phylogenetics of protozoan tubulin with reference to the amitochondriate eukaryotes. Pages 91-108 in Evolutionary Relationships Among Protozoa (G. H. Coombs, K. Vickerman, M. A. Sleigh, and A. Warren, eds.) Chapman & Hall, London.
Edlind, T. D., J. Li, G. S. Visvesvara, M. H. Vodkin, G. L. McLaughlin, and S. K. Katiyar. 1996. Phylogenetic analysis of beta-tubulin sequences from amitochondrial protozoa. Molecular Phylogenetics and Evolution 5:359-367.
Embley T. M. and R. P. Hirt. 1998. Early branching eukaryotes? Curr. Opinion Gen. Dev. 8:624-629.
Fast, N. M., J. M. Logsdon, and W. F. Doolittle. 1999. Phylogenetic analysis of the TATA box binding protein (TBP) gene from Nosema locustae: evidence for a microsporidia-fungi relationship and spliceosomal intron loss. Molecular Biology and Evolution 16:1415-1419.
Forterre, P., N. Benachenhou-Lahfa, F. Confalonieri, M. Duguet, C. Elie and B. Labedan. 1992. The nature of the last universal ancestor and the root of the tree of life, still open questions. Biosystems 28:15-32.
Germot, A., H. Philippe, and H. Le Guyader. 1997. Evidence for loss of mitochondria in Microsporidia from a mitochondrial-type HSP70 in Nosema locustae. Molecular and Biochemical Parasitology 87:159-168.
Gogarten, J. P. 2003. Gene transfer: Gene swapping craze reaches eukaryotes. Curr Biol 13: R53?54.
Golding, G. B. and R. S. Gupta. 1995. Protein-based phylogenies support a chimeric origin for the eukaryotic genome. Molecular Biology and Evolution 12:1-6.
Grosberg, R. K. and ?R. R. Strathmann. 2007. The evolution of multicellularity: a minor major transition? Annual Review of Ecology, Evolution, and Systematics 38:621-654.
Hampl, V., D. S. Horner, P. Dyal, J. Kulda, J. Flegr, P. G. Foster, and T. M. Embley. 2005. Inference of the phylogenetic position of oxymonads based on nine genes: Support for Metamonada and Excavata. Molecular Biology and Evolution 22(12):2508-2518.
Hanson, E. D. 1977. The Origin and Early Evolution of Animals. Wesleyan University Press, Middletown, Conn.
Hashimoto, T., Y. Nakamura, T. Kamaishi, and M. Hasegawa. 1997. Early evolution of eukaryotes inferred from protein phylogenies of translation elongation factors 1 alpha and 2. Archiv f?r Protistenkunde 148:287-295.
Hirt, R. P., B. Healy, C. R. Vossbrinck, E. U. Canning, and T. M. Embley. 1997. A mitochondrial Hsp70 orthologue in Vairimorpha necatrix: Molecular evidence that microsporidia once contained mitochondria. Current Biology 7:995-998.
Hirt, R. P. and D. Horner (eds.) 2004. Organelles, Genomes and Eukaryote Evolution. Taylor & Francis, London.
Hirt, R. P., J. M. Logsdon, Jr., B. Healy, M. W. Dorey, W. F. Doolittle, and T. M. Embley. 1999. Microsporidia are related to fungi: evidence from the largest subunit of RNA polymerase II and other proteins. Proceedings of the National Academy of Sciences (USA) 96:580-585.
Huang, J., Y. Xu, and J. P. Gogarten. 2005. The presence of a haloarchaeal type tyrosyl-tRNA synthetase marks the opisthokonts as monophyletic. Molecular Biology and Evolution 22:2142-2146.
Jenner, R. A. and F. R. Schram. 1999. The grand game of metazoan phylogeny: rules and strategies. Biological Reviews 74:121-142.
Kamaishi, T., T. Hashimoto, Y. Nakamura, F. Nakamura, S. Murata, N. Okada, K. Okamoto, and M. Hasegawa. 1996. Protein phylogeny of translation elongation factor EF-1alpha suggests microsporidians are extremely ancient eukaryotes. Journal of Molecular Evolution 42:257-263.
Katz, L. A. 1998. Changing perspectives on the origin of eukaryotes. Trends Ecol. Evol. 13:493-497.
