After Conway Morris, 1993, and Conway Morris & Caron, 2007
Alternative phylogeny after Aguinaldo et. al. 1997
Alternative phylogeny after Winnepennickyx et al., 1998
Alternative phylogeny after Zrzavý et al., 2001
Alternative phylogeny after Brusca " Brusca, 2002
Alternative phylogeny after Paps et al., 2009
<==o Protostoma (protostomes; alkusuiset) |--o Aschelmithes [Pseudocoelomata] (valeruumiinonteloiset) | |?- Tardigrada Spallanzani, 1777 (water bears; karhukaiset) | |-- Chaetognatha Leuckart, 1854 [incl. †Protoconodonta] (arrow worms; nuolimadot) | `--+--+-- Nematomorpha Vejdovsky, 1886 (horse hair worms, gordian worms; jouhimadot) | | `-- Nemata Gobb, 1919 [Nematoda Diesing, 1861] (nematodes, round worms; sukkulamadot) | `--+-- Gastrotricha Metschnikoff, 1865 (hairybacks; sukaspintaiset) | `--o Syndermata | |== Rotifera Cuvier, 1798 (rotifers, wheel animals; rataseläimet) | `-- Acanthocephala Koelreuter, 1771 (thorny-headed worms, spiny-headed worms; väkäkärsämadot) `--+?-+?- Entoprocta Nitsche, 1870 (entoprocts, goblet worms; pikarimadot) | |?- Bryozoa Ehrenberg, 1831 [Ectoprocta Nitsche, 1869] (moss animals; sammaleläimet) | |?- Cycliophora Funch & Kristensen, 1995 (“Wheel Wearers”) | `?- Mesozoa [see Basal Metazoa] `--+?-o †Cribricyathea | |-- †Conoidocyathida | |-- †Cribricyathida | `-- †Vologdinophyllida |?-o †Machaeridia | |-- †Hercollepadida | `-- †Turrilepadida |--o Ecdysozoa Aguinaldo, Turbeville, Linford, Rivera, Garey, Raff & Lake, 1997 (skinshedders; nahanluojat) | |-- †Facivermis yunnanicus (transional for between pripulid/palaeoscolecid worms and lobopods??) | |--o Arthropodomorpha (arthropods and related animals; niveljalkaismaiset eläimet) | | |?-o †Sprigginida | | | |?- †Bomakellia kelleri Fedonkin 1985 [a rangeomorph frond] | | | `-- †Spriggina floundersi Glaessner, 1958 [also a rangeomorph frond?] | | `--o Aiolopoda Hou & Bergström, 1996 [incl. Arthropoda von Siebold, 1848][Panarthropoda] (water bears, velvet worms, anomalocarids and arthropods; karhukaiset, käsnäjalkaiset, anomalocaridit ja niveljalkaiset) | | |-- Tardigrada Spallanzani, 1777 (water bears; karhukaiset) [considered as Aschelmithes by Hou & Bergström (1997)] | | `==o Loboboda Snodgrass, 1938 [Onychophora sensu lato] [Microdictyoniida; Protonychophora Hutchinson, 1930] (lobopods, velvet worms; käsnäjalkaiset) [paraphyletic grade of earliest arthropods] | | |-- Onychophora Grube, 1853 [Euonychophora Hutchinson, 1930] (modern lobopods, velvet worms; nykykäsnäjalkaiset) | | `--+-- †Radiodonta Collins, 1996 (anomalocaritids and relatives) | | `-- Euarthropoda [Arthropoda sensu stricto] [Schizoramia sensu Hou & Bergström, 1997] | `--o Scalidophora sensu Dong, Donoghue, Cheng & Liu, 2004 | |--+-- †Ancalagon minor (Walcott, 1911) Conway Morris, 1977 [Ottoia minor Walcott, 1911] [Ancalagonidae Conway Morris, 1977] | | `-- †Fieldia lanceolata Walcott, 1912 [Fieldiidae] | `--+--o †Markuelia Val´kov, 1983 [Middle and Late Cambrian protostome embryos] | | |-- †M. hunanensis Dong & Donoghue, in Dong, Donoghue, Cheng & Liu, 2004 | | `-- †M. secunda Val´kov, 1983 | `--o Priapulida Delage & Hérouard, 1897 (non Théel, 1906?) sensu lato [Priapuloidea] [incl. Palaeoscolecida Conway Morris & Robison, 1986 sensu Dong, Donoghue, Cheng & Liu, 2004] (priapulid