Octopodidae

Graneledone boreopacifica, from NOAA/MBARI.

Belongs within: Octopoda.
Contains: Benthoctopus, Octopus.

The Octopodidae is a diverse group of benthic octopuses, including the majority of familiar octopus species. Suckers on the arms are biserial in Octopodinae and Bathypolypodinae but uniserial in the deep-sea Graneledoninae (O'Shea 1999).

Characters (from O'Shea 1999): Body muscular to gelatinous; suckers uniserial or biserial; radula heterodont, occasionally homodont; digestive gland anterior to stomach and caecum; hectocotylised arm not autotomised at maturity; mantle-locking apparatus absent; shell vestige occasionally present as stylets in mantle; ink sac present or absent; light organs absent.

<==Octopodidae [Heteroglossa, Octopodoidea]
    |  i. s.: Megaleledone O99
    |         Hapalochlaena O99
    |         Pteroctopus tetracirrus O99, PP64
    |         Cistopus O99
    |         Vosseledone H88
    |         Scaegurus MG-H11
    |--Bathypolypodinae O99
    |    |--Benthoctopus O99
    |    |--Grimpella thaumastocheir O99
    |    `--Bathypolypus O99
    |         |--B. arcticus O99
    |         |--B. faeroensis (Russell 1909) O99
    |         `--B. spondalis S72
    |--Eledone [Eledoninae] O99
    |    |  i. s.: E. caparti Adam 1950 H88
    |    |         E. cirrhosa (Lamarck 1798) H88
    |    |         E. gaucha Haimovici 1988 H88
    |    |         E. massyae Voss 1964 H88
    |    |         E. palari Lu & Stranks 1991 MG-H11
    |    |         E. thysanophora Voss 1962 H88
    |    |--E. (Eledone) moschata (Lamarck 1798) G20, H88
    |    `--E. (Ocithoe) G20
    |         |--E. (O.) antiquorum G20
    |         `--E. (O.) cranchii G20
    |--Octopodinae O99
    |    |--Octopus O99
    |    |--Ameloctopus Norman 1992 O99
    |    |--Pinnoctopus d’Orbigny 1845 O99 (see below for synonymy)
    |    |    |--*P. cordiformis (Quoy & Gaimard 1832) (see below for synonymy) O99
    |    |    |--‘Robsonella’ australis (Hoyle 1885) P61
    |    |    |--‘Octopus fontanianus’ Robson 1929 (see below for synonymy) O99
    |    |    `--P. kermadecensis (Berry 1914) [=Polypus kermadecensis] O99
    |    `--Enteroctopus Gould 1852 O99
    |         |--*E. megalocyathus Gould 1852 O99
    |         |--E. dofleini [=Octopus dofleini] O99
    |         |--E. magnificus O99
    |         `--E. zealandicus (Benham 1944) [=Paroctopus zealandicus] O99
    `--Graneledoninae [Pareledoninae] O99
         |--Pareledone O99
         |--Bentheledone albida O99
         |--Velodona togata O99
         |--Thaumeledone Robson 1930 O99
         |    |--*T. brevis (Hoyle 1885) [=Eledone brevis] O99
         |    |--T. gunteri O99
         |    |--T. marshalli O’Shea 1999 O99
         |    `--T. zeiss O’Shea 1999 O99
         `--Graneledone Joubin 1918 O99
              |--*G. verrucosa (Verrill 1881) [=Eledone verrucosa, Moschites verrucosa] O99
              |    |--G. v. verrucosa O99
              |    `--G. v. media Joubin 1918 O99
              |--G. antarctica Voss 1976 O99
              |--G. boreopacifica O99
              |--G. challengeri (Berry 1916) (see below for synonymy) O99
              |--G. macrotyla O99
              |--G. pacifica Voss & Pearcy 1990 O99
              `--G. taniwha O’Shea 1999 O99
                   |--G. t. taniwha O99
                   `--G. t. kubodera O’Shea 1999 O99

Graneledone challengeri (Berry 1916) [=Moschites challengeri, Eledone challengeri, M. verrucosa var. challengeri] O99

‘Octopus fontanianus’ Robson 1929 non d’Orbigny 1835 [=*Joubinia fontanianus, *Robsonella fontanianus] O99

Pinnoctopus d’Orbigny 1845 O99 [incl. Callistoctopus O99, Joubinia Robson 1930 (preoc.) O99, Macroctopus Robson 1928 O99, P61, Robsonella Adam 1938 O99]

*Pinnoctopus cordiformis (Quoy & Gaimard 1832) [=Octopus cordiformis; incl. O. (Macroctopus) communis Park 1885, Polypus communis, O. flindersi Cotton 1932, O. maorum Hutton 1880, *Macroctopus maorum, Polypus maorum] O99

*Type species of generic name indicated

REFERENCES

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[H88] Haimovici, M. 1988. Eledone gaucha, a new species of eledonid octopod (Cephalopoda: Octopodidae) from southern Brazil. Nautilus 102 (2): 82–87.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[O99] O’Shea, S. 1999. The marine fauna of New Zealand: Octopoda (Mollusca: Cephalopoda). NIWA Biodiversity Memoir 112: 1–280.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d'Endoume, Bulletin 31 (27): 5–137.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4^th ed. Whitcombe and Tombs Limited: Christchurch.

[S72] Stunkard, H. W. 1972. Clarification of taxonomy in the Mesozoa. Systematic Zoology 21 (2): 210–214.

Last updated: 18 November 2021.

Octopoda

Blanket octopus Tremoctopus violaceus, copyright Cassandra L. Cox.

Belongs within: Neocoleoidea.
Contains: Grimpoteuthididae, Opisthoteuthididae, Octopodidae.

The Octopoda, octopuses, are a group of stout-bodied cephalopods with eight arms and no tentacles. A small internal shell is present in members of the suborder Cirrata which also have a pair of fins on the head and cilia-like cirri associated with the suckers on the arms; all three features are absent in the suborder Incirrata.