Katz, L. A. 1999. The tangled web: gene genealogies and the origin of eukaryotes. American Naturalist 154 (suppl.):S137-S145.
Keeling, P. J. 1998. A kingdom's progress: Archezoa and the origin of eukaryotes. BioEssays 20:87-95.
Keeling, P. J. 2004. Diversity and evolutionary history of plastids and their hosts. American Journal of Botany 91:1481-1493.
Keeling, P. J., G. Burger, D. G. Durnford, B. F. Lang, R. W. Lee, R. E. Pearlman, A. J. Roger and M. W. Gray. 2005. The tree of eukaryotes. Trends in Ecology & Evolution 20(12):670-676.
Keeling, P. J. and W. F. Doolittle. 1996. Alpha-tubulin from early-diverging eukaryotic lineages and the evolution of the tubulin family. Molecular Biology and Evolution 13:1297-1305.
Keeling, P. J., M. A. Luker, and J. D. Palmer. 2000. Evidence from beta-tubulin phylogeny that microsporidia evolved from within the fungi. Molecular Biology and Evolution 17:23-31.
Keeling, P. J. and G. I. McFadden. 1998. Origins of microsporidia. Trends Microbiol. 6:19-23.
Keeling, P. J. and J. D. Palmer. 2000. Phylogeny - Parabasalian flagellates are ancient eukaryotes. Nature 405:635-637.
Kim, J., W. Kim, and C. W. Cunningham. 1999. A new perspective on lower metazoan relationships from 18S rDNA sequences. Molecular Biology and Evolution 16:423-427.
Knoll, A. H. 1992. The early evolution of eukaryotes: a geological perspective. Science 256:622-627.
Kumar, S. and A. Rzhetsky. 1996. Evolutionary relationships of eukaryotic kingdoms. Journal of Molecular Evolution 42:183-193.
Lake, J. A. and M. C. Rivera. 1994. Was the nucleus the first endosymbiont? Proceedings of the National Academy of Sciences (USA) 91:2880-2881.
Lang, B. F., G. Burger, C. J. O'Kelly, R. Cedergren, G. B. Golding, C. Lemieux, D. Sankoff, M. Turmel, and M. W. Gray. 1997. An ancestral mitochondrial DNA resembling a eubacterial genome in miniature. Nature 387:493-497.
Lang, B. F., M. W. Gray, and G. Burger. 1999. Mitochondrial genome evolution and the origin of eukaryotes. Annual Review of Genetics 33:351-397.
Leipe, D., J. H. Gunderson, T. A. Nerad and M. L. Sogin. 1993. Small subunit ribosomal RNA of Hexamita inflata and the quest for the first branch in the eukaryotic tree. Mol. Biochem. Parasitol. 59:41-48.
Li, J., S. K. Katiyar, A. Hamelin, G. S. Visvesvara, and T. D. Edlind. 1996. Tubulin genes from AIDS-associated microsporidia and implications for phylogeny and benzimidazole sensitivity. Molecular and Biochemical Parasitology 78:289-295.
Lipscomb D. L., J. S. Farris, M. Kallersjo and A. Tehler. 1998. Support, ribosomal sequences and the phylogeny of the eukaryotes. Cladistics 14:303-338.
Maldonado, M. 2004. Choanoflagellates, choanocytes, and animal multicellularity. Invertebrate Biology 123:1-22.
Margulis, L. 1970. Origin of Eukaryotic Cells. Yale University Press.
Margulis, L., M. Chapman, R. Guerrero, and J. Hall. 2006. The last eukaryotic common ancestor (LECA): Acquisition of cytoskeletal motility from aerotolerant spirochetes in the Proterozoic Eon. Proceedings of the National Academy of Sciences (USA) 103(35):13080-13085.
Margulis, L., J. O. Corliss, M. Melkonian and D. J. Chapman. 1990. Handbook of Protoctista. Jones and Bartlett Publishers, Boston.
Mendoza, L., J. W. Taylor, and L. Ajello. 2002. The class mesomycetozoea: a heterogeneous group of microorganisms at the animal-fungal boundary. Annual Review of Microbiology 56:315-44.
