Field of Science

Tubificida

Section of genitalic segments of Phreodrilus linnaei, male genitalia to the left and spermathecal duct to the right, from Pinder (2008).


Belongs within: Clitellata.
Contains: Naididae, Enchytraeidae, Haplotaxidae, Phallodrilinae.

The Tubificida are a group of small, mostly aquatic oligochaetes that include many of the best-known freshwater worms.

Characters (from Jamieson 1988): One pair of testes followed by one pair of ovaries; male pores anterior to female pores; male ducts penetrating one intersegment. Sperm with no connectives; acrosome very short.

Tubificida [Tubificina] J88
    |  i. s.: Dorydrilus Piguet 1913 JD86 [Dorydrilidae J88]
    |           |--D. michaelseni Piguet 1913 JD86
    |           |--D. mirabilis Hrabĕ 1936 JD86
    |           `--D. tetrathecus Hrabĕ 1960 JD86
    |         Naididae J88
    |         Opistocystidae J88
    |--Phreodrilidae RP07
    |    |--Phreodrilus Beddard 1891 P08 [incl. Hesperodrilus J31]
    |    |    |--*P. subterraneus Beddard 1891 P08
    |    |    |--P. albus Beddard 1894 G09b
    |    |    |--P. beddardi Benham 1904 G09b
    |    |    |--P. branchiatus Beddard 1894 G09b
    |    |    |--P. kerguelenensis Michaelsen 1902 G09b
    |    |    |--P. linnaei Pinder 2008 P08
    |    |    |--P. mauiensis Benham 1904 G09b
    |    |    |--P. niger Beddard 1904 G09b
    |    |    |--P. pellucidus Beddard 1894 G09b
    |    |    `--P. peniculus Pinder 2003 P08
    |    |--Phreodriloides Benham 1907 G09b
    |    |    `--P. notabilis Benham 1907 ED08
    |    |--Astacopsidrilus Goddard 1909 G09a
    |    |    |--A. edwardi Pinder 2003 P08
    |    |    |--A. fusiformis Goddard 1909 G09a
    |    |    |--A. notabilis Goddard 1909 G09a
    |    |    `--A. novus Jackson 1931 J31
    |    |--Insulodrilus Brnkhurst 1991 P08
    |    |    |--*I. lacustris (Benham 1903) [=Phreodrilus lacustris] P08
    |    |    |--I. angela Pinder 2008 P08
    |    |    |--I. genitalisetifera Martin & Brinkhurst 1994 P08
    |    |    |--I. litoralis Michaelsen 1903 P08
    |    |    `--I. martensi Martin & Giani 1995 P08
    |    `--Antarctodrilus proboscidea (Brinkhurst & Fulton 1979) RP07
    `--+--+--Enchytraeidae RP07
       |  `--+--Haplotaxidae RP07
       |     `--Propappus Michaelsen 1905 RP07, D90 [Propappidae]
       |          `--P. volki Michaelsen 1916 RP07
       `--Tubificidae RP07
            |  i. s.: Tubifex Lamarck 1816 [=Tubifer (l. c.) non Piette 1856] BR05
            |           |--T. davidis Benham 1907 ED08
            |           |--T. flabellisetosus Hrabĕ 1966 JD86
            |           |--T. ignotus KA01
            |           `--T. tubifex KA01
            |         Rhyacodrilus Bretscher 1901 JD86
            |           |--R. amphigenus JD86
            |           |--R. balmensis Juget 1959 JD86
            |           |--R. carsticus Kosel 1980 JD86
            |           |--R. lindbergi Hrabĕ 1963 JD86
            |           |--R. phreaticola JD86
            |           |--R. sketi Karaman 1974 JD86
            |           `--R. subterraneus Hrabĕ 1963 JD86
            |         Psammoryctides Hrabĕ 1964 JD86
            |           |--P. hadzii Karaman 1974 JD86
            |           `--P. hrabei Karaman 1971 JD86
            |         Peloscolex Leidy 1851 JD86
            |           |--P. arganoi Pop 1974 JD86
            |           |--P. boitanii Pop 1974 JD86
            |           |--P. cottarellii Pop 1974 JD86
            |           |--P. latens JD86
            |           `--P. pescei Dumnicka 1981 JD86
            |         Haber Holmquist 1978 JD86
            |           |--H. monfalconensis Hrabĕ 1966 JD86
            |           |--H. turquini Juget & Lafont 1979 JD86
            |           `--H. zavreli Hrabĕ 1942 JD86
            |         Potamothrix Vejdovsky & Mrazek 1902 JD86
            |           |--P. moldaviensis JD86
            |           |    |--P. m. moldaviensis JD86
            |           |    `--P. m. minimus Pop 1971 JD86
            |           `--P. postojnae Karaman 1974 JD86
            |         Epirodrilus Hrabĕ 1930 JD86
            |           `--E. slovenicus Karaman 1976 JD86
            |         Spirospermoides Dumnicka 1983 JD86
            |           `--*S. stocki Dumnicka 1983 JD86
            |         Krenedrilus Dumnicka 1983 JD86
            |           `--*K. papillatus Dumnicka 1983 JD86
            |         Marcusaedrilus P86
            |         Smithsonidrilus P86
            |         Clitellio abjornseni J31
            |         Macquaridrilus bennettae J88
            |         Limnodrilus hoffmeisteri S30, KA01
            |         Heronidrilus gravidus Erséus 1990 RP07
            |         Heterochaeta costata Claparède 1863 RP07
            |         Tubificoides amplivasatus (Erséus 1975) RP07
            |         Pronaidites Kušta 1888 H62
            |           `--*P. carbonarius Kušta 1888 H62
            |--Phallodrilinae P86
            `--Aulodrilinae P86
                 |--Kaketio P86
                 |--Thalassodrilides P86
                 `--Limnodriloides Pierantoni 1904 P86
                      `--L. winckelmanni Michaelsen 1914 P86

*Type species of generic name indicated

REFERENCES

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[D90] Dash, M. C. 1990. Oligochaeta: Enchytraeidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 311–340. John Wiley & Sones: New York.

