Lophotrochozoa

The phylogenetically distinct Limnognathia maerski, first described in 2000 from a cold spring in Greenland. Photo from ZMUC.

Belongs within: Bilateria.
Contains: Bryozoa, Entoprocta, Platyhelminthes, Gastrotricha, Gnathostomulida, Syndermata, Nemertea, Annelida, Brachiozoa, Mollusca.

The Lophotrochozoa is a major clade of animals identified by molecular analyses, named to indicate the association within it of animals bearing a lophophore (bryozoans and brachiozoans) and those with a trochophore-like ciliated larva (annelids and molluscs). Those animals that, at least ancestrally, possess a trochophore-like larva are placed in the clade Trochozoa; as well as the annelids and molluscs, this clade is usually regarded as including the nemerteans and brachiozoans. However, most recent analyses have not supported a direct relationship between the lophophorate brachiozoans and bryozoans; instead, the bryozoans may be placed with the Entoprocta in a clade Polyzoa outside the trochozoans. Molecular analyses have also suppored the association of Platyhelminthes (flatworms) and Syndermata (rotifers and acanthocephalans) with the Lophotrochozoa with at least some analyses placing these two taxa within a clade Platyzoa whose members differ from most other lophotrochozoans in lacking a developed coelom, being either pseudocoelomate or acoelomate.

<==Lophotrochozoa [Acoelomata, Lophozoa, Lophophorata, Molluscoidea, Parenchymia, Procoelomata, Prosomastigozoa]
    |  i. s.: Phlogites [incl. Cheungkongella Shu, Chen et al. 2001] CH03
    |           `--‘*Cheungkongella’ ancestralis Shu, Chen et al. 2001 SC01
    |         Cambrotentacus Zhang, Shu et al. 2001 ZS01
    |           `--*C. sanwuia Zhang, Shu et al. 2001 ZS01
    |--+--Polyzoa HO09
    |  |    |--Bryozoa HO09
    |  |    `--+--Entoprocta HO09
    |  |       `--Symbion [Cycliophora, Symbiidae] HO09
    |  |            |--S. americanus GO06
    |  |            `--S. pandora HO09
    |  `--Platyzoa [Acanthognatha, Gnathifera, Neotrichozoa, Monokonta, Plathelminthomorpha] ZHT01
    |       |--+--Platyhelminthes HO09
    |       |  `--+--Gastrotricha HO09
    |       |     `--Gnathostomulida HO09
    |       `--+--Syndermata HO09
    |          `--Limnognathia Kristensen & Funch 2000 [Limnognathida, Limnognathiidae, Micrognathozoa] KF00
    |               `--*L. maerski Kristensen & Funch 2000 KF00
    `--Trochozoa (see below for synonymy) HO09
         |  i. s.: Lobatocerebrum [Lobatocerebridae, Lobatocerebromorpha] ZHT01
         |           `--L. psammicola Rieger 1980 ZHT01
         |         Miskoia Walcott 1911 [Miskoidae, Miskoiidae] H62
         |           `--*M. preciosa Walcott 1911 H62
         |         Maotianchaeta EL11
         |--Nemertea EL11
         `--+--+--Phragmochaeta canicularis PTV14
            |  `--+--Burgessochaeta setigera (Walcott 1911) PTV14, E-J04 [=Canadia setigera E-J04]
            |     |--Canadia Walcott 1911 PTV14, H62 [Canadidae, Canadiidae]
            |     |    `--*C. spinosa Walcott 1911 H62
            |     `--+--Pygocirrus butyricampum PTV14
            |        `--Annelida EL11
            `--+--Brachiozoa EL11
               `--+--Kimberellomorpha EL11
                  |    |--Solza margarita EL11
                  |    `--Kimberella Wade 1972 CS06, G79 [=Kimberia Glaessner & Wade 1966 non Cotton & Woods 1935 G79]
                  |         `--*K. quadrata (Glaessner & Wade 1966) [=*Kimberia quadrata] G79
                  `--+--Odontogriphidae RE01
                     |    |--Odontogriphus omalus Conway Morris 1976 VVRB08, RE01
                     |    `--Bowengriphus Ritchie & Edgecombe 2001 RE01
                     |         `--*B. perphlegis Ritchie & Edgecombe 2001 RE01
                     `--+--Mollusca EL11
                        `--+--Orthrozanclus SB12
                           `--+--Wiwaxiidae [Wiwaxidae] H62
                              |    |--Wiwaxia Walcott 1911 SB12, P08
                              |    |    `--*W. corrugata (Matthew 1899) [=Orthotheca corrugata] H62
                              |    `--Pollingeria Walcott 1911 H62
                              |         `--*P. grandis Walcott 1911 H62
                              `--Coeloscleritophora P08
                                   |  i. s.: Hippopharangites Bengtson in Bengtson et al. 1990 MP10
                                   |           `--H. dailyi Bengtson in Bengtson et al. 1990 P08
                                   |         Sachites Meshkova 1969 MP10
                                   |           `--S. proboscideus Meshkova 1969 P08
                                   |         Siphonoguchites triangularis Qian 1977 P08
                                   |         Thambetolepis delicata Jell 1981 P08
                                   |         Maikhanella Zhegallo in Voronin et al. 1982 P08 (see below for synonymy)
                                   |         Australohalkieria P08
                                   |           |--A. parva (Conway Morris in Bengtson et al. 1990) P08
                                   |           `--A. superstes Porter 2004 P08
                                   |         Halkieria Poulsen 1967 SB12, VN05 [Diplacophora]
                                   |           |--*H. obliqua Poulsen 1967 VN05
                                   |           |--H. evangelista Conway Morris & Peel 1995 VN05
                                   |           |--H. fordi Landing 1991 L95
                                   |           `--H. mira (Qian & Xiao 1984) P08
                                   |         Aurisella Qian & Xiao 1984 (see below for synonymy) MP10
                                   |         Cambrothyra Qian & Zhang 1983 (see below for synonymy) MP10
                                   |           `--*C. ampulliformis Qian & Zhang 1983 (see below for synonymy) MP10
                                   |         Chancelloriella irregularis Demidenko 2000 MP10
                                   |         Eremactis Bengtson & Conway Morris in Bengtson et al. 1990 MP10
                                   |           |--E. conara Bengtson & Conway Morris in Bengtson et al. 1990 MP10
                                   |           `--E. mawsoni Bengtson & Conway Morris in Bengtson et al. 1990 MP10
                                   |         Monospinites Sayutina in Vasil’eva & Sayutina 1988 MP10
                                   |         Dabashanites Chen 1979 MP10
                                   |         Mirusilites He & Yang 1986 MP10
                                   |         Archicladium Qian & Xiao 1984 MP10
                                   |         Drepanochites EL11
                                   |         Zhiiginites EL11
                                   `--Chancelloriidae MP10
                                        |--Chancelloria racemifundis Bengtson in Bengtson et al. 1990 MP10
                                        |--Allonnia Doré & Reid 1965 [incl. Dimidia Jiang in Luo et al. 1982] MP10
                                        `--Archiasterella Sdzuy 1969 MP10
                                             |--A. fletchergryllus Randell et al. 2005 P08
                                             `--A. hirundo Bengtson in Bengtson et al. 1990 P08