Octopoda [Octopodida]
    |  i. s.: Polypus H09
    |           |--P. octopodia [=Eledone (Polypus) octopodia] G20
    |           |--P. pictus (Brock 1882) [=Octopus pictus] H09
    |           |--P. polyzenia (Gray 1849) [=Octopus polyzenia] H09
    |           `--P. tenebricus (Smith 1884) [=Octopus tenebricus] H09
    |         Amphioctopus marginatus KVF11
    |--Cirrata [Cirroctopoda] O99
    |    |  i. s.: Chunioteuthis gilchristi (Robson 1924) [=Cirrhoteuthis gilchristi] O99
    |    |         Cirroteuthopsis massayae Grimpe 1920 O99
    |    |         Frokenia clara Hoyle 1904 O99
    |    |--Grimpoteuthididae O99
    |    |--Opisthoteuthididae O99
    |    |--Stauroteuthis GO06 [Stauroteuthididae YVD98]
    |    |    `--S. syrtensis GO06
    |    |--Luteuthis O’Shea 1999 [Luteuthididae] O99
    |    |    |--*L. dentatus O’Shea 1999 O99
    |    |    `--‘Grimpoteuthis’ tuftsi Voss & Pearcy 1990 O99
    |    `--Cirroteuthididae O99
    |         |--Cirroteuthis Eschricht 1836 O99
    |         |    `--*C. muelleri Eschricht 1836 O99
    |         `--Cirrothauma Chun 1911 O99
    |              |--*C. murrayi Chun 1911 O99
    |              `--C. magna (Hoyle 1885) [=Cirroteuthis magna] O99
    `--Incirrata O99
         |  i. s.: Palaeoctopodidae D93
         |           |--Palaeoctopus newboldi (Woodward 1896) YVD98, D93
         |           `--Proteroctopus ribeti Fischer & Riou 1982 D93
         |--Octopodidae O99
         |--Vitreledonella Joubin 1918 [Vitreledonellidae] O99
         |    `--*V. richardi Joubin 1918 O99
         |--Amphitretus Hoyle 1885 [Amphitretidae] O99
         |    |--*A. pelagicus Hoyle 1885 [incl. Idioctopus gracilipes Taki 1964] O99
         |    `--A. thielei Robson 1930 [=A. pelagicus thielei] O99
         |--Bolitaenidae [Bathypelagoctopoda, Bolitaenoidea] O99
         |    |--Haliphron atlanticus YVD98, O99 [incl. Alloposus mollis O99]
         |    |--Bolitaena Steenstrup 1859 O99
         |    |    `--*B. microtyla Steenstrup in Hoyle 1886 O99
         |    |--Japetella Hoyle 1885 O99
         |    |    `--J. diaphana (Hoyle 1885) [=Eledonella diaphana, Bolitaena diaphana, Chunella diaphana] O99
         |    `--Eledonella Verrill 1884 O99
         |         `--*E. pygmaea Verrill 1884 (see below for synonymy) O99
         `--Argonautoidea YVD98
              |--Alloposidae YVD98
              `--+--Tremoctopus delle Chiaje 1830 O99 [Tremoctopodidae YVD98]
                 |    |--*T. violaceus delle Chiaje 1830 O99
                 |    |    |--T. v. violaceus O99
                 |    |    `--T. v. gracilis O99
                 |    |--T. gelatus O99
                 |    `--T. robsonianus Kirk 1883 [=T. robsoni (l. c.)] O99
                 `--+--Ocythoe Rafinesque 1814 O99 [Ocythoidae YVD98]
                    |    `--*O. tuberculata Rafinesque 1814 O99
                    `--Argonautidae YVD98
                         |--Obinautilus Kobayashi 1954 D93, K64
                         |    `--*O. pulcher Kobayashi 1954 K64
                         `--Argonauta Linnaeus 1758 O99
                              |--*A. argo Linnaeus 1758 O99 [incl. A. bulleri Kirk 1885 O99, A. papyracea C64]
                              |--A. boettgeri Maltzan 1881 P61
                              |--A. cymbiu Linnaeus 1758 L58
                              |--A. hians Solander 1786 P61
                              |--A. navicula Lightfoot 1786 K65
                              `--A. nodosa Solander 1786 [incl. A. gracilis Kirk 1885, A. tuberculata Shaw 1791] O99

*Eledonella pygmaea Verrill 1884 [incl. E. ijimae Sasaki 1929, Bolitaena ijimai, E. massayae Robson 1924, B. massayae, Japetella prismatica Hoyle 1885, B. massayae purpurea Robson 1932, Eledonella sheardi Allan 1945] O99

*Type species of generic name indicated

REFERENCES

[C64] Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 33: 517–686.

[D93] Doyle, P. 1993. Mollusca: Cephalopoda (Coleoidea). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 229–236. Chapman & Hall: London.

[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.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[K65] Kay, E. A. 1965. The Reverend John Lightfoot, Daniel Solander, and the Portland Catalogue. Nautilus 79 (1): 10–19.

[KVF11] Kröger, B., J. Vinther & D. Fuchs. 2011. Cephalopod origin and evolution: a congruent picture emerging from fossils, development and molecules. Bioessays 33: 602–613.

[K64] Kummel, B. 1964. Nautiloidea—Nautilida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt K. Mollusca 3. Cephalopoda—General Features—Endoceratoidea—Actinoceratoidea—Nautiloidea—Bactritoidea pp. K383–K457. The Geological Society of America and the University of Kansas Press.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[O99] O’Shea, S. 1999. The marine fauna of New Zealand: Octopoda (Mollusca: Cephalopoda). NIWA Biodiversity Memoir 112: 1–280.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4^th ed. Whitcombe and Tombs Limited: Christchurch.

[YVD98] Young, R. E., M. Vecchione & D. T. Donovan. 1998. The evolution of coleoid cephalopods and their present biodiversity and ecology. South African Journal of Marine Science 20: 393–420.

Last updated: 20 November 2021.

Comatulida

Comatula rotalaria, copyright Teresa Zubi.