[ED08] Edgeworth David, T. W. 1908. Geological notes on Kosciusko, with special reference to evidences of glacial action. Part II. Proceedings of the Linnean Society of New South Wales 33: 657–668, pl. 623.

[G09a] Goddard, E. J. 1909a. Contribution to a further knowledge of Australasian Oligochaeta. Part I. Descriptions of two species of a new genus of Phreodrilidae. Proceedings of the Linnean Society of New South Wales 33: 768–793, pls 729–731.

[G09b] Goddard, E. J. 1909b. Contribution to a further knowledge of Australian Oligochaeta. Part II. Description of a Tasmanian phreodrilid. Proceedings of the Linnean Society of New South Wales 33: 845–853.

[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.

[J31] Jackson, A. 1931. The Oligochaeta of south-western Australia. Journal of the Royal Society of Western Australia 17: 71–136.

[J88] Jamieson, B. G. M. 1988. On the phylogeny and higher classification of the Oligochaeta. Cladistics 4: 367–402.

[JD86] Juget, J., & E. Dumnicka. 1986. Oligochaeta (incl. Aphanoneura) des eaux souterraines continentales. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 234–244. E. J. Brill/Dr W. Backhuys: Leiden.

[KA01] Kent, M. L., K. B. Andree, J. L. Bartholomew, M. El-Matbouli, S. S. Desser, R. H. Devlin, S. W. Feist, R. P. Hedrick, R. W. Hoffmann, J. Khattra, S. L. Hallett, R. J. G. Lester, M. Longshaw, O. Palenzuela, M. E. Siddall & C. Xiao. 2001. Recent advances in our knowledge of the Myxozoa. Journal of Eukaryotic Microbiology 48: 395–413.

[P86] Pfannkuche, O. 1986. Oligochaeta from marine and brackish-water subterranean habitats. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 245–249. E. J. Brill/Dr W. Backhuys: Leiden.

[P08] Pinder, A. M. 2008. Phreodrilidae (Clitellata: Annelida) in north-western Australia with descriptions of two new species. Records of the Western Australian Museum 24 (4): 459–468.

[RP07] Rousset, V., F. Pleijel, G. W. Rouse, C. Erséus & M. E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 23: 41–63.

[S30] Stephenson, J. 1930. Oligochaeta collected by Dr. G. S. Carter and Mr. L. C. Beadle on the expedition to Brazil and Paraguay in 1926–27. Proceedings of the Linnean Society of London 141: 19–20.

Cheirolepidiaceae

Frenelopsis hoheneggeri, from the Peabody Museum of Natural History.


Belongs within: Pinopsida.

The Cheirolepidiaceae are a group of fossil conifers known from the Mesozoic. They are characterised by a distinct pollen type (assigned to the form genus Classopollis) with a complex, tectate-columellate wall structure (Axsmith et al. 2004). Shoots were often opposite-decussate or spiral, and leaves (which may be dimorphic) were typically reduced or scale-like, and imbricate (Pole 2000).

<==Cheirolepidiaceae [Frenelopsidaceae]
    |--Alvinia bohemica AKW04
    |--Tomaxellia biforme P00
    |--Otwayia Pole 2000 P00
    |    |--*O. tetragona (Cantrill & Douglas) Pole 2000 [=Geinitzea tetragona Cantrill & Douglas 1988] P00
    |    `--O. cudgeloides Pole 2000 P00
    |--Classostrobus Alvin, Spicer & Watson 1978 AKW04 [incl. Pseudofrenelopsis AKW04, Suturovagina GM-C02]
    |    |--*C. comptonensis Alvin, Spicer & Watson 1994 AKW04 (see below for synonymy)
    |    |--C. arkansensis Axsmith, Krings & Waselkov 2004 AKW04
    |    |--C. cathayanus [incl. Pseudofrenelopsis intermedia] AKW04
    |    |--C. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
    |    `--C. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
    |--Classopollis Pflug 1953 AKW04
    |    |--*C. classoides Pflug 1953 GM-C02
    |    |--C. brasiliensis Herngreen 1975 I02
    |    |--C. mirabilis Reyre 1970 GM-C02
    |    |--C. noeli Reyre 1970 GM-C02
    |    `--C. obidosensis Groot & Groot 1962 GM-C02
    |--Cupressinocladus Seward 1919 AKW04 [incl. Watsoniocladus GM-C02]
    |    |--C. asferensis Barale & Ouaja 2002 BO02
    |    |--C. itieri (Saporta) Barale 1981 BO02
    |    |--C. leptocladoides (Berry) Pons 1990 BO02
    |    |--C. micromerum (Heer) Pais 1974 BO02
    |    |--C. pompeckji (Salfeld) Pons 1990 BO02
    |    |--C. pseudoexpansum Barnard & Miller 1976 BO02
    |    `--C. soltanensis Barale, Ouaja & Philippe 2000 B)2
    `--Frenelopsis Schenck 1869 AKW04, GM-C02
         |--*F. hoheneggeri (Ettingshausen) Schenck 1869 [=Thuites hoheneggeri Ettinghausen 1852] GM-C02
         |--F. alata Feistmantel 1881 AKW04
         |--F. choshiensis GM-C02
         |--F. elegans GM-C02
         |--F. harrisii Doludenko & Reymanówna 1978 GT02
         |--F. kaneviensis Barale & Doludenko 1985 GM-C02
         |--F. occidentalis GM-C02
         |--F. oligostomata AKW04 [incl. Classoidites glandis GM-C02]
         |--F. pombetsuensis Saiki 1997 GM-C02
         |--F. ramosissima GM-C02
         |--F. rubiesensis Barale 1973 GM-C02
         |--F. sifloana Watson 1983 GM-C02
         |--F. teixeirae GM-C02
         |--F. turolensis Gomez, Martín-Closas et al. 2002 GM-C02
         |--F. ugnaensis Gomez, Martín-Closas et al. 2002 GM-C02
         `--F. veneta Gomez, Thévenard et al. 2002 GT02