worms; makkaramadot) | |== †Palaeoscolecida Conway Morris & Robison, 1986 [paraphyletic basal Scalidophorans?] | `--+-- Kinorhyncha Reinhard, 1881 (mud dragons; okakaulamadot) | `--+-- Priapulida Delage & Hérouard, 1897 (non Théel, 1906?) (s.s.) [Priapuloidea] (priapulid worms, penis worms; makkaramadot) | `-- Loricifera Kristensen, 1983 (Brush heads) `--o Coeloscleritophora Bengtson & Missarzhevsky, 1981 [paraphyletic] `--o Lophotrochozoa Halanych, Bacheller, Liva, Aguinaldo,, Hillis & Lake, 1995 |--o “Molluscamorpha” [Mollusca sensu lato] | |-- †Kimberella quadrata (Glaessner & Wade, 1966) Wade, 1972 [Kimberia quadrata Glaessner & Wade, 1966] | `--+-- †Odontogriphus omalus Conway Morris, 1976 [Odontogriphidae Conway Morris, 1976] | |?= †Halwaxida Conway Morris & Caron, 2007 [incl. Siphogonuchitida; Halkieriida; Wiwaxiida; ?Sachitida; ?Chancelloriida] [paraphyletic basal Aculiferas sensu Vinther, 2015] | `--+?- †Tullimonstrum gregarium Richardson, Jr., 1966 [Tullimonsterida] [has been reidentified as hyperoartian vertebrate by McCoy et al. (2016)] | `--+-- Mollusca Cuvier 1795 (molluscs; nilviäiset) | `--+?- †Hyolitha Marek, 1963 (hyoliths, hyoliitit) | `-- Sipuncula Rafinesque, 1814 [Sipunculoidea] (sipunculid worms, peanut worms; ruiskumadot) `--+--o “Annelidamorpha” | |-- †Canadia spinosa Walcott, 1911 | |--+-- †Amiskwia sagittiformis Walcott,1911 [Amiskwiida] | | `-- Nemertea Schultze, 1851 [Nemertinea] (ribbon worms, proboscis worms; limamadot) | `--o Annelida sensu lato | |-- Lobatocerebrum psammicola Rieger, 1980 [Lobatocerebrida; Lobatocerebridae Rieger, 1980] | `--+--+?- †Banffia constricta [Banffozoa: Banffida: Banffidae] | | `-- Echiura Newby, 1940 [Echiuroidea] (spoon worms; tähtimadot) | `-- Annelida Lamarck, 1809 [incl. Vestimentifera & Pogonophora sensu Rouse, 2001] (annelids, segmented worms; nivelmadot) `--o Lophophorata Hyman, 1959 ex Emig, 1977 |-- Phoronida Hatschek, 1888 (phoronids, horseshoe worms; tupsumadot) `--o “Pan-Brachiopoda” |== †Tommotiida Missarzhevsky, 1970 (tommotians) [paraphyletic stem-brachiopods] |?- †Hyolitha Marek, 1963 sensu Moysiuk, Smith & Caron, 2017 (hyoliths, hyoliitit) `-- Brachiopoda Duméril, 1806 (brachiopods, lampshells; lonkerojalkaiset)
After Aguinaldo, Turbeville, Linford, Rivera, Garey, Raff & Lake, 1997 and Dong, Donoghue, Cheng & Liu, 2004<==o Protostoma |?- Myzostomida |?- Annelida |?- Mollusca |?- Sipuncula |--o Lophotrochozoa | |?- Mesozoa | |?- Entoprocta | |?- Bryozoa [Ectoprocta] | |?- Cycliophora | |?- Gastrotricha | |?- Rotifera | `?- Acanthocephala `--o Ecdysozoa |?- Chaetognatha |--o Introverta | |--+-- Nemata | | `-- Nematomorpha | `--o Scelidophora | |--+-- †Ancalagon | | `-- †Fieldia | `--+--o †Markuelia | | |-- †M. hunanensis | | `-- †M. secunda | `--+-- Kinorhyncha | `--+-- Loricifera | `-- Priapulida `--o Euarthropoda |-- Loboboda |-- Tardigrada |-- Crustaceomorpha |-- Insecta `--+-- Myriapoda `-- Chelicerata Reference(s):
|
