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Animals

Metazoa


     =========================================== Porifera  (sponges) 
     |
     |        ================================== Cnidaria  (jellyfish, anemones, corals, etc.) 
     |        |
     |        |  =============================== Ctenophora  (comb-jellies) 
     |        |  |
     |        |  |                        ====== Arthropoda  (insects, spiders, crabs, etc.) 
     |        |  |                        |
     |        |  |                     ===|===== Onychophora  (velvet worms) 
     |        |  |                     |  |
     |  ======|  |               ======|  ====== Tardigrada  (water bears) 
     |  |     |  |               |     |
     |  |     |  |               |     |  ====== Annelida  (segmented worms) 
     |  |     |  |               |     ===|
<<===|  |     |  |               |        |  === Pogonophora 
     |  |     |  |               |        ===|
     |  |     |  |            ===|           === Vestimentifera 
     |  |     |  |            |  |
     |  |     ===|            |  |============== Echiura 
     |  |        |            |  |
     |  |        |         ===|  |============== Mollusca  (snails, clams, squids, etc.) 
     |  |        |         |  |  |
     |  |        |         |  |  =============== Sipuncula 
     |  |        |     ==P=|  |
     |  |        |     |   |  ================== Nemertea  (ribbon worms) 
     |  |        |     |   |
     |  |        |     |   ===================== Platyhelminthes  (flatworms) 
     |  |        |     |
     ===|        |  ===|            ============ Chordata  (vertebrates and relatives) 
        |        |  |  |         ===|
        |        |  |  |      ===|  ============ Hemichordata 
        |        |  |  |      |  |
        |        |  |  |   ===|  =============== lophophorates 
        |        ===|  |   |  |
        |           |  ==D=|  ================== Chaetognatha 
        |           |      |
        |           |      ===================== Echinodermata  (starfish, urchins, sea cucumbers, etc.) 
        |           |
        |           ============================ pseudocoelomates 
        |
        |======================================= Placozoa 
        |
        |======================================= Monoblastozoa 
        |
        |======================================= Rhomobozoa 
        |
        ======================================== Orthonectida 
Tree modified from Brusca and Brusca (1990).
Note that this is a fairly traditional view of animal phylogeny, and other recent papers have substantially different arrangements. We will soon present an up to date discussion of metazoan relationships on this page. For the time being, please refer to the papers listed in the References section

In the tree, the descendents of the branch labelled "P" are the protostomes, of that labelled "D" are the deuterostomes.

Containing clade(s): Eukaryotes


Table of Contents

References

References

Adoutte, A., G. Balavoine, N. Lartillot, O. Lespinet, B. Prud'homme, 
          and R. de Rosa.  2000.  The new animal phylogeny: Reliability 
          and implications.  Proceedings of the National Academy 
          of Sciences (U.S.A.) 97:4453-4456.  

Adoutte, A., G. Balavoine, N. Lartillot, and R. de Rosa.  1999. 
          Animal evolution, the end of the intermediate taxa? 
          Trends in Genetics 15:104-108.

Aguinaldo, A. M. A., J. M. Turbeville, L. S. Linford, M. C. Rivera, 
          J. R. Garey, R. A. Raff, and J. A. Lake.  1997.  Evidence 
          for a clade of nematodes, arthropods and other moulting 
          animals.  Nature 387:489-493.

Aleshin, V. V., I. A. Milyutina, O. S. Kedrova, N. S. Vladychenskaya, 
          and N. B. Petrov.  1998.  Phylogeny of Nematoda and 
          Cephalorhyncha derived from 18S rDNA.  J. Mol. Evol. 
          47:597-605.

Anderson, C. L.  1998.  Phylogenetic relationships of the Myxozoa. 
          Pages 341-350 in Evolutionary Relationships among Protozoa 
          (G.H. Coombs, K. Vickerman, M.A. Sleigh, and A. Warren, 
          eds.)  Chapman & Hall, London.

Anderson, C. L., E. U. Canning, and B. Okamura.  1998.  A triploblast 
          origin for Myxozoa?  Nature 392:346-347.