*Classostrobus comptonensis Alvin, Spicer & Watson 1994 AKW04 [incl. Classopollis martinottii GM-C02, Pseudofrenelopsis parceramosa (Fontaine) Watson 1977 AKW04]

*Type species of generic name indicated

REFERENCES

[AKW04] Axsmith, B. J., M. Krings & K. Waselkov. 2004. Conifer pollen cones from the Cretaceous of Arkansas: implications for diversity and reproduction in the Cheirolepidiaceae. Journal of Paleontology 78: 402–409.

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

[GM-C02] Gomez, B., C. Martín-Closas, G. Barale, N. Solé de Porta, F. Thévenard & G. Guignard. 2002. Frenelopsis (Coniferales: Cheirolepidiaceae) and related male organ genera from the Lower Cretaceous of Spain. Palaeontology 45 (5): 997–1036.

[GT02] Gomez, B., F. Thévenard, M. Fantin & L. Guisberti. 2002. Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy. Cretaceous Research 23: 671–685.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775–788.

[P00] Pole, M. S. 2000. Mid-Cretaceous conifers from the Eromanga Basin, Australia. Australian Systematic Botany 13 (2): 153–197.

Myzostomatida

Myzostoma cirriferum, from Christoph Bleidorn.


Belongs within: Aciculata.

The Myzostomatida are a group of small marine worms that live in close association with echinoderms, either as commensals or parasites (Lanterbecq et al. 2006).

Characters (from Lanterbecq et al. 2006): Anterior introvert present. Trunk with five pairs of conical locomotory organs (parapodia) with distal hook-shaped chaeta and basal parapodial fold, alternating with four pairs of lateral sense organs; parapodia and lateral organs may be reduced or lost in parasitic forms.

Myzostomatida [Endomyzostomatidae, Myzostomida]
    |  i. s.: Stelechopus [Stelechopidae] LR06
    |           `--S. hyocrini LR06
    |         Mycomyzostoma calcidicola LR06
    |--Pharyngidea LR06
    |    |--Pulvinomyzostomum [Pulvinomyzostomatidae] LR06
    |    |    `--P. pulvinar (Graff 1884) LR06 [=Myzostoma pulvinar BK77]
    |    `--Endomyzostoma LR06
    |         |--E. cysticolum (Graff 1883) LR06
    |         `--+--E. deformator (Graff 1884) LR06
    |            `--+--E. clarki (McClendon 1906) LR06
    |               `--E. tenuispinum (Graff 1884) LR06
    `--Myzostoma LR06 [=Myzostomum Leuckart 1827 BK77, H62; Myzostomatidae]
         |  i. s.: *Myzostomum’ costatum Leuckart 1827 H62
         |         M. agassizii von Graff 1883 C50
         |         ‘Myzostomum’ brevipes C50
         |         M. carpenteri [=Myzostomum carpenteri] BK77
         |         M. fimbriatum [=Myzostomum fimbriatum] BK77
         |         M. giganteum [=Myzostomum giganteum] BK77
         |         M. gigas [=Myzostomum gigas] BK77
         |         M. graffi [=Myzostomum graffi] BK77
         |         ‘Myzostomum’ parasiticum [incl. Myzostoma tuberculosum Semper 1858] BK77
         |         ‘Myzostomum’ pictum von Graff 1884 C50
         |         M. seymourcollegiorum DH08
         |--+--M. cirriferum (Leuckart 1827) LR06 (see below for synonymy)
         |  `--+--M. alatum (Graff 1884) LR06 [=Myzostomum alatum BK77]
         |     `--M. glabrum (Leuckart 1842) LR06
         `--+--+--+--Notopharyngoides aruensis (Remscheid 1918) LR06
            |  |  `--+--M. cuniculus (Eeckhaut, Grygier & Deheyn 1998) LR06
            |  |     `--M. pseudocuniculus (Lanterbecq & Eeckhaut 2003) LR06
            |  `--+--Contramyzostoma sphaera (Eeckhaut, Grygier & Deheyn 1998) LR06
            |     `--+--M. capitocutis (Eeckhaut, VandenSpiegel & Grygier 1994) LR06
            |        `--+--M. coriaceum (Graff 1884) LR06
            |           `--M. nigromaculatum (Eeckhaut, Grygier & Deheyn 1998) LR06
            `--+--+--+--M. polycyclus (Atkins 1927) LR06
               |  |  `--M. toliarense (Lanterbecq & Eeckhaut 2003) LR06
               |  `--+--M. ambiguum (Graff 1887) LR06
               |     `--+--M. mortenseni (Jägersten 1842) LR06
               |        `--+--Asteriomyzostomum [Asteriomyzostomatidae] LR06
               |           `--Asteromyzostomum [Asteromyzostomatidae] LR06
               `--+--+--+--M. fissum (Graff 1884) LR06
                  |  |  `--M. furcatum (Graff 1887) LR06
                  |  `--Hypomyzostoma LR06
                  |       |--H. crosslandi (Boulenger 1913) LR06
                  |       |--H. dodecephalcis (Grygier 1992) LR06
                  |       `--H. fasciatum (Remscheid 1918) LR06
                  `--+--M. laingense (Eeckhaut, Grygier & Deheyn 1998) LR06
                     `--Mesomyzostoma [incl. Protomyzostomum; Mesomyzostomatidae, Protomyzostomatidae] LR06
                          |  i. s.: ‘Protomyzostomum’ sagamiense BK77
                          |--‘Protomyzostomum’ polynephris (Fedotov 1912) LR06 [=Protomyzostoma polynephris BK77]
                          `--+--M. reichenspergi (Remscheid 1918) LR06
                             `--+--‘Protomyzostomum’ glanduliferum (Bartsch 1995) LR06
                                `--M. katoi (Okada 1933) LR06

