Metazoa phylogeny

From Palaeos

Choanomonada
`--Metazoa
   `==Porifera (paraphyletic?)
      `==Radiata (paraphyletic?)
         `--Bilateria
            `==Acoelomorpha (paraphyletic?)
               |--Protostomia
               |  |--Ecdysozoa
               |  `--Lophotrochozoa
               `--Deuterostomia

Note that the phylogeny given below differs in some details from that given on the main page for Metazoa. It is almost universally agreed that the presence of tissue systems in Cnidaria, Ctenophora and Bilateria indicates that they form a clade to the exclusion of the less organised sponges (Porifera). These days this clade is generally known as Eumetazoa, but in older textbooks you may find this clade referred to as Metazoa, with sponges excluded as Parazoa.

Within Eumetazoa, Ctenophora have often been regarded as closer to Bilateria than Cnidaria (as shown on the main page) due to the former two having an anterior-posterior body axis and muscle blocks (old classificatory schemes united Cnidaria and Ctenophora as Coelenterata - however, this was based solely on plesiomorphic characters of diploblasty and radial symmetry). In contrast, molecular phylogenies suggest that Ctenophora is further away from Bilateria (as shown below), which would require shared characters between the two to be homoplastic. While this may seem unlikely at first glance, fossil evidence has suggested that Ctenophora may be derived from sessile Ediacaran ancestors, making convergence with Bilateria more likely.

Another notable point of contention is the position of Trichoplax (Placozoa). This simple animal has been thought to be morphologically intermediate between Porifera and Eumetazoa (main page). However, Cavalier-Smith and Chao (2003) found Trichoplax to be nested within Cnidaria in their molecular phylogeny, and it has been suggested that Trichoplax is derived from a neotenous cnidarian larva.