Belongs within: Articulata.
Contains: Tropiometracea, Antedonacea, Solanocrinitacea, Comasterinae, Capillasterinae.

The Comatulida, feather stars, are a group of crinoids in the larval column has been reduced to a single cirriferous centrodorsal attached to the calyx (Wienberg Rasmussen 1978). They are the most prominent crinoid group in the modern fauna.

<==Comatulida [Comatuladae, Comatulidae, Comatulidia, Macrophreata, Macrophreatina, Oligophreata]
    |  i. s.: Pachyantedon Jaekel 1891 (n. d.) WR78
    |           |--*P. beyrichi Jaekel 1891 WR78
    |           `--P. gracilis (Walther 1886) [=Solanocrinus gracilis, Antedon gracilis] C50
    |--Paracomatulacea UL78
    |    |--Paracomatula Hess 1951 [Paracomatulidae] WR78
    |    |    `--*P. helvetica Hess 1951 WR78
    |    `--Atelecrinidae [Atelecrininae] WR78
    |         |--Atopocrinus Clark 1912 WR78
    |         |    `--*A. sibogae Clark 1912 WR78
    |         |--Jaekelometra Gislén 1924 WR78
    |         |    |--J. belgica (Jaekel 1902) [=Atelecrinus belgicus] WR78
    |         |    |--J. columnaris Gislén 1924 E01
    |         |    |--J. concava (Schlüter 1878) E01
    |         |    `--J. meijeri Rasmussen 1961 E01
    |         |--Sibogacrinus Clark in Clark & Clark 1967 WR78
    |         |    `--*S. anomalus (Clark 1912) [=Atelecrinus anomalus] WR78
    |         `--Atelecrinus Carpenter 1881 WR78
    |              |--*A. balanoides Carpenter 1881 (see below for synonymy) WR78
    |              `--A. conifer WR78
    |--+--Tropiometracea UL78
    |  `--Antedonacea UL78
    `--+--Solanocrinitacea UL78
       `--Comasteridae [Actinometridae, Actinometrinae, Comasteracea, Comasterida, Comasterina] UL78
            |  i. s.: Clarkcomanthus MG-H11
            |           |--C. albinotus (Rowe, Hogget et al. 1986) LR06
            |           |--C. littoralis (Carpenter 1888) MG-H11
            |           `--C. luteofuscum (Clark 1915) MG-H11
            |         Oxycomanthus MG-H11
            |           |--O. bennetti (Müller 1841) LR06
            |           |--O. japonica (Müller 1841) LR06
            |           `--O. plectrophorum (Clark 1916) MG-H11
            |--Comasterinae WR78
            |--Capillasterinae WR78
            `--Comactiniinae WR78
                 |--Comatulella Clark 1911 WR78
                 |    `--*C. brachiolata (Lamarck 1816) [=Comatula brachiolata; incl. Alecto rosea Müller 1841] WR78
                 |--Comatulides Clark 1918 [=Cominia Clark 1909 non Brown 1844] WR78
                 |    `--*C. decameros (Clark 1908) [=Comanthus decameros] WR78
                 |--Comactinia Clark 1909 WR78
                 |    |--*C. echinoptera (Müller 1841) [=Alecto echinoptera] WR78
                 |    `--C. meridionalis BLWR78 [=C. echinoptera var. meridionalis MMR78]
                 `--Comatula Lamarck 1816 (see below for synonymy) WR78
                      |--*C. solaris Lamarck 1816 [incl. *Actinometra imperialis Müller 1841] WR78
                      |--C. pectinata (Linnaeus 1758) MG-H11
                      |--C. rotalaria WR78
                      `--C. tenuicirra Clark 1912 MG-H11

*Atelecrinus balanoides Carpenter 1881 [incl. A. cubensis Carpenter 1881, A. pourtalesi Clark 1907, A. helgae Clark 1913] WR78

Comatula Lamarck 1816 [incl. Actinometra Müller 1841, Actinomedra (l. c.), Actinomerta (l. c.), Actinometer (l. c.), Actynometra (l. c.), Aetinometra (l. c.)] WR78

*Type species of generic name indicated

REFERENCES

[BLWR78] Brower, J. C., N. G. Lane & H. Wienberg Rasmussen. 1978. Postlarval ontogeny of fossil crinoids. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T244–T274. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[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.

[E01] Eagle, M. K. 2001. New fossil crinoids (Articulata: Comatulida) from the late Oligocene of Waitete Bay, northern Coromandel Peninsula, New Zealand. Records of the Auckland Museum 37: 81–92.

[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.

[MMR78] Macurda, D. B., Jr., D. L. Meyer & M. Roux. 1978. The crinoid stereom. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T217–T228. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[UL78] Ubaghs, G., N. G. Lane, H. Wienberg Rasmussen & H. L. Strimple. 1978. Evolution. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 2 pp. T275–T316. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Sepiolidae

Sepiola atlantica, copyright Mick Otten.

Belongs within: Coleoidea.

The Sepiolidae, bobtail squid, are a group of small (1–10 cm long) cephalopods with a broad body, round posterior mantle and ear-like fins. They bear a funnel locking-apparatus with a single depression (Young 1996).

<==Sepiolidae MG-H11
    |--Heteroteuthis hawaiiensis GO06
    `--+--Rossia palpebrosa GO06
       `--Sepiola GO06
            |--S. affinis GO06
            |--S. atlantica NN03
            |--S. birostrata V62
            |--S. intermedia NN03
            |--S. ligulata NN03
            |--S. robusta NN03
            |--S. rondeleti NN03
            `--S. trirostrata Voss 1962 V62

Sepiolidae incertae sedis:
  Euprymna MG-H11
    |--E. albatrossae Voss 1962 V62
    |--E. berryi NN03
    |--E. hyllebergi NN03
    |--E. morsei NN03
    |--E. phenax Voss 1962 V62
    |--E. scolopes DH08
    `--E. tasmanica (Pfeffer 1884) MG-H11
  Austrorossia australis (Berry 1918) MG-H11
  Rondeletiola minor MG-H11, NN03
  Sepiolina nipponensis (Berry 1911) MG-H11

*Type species of generic name indicated

REFERENCES

[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.