Myzostoma cirriferum (Leuckart 1827) LR06 [=Myxostoma (l. c.) cirriferum BK77, Myzostomum cirriferum BK77; incl. Myzostomum schultzeanum Diesing 1858 BK77, Cyclocirra thompsonii Müller 1841 BK77, Myzostomum thompsoni BK77]

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[C50] Clark, A. H. 1950. A monograph of the existing crinoids. Vol. 1. The comatulids. Part 4c. Superfamily Tropiometrida (the families Thalassometridae and Charitometridae). Bulletin of the United States National Museum 82 vol. 1 (4c): 1–383.

[DH08] Dunn, C. W., A. Hejnol, D. Q. Matus, K. Pang, W. E. Browne, S. A. Smith, E. Seaver, G. W. Rouse, M. Obst, G. D. Edgecombe, M. V. Sørensen, S. H. D. Haddock, A. Schmidt-Rhaesa, A. Okusu, R. M. Kristensen, W. C. Wheeler, M. Q. Martindale & G. Giribet. 2008. Broad phylogenomic sampling improves resolution of the animal tree of life. Nature 452: 745–749.

[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.

[LR06] Lanterbecq, D., G. W. Rouse, M. C. Milinkovitch & I. Eeckhaut. 2006. Molecular phylogenetic analyses indicate multiple independent emergences of parasitism in Myzostomida (Protostomia). Systematic Biology 55 (2): 208–227.

Deuterostomia

Specimen of Vetulicola cuneata (scale bar 5 mm), with close-up of lateral groove structure (scale bar 1 mm), from Donoghue & Purnell (2009).


Belongs within: Bilateria.
Contains: Hemichordata, Echinodermata, Chordata.

The Deuterostomia are a group of animals characterised by features of embryonic development including radial cleavage, and development of the blastopore into the anus. Possible synapomorphies of this group (albeit lost or modified in many subgroups) also include the presence of gill slits and the endostyle (Shu et al. 2001).

Deuterostomia [Cristozoa, Dexiothetica]
    |--Vetulicolia SCM04
    |    |  i. s.: Yuyuanozoon BL05
    |    |--Heteromorphida EL11
    |    |    |--Heteromorphus EL11
    |    |    `--Banffia Walcott 1911 EL11, H62
    |    |         |--*B. constricta Walcott 1911 H62
    |    |         `--B. confusa Chen & Zhou 1997 BL05
    |    |--Yunnanozoa [Yunnanozoida, Yunnanozoonidae] SCM03
    |    |    |--*Yunnanozoon lividum Hou et al. 1991 SCM03, D95
    |    |    `--Haikouella Chen, Huang & Li 1999 SCM03
    |    |         |--*H. lanceolata SCM03
    |    |         `--H. jianshanensis Shu, Conway Morris et al. 2003 SCM03
    |    `--Vetulicolida BL05
    |         |--Pomatrum ventralis SCM01
    |         |--Xidazoon Shu, Conway Morris & Zhang in Shu, Conway Morris et al. 1999 SCM01, SCM99
    |         |    `--*X. stephanus Shu, Conway Morris & Zhang in Shu, Conway Morris et al. 1999 SCM99
    |         |--Didazoon Shu & Han in Shu, Conway Morris et al. 2001 SCM01
    |         |    `--*D. haoae Shu & Han in Shu, Conway Morris et al. 2001 SCM01
    |         |--Skeemella Briggs, Lieberman et al. 2005 BL05
    |         |    `--*S. clavula Briggs, Lieberman et al. 2005 BL05
    |         `--Vetulicola BL05
    |              |--V. cuneata Hou 1987 [=V. cuneatus] BL05
    |              `--V. rectangulata SCM01
    |--+--Eldonia ludwigi EL11, CM98
    |  `--Herpetogaster EL11
    `--+--Ambulacraria [Coelomopora] SCM04
       |    |--Hemichordata SCM04
       |    `--+--Echinodermata SCM04
       |       `--Vetulocystidae [Vetulocystida] SCM04
       |            |--Vetulocystis Shu, Conway Morris et al. 2004 SCM04
       |            |    `--*V. catenata Shu, Conway Morris et al. 2004 SCM04
       |            `--Dianchicystis Shu, Conway Morris et al. 2004 SCM04
       |                 `--*D. jianshanensis Shu, Conway Morris et al. 2004 SCM04
       `--Chordata SCM04

*Type species of generic name indicated

REFERENCES

[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.