After Winnepennickyx et al., 1998<==o Protostoma |?- Mesozoa |?- Sipuncula |?- Myzostomida |?- Loricifera |?- Chaetognatha |?- Kinorhyncha |?- Nematomorpha |-- Mollusca |-- Annelida |--o Lophophorata | |-- Phoronida | `-- Brachiopoda |-- Entoprocta |-- Bryozoa [[Ectoprocta] |-- Gastrotricha |--+?- Cycliophora | `--+?- Rotifera | `?- Acanthocephala |-- Lobopoda |-- Nemata `--+-- Priapulida `--+?- Tardigrada |?- Myriapoda |?- Chelicerata `--+-- Crustaceomorpha `-- Insecta Note:This tree is from molecular phylogenetic study made by Winnepennickyx, Backeljau & Kristensen (1998). This doesn't agree with a study by Aguinaldo et al. (1997), although it too contains a clade made by priapulids and arthropods. Aguinaldo et al. also contains clade made by phoronids and brachiopods. Several Protostome-phylums were apparently untreated, I have placed them here too, marked as unsure relationship. [ |?- ] Reference(s):
|
After Zrzavý et al., 2001, with fossil phyla from Conway Morris, 1993<==o PROTOSTOMA |--o Aschelmithes | |-- Xenoturbellida | |-- Platyhelmithes | `--o Acanthognatha | |--o Neotrichozoa | | |-- Gastrotricha | | `-- Gnathostomulida | `--o Prosomastigozoa | |-- Myzostomida | `--+-- Cycliophora | `--o Syndermata | `==o Rotifera | |-- Monogononta | `--o Lemniscea | |-- Bdelloida | `--+-- Seisonida | `-- Acanthocephala `--+--o Arthropodomorpha | |-- Cephalorhyncha | |--+-- Chaetognatha | | `--+-- Nematomorpha | | `-- Nemata | `--+--o †Sprigginida | | |?- †Bomakellia | | `-- †Spriggina floundersi | |?- Tardigrada | `-- Arthropoda `--+-- Entoprocta |-- Bryozoa [Ectoprocta] |--+-- †Chancelloriida | `-- †Sachitida `--+-- †Siphogunuchitida `--+--o †Halkieriida | `-- †Halkeria evangelista `--+--o “Molluscamorpha” | `--+?- †“Tullimonsterida” | `--+-- Mollusca | `--+-- †Hyolitha | `-- Sipuncula `--+--o “Annelidamorpha” | |?- †Wiwaxia corrugata | `--+?- †Canadia spinosa | |--+-- †Amiskwia sagittiformis | | `-- Nemertea | `--+-- Lobatocerebrida | `--+--+-- †Banffida | | `-- Echiura | `-- Annelida `--+--o Lophophorata | |-- Phoronida | `--+== †Tommotiida | `-- Brachiopoda `--+-- †Palaeoscolecida `--+-- Kinorhyncha `--+-- Loricifera `-- Priapulida Reference(s):
|
After Brusca & Brusca, 2002<==o Bilateralia |--o Deuterostomia | |-- Chaetognatha | |-- Ectoprocta | |-- Phoronida | |-- Brachiopoda | |-- Echinodermata | `--+--o Hemichordata | | |-- Pterobranchia | | `-- Enteropneusta | `--o Chordata | |-- Urochordata | `--+-- Cephalochordata | `-- Vertebrata `--o Protostomia |-- Platyhelmithes `--+-- Nemertea |--+-- Sipuncula | `-- Mollusca |-- Echiura |--o Articulata | |-- Annelida | `--+-- Onychophora | `--+-- Tardigrada | `-- Arthropoda |-- Gnathostomulida |--+-- Entoprocta | `-- Cycliophora |--+-- Rotifera | `-- Acanthocephala `--o Cycloneuralia |-- Gastrotricha |-- Nemata |-- Nematomorpha `--+-- Priapulida `--+-- Kinorhyncha `-- Loricifera Reference(s):
|
After Paps, Bagañà & Riutort, 2009<==o Protostoma |--o Ecdysozoa (nahanluojat) | |?- Chaetognatha | |--+-- Nematomorpha | | `-- Nemata | `?-+--+-- Kinorhyncha | | `-- Priapulida | `--+-- Tardigrada | `-- Arthropoda `--o Lophotrochozoa sensu lato |-- Gnathostomulida `--+-- Gastrotricha `--+--o Gnathifera | |-- Limnognathia maerski [Micrognathozoa] | `--+== Rotifera | `-- Acanthocephala `--o Lophotrochozoa sensu stricto |--+-- Platyhelmithes | `--+-- Bryozoa [Ectoprocta] | `--+-- Cycliophora | `-- Entoprocta `--+-- Nemertea `--+--+-- Echiura | `--+-- Sipuncula | `-- Annelida `--+-- Mollusca `--o Brachiozoa |-- Phoronida `-- Brachiopoda Reference(s):
|
Reference(s):
- Caron, J.-B., 2005: Banffia constricta, a putative vetulicolid from the Middle Cambrian Burgess Shale.