<==Animalia [Diploblastica, Metazoa, Parazoa, Porifera, Radiata, Spongiaria, Vendobionta]
   |  i. s.: Namapoikia Wood, Grotzinger & Dickson 2002
   |           `--*N. rietoogensis Wood, Grotzinger & Dickson 2002
   |         Yaworipora
   |         Labyrinthus
   |         Rosellatana
   |         Limnea luteola
   |         Coleoloides typicalis Walcott 1889
   |         Purella Missarzhevsky 1974 [Purellidae]
   |           `--*P. cristata Missarzhevsky 1974
   |         Maikhanella Zhegallo 1982
   |         Albinaria coerulea
   |         Ventogyrus
   |         Podolimirus
   |         Swartpuntia
   |         Calyptrina
   |         Kimberella
   |         Diagramma labiosum
   |         Neotricula aperta
   |         Tricula
   |         Indoplanorbis exustus
   |         Radix peregra
   |         Uroteuthis noctiluca
   |         Rondeletiola minor
   |         Sepiolina nipponensis
   |         Cleisopus gloriamaris
   |         Nowakia
   |           |--N. barrandei Boucek & Prantl 1959
   |           `--N. otomari
   |         Ainiktozoon
   |         Amiella ornata Walcott 1911
   |         Hirnantia Lamont 1935
   |         Foliomena Havlicek 1952
   |         Caveasphaera costata
   |         Megaclonophycus onustus
   |         Sinocyclocyclicus guizhouensis
   |         Spiralicellula bulbifera
   |         Trimerotropis citrina
   |         Eurytholia Sutton, Holmer & Cherns 2001
   |           |--*E. prattensis Sutton, Holmer & Cherns 2001
   |           `--E. elibata Sutton, Holmer & Cherns 2001
   |         Dimorphoconus granulatus Donovan & Paul 1985
   |         Stromatoporoidea
   |           |--Keega
   |           `--Clathrocoilona spissa
   |         Dinomischus isolatus
   |         Eldonia ludwigi
   |         Spriggina floundersi
   |         Cervulus
   |           |--C. reevesii
   |           `--C. vaginalis
   |         Jennaria pulchra Rieger 1991
   |         Amauroceras ferrugineum
   |         Declinognathodus noduliferus (Ellison & Graves 1941)
   |         Pseudomytiloides dubius
   |         Asbestopluma Norman 1882 [Cladorhizidae]
   |           `--A. lycopodium (Levinsen 1886)
   |         Acarnidae
   |           |--Acarnus
   |           |--Cornulum dubium Hentschel 1912
   |           `--Damiria simplex Keller 1891 [incl. D. simplex fistulata Hentschel 1912]
   |         Actinostromaria
   |         Parastromatopora
   |         Gilletia
   |         Sestromella
   |         Chaetetopsis
   |         Ellipsactinia
   |         Sphaeractinia
   |         Doryderma
   |         Elasmoiera
   |         Tethya Lamarck 1814 [incl. Donatia Nardo 1833; Tethyidae]
   |           |--T. aurantium (Pallas 1766) [incl. T. limski Müller & Zahn 1968]
   |           |--T. clavigera Hentschel 1912
   |           |--T. lyncurium
   |           |--T. maza Selenka 1893
   |           `--‘Donatia’ tylota Hentschel 1912
   |         *Crambe crambe
   |         Ceratoporella nicolsoni
   |         Disyringa dissimilis
   |         Poterion neptuni
   |         Callyspongiidae
   |           |--Callyspongia
   |           |    |--C. plicifera
   |           |    `--C. vaginalis
   |           `--Siphonochalina
   |                |--S. fascigera Hentschel 1912
   |                `--S. siphonella Lévi 1965
   |         Holopsamma turbo Carter 1885 [=Sigmatella turbo]
   |         Plakina [Plakinidae]
   |           |--P. monolopha
   |           `--P. trilopha
   |                |--P. t. trilopha
   |                `--P. t. antarctica von Lendenfeld 1907
   |         Psammoclemma Marshall 1880 [incl. Psammopemma Marshall 1880; Chondropsidae]
   |           |--‘Psammopemma’ densum Marshall 1881 [incl. Holopsamma laevis Carter 1885]
   |           |--‘Psammopemma’ durissimum
   |           |    |--P. d. durissimum
   |           |    `--P. d. grisea Hentschel 1912
   |           `--‘Psammopemma’ porosum Polejaff 1884
   |         Dysidea Johnston 1842 [incl. Spongelia Schmidt 1862; Dysideidae]
   |           |--‘Spongelia’ fragilis
   |           |    |--S. f. fragilis
   |           |    `--S. f. clathrata Hentschel 1912
   |           |--‘Sarcocornea’ nodosa [=Spongelia nodosa]
   |           `--‘Spongelia’ spinifera
   |                |--S. s. spinifera
   |                `--S. s. australis Hentschel 1912
   |         Ochridaspongia
   |         Microcordyla asteriae
   |         Spheciospongia vesparium (Lamarck 1814)
   |         Ircinia Nardo 1833 [incl. Hircinia Nardo 1834]
   |           |--‘Hircinia’ aruensis Hentschel 1912
   |           |--I. felix Duchassaing & Michelotti 1864
   |           |--‘Hircinia’ pinna Hentschel 1912
   |           |--‘Hircinia’ spiculosa Hentschel 1912
   |           `--I. strobilina Lamarck 1816
   |         Stematumenia
   |           |--S. foetida
   |           |--S. strobilina
   |           `--S. variabilis
   |         Astrosclera willeyana
   |         Solenoporaceae [Solenoporeae, Solenoporideae]
   |           |--*Solenopora spongioides Dybowsky 1877
   |           `--Parachaetetes Deninger 1906
   |         Hemectyon ferox
   |         Vauxia gracilenta
   |         Choia carteri
   |         Pirania muricata
   |         Eiffelia globosa
   |         Acanthella [Dictyonellidae]
   |           `--A. euctimena Hentschel 1912
   |         Archaeocyatha
   |         Speophilosoma
   |           |--S. koyama
   |           `--S. tottoriense
   |         Lycocerus suturellus
   |         Araragi enthea
   |         Buergeria buergeri
   |         Parechthistatus gibber
   |         Parapodisma setouchiensis
   |         Arcuphantes
   |           |--A. hibanus
   |           |--A. iharai
   |           |--A. saitoi
   |           `--A. tsurusakii
   |         Cynops pyrrhogaster
   |         ‘Leptophyllus’ Quenstedt 1876
   |         Niphatidae
   |           |--Pachychalina diffusa
   |           |    |--P. d. diffusa
   |           |    `--P. d. affinis Hentschel 1912
   |           `--Gelliodes
   |                |--G. gracilis Hentschel 1912
   |                |--G. hamata Thiele 1903