Balavoine, G. and A. Adoutte.  1998.  One or three cambrian radiations? 
          Science 280:397-398

Balavoine, G.  1997. The early emergence of platyhelminths is 
          contradicted by the agreement between 18S rRNA and 
          Hox genes data. C.R. Acad. Sci. Paris, Sciences de 
          la vie/Life Sciences 320:83-94.

Balavoine, G.  1998.  Are Platyhelminthes coelomates without 
          a coelom?  An argument based on the evolution of Hox 
          genes.  American Zoologist 38:843-858.

Boore, J. L. and W. M. Brown.  2000.  Mitochondrial genomes of 
          Galathealinum, Helobdella, and Platynereis: Sequence 
          and gene arrangement comparisons indicate that Pogonophora 
          is not a phylum and Annelida and Arthropoda are not 
          sister taxa.  Molecular Biology and Evolution 17:87-106.

Briggs, D. E. G., D. H. Erwin, and F. J. Collier.  1994. The 
          Fossils of the Burgess Shale. Smithsonian Institution 
          Press, Washingthon, D.C.

Brooke, M. D.  1999.  How old are animals?  Trends in Ecology 
          and Evolution 14: 211-212.

Brusca, R. C., and G. J. Brusca.  1990. Invertebrates. Sinauer 
          Associates, Sunderland, Massachusetts.

Carranza, S., J. Baguña, and M. Riutort.  1997.  Are the platyhelminthes 
          a monophyletic primitive group?  An assessment using 
          18S rDNA sequences.  Mol. Biol. Evol. 14:485-497.

Cavalier-Smith, T., M. T. E. P. Allsopp, E. E. Chao, N. Boury-Esnault, 
          and J. Vacelet.  1996.  Sponge phylogeny, animal monophyly, 
          and the origin of the nervous system: 18S rRNA evidence. 
          Can. J. Zool. 74:2031-2045.

Chen, J.-Y., P. Oliveri, C.-W. Li, G.-Q. Zhou, F. Gao,  J. W. 
          Hagadorn, K. J. Peterson, and E. H. Davidson.  2000. 
          Precambrian animal diversity: Putative phosphatized 
          embryos from the Doushantuo Formation of China. Proceedings 
          of the National Academy of Sciences (U.S.A.) 97:4457-4462.

Collins, A. G.  1998.  Evaluating multiple alternative hypotheses 
          for the origin of Bilateria: An analysis of 18S rRNA 
          molecular evidence.  Proceedings of the National Academy 
          of Sciences (U.S.A.) 95:15458-15463.

de Rosa R., J. K. Grenier, T. Andreeva, C. E. Cook, A. Adoutte, 
          M. Akam, S. B. Carroll, and G. Balavoine.  1999.  Hox 
          genes in brachiopods and priapulids and protostome 
          evolution.  Nature 399:772-776.

Dewel, R. A.  2000.  Colonial origin for Eumetazoa: Major morphological 
          transitions and the origin of Bilaterian complexity. 
          Journal of Morphology 243:35-74.

Dewel, R. A. and W. C. Dewel.  1998. The place of tardigrades 
          in arthropod evolution. Pages 109-123 in Arthropod 
          Relationships (R.A. Fortey and R.H. Thomas, eds.) The 
          Systematics Association and Chapman and Hall, London.

Eernisse, D. J. 1998. Arthropod and annelid relationships re-examined. 
          Pages 43-56 in Arthropod Relationships (R.A. Fortey 
          and R.H. Thomas, eds.) The Systematics Association 
          and Chapman and Hall, London.

Eernisse, D. J., J. S. Albert, F. E. Anderson.  1992.  Annelida 
          and Arthropoda are not sister taxa.  A phylogenetic 
          analysis of spiralian metazoan morphology.  Systematic 
          Biology 41:305-330.

Emig, C. C.  1997.  Are the Lophophorata a phylum?  Bulletin 
          de la Societé Zoologique de France 122:279-288.

Erwin, D. H.  1993.  The origin of metazoan development: A palaeobiological 
          perspective. Biological Journal of the Linnean Society 
          50: 255-274.

Fauchald, K. and G. W. Rouse.  1997.  Polychaete systematics: 
          Past and present.  Zool. Scr. 26:71-138.