Pseudofossils: Cambrothyra miliaria Duan & Cao 1989 MP10
               Situlitesta antiquata Duan & Cao 1989 MP10
               Situlitesta cyclocarpa Duan & Cao 1989 MP10

Aurisella Qian & Xiao 1984 [incl. Ninella Missarzhevsky in Missarzhevsky & Mambetov 1981 nec Gray 1850 nec Malakhova 1975] MP10

Cambrothyra Qian & Zhang 1983 [incl. Acatomus Duan 1986, Clinopa Duan 1986, Cyphinites Duan, Cao & Zhang 1993, Globifructus Geng & Zhang 1987, Horridomus Duan 1986, Hubeitesta Duan 1986, Lapistamnia Duan, Cao & Zhang 1993, Laxicavia Duan, Cao & Zhang 1993, Mirabichitina Yang, He & Deng 1983 (n. n.), Mirabifolliculus Yang & He 1984, Nanjiangochitina Yang, He & Deng 1983 (n. n.), Nanjiangofolliculus Yang & He 1984, Ovitesta Duan 1986, Parahorridomus Duan, Cao & Zhang 1993, Pollofructus Geng & Zhang 1987, Situlitesta Duan 1986, Trymitesta Duan 1986, Turbinella Duan, Cao & Zhang 1993 non Lamarck 1799, Tympanites Duan, Cao & Zhang 1993] MP10

*Cambrothyra ampulliformis Qian & Zhang 1983 [=Cambrotyca (l. c.) ampulliformis; incl. Situlitesta afflata Duan 1986, Hubeitesta amblybasis Duan 1986, Situlitesta baccata Duan 1986, Parahorridomus biarmillaris Duan, Cao & Zhang 1993, Acatomus bursoides Duan 1986, Anterosculum cambricum Duan, Cao & Zhang 1993, Situlitesta cordata Duan 1986, Laxicavia crassa Duan, Cao & Zhang 1993, Cambrothyra cymatoidea Duan 1986, Lax. debilis Duan, Cao & Zhang 1993, Trymitesta declinata Duan 1986, Tr. ellipsoidalis Duan 1986, Clinopa gibberosa Duan 1986, Cambrothyra glandiformis Duan 1986, Trymitesta globularis Zhang in Zhao et al. 1988, Ca. granosa Geng & Zhang 1987, Acatomus lacrimiformis Duan 1986, Tympanites lebericus Duan, Cao & Zhang 1993, Globifructus meloniformis Geng & Zhang 1987, Cyphinites nidificus Duan, Cao & Zhang 1993, Pollofructus nodus Geng & Zhang 1987, Clinopa nucleola Duan 1986, Cyphinites ovalis Duan, Cao & Zhang 1993, Situlitesta rostellata Duan 1986, Globifructus scutatus Geng & Zhang 1987, Lapistamnia septalis Duan, Cao & Zhang 1993, Horridomus stephanoideus Duan 1986, Turbinella trochalis Duan, Cao & Zhang 1993, Ovitesta truncata Duan 1986, Cambrothyra truncata, Ca. vasiformis Duan 1986, H. vemetovatus Duan 1986, Clinopa verruculata Duan 1986, Cl. xihaopingensis Duan 1986] MP10