[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.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[NN03] Nishiguchi, M. K., & V. S. Nair. 2003. Evolution of symbiosis in the Vibrionaceae: a combined approach using molecules and physiology. International Journal of Systematic and Evolutionary Microbiology 53: 2019–2026.

[V62] Voss, G. L. 1962. Six new species and two new subspecies of cephalopods from the Philippine Islands. Proceedings of the Biological Society of Washington 75: 169–176.

Antedonacea

Florometra serratissima, from Ocean Spaces.

Belongs within: Comatulidia.
Contains: Antedoninae, Zenometrinae, Bathymetrinae.

The Antedonacea are a group of feather stars with a relatively large internal cavity to the calyx.

Characters (from Wienberg Rasmussen 1978): Centrodorsal variable, discoidal or hemispherical to rounded subconical, conical or columnar. Cirrus sockets generally small and numerous, closely placed, often closely alternating in several circles, may tend to form several indistinct columns, in Zenometrinae forming 10 to 20 distinct columns. Sockets without distinct ornament. Basals often transformed to rosette. Rod-shaped basals usually reduced or absent, sometimes present and commonly exposed. Centrodorsal cavity moderate, about 0.3 of centrodorsal diameter in most Antedoninae, Thysanometrinae, Heliometrinae and Isometrinae, and rather large to very large in Perometrinae, Bathymetrinae, Zenometrinae and Pentametrocrinidae. Radials 10 (5 radials and 5 pararadials) in Promachocrinus and Thaumatocrinus, but five in all other Antedonacea. Ventral muscular fossae high and almost vertical on thin, muscular flange of radial, and meeting in midradial line at angle of about 90 degrees. Radial cavity narrow or funnel shaped, without calcareous filling (no central plug). Arms undivided in Pentametrocrinidae, divided at primibrachs 2 in Antedonidae. Synarthry at brachials 2–2 often embayed. Syzygy at brachials 3–4 and generally at brachials 9–10, and distal with short intervals. Distal brachials strongly wedge shaped. Pinnules cylindrical to flattened, not carinate. Mouth central.

Antedonacea [Antedonida]
    |--Pentametrocrinidae [Decametrocrinidae, Thaumatocrinida, Thaumatocrinidae, Thaumatocrininae] WR78
    |    |--Thaumatocrinus Carpenter 1884 [incl. Decametrocrinus Minckert 1905] WR78
    |    |    |--*T. renovatus Carpenter 1884 (see below for synonymy) WR78
    |    |    `--T. naresi WR78
    |    `--Pentametrocrinus Clark 1908 WR78
    |         |--*P. japonicus (Carpenter 1882) [=Eudiocrinus japonicus] WR78
    |         |--P. semperi (Carpenter 1882) MG-H11
    |         `--P. varians (Carpenter 1882) MG-H11
    `--Antedonidae WR78
         |--Antedoninae WR78
         |--Zenometrinae WR78
         |--Bathymetrinae WR78
         |--Isometra Clark 1908 [Isometrinae] WR78
         |    |--*I. challengeri (Clark 1907) (see below for synonymy) WR78
         |    `--I. vivipara WR78
         |--Thysanometrinae WR78
         |    |--Thysanometra Clark 1907 WR78
         |    |    `--*T. tenelloides (Clark 1907) [=Antedon tenelloides] WR78
         |    `--Coccometra Clark 1908 WR78
         |         `--*C. hagenii (Pourtalès 1868) [=Comatula hagenii] WR78
         |--Perometrinae WR78
         |    |--Nanometra Clark 1907 WR78
         |    |    `--N. bowersi (Clark 1907) (see below for synonymy) WR78
         |    |--Perometra Clark 1907 WR78
         |    |    `--*P. diomedeae (Clark 1907) [=Antedon diomedeae] WR78
         |    |--Erythrometra Clark 1908 WR78
         |    |    `--*E. rubra (Clark 1907) [=Antedon ruber] WR78
         |    `--Hypalometra Clark 1908 WR78
         |         `--*H. defecta (Carpenter 1888) [=Antedon defecta] WR78
         `--Heliometrinae [Promachocrininae] WR78
              |--Promachocrinus Carpenter 1879 WR78
              |    `--*P. kerguelensis Carpenter 1881 (see below for synonymy) WR78
              |--Solanometra Clark 1911 WR78
              |    `--*S. antarctica (Carpenter 1881) (see below for synonymy) WR78
              |--Hertha von Hagenow 1840 WR78
              |    `--*H. mystica von Hagenow 1840 WR78
              |--Allionia Michelotti 1861 WR78
              |    `--*A. oblita Michelotti 1861 WR78
              |--Roiometra Clark 1944 WR78
              |    `--*R. columbiana Clark 1944 WR78
              |--Anthometra Clark 1913 WR78
              |    `--*A. adriani (Bell 1908) [=Antedon adriani] WR78
              |--Heliometra Clark 1907 WR78
              |    `--H. glacialis (Leach ex Owen 1833) WR78 (see below for synonymy)
              |         |--H. g. glacialis BK77
              |         `--H. g. maxima BK77
              `--Florometra Clark 1913 WR78
                   |--*F. magellanica (Bell 1882) [=Antedon eschrichti var. magellanica] WR78
                   |--F. asperrima WR78
                   `--F. serratissima HF04

Heliometra glacialis (Leach ex Owen 1833) WR78 [=Alecto glacialis WR78; incl. Antedon arctica Clark 1907 WR78, An. bartensi Carpenter 1886 WR78, Alecto eschrichtii Müller 1841 WR78, An. eschrichtii BK77, *Heliometra eschrichtii WR78, H. juvenalis Clark 1908 WR78, Antedon quadrata Carpenter 1884 WR78]

*Isometra challengeri (Clark 1907) [=Antedon challengeri Clark 1907, A. lineata Carpenter 1888 non Pomel 1887] WR78

Nanometra bowersi (Clark 1907) [=Antedon bowersi; incl. A. minor Clark 1907 non A. avenionensis var. minor Nicolas 1898, *Nanometra minor, N. minckerti Clark 1907, Antedon orientalis Clark 1907] WR78

*Promachocrinus kerguelensis Carpenter 1881 [incl. P. joubini Vaney 1910, P. vanhoeffenianus Minckert 1905] WR78

*Solanometra antarctica (Carpenter 1881) [=Antedon antarctica; incl. A. australis Carpenter 1888 non Carpenter 1882, Heliometra glabra Clark 1907] WR78

*Thaumatocrinus renovatus Carpenter 1884 [incl. Promachocrinus abyssorum Carpenter 1888, *Decametrocrinus abyssorum] WR78

*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.