   |                |--G. macrosigma Hentschel 1912
   |                |--G. obtusa Hentschel 1912
   |                `--G. porosa Thiele 1903
   |         Periphragella [Euretidae]
   |           `--P. antarctica Janussen, Tabachnick & Tendal 2004
   |         Rossellidae
   |           |--Schaudinnia rosea (Fristedt 1887) [incl. S. arctica Schulze 1900]
   |           |--*Trichasterina borealis Schulze 1900
   |           `--Caulophacus
   |                |--C. (Caulodiscus) brandti Janussen, Tabachnick & Tendal 2004
   |                |--C. valdiviae Schulze 1904
   |                `--C. (*Oxydiscus) weddelli Janussen, Tabachnick & Tendal 2004
   |         Raspailiidae
   |         Aplysina [Aplysinidae]
   |           `--A. mollis
   |                |--A. m. mollis
   |                `--A. m. aruensis Hentschel 1912
   |         Desmacellidae
   |           |--Tylodesma microstrongyla Hentschel 1912
   |           `--Biemna
   |                |--B. aruensis Hentschel 1912
   |                |--B. humilis Thiele 1903
   |                |--B. megalosigma Hentschel 1912
   |                |    |--B. m. megalosigma
   |                |    `--B. m. liposphaera Hentschel 1912
   |                `--B. truncata Hentschel 1912
   |         Terpios fugax Duchassaing & Michelotti 1864
   |         Petrosia
   |         Ancorinidae
   |         Iotrochota
   |           |--I. baculifera Ridley 1884
   |           `--I. purpurea (Bowerbank 1875)
   |         Cerbaris Topsent 1898 [incl. Rhabdoploca Topsent 1904; Bubaridae]
   |           `--‘Rhabdoploca’ topsenti Hentschel 1912
   |         Darwinellidae
   |           |--Darwinella australiensis Carter 1885
   |           `--Dendrilla
   |                |--D. lacunosa Hentschel 1912
   |                |--D. lendenfeldi Hentschel 1912
   |                `--D. mertoni Hentschel 1912
   |         Thorectidae
   |           |--Aplysinopsis reticulata Hentschel 1912
   |           |--Hyrtios Duchassaing & Michelotti 1864 [incl. Dysideopsis von Lendenfeld 1888]
   |           |    `--‘Dysideopsis’ topsenti Hentschel 1912
   |           `--Fasciospongia Burton 1934 [incl. Stelospongia Schulze 1879 (nom. inv.)]
   |                |--‘Stelospongia’ cava Hentschel 1912
   |                `--‘Stelospongia’ euplectella Hentschel 1912
   |         Monorhaphis [Monorhaphididae]
   |           `--*M. chuni Schulze 1904 [incl. M. dives Schulze 1904]
   |         Pachastrellidae
   |           |--Pachastrella
   |           |    |--P. chuni von Lendenfeld 1907
   |           |    `--P. tenuipilosa von Lendenfeld 1907
   |           `--Thenea
   |                |--T. megaspina von Lendenfeld 1907
   |                |--T. multiformis von Lendenfeld 1907
   |                |--T. pendula von Lendenfeld 1907
   |                `--T. valdiviae von Lendenfeld 1907
   |         Theonellidae
   |           |--Discodermia claviformis Kieschnick 1896 (n. d.)
   |           `--Theonella
   |                |--T. conica (Kieschnick 1896) [=Discodermia conica]
   |                |--T. incerta Thiele 1900
   |                `--T. lacerata von Lendenfeld 1907
   |         Chondrillidae
   |           |--Chondrosia
   |           |    |--C. corticata Thiele 1900
   |           |    `--C. debilis Thiele 1900
   |           `--Chondrilla
   |                |--C. australiensis Carter 1873
   |                |--C. grandistellata Thiele 1900
   |                |--C. jinensis Hentschel 1912
   |                |--C. media Hentschel 1912
   |                `--C. ternatensis Thiele 1900
   |         Hymedesmiidae
   |           |--Hymedesmia mertoni Hentschel 1912
   |           `--Hamigera [incl. Amoibodictya]
   |                `--*H. hamigera [incl. *Amoibodictya forsteri Zahn, Müller & Müller 1977]
   |         Desmanthus [incl. Lophacanthus non Stock 1880; Desmanthidae]
   |           |--D. rhabdophorus (Hentschel 1912) [=*Lophacanthus rhabdophorus]
   |           `--D. topsenti Hentschel 1912
   |         Placospongia [Placospongiidae]
   |           `--P. mixta Thiele 1900
   |         Dendropsis [Desmoxyidae]
   |           `--D. mixta Hentschel 1912
   |         Scleritoderma [Scleritodermatidae]
   |           `--S. nodosum Thiele 1900
   |         *Platylistrum [Pheronematidae]
   |           `--P. platessa Schulze 1904
   |         Holascus [Euplectellidae]
   |           `--H. pseudostellatus Janussen, Tabachnick & Tendal 2004
   |         Timea [Timeidae]
   |           `--T. tetractis Hentschel 1912
   |         Sarostegia [incl. Ramella; Farreidae]
   |           `--S. occulata Topsent 1904 [incl. *Ramella tubulosa Schulze 1904]
   |         Paracharnia dengyingensis
   |         Sinotubulites baimatuoensis
   |         Cloudina hartmannae
   |         Pararenicola Wang 1982 [incl. Paleorhynchus Wang 1982]
   |           `--*P. huaiyuanensis Wang 1982 (see below for synonymy)
   |         Protoarenicola Wang 1982
   |           `--*P. baiguashanensis Wang 1982
   |         Gunnarites antarcticus
   |         Oscarella carmela
   |         Clausocalanus laticeps
   |--+--Calcarea
   |  `--Eumetazoa [Agnotozoa, Coelenterata]
   |       |--Trichoplax [Placozoa]
   |       |    `--T. adhaerens
   |       `--+--Ctenophora
   |          `--+--Cnidaria
   |             `--Bilateria
   `--Hyalospongiae
        |  i. s.: Microcionidae
        |--Demospongiae
        `--Hexactinellida
             |--Oopsacas minuta
             |--Rhabdocalyptus dawsoni
             |--Semperella schulzei
             `--Hyalonema [Hyalonematidae]
                  |--H. hozawai
                  |--H. molle Schulze 1904
                  |--H. proximum Schulze 1904
                  |--H. simile Schulze 1904
                  `--H. valdiviae Schulze 1904

*Pararenicola huaiyuanensis Wang 1982 [incl. Paleorhynchus anhuiensis Wang 1982, Ruedemannella minuta Wang 1982, Paleolina tortuosa Wang 1982]

* Type species of generic name indicated

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[edit] Credits

introduction CKT061026; phylogeny CKT080404