Mereschkowsky, C. 1910. Theorie der zwei Plasmaarten als Grundlage der Symbiogenesis. Eine neue Lehre von der Entstehung der Organismen. Biologisches Centralblatt 30: 278-303, 321-347, 353-367.
Moreira, D., H. Le Guyader, and H. Philippe. 2000. The origin of red algae and the evolution of chloroplasts. Nature 405:69-72.
Moreira, D., S. von der Heyden, D. Bass, P. L?pez-Garc?a, E. Chao, and T. Cavalier-Smith. 2007. Global eukaryote phylogeny: Combined small- and large-subunit ribosomal DNA trees support monophyly of Rhizaria, Retaria and Excavata. Molecular Phylogenetics and Evolution 44(1):255-266.
Morin, L. 2000. Long branch attraction effects and the status of "basal eukaryotes": Phylogeny and structural analysis of the ribosomal RNA gene cluster of the free-living diplomonad Trepomonas agilis. Journal of Eukaryotic Microbiology 47:167-177.
Morris, P. J. 1993. The developmental role of the extracellular matrix suggests a monophyletic origin of the Kingdom Animalia. Evolution 47:152-165.
Narbonne, G. M. 2004. Modular construction of early Ediacaran complex life forms. Science 305(5687):1141-1144.
Nozaki, H., M. Matsuzaki, M. Takahara, O. Misumi, H. Kuroiwa, M. Hasegawa, T. Shin-i, Y. Kohara, N. Ogasawara, and T. Kuroiwa. 2003. The phylogenetic position of red algae revealed by multiple nuclear genes from mitochondria-containing eukaryotes and an alternative hypothesis on the origin of plastids. Journal of Molecular Evolution 56(4):485-497.
Nozaki, H., M. Iseki, M. Hasegawa, K. Misawa, T. Nakada, N. Sasaki, and M. Watanabe. 2007. Phylogeny of primary photosynthetic eukaryotes as deduced from slowly evolving nuclear genes. Molecular Biology and Evolution 24:1592-1595.
Patterson, D. J. 1994. Protozoa: Evolution and Systematics. Pages 1-14 in Progress in Protozoology. Proceedings of the IX International Congress of Protozoology, Berlin 1993. (K. Hausmann and N. H?lsmann, eds.) Gustav Fischer Verlag, Stuttgart, Jena, New York.
Patterson, D. J. 1999. The diversity of eukaryotes. American Naturalist 154 (suppl.):S96-S124.
Patterson, D. J. and M. L. Sogin. 1992. Eukaryote origins and protistan diversity. Pages 13-46 in The Origin and Evolution of Prokaryotic and Eukaryotic Cells (H. Hartman and K. Matsuno, eds.) World Scientific Pub. Co. NJ.
Philip, G. K., C. J. Creevey, and J. O. McInerney. 2005. The Opisthokonta and the Ecdysozoa may not be clades: Stronger support for the grouping of plant and animal than for animal and fungi and stronger support for the Coelomata than Ecdysozoa. Molecular Biology and Evolution 22(5):1175?1184.
Philippe, H. and A. Adoutte. 1998. The molecular phylogeny of Eukaryota: solid facts and uncertainties. Pages 25-56 in Evolutionary Relationships among Protozoa (G. H. Coombs, K. Vickerman, M. A. Sleigh, and A. Warren, eds.) Chapman & Hall, London.
Philippe, H. and A. Germot. 2000. Phylogeny of eukaryotes based on ribosomal RNA: Long-branch attraction and models of sequence evolution. Molecular Biology and Evolution 17:830-834.
Philippe, H., P. Lopez, H. Brinkmann, K. Budin, A. Germot, J. Laurent, D. Moreira, M. Muller, and H. Le Guyader. 2000. Early-branching or fast-evolving eukaryotes? An answer based on slowly evolving positions. Proceedings of the Royal Society of London Series B 267:1213-1221.
Philippe, H., E. A. Snell, E. Bapteste, P. Lopez, P. W. H. Holland, and D. Casane. 2004. Phylogenomics of eukaryotes: impact of missing data on large alignments. Molecular Biology and Evolution 21(9):1740-1752.
Ragan, M. A. and R. R. Gutell. 1995. Are red algae plants? Botanical Journal of the Linnean Society 118:81-105.
