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The New View of Animal Phylogeny
Abstract
▪ Abstract Molecular tools have profoundly rearranged our understanding of metazoan phylogeny. Initially based on the nuclear small ribosomal subunit (SSU or 18S) gene, recent hypotheses have been corroborated by several sources of data (including the nuclear large ribosomal subunit, Hox genes, mitochondrial gene order, concatenated mitochondrial genes, and the myosin II heavy chain gene). Herein, the evidence supporting our current understanding is discussed on a clade by clade basis. Bilaterian animals consist of three clades: Deuterostomia, Lophotrochozoa, and Ecdysozoa. Each clade is supported by molecular and morphological data. Deuterostomia is smaller than traditionally recognized, consisting of hemichordates, echinoderms, chordates, and Xenoturbella (an enigmatic worm-like animal). Lophotrochozoa groups animals with a lophophore feeding apparatus (Brachiopoda, Bryozoa, and Phoronida) and trochophore larvae (e.g., annelids and mollusk), as well as several other recognized phyla (e.g., platyhelmin thes, sipunculans, nemerteans). Ecdysozoa comprises molting animals (e.g., arthropods, nematodes, tardigrades, priapulids), grouping together two major model organisms (Drosophila and Caenorhabditis) in the same lineage. Platyhelminthes do not appear to be monophyletic, with Acoelomorpha holding a basal position in Bilateria. Before the emergence of bilateral animals, sponges, ctenophorans, cnidarians, and placozoans split from the main animal lineage, but order of divergence is less than certain. Many questions persist concerning relationships within Ecdysozoa and Lophotrochozoa, poriferan monophyly, and the placement of many less-studied taxa (e.g., kinorhynchs, gastrotrichs, gnathostomulids, and entoprocts).
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“These have been interpreted as grasping spines of the Chaetognatha, a protostomian phylum close to the Lophotrochozoa, but most likely its sister-group (Halanych, 2004; Dunn et al . 2008).”
From Chronology of early Cambrian biomineralization (2012) · cites this paper as Halanych, 2004
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Cites
PHYLOGENIES FROM MOLECULAR SEQUENCES: INFERENCE AND RELIABILITY
Felsenstein, J. (1988) · Annual Review of Genetics · doi:10.1146/annurev.ge.22.120188.002513 · Cited by 2,159
Evidence for a clade of nematodes, arthropods and other moulting animals
Aguinaldo, A. M. A., Turbeville, J. M., Linford, L. S., Rivera, M. C., Garey, J. R., Raff, R. A., & Lake, J. A. (1997) · Nature · doi:10.1038/387489a0 · Cited by 1,609
A revised six-kingdom system of life
Cavalier‐Smith, T. (1998) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1017/s0006323198005167 · Cited by 1,213
Animal evolution: interrelationships of the living phyla
(2012) · Choice Reviews Online · doi:10.5860/choice.50-1469 · Cited by 1,041
Early animal evolution: emerging views from comparative biology and geology
Knoll, A. H., & Carroll, S. B. (1999) · Science · doi:10.1126/science.284.5423.2129 · Cited by 821
Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorite
Xiao, S., Zhang, Y., & Knoll, A. H. (1998) · Nature · doi:10.1038/35318 · Cited by 783
Molecular Phylogeny of the Animal Kingdom
Field, K. G., Olsen, G. J., Lane, D. J., Giovannoni, S. J., Ghiselin, M. T., Raff, E. C., Pace, N. R., & Raff, R. A. (1988) · Science · doi:10.1126/science.3277277 · Cited by 749
Biostratigraphic and Geochronologic Constraints on Early Animal Evolution
Grotzinger, J. P., Bowring, S. A., Saylor, B. Z., & Kaufman, A. J. (1995) · Science · doi:10.1126/science.270.5236.598 · Cited by 701
Rouse, G. W., & Fauchald, K. (1997) · Zoologica Scripta · doi:10.1111/j.1463-6409.1997.tb00412.x · Cited by 683
Reading the entrails of chickens: molecular timescales of evolution and the illusion of precision
Graur, D., & Martin, W. (2004) · Trends in Genetics · doi:10.1016/j.tig.2003.12.003 · Cited by 669
Research works printed but not linked: 31
- Abele LG, 1989, Mol. Biol. Evol., 6, 685
- Aleshin VV, 2002, Zh. Obshch. Biol., 63, 195
- Bridge D, 1995, Mol. Biol. Evol., 12, 679
- Brusca RC, Brusca GJ. 2003.Invertebrates. Sunderland, MA: Sinauer. 936 pp. 2nd ed.