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

[D95] Dzik, J. 1995. Yunnanozoon and the ancestry of chordates. Acta Palaeontologica Polonica 40: 341–360.

[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.

[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.

[SCM99] Shu, D., S. Conway Morris, X.-L. Zhang, L. Chen, Y. Li & J. Han. 1999. A pipiscid-like fossil from the Lower Cambrian of south China. Nature 400: 746–749.

[SCM01] Shu, D.-G., S. Conway Morris, J. Han, L. Chen, X.-L. Zhang, Z.-F. Zhang, H.-Q. Liu, Y. Li & J.-N. Liu. 2001. Primitive deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China). Nature 414: 419–424.

[SCM04] Shu, D.-G., S. Conway Morris, J. Han, Z.-F. Zhang & J.-N. Liu. 2004. Ancestral echinoderms from the Chengjiang deposits of China. Nature 430: 422–428.

[SCM03] Shu, D., S. Conway Morris, Z. F. Zhang, J. N. Liu, J. Han, L. Chen, X. L. Zhang, K. Yasui & Y. Li. 2003. A new species of yunnanozoan with implications for deuterostome evolution. Science 299: 1380–1384.

Rhabdocoela

Dalyellia viridis, from the Vernal Pool Association.


Belongs within: Neoophora.
Contains: Typhloplanoida.

The Rhabdocoela are a group of flatworms with a straight, simple gut and an anterior mouth.

Rhabdocoela [Eleucithophora]
    `--Neorhabdocoela S86
         |  i. s.: Mesostoma C81
         |           |--M. californicum C81
         |           `--M. craci C81
         |--Typhloplanoida S86
         |--Dalyellida [Dalyellioida] S86
         |    |--Balgetia Luther 1962 S86
         |    |    `--B. papii Kolasa 1976 S86
         |    |--Provortex LOL03
         |    |    |--P. psammophilus LOL03
         |    |    `--P. tubiferus LOL03
         |    |--Pterastericola LOL03
         |    |    |--P. australis LOL03
         |    |    `--P. fedotovi BK77
         |    `--Dalyellia S86
         |         |--D. rossi H62
         |         |--D. tatrica S86
         |         `--D. viridis H62
         `--Kalyptorhynchia S86
              |--Macrorhynchus crocea (Fabricius 1826) S03
              |--Amphirhynchus caudatus K92
              |--Cheliplana boadeni K92
              |--Cicerina brevicirrus K92
              |--Mesorhynchus terminostylus K92
              |--Proschizorhynchus gullmarensis K92
              |--Schizochilus choriurus K92
              |--Gnathorhynchus Meixner 1929 S86
              |    `--G. aprosmeno An der Lan 1967 S86
              |--Uncinorhynchus Karling 1947 S86
              |    `--U. karlingi Kolasa 1977 S86
              `--Ethmorhynchus Meixner 1938 S86
                   |--E. anophthalmus K92
                   `--E. youngi Kolasa 1977 S86

Rhabdocoela incertae sedis:
  Bothromesostoma GD00
  Mesocastrada GD00
  Anoplodium H01
    |--A. chiridotae BK77
    |--A. clypeastris BK77
    |--A. gracile BK77
    |--A. myriotrochi BK77
    |--A. stichopi BK77
    `--A. tubiferum BK77
  Astrorhynchus H01
  Cheliplanilla H01
  Jensenia H01
  Nygulgus H01
  Promesostoma H01
  Temnocephalida LOL03
    |--Didymorchis LOL03
    `--Temnosewellia minor LOL03
  Umagilla forskalensis BK77
  Wahlia macrostylifera BK77
  Anoplodiopsis gracilis BK77
  Anoplodiera voluta BK77
  Ozametra elegans (Westblad 1926) [=Umagilla elegans] BK77
  Syndesmis echinorum BK77
  Marcusella BK77
    |--M. atriovillosa BK77
    `--M. pallida BK77
  Desmote vorax BK77
  Triloborhynchus astropectinis BK77

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[C81] Collins, F. H. 1981. Predation as a major component of mortality among immature mosquitoes in rice fields, flooded pastures, and vernal pools in northern California. Dissertation Abstracts International B 41: 4375.

[GD00] Giribet, G., D. L. Distel, M. Polz, W. Sterrer & W. C. Wheeler. 2000. Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: a combined approach of 18S rDNA sequences and morphology. Systematic Biology 49: 539–562.

[H62] Hass, W. H. 1962. Conodonts. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W3–W69. Geological Society of America, and University of Kansas Press.

[H01] Hooge, M. D. 2001. Evolution of body-wall musculature in the Platyhelminthes (Acoelomorpha, Catenulida, Rhabditophora). Journal of Morphology 249: 171–194.

[K92] Kozloff, E. N. 1992. The genera of the phylum Orthonectida. Cahiers de Biologie Marine 33: 377–406.

[LOL03] Lockyer, A. E., P. D. Olson & D. T. J. Littlewood. 2003. Utility of complete large and small subunit rRNA genes in resolving the phylogeny of the Neodermata (Platyhelminthes): implications and a review of the cercomer theory. Biological Journal of the Linnean Society 78: 155–171.