Field, K. G., G. J. Olsen, D. J. Lane, S. J. Giovannoni, M. T. 
          Ghiselin, E. C. Raff, N. R. Pace, and R. A. Raff.  
          1988. Molecular phylogeny of the animal kingdom.  Science, 
          239:748-753.

Funch, P. and R. M. Kristensen.  1995.  Cycliophora is a new 
          phylum with affinities to Entoprocta and Ectoprocta. 
          Nature 378:711-714.

Garey, J. R., M. Krotec, D. R. Nelson, and J. Brooks.  1996. 
          Molecular analysis supports a tardigrade-arthropod 
          association.  Invertebrate Biology 115:79-88.

Garey, J. R., T. J. Near, M. R. Nonnemacher, and S. A. Nadler. 
          1996.  Molecular evidence for Acanthocephala as a subtaxon 
          of Rotifera.  J. Mol. Evol. 43:287-292.

Garey, J. R., A. Schmidt-Rhaesa, T. J. Near, and S. A. Nadler. 
          1998.  The evolutionary relationships of rotifers and 
          acanthocephalans. Hydrobiologia 387:83-91.

Garey, J. R. and A. Schmidt-Rhaesa.  1998.  The essential role 
          of "minor" phyla in molecular studies of animal evolution. 
          American Zoologist 38:907-917.

Giribet, G., S. Carranza, J. Baguna, M. Riutort, C. Ribera.  
          1996.  First molecular evidence for the existence of 
          a Tardigrada plus arthropoda clade.  Molecular Biology 
          and Evolution 13:76-84. 

Giribet, G., D. L. Distel, M. Polz, and W. Sterrer.  2000.  Triploblastic 
          relationships with emphasis on the acoelomates and 
          the position of Gnathostomulida, Cycliophora, Plathelminthes, 
          and Chaetognatha: a combined approach of 18S rDNA sequences 
          and morphology.  Systematic Biology 49:539-562.   

Giribet, G. and C. Ribera.  1998.  The position of arthropods 
          in the animal kingdom: A search for a reliable outgroup 
          for internal arthropod phylogeny.  Molecular Phylogenetics 
          and Evolution 9:481-488.

Halanych, K. M., J. D. Bacheller, A. M. A. Aguinaldo, S. M. Liva, 
          D. M. Hillis, and J. A. Lake. 1995. Evidence from 18S 
          ribosomal DNA that the lophophorates are protostome 
          animals. Science 267: 1641-1643.

Halanych, K. M.  1996. Testing hypotheses of Chaetognath origins: 
          long branches revealed by 18S ribosomal DNA. Systematic 
          Biology 45:223-246.

Halanych, K. M., R. A. Lutz, and R. C. Vrijenhoek.  1998.  Evolutionary 
          origins and age of vestimentiferan tube worms.  Cahiers 
          de Biologie Marine 39:355-358.

Hanelt, B., D. VanSchyndel, C. M. Adema, L. A. Lewis, E. S. Loker. 
          1996.  The phylogenetic position of Rhopalura ophiocomae 
          (Orthonectida) based on 18S ribosomal DNA sequence 
          analysis. Molecular Biology and Evolution 13:1187-1191.

Jenner, R. A. and F. R. Schram. 1999.  The grand game of metazoan 
          phylogeny: rules and strategies.  Biological Reviews 
          74:121-142.

Katayama, T., H. Wada, H. Furuya, N. Satoh, and M. Yamamoto. 
          1995.  Phylogenetic position of the dycyemid Mesozoa 
          inferred from 18S rDNA sequences.  Biological Bulletin 
          189:81-90.

Kim, J., W. Kim, and C. W. Cunningham.  1999.  A new perspective 
          on lower metazoan relationships from 18S rDNA sequences. 
          Mol. Biol. Evol. 16:423-427.

Knoll, A. H. and S. B. Carroll.  1999. Early animal evolution: 
          emerging views from comparative biology and geology. 
          Science 284:2129-2137.

Kobayashi, M., H. Furuya, and P. W. H. Holland.  1999.  Dicyemids 
          are higher animals.  Nature 401:762.