- Caullery M, 1914, C.R. Acad. Sci., 158, 2014
- Chen J-Y, 1997, The Cambrian Explosion and the Fossil Record, 10, 11
- Collins AG, 2000, Sci. Mar., 4, 1
- 10.2307/2992569 · DOI
- Ehlers U, 1996, Verh. Dtsch. Zool. Ges., 89, 8
- 10.1007/PL00006088 · DOI
- 10.1006/clad.1999.0128 · DOI
- 10.2307/2413616 · DOI
- Halanych KM, 1998, Cah. Biol. Mar., 39, 355
- Haszprunar G, 2000, Am. Malacol. Bull., 15, 115
- 10.1093/molbev/msh011 · DOI
- Jenner RA, 1999, Belg. J. Zool., 129, 245
- 10.2307/2640769 · DOI
- Malakhov VV, 1980, Zool. Zh., 59, 485
- 10.1139/cjz-78-11-1873 · DOI
- Metschnikoff VE, 1881, Zool. Anz., 4, 139
- 10.1006/bijl.1996.0023 · DOI · Paper page
- 10.1006/zjls.2000.0263 · DOI
- Seaver EC, 2003, Int. J. Dev. Biol., 47, 583
- Sørensen MV, 2000, Zool. Anz., 239, 297
- Telford MJ, 1993, Mol. Biol. Evol., 10, 660
- Turbeville JM, 1992, Mol. Biol. Evol., 9, 235
- Turbeville JM, 1994, Mol. Biol. Evol., 11, 648
- van der Land J, 1977, Z. Zool. Syst. Evol., 1975, 85
- Winnepenninckx B, 1995, Mol. Biol. Evol., 12, 1132
- Zimmer RL, 1973, Living and Fossil Bryozoa, 593
- 10.1006/clad.1998.0070 · DOI
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Metadata
- Authors
- Halanych, K. M.
- Affiliations
- Auburn University (US)
- Corresponding author
- Halanych, Kenneth M.
- ORCID
- Halanych, Kenneth M.
- Venue
- Annual Review of Ecology, Evolution, and Systematics
- Publisher
- Annual Reviews
- Published
- 2004-12-15
- Volume
- 35
- Issue
- 1
- Pages
- 229-256
- ISSN
- 1543-592X (ISSN-L)
- 1545-2069
- OpenAlex
- W2114395497
- Cited by
- 489
- References
- 204
- Citations by year
- 2026: 8 · 2025: 12 · 2024: 7 · 2023: 3 · 2022: 11 · 2021: 13 · 2020: 8 · 2019: 7 · 2018: 5 · 2017: 10 · 2016: 16 · 2015: 20 · 2014: 22 · 2013: 15 · 2012: 22
- FWCI
- 6.04
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Annual Review of Ecology Evolution and Systematics (publishedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Marine Invertebrate Physiology and Ecology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Invertebrate Taxonomy and Ecology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Protist diversity and phylogeny
- Keywords
- animal phylogeny, metazoan phylogeny, deuterostomes, Lophotrochozoa, Ecdysozoa, Acoelomorpha, Cnidaria, placozoans, Hox genes, mitochondrial gene order, myosin II heavy chain, Xenoturbella, hemichordates, Echinodermata, Chordata
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