Reconstruction of Isoxys, from Cambrian Cafe.

Belongs within: Cephalata.
Contains: Euarthropoda, Leanchoiliida.

The Arthropoda are a major clade of animals characterised by the presence of sclerotised, jointed limbs. The arthropods have been one of the most successful of all animal groups and dominate many modern environments. As defined herein, this taxon includes the Anomalocarida, a Cambrian to Devonian group of nektonic animals with large grasping appendages at the front of the head and a mouth surrounded by a radial array of sclerotised, overlapping plates (Vinther et al. 2014). A number of Cambrian forms may also represent lineages outside the arthropod crown group (Euarthropoda). Examples of these forms include the genus Isoxys, members of which had a large carapace covering most of the body, with a more or less well-developed spine at the anterior end of the carapace.

Arthropoda (see below for synonymy) LSE13
    |--Anomalocarida [Anomalocaridata, Radiodonta] VS14
    |    |--+--‘Anomalocaris’ pennsylvanica VS14
    |    |  `--+--Paranomalocaris multisegmentalis VS14
    |    |     `--+--Anomalocaris Whiteaves 1892 VS14, B95 [Anomalocarididae]
    |    |        |    `--*A. canadensis Whiteaves 1892 WB85
    |    |        `--Amplectobeluidae VS14
    |    |             |--‘Anomalocaris’ saron VS14
    |    |             `--Amplectobelua VS14
    |    |                  |--A. kunmingensis VS14
    |    |                  `--+--A. stephenensis VS14
    |    |                     `--A. symbrachiata B14
    |    `--+--Cetiocaridae VS14
    |       |    |--‘Anomalocaris’ briggsi VS14
    |       |    `--Tamisiocaris borealis VS14
    |       `--Hurdiidae VS14
    |            |--+--Schinderhannes Kühl, Briggs & Rust 2009 VS14, KBR09
    |            |  |    `--*S. bartelsi Kühl, Briggs & Rust 2009 KBR09
    |            |  `--+--Stanleycaris hirpex VS14
    |            |     `--Hurdia Walcott 1912 VS14, B95
    |            |          `--*H. victoria Walcott 1912 (see below for synonymy) DB09
    |            `--+--Peytoia Walcott 1911 VS14, WB85
    |               |    `--*P. nathorsti Walcott 1911 [=Anomalocaris nathorsti] WB85
    |               `--Laggania Walcott 1911 KBR09, WB85
    |                    `--*L. cambria Walcott 1911 WB85
    `--+--Isoxys LSE13
       |    |--I. auritus LSE13
       |    `--+--I. acutangulus LSE13
       |       |--I. curvirostrus LSE13
       |       `--I. volucris LSE13
       `--+--Nereocaris LSE13
          |    |--N. briggsi LSE13
          |    `--N. exilis LSE13
          `--+--+--Jugatacaris LSE13
             |  `--Pectocaris LSE13
             |       |--P. euryptelata LSE13
             |       `--P. spatiosa LSE13
             `--+--Odaraia Walcott 1912 LSE13, B95 [Odaraiata, Odaraiida]
                |    `--O. alata CM98
                `--+--Protocaris Walcott 1884 LSE13, G89
                   `--+--‘Canadaspis’ laevigata LSE13
                      |--+--Branchiocaris Briggs 1976 LSE13, B95 [Branchiocarata, Branchiocarida]
                      |  |    `--B. pretiosa (Resser 1929) [=Protocaris pretiosa] G89
                      |  `--Loricicaris LSE13
                      `--+--Canadaspidida LSE13
                         |    |--Canadaspis Novoshilov 1960 B95
                         |    |    `--C. perfecta (Walcott 1912) RL02
                         |    `--Perspicaris Briggs 1977 LSE13, B95
                         |         |--P. dictynna LSE13
                         |         `--P. recondita LSE13
                         `--+--Fuxianhuiida [Fuxianhuiata] LSE13
                            |    |--Shankouia LSE13
                            |    `--+--Liangwangshania LSE13
                            |       `--+--Chengjiangocaris LSE13
                            |          |    |--C. kunmingensis Yang, Ortega-Hernández et al. 2013 YO-H13
                            |          |    `--C. longiformis Hou & Bergström 1991 B95
                            |          `--+--Guangweicaris LSE13
                            |             `--Fuxianhuia LSE13
                            |                  |--F. protensa Hou 1987 B95
                            |                  `--F. xiaoshibaensis Yang, Ortega-Hernández et al. 2013 YO-H13
                            `--+--Kootenichelidae LSE13
                               |    |--Worthenella Walcott 1911 LSE13, H62
                               |    |    `--*W. cambria Walcott 1911 H62
                               |    `--Kootenichela LSE13
                               `--+--Fortiforceps foliosa Hou & Bergström 1997 LSE13, CB04
                                  `--+--Jiangfengia Hou 1987 LSE13, B95 [Jianfengiida]
                                     |    `--J. multisegmentalis Hou 1987 CB04
                                     `--+--+--Euarthropoda LSE13
                                        |  `--Pseudonotostraca G89
                                        |       |--Waptia fieldensis G89, CM98
                                        |       `--‘Burgessia’ Walcott 1912 non McCook 1894 CB04, B95 [Burgessiida]
                                        |            `--B. bella Walcott 1912 CB04
                                        `--Megacheira LSE13
                                             |--Yohoia Walcott 1912 LSE13, B95 [Yohoiida]
                                             |    `--Y. tenuis Walcott 1912 CB04
                                             |--Parapeytoia yunnanensis LSE13, KBR09
                                             `--+--Haikoucaris LSE13
                                                `--Leanchoiliida LSE13