Lophotrochozoa [Acoelomata, Lophozoa, Lophophorata, Molluscoidea, Parenchymia, Procoelomata, Prosomastigozoa, Trochozoa]

Maikhanella Zhegallo in Voronin et al. 1982 P08 [=Majkhanella (l. c.) BR05; Maikhanellidae, Maikhanellinae, Majkhanellidae]

Trochozoa [Agama, Canadiacea, Coelhelminthes, Conchozoa, Eutrochozoa, Halkierida, Kryptrochozoa, Miskoida, Miskoiida, Neotrochozoa, Vermizoa] HO09

*Type species of generic name indicated

REFERENCES

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[CS06] Caron, J.-B., A. Scheltema, C. Schander & D. Rudkin. 2006. A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale. Nature 442: 159–163.

[CH03] Chen, J.-Y., D.-Y. Huang, Q.-Q. Peng, H.-M. Chi, Z.-Q. Wang & M. Feng. 2003. The first tunicate from the Early Cambrian of South China. Proceedings of the National Academy of Sciences of the USA 100 (14): 8314–8318.

[E-J04] Eibye-Jacobsen, D. 2004. A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale. Lethaia 37: 317–335.

[EL11] Erwin, D. H., M. Laflamme, S. M. Tweedt, E. A. Sperling, D. Pisani & K. J. Peterson. 2011. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334: 1091–1097.

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[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).

[HO09] Hejnol, A., M. Obst, A. Stamatakis, M. Ott, G. W. Rouse, G. D. Edgecombe, P. Martinez, J. Baguña, X. Bailly, U. Jondelius, M. Wiens, W. E. G. Müller, E. Seaver, W. C. Wheeler, M. Q. Martindale, G. Giribet & C. W. Dunn. 2009. Assessing the root of bilaterian animals with scalable phylogenomic methods. Proceedings of the Royal Society of London Series B—Biological Sciences 276 (1677): 4261–4270.

[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.

[KF00] Kristensen, R. M., & P. Funch. 2000. Micrognathozoa: a new class with complicated jaws like those of Rotifera and Gnathostomulida. Journal of Morphology 246: 1–49.

[L95] Landing, E. 1995. Upper Placentian–Branchian series of mainland Nova Scotia (middle–upper Lower Cambrian): faunas, paleoenvironments, and stratigraphic revision. Journal of Paleontology 69 (3): 475–495.

[MP10] Moore, J. L., S. M. Porter, M. Steiner & G. Li. 2010. Cambrothyra ampulliformis, an unusual coeloscleritophoran from the Lower Cambrian of Shaanxi Province, China. Journal of Paleontology 84 (6): 1040–1060.

[PTV14] Parry, L., A. Tanner & J. Vinther. 2014. The origin of annelids. Palaeontology 57 (6): 1091–1103.

Passamaneck, Y., & K. M. Halanych. 2006. Lophotrochozoan phylogeny assessed with LSU and SSU data: evidence of lophophorate polyphyly. Molecular Phylogenetics and Evolution 40 (1): 20–28.

[P08] Porter, S. M. 2008. Skeletal microstructure indicates chancelloriids and halkieriids are closely related. Palaeontology 51 (4): 865–879.

[RE01] Ritchie, A., & G. D. Edgecombe. 2001. An odontogriphid from the Upper Permian of Australia. Palaeontology 44 (5): 861–874.

[SC01] Shu, D.-G., L. Chen, J. Han & X.-L. Zhang. 2001. An Early Cambrian tunicate from China. Nature 411: 472–473.

[SB12] Sutton, M. D., D. E. G. Briggs, D. J. Siveter, D. J. Siveter & J. D. Sigwart. 2012. A Silurian armoured aplacophoran and implications for molluscan phylogeny. Nature 490: 94–97.

[VN05] Vinther, J., & C. Nielsen. 2005. The Early Cambrian Halkieria is a mollusc. Zoologica Scripta 34: 81–89.

[VVRB08] Vinther, J., P. Van Roy & D. E. G. Briggs. 2008. Machaeridians are Palaeozoic armoured annelids. Nature 451 (7175): 185–188.

[ZS01] Zhang, X., D. Shu, Y. Li & J. Han. 2001. New sites of Chenjiang fossils: crucial windows on the Cambrian explosion. Journal of the Geological Society 158: 211–218.

[ZHT01] Zrzavý, J., V. Hypša & D. F. Tietz. 2001. Myzostomida are not annelids: molecular and morphological support for a clade of animals with anterior sperm flagella. Cladistics 17: 170–198.