[S86] Schwank, P. 1986. Microturbellaria from subterranean freshwater habitats. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 47–56. E. J. Brill / Dr. W. Backhuys: Leiden.

[S03] Slyusarev, G. S. 2003. The fine structure of the muscle system in the female of the orthonectid Intoshia variabilis (Orthonectida). Acta Zoologica 84: 107–111.

Graptolithina

Fragments of Stolonodendrum, from Bulman (1970).


Belongs within: Hemichordata.
Contains: Graptoblasti, Dendroidea, Graptoloida, Wimanicrustidae.

The Graptolithina, graptolites, are a group of colonial organisms known from the Middle Cambrian to the Carboniferous that produced linear, branched or encrusting colonies.

Characters (from Bulman 1970): Colonial organisms secreting a sclerotised exoskeleton with characteristic growth bands (fuselli) and growth lines. Thecae usually arranged in single or double row along branches (stipes) of colony (rhabdosome), rarely in irregular aggregates; thecae may be polymorphic, may be related to internal sclerotised stolon system. Rhabdosomes originating by single bud from initial zooid housed in conical sicula, producing simple, branched or rarely encrusting colonies. Sessile or pelagic.

Graptolithina
    |--+--Dendroidea B70
    |  `--Graptoloida B70
    |--Mastigograptida [Mastigograptidae] BU02
    |    |--Micrograptus Eisenack 1974 BU02
    |    `--Mastigograptus Ruedemann 1908 BU02
    |         |--*M. tenuiramosus (Walcott 1879) [=Dendrograptus tenuiramosus] BU02
    |         `--M. gracillimus (Lesquereux 1878) BU02
    |--Crustoidea B70
    |    |--Hormograptus Öpik 1930 [=Thallograptus Öpik 1928 non Ruedemann 1925; Hormograptidae] B70
    |    |    `--*H. sphaericola (Öpik 1928) [=*Thallograptus sphaericola] B70
    |    `--Wimanicrustidae B70
    `--Stolonoidea B70
         |--Stolonofolliculis Zessin & Puttkamer 1994 [Stolonofolliculidae] MU04
         |    `--*S. signum (Öpik 1930) (n. d.) [=Melanostrophus signum] MU04
         `--Stolonodendrum Kozłowski 1948 [Stolonodendridae] B70
              |--*S. uniramosum Kozłowski 1948 B70
              `--S. parasiticum Kozłowski 1948 K48

Graptolithina incertae sedis:
  Protistograptus McLearn 1915 (n. d.) B70
  Haplograptus Ruedemann 1933 B70
    `--*H. wisconsinensis Ruedemann 1933 B70
  Graptovermis Kozłowski 1949 [Graptovermida] MU04
    |--*G. spiralis Kozłowski 1948 B70
    `--G. intestinalis Kozłowski 1948 K48
  Graptoblasti B70
  Aellograptus Obut 1964 B70
    `--*A. savitskyi Obut 1964 B70
  Archaeocryptolaria Chapman 1919 B70
    `--*A. skeatsi Chapman 1919 B70
  Ascograptus Ruedemann 1925 B70
    `--*A. similis Ruedemann 1925 B70
  Cactograptus Ruedemann 1908 B70
    `--*C. crassus Ruedemann 1908 B70
  Ceramograptus Hudson 1915 B70
    `--*C. ruedemanni Hudson 1915 B70
  Coelograptus Ruedemann 1947 B70
    `--*C. problematicus (Spencer 1878) [=Inocaulis problematica] B70
  Crinocaulis Obut 1960 B70
    `--*C. flosculus Obut 1960 B70
  Diplospirograptus Ruedemann 1925 B70
    `--*D. goldringae Ruedemann 1925 B70
  Dithecodendrum Obut 1964 B70
    `--*D. sibericum Obut 1964 B70
  Estoniocaulis Obut & Rytsk 1958 B70
    `--*E. jaervensis (Rosenstein in Obut & Rytsk 1958) [=Inocaulis jaervensis] B70
  Leveillites Foerste 1923 B70
    `--*L. hartnageli Foerste 1923 B70
  Medusaegraptus Ruedemann 1925 B70
    `--*M. mirabilis Ruedemann 1925 B70
  Palmatophycus Bouček 1941 B70
    `--*P. kettneri Bouček 1957 B70
  Polygonograptus Bouček 1957 B70
    `--*P. sokolowi (Obut 1953) [=Palaeodictyota sokolowi] B70
  Protohalecium Chapman & Thomas 1936 B70
    `--*P. hallianum Chapman & Thomas 1936 B70
  Rhadinograptus Obut 1960 B70
    `--*R. jurgensonae Obut 1960 B70
  Ruedemannograptus Termier & Termier 1948 [=Streptograptus Ruedemann 1947 non Yin 1937] B70
    `--*R. tenuis (Ruedemann 1947) [=*Streptograptus tenuis] B70
  Siberiodendrum Obut 1964 B70
    `--*S. robustum Obut 1964 B70
  Siberiograptus Obut 1964 B70
    `--*S. kotujensis Obut 1964 B70
  Sphenoecium Chapman & Thomas 1936 [=Sphenothallus Chapman 1917 non Hall 1848] B70
    `--*S. filicoides (Chapman 1917) [=Sphenothallus filicoides] B70
  Birastrites Geinitz 1866 (n. d.) B70
  Buthograptus Hall 1861 (n. d.) B70
  Cameragraptus Hundt 1951 (n. d.) B70
  Cardograptus Hundt 1965 (n. d.) B70
  Conograptus Ruedemann 1947 (n. d.) B70
  Ctenograptus Nicholson 1876 (n. d.) B70
  Cystoturriculograptus Hundt 1952 (n. d.) B70
  Demicystograptus Hundt 1942 (n. d.) B70
  Dibranchiograptus Hundt 1949 (n. d.) B70
  Didymograptoides Hundt 1951 (n. d.) B70
  Eiseligraptus Hundt 1965 (n. d.) B70
  Geminograptus Hundt 1951 (n. d.) B70
  Labrumograptus Hundt 1952 (n. d.) B70
  Limpidograptus Khaletskaya 1962 (n. d.) B70
  Nereitograptus Hundt 1951 (n. d.) B70
  Paradimorphograptus Hundt 1951 (n. d.) B70
  Phycograptus Gurley 1896 (n. d.) B70
  Planktograptus Yakovlev 1933 (n. d.) B70
  Procrytograptus Poulsen 1943 (n. d.) B70
  Protograptus Matthew 1886 (n. d.) B70
  Spinosidiplograptus Hundt 1951 (n. d.) B70
  Stelechograptus Ruedemann 1947 (n. d.) B70
  Strophograptus Ruedemann 1904 (n. d.) B70
  Thecocystograptus Hundt 1947 (n. d.) B70
  Thuringiagraptus Hundt 1935 (n. d.) B70
  ‘Triplograptus’ Hundt 1965 non Richter 1871 (n. d.) B70
  Undograptus Hundt 1949 (n. d.) B70
  Triramograptus atavus FEZ05
  Parakidograptus acuminatus P98
  Amansites Brongniart 1849 (n. d.) H62
    `--*A. dentatus (Brongniart 1828) (n. d.) [=Fucoides dentatus] H62
  Stimulograptus sedgwicki K-D04
  Lituigraptus convolutus K-D04