Arthropoda incertae sedis:
  Dictyocaris Salter 1860 LS02
  Tontoia Walcott 1912 B95
  Corcorania Jell 1980 B95
  Serracaris Briggs 1978 B95
    `--S. lineata [=Anomalocaris lineata] WB85
  Cambropodus gracilis BL05
  Achanarraspis Anderson et al. 2000 N02
  Magnoculocaris blindi CB04
  ‘Leuconoe’ Bogachev 1930 non Boie 1830 (n. d.) H75
    `--*L. paradoxa Bogachev 1930 (n. d.) H75
  Velancorina Pflug 1966 G79
    `--*V. martina Pflug 1966 G79
  Occacaris B02
  Clypecaris B02
  Caryosyntrips serratus VS14

Inorganic: Hurdia davidi Chapman 1926 H75

Arthropoda [Annulosa, Brachiocaridea, Burgessidea, Canadaspididea, Cheliceromorpha, Euthycarcinomorpha, Merostomoidea, Schizoramia, Trilobitoidea, Yohoiidacea, Yohoiidea] LSE13

*Hurdia victoria Walcott 1912 [incl. Proboscicaris agnosta Rolfe 1962, H. dentata Simonetta & Della Cave 1975, P. hospes Chlupáč & Kordule 2002, Liantuoia inflata Cui & Huo 1990, P. ingens Rolfe 1962, P. obtusa Simonetta & Della Cave 1975, Amiella ornata Walcott 1911, Sidneyia ornata, Hurdia triangulata Walcott 1912, Huangshandongia yichangensis Cui & Huo 1990] DB09

*Type species of generic name indicated


[B95] Bousfield, E. L. 1995. A contribution to the natural classification of Lower and Middle Cambrian arthropods: food-gathering and feeding mechanisms. Amphipacifica 2: 3–34.

[BL05] Briggs, D. E. G., B. S. Lieberman, S. L. Halgedahl & R. D. Jarrard. 2005. A new metazoan from the Middle Cambrian of Utah and the nature of the Vetulicolia. Palaeontology 48 (4): 681–686.

[B02] Budd, G. E. 2002. A palaeontological solution to the arthropod head problem. Nature 417: 271–275.

[CM98] Conway Morris, S. 1998. The Crucible of Creation. Oxford University Press: Oxford.

[CB04] Cotton, T. J., & S. J. Braddy. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 169–193.

[DB09] Daley, A. C., G. E. Budd, J.-B. Caron, G. D. Edgecombe & D. Collins. 2009. The Burgess Shale anomalocaridid Hurdia and its significance for early euarthropod evolution. Science 323: 1597–1600.

[G79] Glaessner, M. F. 1979. Precambrian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A79–A118. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[G89] Gould, S. J. 1989. Wonderful Life. Vintage: London.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[H62] Howell, B. F. 1962. Worms. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W144–W177. Geological Society of America, and University of Kansas Press.

[KBR09] Kühl, G., D. E. G. Briggs & J. Rust. 2009. A great-appendage arthropod with a radial mouth from the Lower Devonian Hunsrück Slate, Germany. Science 323: 771–773.

[LS02] Lange, S., & F. R. Schram. 2002. Possible lattice organs in Cretaceous Thylacocephala. Contributions to Zoology 71 (4): 159–169.

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4: 2485.

[N02] Newman, M. J. 2002. A new naked jawless vertebrate from the Middle Devonian of Scotland. Palaeontology 45 (5): 933–941.

[RL02] Rode, A. L., & B. S. Lieberman. 2002. Phylogenetic and biogeographic analysis of Devonian phyllocarid crustaceans. Journal of Paleontology 76 (2): 271–286.

[VS14] Vinther, J., M. Stein, N. R. Longrich & D. A. T. Harper. 2014. A suspension-feeding anomalocarid from the Early Cambrian. Nature 507: 496–499.

[WB85] Whittington, H. B., & D. E. G. Briggs. 1985. The largest Cambrian animal, Anomalocaris, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London 309: 569–609.

[YO-H13] Yang, J., J. Ortega-Hernández, N. J. Butterfield & X.-G. Zhang. 2013. Specialized appendages in fuxianhuiids and the head organization of early euarthropods. Nature 494: 468–471.

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