Kojima, S.,  T. Hashimoto, M. Hasegawa,  S. Murata,  S. Ohta, 
          H. Seki, N. Okada.  1993.  Close phylogenetic relationship 
          between Vestimentifera tube worms and Annelida revealed 
          by the amino acid sequence of elongation factor-1-alpha. 
          Journal of Molecular Evolution 37: 66-70.

Littlewood, D. T. J., M. J. Telford, K. A. Clough, K. Rohde. 
          1998.  Gnathostomulida - an enigmatic metazoan phylum 
          from both morphological and molecular perspectives. 
          Molecular Phylogenetics and Evolution 9:72-79.

Luter, C.  1997.  The phylogenetic position of Brachiopoda - 
          a comparison of morphological and molecular data.  
          Zoologica Scripta 26:245-253.

Mackey, L. Y., B. Winnepenninckx, R. DeWachter, T. Backeljau, 
          P. Emschermann, and J. R. Garey.  1996.  18S rRNA suggests 
          that Entoprocta are protostomes, unrelated to Ectoprocta. 
          Journal of Molecular Evolution 42:552-559.

McHugh, D.  1997.  Molecular evidence that echiurans and pogonophorans 
          are derived annelids.  Proc. Natl. Acad. Sci. USA 94:8006-8009.

McHugh, D. and G. W. Rouse.  1998.  Life history evolution of 
          marine invertebrates: new views from phylogenetic systematics. 
          Trends in Ecology and Evolution 13: 182-186. 

Morris, S. C.  1992.  Early metazoan evolution: First steps to 
          an integration of molecular and morphological data. 
          Pp. 450-459 in Bengtson, S. (ed.).  Early Life on Earth; 
          84th Nobel Symposium, Karlskoga, Sweden, May 16, 1992. 
          Columbia University Press: New York, New York, USA.

Nielsen, C.  1995.  Animal Evolution: Interrelationships of the 
          living phyla.  Oxford University Press.

Nielsen, C.  1997.  The phylogenetic position of the Arthropoda. 
          Pages 11-22 in Arthropod Relationships (R. A. Fortey 
          and R. H. Thomas, eds.)  Systematics Association Special 
          Volume Series 55.  Chapman & Hall, London.

Nielsen, C., N. Scharff, and D. Eibye-Jacobsen.  1996.  Cladistic 
          analyses of the animal kingdom.  Biol. J. Linn. Soc. 
          57:385-410.

Pawlowski, J., J. I. MontoyaBurgos, J. F. Fahrni, J. Wuest, and 
          L. Zaninetti.  1996.  Origin of the Mesozoa inferred 
          from 18S rRNA gene sequences.  Molecular Biology and 
          Evolution 13:1128-1132.

Ramsköld, J. and J.-Y. Chen.  1998.  Cambrian lobopodians:  morphology 
          and phylogeny.  Pages 107-150 in Arthropod Fossils 
          and Phylogeny (G. D. Edgecombe, ed.)  Columbia University 
          Press, New York.

Rouse, G. W. and K. Fauchald.  1998.  Recent views on the status, 
          delineation and classification of the Annelida.  American 
          Zoologist 38:953-964.

Ruiz-Trillo, I., M. Riutort, D. T. J. Littlewood, E. A. Herniou, 
          J. Baguña.  1999.  Acoel flatworms: earliest extant 
          bilaterian metzoans, not members of platyhelminthes. 
          Science 283:1919-1923.

Schlegel, M., J. Lom, A. Stechmann, D. Bernhard, D. Leipe, I. 
          Dykova, and M. L. Sogin.  1996.  Phylogenetic analysis 
          of complete small subunit ribosomal RNA coding region 
          of Myxidium lieberkuehni: Evidence that Myxozoa are 
          Metazoa and related to the Bilateria. Archiv für Protistenkunde 
          147:1-9.

Schmidt-Rhaesa, A., T. Bartolomaeus, C. Lemburg, U. Ehlers, and 
          J. R. Garey.  1998.  The position of the Arthropoda 
          in the phylogenetic system.  Journal of Morphology 
          238:263-285.

Schmidt-Rhaesa, A.  1998.  Phylogenetic relationships of the 
          nematomorpha - a discussion of current hypotheses. 
          Zoologischer Anzeiger 236:203-216.