*Type species of generic name indicated

REFERENCES

[BU02] Bates, D. E. B., & A. Urbanek. 2002. The ultrastructure, development, and systematic position of the graptolite genus Mastigograptus. Acta Palaeontologica Polonica 47 (3): 445–458.

[B70] Bulman, O. M. B. 1970. Graptolithina with sections on Enteropneusta and Pterobranchia. In: Teichert, C. (ed.) Treatise on Invertebrate Paleontology pt V 2nd ed. pp. V1–V149. The Geological Society of America, Inc.: Boulder (Colorado), and the University of Kansas: Lawrence (Kansas).

[FEZ05] Feng, H., B.-D. Erdtmann & Y. Zhang. 2005. Taxonomy and phylogeny of Staurograptus Emmons from the base of the Ordovician. Palaeontology 48 (5): 1007–1019.

[H62] Häntzschel, W. 1962. Trace fossils and problematica. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W177–W245. Geological Society of America, and University of Kansas Press.

[K-D04] Kozłowska-Dawidziuk, A. 2004. Evolution of retiolitid graptolites—a synopsis. Acta Palaeontologica Polonica 49 (4): 505–518.

[K48] Kozłowski, R. 1948. Les graptolithes et quelques nouveaux groupes d’animaux du Tremadoc de la Pologne. Palaeontologica Polonica 3: i–xii, 1–235.

[MU04] Mierzejewski, P., & A. Urbanek. 2004. The morphology and fine structure of the Ordovician Cephalodiscus-like genus Melanostrophus. Acta Palaeontologica Polonica 49 (4): 519–528.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

Graptoloida

Rhabdinopora flabelliformis flabelliformis, copyright Institute of Geology at TUT.


Belongs within: Graptolithina.
Contains: Eugraptoloida.

The Graptoloida is the clade including the planktic graptolites, characterised by the possession of a nematophorous sicula with a free nema in the adult stage (Maletz et al. 2009).

<==Graptoloida (see below for synonymy)
    |  i. s.: Apiograptus K79
    |         Neolobograptus ZL97
    |           |--N. ariculatus ZL97
    |           `--N. nilssoni K-D04
    |         Calyxdendrum Kozłowski 1960 B70
    |           `--*C. graptoloides Kozłowski 1960 B70
    |         Psigraptus Jackson 1967 B70
    |           `--*P. arcticus Jackson 1967 B70
    |--Radiograptus Lapworth ex Bulman 1950 FC86, B70
    |    `--*R. rosieranus Bulman 1950 B70
    `--+--Staurograptus Emmons 1855 FC86, FEZ05 (see below for synonymy)
       |    |--*S. dichotomus Emmons 1855 (see below for synonymy) FEZ05
       |    |--S. apertus Ruedemann 1904 [=S. dichotomus var. apertus] FEZ05
       |    `--S. hyperboreus (Obut & Sobolevskaya 1962) (see below for synonymy) FEZ05
       `--+--Rhabdinopora Eichwald 1855 FC86, B70 (see below for synonymy)
          |    |--*R. flabelliformis (Eichwald 1840) B70 (see below for synonymy)
          |    |    |--R. f. flabelliformis FEZ05
          |    |    |--R. f. anglica FEZ05
          |    |    |--‘Dictyonema flabelliforme’ norvegicum B70
          |    |    |--R. f. parabola [incl. Staurograptus dichotomus priscus Lin 1986] FEZ05
          |    |    `--‘Dictyonema flabelliforme’ sociale (Salter 1858) [=*Graptopora socialis] B70
          |    |--*Ceramites’ hisingeri Liebmann in Forchhammer 1845 H62
          |    `--R. praeparabola FEZ05
          `--+--Anisograptinae FC86
             |    |--Triograptus Monsen 1925 FC86, B70
             |    |    |--*T. osloensis Monsen 1925 B70
             |    |    `--T. canadensis B70
             |    `--Anisograptus Ruedemann 1937 B70
             |         |--*A. matanensis Ruedemann 1937 B70
             |         |    |--A. m. matanensis FEZ05
             |         |    `--A. m. tetragraptoides FEZ05
             |         |--A. flexuosus B70
             |         `--A. richardsoni FEZ05
             `--Adelograptinae [Didymograpta, Didymograptina] FC86
                  |--Eugraptoloida FC86
                  `--+--Kiaerograptus Spjeldnaes 1963 FC86, B70
                     |    `--*K. kiaeri (Monsen 1925) [=Didymograptus kiaeri] B70
                     `--Adelograptus Bulman 1941 FC86, B70 [incl. Choristograptus Legrand 1963 B70]
                          |--*A. hunnebergensis (Moberg 1892) [=Bryograptus hunnebergensis] B70
                          |--A. clarki C68
                          |--A. lapworthi B70
                          |--*Choristograptus’ louhai Legrand 1963 B70
                          |--A. tenellus (Linnarsson 1871) MCM09
                          `--A. victoriae C68