[HF04] Helfenbein, K. G., H. M. Fourcade, R. G. Vanjani & J. L. Boore. 2004. The mitochondrial genome of Paraspadella gotoi is highly reduced and reveals that chaetognaths are a sister group to protostomes. Proceedings of the National Academy of Sciences of the USA 101 (29): 10639-10643.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1-191.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In Treatise on Invertebrate Paleontology pt. T. Echinodermata 2. Crinoidea (R. C. Moore & C. Teichert, eds) vol. 3 pp. T813-T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Tropiometracea

Fossil of Pterocoma pennata in the Paläontologische Museum München, copyright Ghedoghedo.

Belongs within: Comatulida.
Contains: Charitometridae, Notocrinacea, Conometridae, Thalassometridae.

The Tropiometracea are a group of feather stars in which the pinnules are prismatic in shape with a triangular cross-section (Wienberg Rasmussen 1978).

Characters (from Wienberg Rasmussen 1978): Centrodorsal hemispherical or conical to truncated conical, columnar or discoidal. Adult specimens often with flattened, cirrus-free dorsal area, which may be rugose or tuberculate. With or without a radial dorsal star, radial pits and subradial cleft. Cirrus sockets large, without distinct ornament or with articular tubercles, occasionally with marginal crenellae. Sockets arranged in (five or) 10 distinct columns or irregular and crowded, tending to form 10, 15 or 20 lateral columns. Cirrals with or without dorsal spine. Centrodorsal cavity narrow to moderate, generally less than three-tenths of centrodorsal diameter except for very small specimens. Rod-shaped basals present, generally exposed in interradial points. Surface of radials variable, often low or concealed, but may be moderate to large and recumbent or overhanging. Articular face of radials gently sloping to steep, almost vertical and often high. Ventral muscular fossae low and wide to high and wide or narrow. Radial cavity narrow to wide. Arms divided at primibranchs 2 and commonly at some or all secundibranchs 2 and tertibrachs 2, seldom quartibrachs 2, occasionally secundibrachs 4 and tertibrachs 4. Synarthry at brachials 1–2 may be flat or embayed, often narrow. Syzygy at brachials 3–4, seldom at brachials 1–2 or distal branches. Further syzygy with variable intervals through distal branches. Pinnulation complete; pinnules distinctly prismatic, triangular in section with dorsal keel (unknown in most fossil species). Distal pinnules extend beyond tip of arm. Ambulacral covering plates often distinct along pinnules. Mouth central.

<==Tropiometracea [Tropiometrida, Thalassometrida, Thalassometrina, Thalassometroida]
    |--+--Notocrinacea UL78
    |  `--Asterometridae UL78
    |       |--Asterometra Clark 1907 WR78
    |       |    |--*A. macropoda (Clark 1907) [=Antedon macropoda] WR78
    |       |    `--A. anthus (Clark 1907) MG-H11
    |       `--Pterometra Clark 1909 WR78
    |            |--*P. trichopoda (Clark 1908) [=Ptilometra trichopoda] WR78
    |            `--P. pulcherrima WR78
    `--+--+--Conometridae UL78
       |  `--Thalassometridae UL78
       `--Pterocomidae UL78
            |--Pterocoma Agassiz 1836 [=Comathulites von Schlotheim 1823, Comaturella Münster 1839] WR78
            |    |--*P. pennata (von Schlotheim 1813) (see below for synonymy) WR78
            |    `--‘*Comaturella’ wagneri Münster 1839 WR78
            `--Placometra Gislén 1924 WR78
                 |--*P. mortenseni Gislén 1924 WR78
                 `--P. laticirra WR78

Tropiometracea incertae sedis:
  Tropiometra Clark 1907 [Tropiometridae] WR78
    `--*T. carinata (Leach 1815) (see below for synonymy) WR78
  Ptilometra Clark 1907 [Ptilometridae] WR78
    `--*P. macronema (Müller 1846) [=Comatula macronema, Alecto macronema] WR78
  Calometridae WR78
    |  i. s.: ‘Antedon’ discoidea C50
    |--Calometra Clark 1907 WR78
    |    `--*C. callista (Clark 1907) [=Antedon callista] WR78
    |--Gephyrometra Clark 1912 WR78
    |    `--*G. versicolor (Clark 1907) [=Antedon versicolor] WR78
    |--Pectinometra Clark 1912 [=Pectiometra (l. c.)] WR78
    |    `--*P. flavopurpurea (Clark 1907) [=Antedon flavopurpurea] WR78
    |--Reometra Clark 1934 [=Oreometra Clark 1912 non Aurivillius 1910] WR78
    |    `--*R. mariae (Clark 1912) [=*Oreometra mariae] WR78
    `--Neometra Clark 1912 WR78
         |--*N. multicolor (Clark 1907) [=Antedon multicolor] WR78
         |--N. conaminis Clark 1914 MG-H11
         |--N. gorgonia Clark 1915 MG-H11
         `--N. spinosissima U78
  Charitometridae C50
  Pseudoconometridae E01
    |--Pseudoconometra Eagle 2001 E01
    |    `--*P. coromandelensis Eagle 2001 E01
    `--Moanametra Eagle 2001 E01
         `--*M. torehinaensis Eagle 2001 E01

*Pterocoma pennata (von Schlotheim 1813) [=Asteriatites pennatus, *Comathulites mediterraneaeformis von Schlotheim 1823, Geocoma pennata, Ophiurites pennatus von Schlotheim 1820; incl. Comatula pinnata Goldfuss 1831, *Pterocoma pinnata] WR78

*Tropiometra carinata (Leach 1815) [=Alecto carinata, Antedon carinata, Comatula carinata Lamarck 1816; incl. T. picta] WR78

*Type species of generic name indicated

REFERENCES

[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.