–Earth and Environmental Science Transactions of the Royal Society of Edinburgh: Vol. 96, #02, pp. 95-111 [doi: doi:10.1017/S0263593300001255] - Conway Morris, S., 1993: The fossil record and early evolution of the Metazoa.
–Nature, vol. 361, 21 January, pp. 219-225 - Conway Morris, S. & Caron, J.-B., 2007: Halwaxiids and the early evolution of the Lophotrochozoans.
–Science: Vol. 315, pp. 1255-1258 [doi: 10.1126/science.1137187] - Conway Morris, S. & Peel, J. S., 2010: New palaeoscolecidan worms from Lower Cambrian: Sirius Passet, Latham Shale and Kinzers Shale.
–Acta Palaeontologica Polonica: Vol. 55, #1, pp. 141-156 [doi: 10.4202/app.2009.0058] - Dong, X.-p., Donoghue, P. C. J., Cheng, H. & Liu, J.-b., 2004: Fossil embryos from the Middle and Late Cambrian period of Hunan, south China.
–Nature: Vol. 427, pp. 237-240 - Funch, P. & Kristensen, R. M., 1995: Cycliophora is a new phylum with affinities to Endoprocta and Ectoprocta.
–Nature: Vol. 378, 14 December, pp. 711-714 - Hou, X. & Bergström, J., 1997: Arthropods of the Lower Cambrian Chengjiang fauna, southwest China.
–Fossils & Strata, Number 45, 22nd December 1997, pp. 1-116 - –iNet: Tree of Life: http://tolweb.org/tree/
- Kazlev, M. A., 2002-: The "Procoelomates"
–iNet: The Palaeos: http://www.palaeos.com/Invertebrates/Procoelomates/Procoelomates.html - McCoy, V. E., Saupe, E. E., Lamsdell, J. C., Tarhan, L. G., McMahon, S., Lidgard, S., Mayer, P., Whalen, C. D., Soriano, C., Finney, L., Vogt, S., Clark, E. G., Anderson, R. P., Petermann, H., Locatelli, E. R. & Briggs, D. E. G., 2016: The ‘Tully monster’ is a vertebrate.
–Nature: Vol. 532, #7600, pp. 496-499 [doi: 10.1038/nature16992] - Moysiuk, J., Smith, M. R. & Caron, J.-B., 2017: Hyoliths are Palaeozoic lophophorates.
–Nature: in press [doi: 10.1038/nature20804] - Paps, J., Bagañà, J. & Riutort, M., 2009: Lophotrochozoa internal phylogeny: new insights from an up-to-date analysis of nuclear ribosomal genes.
–Proceedings of the Royal Society of London: Biological series: Vol. 276, pp. 1245-1254 [doi: 10.1098/rspb.2008.1574] - Parker, S. P. (ed.), 1982: Synopsis and classification of living organisms. Vols. 1 & 2
–McGrew-Hill Book Company - Porter, S. M., 2008: Skeletal microstructure indicates chancelloriid and halkieriids are closely related.
–Palaeontology: Vol. 51, #4, pp. 865-879 [doi: 10.1111/j.1475-4983.2008.00792.x] - Rieger, R. M., 1980: A new group of interstitial worms, Lobatocerebridae nov. fam. (Annelida) and its significance for metazoan phylogeny.
–Zoomorphologie: Vol. 95, #1, pp. 41-84 [doi: 10.1007/BF01342233] - Szaniawski, H., 2002: New evidence for the protoconodont origin of chaetognaths.
–Acta Palaeontologica Polonica: Vol. 47, #3, pp. 405-419