Graptoloida [Anisograptidae, Graptodendroidina, Graptolitidae, Graptoloidea, Retioloidea, Rhabdinoporinae, Rhabdophora, Tetragrapti]

Rhabdinopora Eichwald 1855 FC86, B70 [=Dictyograptus Westergård 1909 non Hopkinson 1875 B70, Phyllograpta Angelin 1854 B70; incl. Ceramites Liebmann in Forchhammer 1845 H62, Graptopora Salter 1858 B70]

*Rhabdinopora flabelliformis (Eichwald 1840) B70 [=Gorgonia flabelliformis B70, *Dictyograptus flabelliformis B70, Dictyonema flabelliforme FC86, *Phyllograpta flabelliformis B70; incl. Dictyonema flabelliforme var. conferta K48]

Staurograptus Emmons 1855 FC86, FEZ05 [=Staurograpsus FEZ05; incl. Aletograptus Obut & Sobolevskaya 1962 FEZ05; Staurograptinae]

*Staurograptus dichotomus Emmons 1855 [incl. S. borealis Zhao 1983, S. dasycladus Zhao 1983, Radiograptus rosieranus flexibilis Fortey in Fortey et al. 1982, S. (Radiograptus) flexibilis, S. immaturus Wang, Liu & Zhou 1979, S. dichotomus immaturus, S. magnus Yu, Liu & Fang 1984, S. minimus Mu in Mu & Lin 1984, S. orientalis Mu in Mu & Lin 1984, S. dichotomus sinicus Mu in Mu & Lin 1984, S. taishanensis Wang, Liu & Zhou 1979, S. dichotomus taishanensis, S. tenuis tangpanensis Yu, Liu & Fang 1984, S. tenuis Jackson 1974] FEZ05

Staurograptus hyperboreus (Obut & Sobolevskaya 1962) [=*Aletograptus hyperboreus; incl. A. flexibilis Yu, Liu & Fang 1984, A. incertus Yu, Liu & Fang 1984, Triograptus rigidus Yu, Liu & Fang 1984] FEZ05

*Type species of generic name indicated

REFERENCES

[B70] Bulman, O. M. B. 1970. Graptolithina with sections on Enteropneusta and Pterobranchia. In: Teichert, C. (ed.) Treatise on Invertebrate Paleontology Part V 2nd ed. pp. V1–V149. The Geological Society of America, Inc.: Boulder (Colorado), and the University of Kansas: Lawrence (Kansas).

[C68] Cooper, R. A. 1968. Lower and Middle Paleozoic fossil localities of north-west Nelson. Transactions of the Royal Society of New Zealand, Geology 6 (7): 75–89.

[FEZ05] Feng, H., B.-D. Erdtmann & Y. Zhang. 2005. Taxonomy and phylogeny of Staurograptus Emmons from the base of the Ordovician. Palaeontology 48 (5): 1007–1019.

[FC86] Fortey, R. A., & R. A. Cooper. 1986. A phylogenetic classification of the graptoloids. Palaeontology 29: 631–654.

[H62] Häntzschel, W. 1962. Trace fossils and problematica. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W177–W245. Geological Society of America, and University of Kansas Press.

[K79] Kirk, N. H. 1979. Thoughts on coloniality in the graptolites. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 411–432. Academic Press: London.

[K-D04] Kozłowska-Dawidziuk, A. 2004. Evolution of retiolitid graptolites—a synopsis. Acta Palaeontologica Polonica 49 (4): 505–518.

[K48] Kozłowski, R. 1948. Les graptolithes et quelques nouveaux groupes d’animaux du Tremadoc de la Pologne. Palaeontologica Polonica 3:i–xii, 1–235.

[MCM09] Maletz, J., J. Carlucci & C. E. Mitchell. 2009. Graptoloid cladistics, taxonomy and phylogeny. Bulletin of Geosciences 84 (1): 7–19.

[ZL97] Zhang Y. & A. C. Lenz. 1997. Uppermost Wenlock and Ludlow graptolites from southern Yunnan, China. Canadian Journal of Earth Sciences 34: 1220–1238.