[E01] Eagle, M. K. 2001. New fossil crinoids (Articulata: Comatulida) from the late Oligocene of Waitete Bay, northern Coromandel Peninsula, New Zealand. Records of the Auckland Museum 37: 81–92.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[U78] Ubaghs, G. 1978. Skeletal morphology of fossil crinoids. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T58–T216. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[UL78] Ubaghs, G., N. G. Lane, H. Wienberg Rasmussen & H. L. Strimple. 1978. Evolution. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 2 pp. T275–T316. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Bathymetrinae

Phrixometra nutrix, copyright Pablo E. Penchaszadeh.

Belongs within: Antedonacea.

The Bathymetrinae are a group of feather stars with moderate-sized cirri arising from small, closely placed sockets, and arms branching once only.

Characters (from Wienberg Rasmussen 1978): Centrodorsal conical or rounded subconical to low hemispherical. Cirrus sockets rather small, closely placed, often numerous (25 to more than 100), covering almost completely sides of centrodorsal, rarely few (10-15) with mostly smooth cirrus-free doral area; cirri closely alternate, generally without distinct arrangement, often interpreted as alternating circles, but sometimes with tendency to form columns, up to 30 in number. Size of sockets may increase slightly upward. Length of cirri moderate, apical cirri rather short. Cirrals laterally compressed and generally long, length one to six times width. Distal cirrals often dorsally carinate, may have dorsal spine. Centrodorsal cavity large. Basal rosette, no rod-shaped basals. Radials may have smooth, rather large, free surface in small species, but generally very low and often concealed in midradial area of large species. Articular face of radials high and rather steep. Interarticular ligament fossae triangular or low and narrow ridge separating interarticular from ventral fossae may be almost horizontal, straight or curved. Ventral muscular fossae large and high, rounded subtriangular with rounded ventral edge, separated by low and narrow median ridge and small notch, forming angle of almost 90 degrees with each other. Radial cavity narrow. Arms divided at primibrachs 2 only. Synarthrial articulation at primibrachs 1-2 often embayed. Primibrachs 1 usually with parallel or converging sides, commonly laterally overhung by primibrachs 2. First pinnule stiff and slender with elongated pinnulars.

<==Bathymetrinae
    |--Boleometra Clark 1936 WR78
    |    `--*B. clio (Clark 1907) [=Antedon clio] WR78
    |--Fariometra Clark 1917 WR78
    |    `--*F. explicata (Clark 1908) [=Trichometra explicata] WR78
    |--Nepiometra Clark 1917 WR78
    |    `--*N. laevis (Carpenter 1888) [=Antedon laevis] WR78
    |--Orthometra Clark 1917 WR78
    |    `--*O. hibernica (Clark 1913) [=Trichometra hibernica] WR78
    |--Retiometra Clark 1936 WR78
    |    `--*R. alascana Clark 1936 WR78
    |--Thaumatometra Clark 1908 WR78
    |    `--T. tenuis (Clark 1907) [=Antedon tenuis; incl. A. ciliata Clark 1907, *Thaumatometra ciliata] WR78
    |--Tonrometra Clark 1917 WR78
    |    `--*T. remota (Carpenter 1888) [=Antedon remota] WR78
    |--Bathymetra Clark 1908 WR78
    |    |--*B. abyssicola (Carpenter 1888) [=Antedon abyssicola] WR78
    |    `--B. carpenteri Clark 1908 MG-H11
    |--Hathrometra Clark 1908 WR78
    |    |--H. tenella (Retzius 1783) WR78 (see below for synonymy)
    |    `--H. sarsi [=Antedon sarsi] BK77
    |--Trichometra Clark 1908 WR78
    |    |--T. cubensis (Pourtalès 1869) [=Antedon cubensis; incl. *T. aspera Clark 1908] WR78
    |    `--T. delicata BK77
    `--Phrixometra Clark 1917 WR78
         |--*P. longipinna (Carpenter 1888) [=Antedon longipinna] WR78
         |--P. exigua WR78
         `--P. nutrix B78

Hathrometra tenella (Retzius 1783) WR78 [=Asterias tenella WR78; incl. Alectro dentata Say 1825 WR78, Antedon dentata BK77, *Hathrometra dentata WR78]

*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.

[B78] Breimer, A. 1978. General morphology: recent crinoids. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T9–T58. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Capillasterinae

Nemaster grandis, copyright Edgardo Ochoa.

Belongs within: Comatulidia.

The Capillasterinae are a group of feather stars with a low centrodorsal and usually numerous cirri, with dorsal spines or tubercles on the distal cirri (Wienberg Rasmussen 1978).

Characters (from Wienberg Rasmussen 1978): Centrodorsal rather low discoidal. Cirri rather numerous, in one to three irregular circles. Distal cirrals with dorsal spine or tubercle. Centrodorsal reduced and cirrus-free only in a single species of Capillaster. Articular face of radials with interarticular ligament fossae much wider than high. Border between interarticular and ventral fossae horrizontal or sloping toward interradial edge. Ventral muscular fossae more or less extended outward at interradial margin. Arms divided at primibrachs 2 and commonly further divided at secundibrachs 2, seldom at secundibrachs 1,3, or 4 and tertibrachs 2, 3, or 4. Synarthrial articulation at primibrachs 1 to 2, secundibrachs 1 to 2 and often tertibrachs 1 to 2. Mouth generally excentric to marginal, occasionally central.