Ribeiro, S. and G. B. Golding. 1998. The mosaic nature of the eukaryotic nucleus. Molecular Biology and Evolution 15:779-788.
Richards, T. A. and M. van der Giezen. 2006. Evolution of the Isd11-IscS complex reveals a single alpha-proteobacterial endosymbiosis for all eukaryotes. Molecular Biology and Evolution 23:1341-1344.
Roger, A. J. 1999. Reconstructing early events in eukaryotic evolution. American Naturalist 154 (suppl.):S146-S163.
Roger, A. J., O. Sandblom, W. F. Doolittle, and H. Philippe. 1999. An evaluation of elongation factor 1 alpha as a phylogenetic marker for eukaryotes. Molecular Biology and Evolution 16:218-233.
Schlegel, M. 2003. Phylogeny of Eukaryotes recovered with molecular data: highlights and pitfalls. European Journal of Protistology 39:113-122.
Sch?tze J., A. Krasko, M. R. Custodio, S. M. Efremova, I. M. M?ller and W. E. G. M?ller. 1999. Evolutionary relationships of Metazoa within the eukaryotes based on molecular data from Porifera. Proceedings of the Royal Society of London Series B 266:63-73.
Siddall, M. E., D. S. Martin, D. Bridge, S. S. Desser, and D.K. Cone. 1995. The demise of a phylum of protists: phylogeny of Myxozoa and other parasitic Cnidaria. J. Parasitol. 81:961-967.
Simpson, A. G. B., Y. Inagaki, and A. J. Roger. 2006. Comprehensive multigene phylogenies of excavate protists reveal the evolutionary positions of "primitive" eukaryotes. Molecular Biology and Evolution 23(3):615-625.
Smothers, J. F., C. D. van Dohlen, L. H. Smith, and R. D. Spall. 1994. Molecular evidence that the myxozoan protists are metazoans. Science 265:1719-1721.
Sogin, M. L. 1991. Early evolution and the origin of eukaryotes. Current Opinion in Genetics and Development 1:457-463.
Sogin, M. L., H. G. Morrison, G. Hinkle and J. D. Silberman. 1996. Ancestral relationships of the major eukaryotic lineages. Microbiologia SEM 12:17-28.
Sogin, M. L. and J. D. Silberman. 1998. Evolution of the protists and protistan parasites from the perspective of molecular systematics. International Journal of Parasitology 28:11-20.
Stechmann, A. and T. Cavalier-Smith. 2002. Rooting the eukaryote tree by using a derived gene fusion. Science 297:89-91.
Steenkamp, E. T., J. Wright, and S. L. Baldauf. 2006. The protistan origins of animals and fungi. Molecular Biology and Evolution 23:93-106.
Stiller, J. W., E. C. S. Duffield and B. D. Hall. 1998. Amitochondriate amoebae and the evolution of DNA-dependent RNA polymerase II. Proceedings of the National Academy of Sciences (USA) 95:11769-11774.
Stiller, J. W. and B. D. Hall. 1997. The origin of red algae: Implications for plastid evolution. Proceedings of the National Academy of Sciences (USA) 94:4520-4525.
Stiller, J. W. and B. D. Hall. 1999. Long-branch attraction and the rDNA model of early eukaryotic evolution. Molecular Biology and Evolution 16:1270-1279.
Stiller, J. W, J. Riley, and B. D. Hall. 2001. Are red algae plants? A critical evaluation of three key molecular data sets. Journal of Molecular Evolution 52(6):527-539.
Taylor F. J. R. 1999. Ultrastructure as a control for protistan molecular phylogeny. American Naturalist 154(suppl.):S125-S136.
Van de Peer, Y., A. Ben Ali, and A. Meyer. 2000. Microsporidia: accumulating molecular evidence that a group of amitochondriate and suspectedly primitive eukaryotes are just curious fungi. Gene 246:1-8.
Van de Peer, Y. and R. de Wachter. 1997. Evolutionary relationships among the eukaryotic crown taxa taking into account site-to-site rate variation in 18S rRNA. Journal of Molecular Evolution 45:619-630.
Van de Peer, Y., G. Van der Auwera and R. DeWachter. 1996. The evolution of stramenopiles and alveolates as derived by 'substitution rate calibration' of small ribosomal subunit RNA. Journal of Molecular Evolution 42:201-210.