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. 
          Journal of Parasitology 81:961-967.

Smothers, J. F.,  C. D. von Dohlen,  L. H. Smith, Jr.,  and R. 
          D. Spall.  1994.   Molecular evidence that the myxozoan 
          protists are metazoans.  Science 265:1719-1721.

Stechmann, A. and M. Schlegel.  1999.  Analysis of the complete 
          mitochondrial DNA sequence of the brachiopod Terebratulina 
          retusa places Brachiopoda within the protostomes.  
          Proc. R. Soc. Lond. Ser. B 266:2043-2052.

Telford, M. J., and P. W. H. Holland.  1993.  The phylogenetic 
          affinities of the chaetognaths:  A molecular analysis. 
          Molecular Biology and Evolution 10:660-676.

Turbeville, J. M., K. G. Field, and R. A. Raff.  1992.  Phylogenetic 
          position of phylum Nemertini, inferred from 18s rRNA 
          sequences: Molecular data as a test of morphological 
          character homology.  Molecular Biology and Evolution 
          9:235-249.

Turbeville, J. M., D. M. Pfeifer, K. G. Field,  and R. A. Raff. 
          1991.  The phylogenetic status of arthropods as inferred 
          from 18s ribosomal RNA sequences.  Molecular Biology 
          and Evolution 8:669-686.

Valentine, J. W.  1989.   Bilaterians of the Precambrian-Cambrian 
          transition and the Annelid-Arthropod relationship. 
          Proceedings of the National Academy of Sciences (U.S.A.) 
          86:2272-2275.

Valentine, J. W.  1994.  Late Precambrian bilaterians: grades 
          and clades. Proceedings of the National Academy of 
          Sciences (U.S.A.) 91:6751-6757.

Valentine, J. W.  1997.  Cleavage patterns and the topology of 
          the metazoan tree of life.  Proceedings of the National 
          Academy of Sciences (U.S.A.) 94:8001-8005.

Valentine, J. W., D. Jablonski, D. H. Erwin.  1999.  Fossils, 
          molecules and embryos: new perspectives on the Cambrian 
          explosion.  Development 126:851-859.

Wada, H., and N. Satoh.  1994.  Details of the evolutionary history 
          from invertebrates to vertebrates, as deduced from 
          the sequences of 18S rDNA.  Proceedings of the National 
          Academy of Sciences (U.S.A.) 91:1801-1804.

Waggoner, B. M.  1998.  Interpreting the earliest metazoan fossils: 
          What can we learn?  American Zoologist 38:975-982.

Wang, D. Y.-C., S. Kumar, and S. B. Hedges.  1999.  Divergence 
          time estimates for the early history of animal phyla 
          and the origin of plants, animals, and fungi.  Proceedings 
          of the Royal Society London Series B 266:163-171.

Westheide, W., D. McHugh, G. Purschke, and G. Rouse.  1999.  
          Systematization of the Annelida: different approaches. 
          Hydrobiologia 402: 291-307.

Winnepenninckx, B., T. Backeljau, and R. M. Kristensen.  1998. 
          Relations of the new phylum Cycliophora.  Nature 393:636-638.

Winnepenninckx, B., T.  Backeljau, L. Y. Mackey, J. M. Brooks, 
          R. de Wachter, S. Kumar, and J. R. Garey.  1995.  18S 
          rRNA data indicate that aschelminthes are polyphyletic 
          in origin and consist of at least three distinct clades. 
          Molecular Biology and Evolution 12:1132-1137.

Winnepenninckx, B., T.  Backeljau, and R. de Wachter.  1995. 
          Phylogeny of protostome worms derived from 18S rRNA 
          sequences.  Molecular Biology and Evolution 12:641-649.

Winnepenninckx, B., Y. van de Peer, and T. Backeljau.  1998. 
          Metazoan relationships on the basis of 18S rRNA sequences: 
          a few years later... American Zoologist 38:888-906.

Zrzavy, J., S. Mihulka, P. Kepka, A. Bezdek, and D. Tietz.  1998. 
          Phylogeny of the Metazoa based on morphological and 
          18S ribosomal DNA evidence.  Cladistics 14:249-285.

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