Capillasterinae
    |--Comatilia Clark 1909 WR78
    |    `--*C. iridometriformis Clark 1909 WR78
    |--Comanthoides Clark 1931 WR78
    |    `--*C. spanochistum (Clark 1916) [=Comanthus spanochistum] WR78
    |--Comatonia Clark 1916 WR78
    |    `--*C. cristata (Hartlaub 1912) [=Actinometra cristata] WR78
    |--Comissia Clark 1909 WR78
    |    `--*C. luetkeni Clark 1909 WR78
    |--Ctenantedon Meyer 1972 WR78
    |    `--*C. kinziei Meyer 1972 WR78
    |--Leptonemaster Clark 1909 WR78
    |    `--*L. venutus Clark 1909 WR78
    |--Microcomatula Clark 1931 WR78
    |    `--*M. mortenseni Clark 1931 WR78
    |--Palaeocomatella Clark 1912 WR78
    |    `--*P. difficilis (Carpenter 1888) [=Actinometra difficilis] WR78
    |--Capillaster Clark 1909 WR78
    |    |--*C. sentosus (Carpenter 1888) [=Actinometra sentosa] WR78
    |    `--C. multiradiata (Linnaeus 1758) MG-H11
    |--Neocomatella Clark 1909 [=Neomatella (l. c.)] WR78
    |    |--*N. alata (Pourtalès 1878) [=Antedon alata] WR78
    |    `--N. pulchella WR78 [=Actinometra pulchella C50]
    |--Comatella Clark 1908 WR78
    |    |--*C. nigra (Carpenter 1888) [=Actinometra nigra] WR78
    |    |--C. maculata WR78
    |    `--C. stelligera (Carpenter 1888) MG-H11
    `--Nemaster Clark 1909 WR78
         |--*N. grandis Clark 1909 WR78
         |--N. discoidea WR78
         `--N. rubiginosa W79

*Type species of generic name indicated

REFERENCES

[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.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[W79] Warner, G. F. 1979. Aggregation in echinoderms. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 375–396. Academic Press: London.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Comasterinae

Underside of Comanthus mirabilis, copyright Lyle Vail and Anne Hoggett.

Belongs within: Comatulida.

The Comasterinae are a group of feather stars with a marginal or submarginal mouth and much-divided arms.

Characters (from Weinberg Rasmussen 1978): Centrodorsal discoidal, moderate to low, with or without a central depression. Cirrus sockets in one to three irregular circles, or absent. Cirri well developed and stout, or small and weak or reduced. Some or all cirri with spine or tubercle on distal cirrals. Centrodorsal in adult specimens sometimes reduced to a small, stellate plate, slightly or not projecting below radial circlet, and with few, commonly reduced cirrus sockets or without sockets. Radials with low, free surface or concealed in specimens with unreduced centrodorsal; overhanging and with trapezoidal surface when centrodorsal is reduced. Articular face of radials with interarticular ligament fossae high and wide. Ventral muscular fossae low, not projecting at interradial edge, forming narrow bands along almost horizontal ventral edge and slightly curved downward along wide midradial furrow. Arms divided at primibrachs 2 and always further divided, often several times, with intervals of four or, variably, four and two. Brachials 1 to 2 synarthrial except in Comaster, where juvenile cryptosynarthry is commonly changed during growth to a smooth articular face or a syzygy with radiating ridges or a concentric pattern of tubercles. Pinnular comb present not only in oral pinnules but also in some genital pinnules. Mouth marginal or submarginal.

<==Comasterinae
    |--Comantheria Clark 1909 WR78
    |    `--*C. briareus (Bell 1884) [=Antedon briareus] WR78
    |--Comanthina Clark 1909 WR78
    |    `--C. schlegelii Carpenter 1881 (see below for synonymy) WR78
    |--Comaster Agassiz 1836 [incl. Phanogenia Lovén 1866] WR78
    |    |--*C. multiradiata (Lamarck 1816) [=Comatula multiradiata; incl. Alecto multifida Müller 1841] WR78
    |    |--C. brevicirrus (Bell 1894) WR78, MG-H11 [=Phanogenia brevicirra MG-H11]
    |    |--C. formae WR78
    |    |--C. fruticosus WR78
    |    |--‘Phanogenia’ gracilis (Hartlaub 1893) MG-H11
    |    |--‘Phanogenia’ schoenovi (Clark 1909) MG-H11
    |    |--C. seranensis WR78
    |    `--‘*Phanogenia’ typica Lovén 1866 WR78
    `--Comanthus Clark 1908 (see below for synonymy) WR78
         |--C. parvicirrus (Müller 1841) [=Alecto parvicirra; incl. *C. intricata Clark 1908] WR78
         |--‘Alecto’ benneti Müller 1841 [=*Bennetia benneti] WR78
         |--C. gisleni Rowe, Hoggett et al. 1986 MG-H11
         |--C. japonicus BL78
         |--C. mirabilis (Rowe, Hogget et al. 1986) LR06
         |--‘Goldfussia’ multiradiata Norman 1891 WR78
         |--C. schlegelii (Carpenter 1881) LR06
         |--‘Comatula’ trichoptera Müler 1846 [=*Cenolia trichoptera] WR78
         `--C. wahlbergii (Müller 1843) MG-H11

Comanthina schlegelii Carpenter 1881 [=Actinometra schlegelii; incl. A. nobilis Carpenter 1888, *Comanthina nobilis] WR78

Comanthus Clark 1908 [incl. Bennetia Clark 1909, Cenolia Clark 1916, Goldfussia Norman 1891 non Castelnau 1843, Validia Clark 1909, Vania Clark 1911] WR78

*Type species of generic name indicated

REFERENCES

[BL78] Breimer, A., & N. G. Lane. 1978. Ecology and paleoecology. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T316–T347. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[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.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Mariametracea

Heterometra savignyi, copyright Igor Kot.

Belongs within: Solanocrinitacea.