Vellai T. and G. Vida. 1999. The origin of eukaryotes: the difference between prokaryotic and eukaryotic cells. Proceedings of the Royal Society of London Series B 266:1571-1577.
Wainright, P. O., G. Hinkle, M. L. Sogin, and S. K. Stickel. 1993. Monophyletic origin of the Metazoa: an evolutionary link with fungi. Science 260:340-342.
Wainright, P. O., D. J. Patterson, and M. L. Sogin. 1994. Monophyletic origin of animals: a shared ancestry with fungi. Pages 39-53 in Molecular Evolution of Physiological Processes. Society of General Physiologists Series No. 49. (D. M. Farmborough, ed.) Rockefeller Press, New York.
Willmer, P. 1990. Invertebrate Relationships: Patterns in Animal Evolution. Cambridge University Press, Cambridge, UK.
Yang, D., Y. Oyaizu, H. Oyaizu, G. J. Olsen, and C. R. Woese. 1985. Mitochondrial origins. Proceedings of the National Academy of Sciences (USA) 82:4443-4447.
Yoon, H. S., J. Grant, Y. I. Tekle, M. Wu, B. C. Chaon, J. C. Cole, J. M. Logsdon, D. J. Patterson, D. Bhattacharya and L. A. Katz. 2008. Broadly sampled multigene trees of eukaryotes. BMC Evolutionary Biology 8:14. doi:10.1186/1471-2148-8-14
- Eu-Tree. Assembling the Tree of Eukaryotic Diversity.
- Protsville. Protist Research Laboratory, University of Sydney, Australia.
- Protist Information Server. Japan Science and Technology Corporation.
- Eukaryota: Systematics. Museum of Paleontology, University of California, Berkeley, USA.
- Malaria, Algae, Amoeba and You: Unravelling Eukaryotic Relationships. Joel B. Dacks. ActionBioScience.org
- Exploring Early Eukaryotic Evolution: Diversity and Relationships Among Novel Deep-Branching Lineages . Virginia Edgcomb, Andrew Roger, Alastair G.B. Simpson, Jeffrey Silberman and Mitchell Sogin, Marine Biological Laboratory, Woods Hole, USA.
- Microbial Life - Educational Resources. Teaching and learning about the diversity, ecology and evolution of the microbial world; discover the connections between microbial life, the history of the earth and our dependence on micro-organisms.
- Eukaryotes in extreme environments. Dave Roberts, the Natural History Museum, London, UK.
- The Homeobox Page. Thomas R. B?rglin's page about the homeobox genes which play important roles in the development of multicellular organisms.
- Protist Image Data. Molecular Evolution and Organelle Genomics program at the University of Montreal, Canada.
This page is being developed as part of the Tree of Life Web Project Protist Diversity Workshop, co-sponsored by the Canadian Institute for Advanced Research (CIFAR) program in Integrated Microbial Biodiversity and the Tula Foundation.
The University of British Columbia, Vancouver, British Columbia, Canada
Brian S. Leander
The University of British Columbia, Vancouver, British Columbia, Canada
Dalhousie University, Halifax, Nova Scotia, Canada
Correspondence regarding this page should be directed to Patrick Keeling at , Brian S. Leander at , and Alastair Simpson at
Page: Tree of Life Eukaryotes. Eukaryota, Organisms with nucleated cells. Authored by Patrick Keeling, Brian S. Leander, and Alastair Simpson. The TEXT of this page is licensed under the Creative Commons Attribution-NonCommercial License - Version 3.0. Note that images and other media featured on this page are each governed by their own license, and they may or may not be available for reuse. Click on an image or a media link to access the media data window, which provides the relevant licensing information. For the general terms and conditions of ToL material reuse and redistribution, please see the Tree of Life Copyright Policies.
- First online 08 September 2000
- Content changed 02 September 2008
Citing this page:
Keeling, Patrick, Leander, Brian S., and Simpson, Alastair. 2008. Eukaryotes. Eukaryota, Organisms with nucleated cells. Version 02 September 2008 (under construction). http://tolweb.org/Eukaryotes/3/2008.09.02 in The Tree of Life Web Project, http://tolweb.org/