The Mariametracea are a group of feather stars with no aboral star and largely unornamented cirrus sockets.

Characters (from Weinberg Rasmussen 1978): Centrodorsal low hemispherical to discoidal with cirrus-free, flattened to concave dorsal side, which is smooth or tuberculate, seldom rugose or with traces of obliterated cirrus sockets. No dorsal star. Cirrus sockets without distinct ornament or with slightly elevated rim around axial canal closely placed in one to three, seldom four, irregular marginal circles. Dorsal side of cirrals rounded or carinate, with or without dorsal spines. Ventral side of centrodorsal with interradial ridges. Centrodorsal cavity narrow to moderate, less than 0.3 of centrodorsal diameter. Basal rosette but usually no rod-shaped basals. Radials with a low, free surface, commonly concealed in midradial area, very seldom moderate to high. Articular face of radials often rather flat, moderately sloping to almost vertical, and often separated by narrow interradial margins. Interarticular ligament fossae large, high and broad. Ventral muscular fossae generally small, commonly forming a narrow crescentic band along ventral edge and midradial furrow. Wide midradial furrow usually present with or without median ridge, except in Cyllometra which has only a narrow midradial ridge. Radial cavity moderate to large, in juvenile specimens often very large, with spongy calcareous filling. Arms usually dividing at primibrach 2 and further divided with intervals of two or four, often different in inner and outer branches; Eudiocrinus with five arms undivided with first pinnule on brachials 2 and second pinnule on brachials 4, but no pinnule on brachials 3. Synarthrial, flat articulations at brachials 1–2 except for primibrachs 1–2 of Zygometridae and Eudiocrinidae. Syzygial articulations at brachials 3–4 and with variable, often large intervals in distal branches, in Zygometridae and Eudiocrinidae syzygy also at primibrachs 1–2. Oral pinnules may be more or less carinate. One or more proximal pinnules, generally the first inner pinnule (P_a), absent in Colobometridae. Genital and distal pinnules not carinate except for trace of carination in the most proximal pinnulars. Ambulacral covering plates inconspicuous or absent. Mouth central.

Mariametracea [Mariametrida]
    |--Eudiocrinus Carpenter 1882 (see below for synonymy) WR78
    |    |--*E. indivisus (Semper 1868) [=Ophiocrinus indivisus; incl. E. granulatus Bell 1894] WR78
    |    |--E. ornatus WR78
    |    `--E. pinnatus WR78
    |--Zygometridae WR78
    |    |--Catoptometra Clark 1908 WR78
    |    |    `--*C. hartlaubi (Clark 1907) [=Antedon hartlaubi] WR78
    |    `--Zygometra Clark 1907 [incl. Hyponome Lovén 1869] WR78
    |         |--*Z. microdiscus (Bell 1884) WR78 (see below for synonymy)
    |         |--Z. andromeda Clark 1912 MG-H11
    |         |--Z. comata WR78
    |         `--Z. elegans (Bell 1882) MG-H11
    |--Mariametridae [Mariametrinae, Stephanometridae, Stephanometrinae] WR78
    |    |--Dichrometra Clark 1909 WR78
    |    |    `--*D. flagellata (Müller 1841) [=Alecto flagellata; incl. A. elongata Müller 1841] WR78
    |    |--Pelometra Clark 1941 WR78
    |    |    `--*P. ambonensis Clark 1941 WR78
    |    |--Mariametra Clark 1909 WR78
    |    |    |--*M. subcarinata (Clark 1908) [=Himerometra subcarinata] WR78
    |    |    `--M. vicaria (Bell 1894) MG-H11
    |    |--Liparometra Clark 1913 WR78
    |    |    |--*L. grandis (Clark 1908) [=Himerometra grandis] WR78
    |    |    `--L. articulata (Müller 1849) LR06
    |    |--Lamprometra Clark 1913 WR78
    |    |    |--*L. imparipinna (Carpenter 1882) [=Antedon imparipinna] WR78
    |    |    `--L. klunzingeri BL78
    |    |--Oxymetra Clark 1909 [incl. Selenemetra Clark 1911] WR78
    |    |    |--*O. erinacea (Hartlaub 1890) [=Antedon erinacea] WR78
    |    |    `--‘Antedon’ finschii Hartlaub 1890 [=*Selenemetra finschii] WR78
    |    `--Stephanometra Clark 1909 WR78
    |         |--‘Antedon’ protectus Lütken in Carpenter 1879 (see below for synonymy) WR78
    |         |--S. oxyacantha (Hartlaub 1890) LR06
    |         `--S. spinipinna (Hartlaub 1890) LR06
    |--Himerometridae WR78
    |    |--Discometra Gislén 1924 WR78
    |    |    `--*D. rhodanica (Fontannes 1877) (see below for synonymy) WR78
    |    |--Craspedometra Clark 1909 WR78
    |    |    `--*C. acuticirra (Carpenter 1882) (see below for synonymy) WR78
    |    |--Homalometra Clark 1918 WR78
    |    |    `--*H. denticulata (Carpenter 1888) [=Antedon denticulata] WR78
    |    |--Amphimetra Clark 1909 WR78
    |    |    |--‘Alecto’ tesselata Müller 1841 (see below for synonymy) WR78
    |    |    |--A. discoidea C50
    |    |    `--A. ensifer WR78
    |    |--Himerometra Clark 1907 WR78
    |    |    |--H. robustipinna (Carpenter 1881) LR06 (see below for synonymy)
    |    |    |--H. bassleri WR78
    |    |    |--H. grippae WR78
    |    |    |--H. magnipinna (Clark 1908) LR06
    |    |    `--H. martensi WR78
    |    `--Heterometra Clark 1909 [incl. Alecto Leach 1815 (n. d.)] WR78
    |         |--*H. quinquedublicava (Carpenter 1888) (see below for synonymy) WR78
    |         |--H. crenulata (Carpenter 1882) MG-H11
    |         |--‘*Alecto’ horrida Leach 1815 (n. d.) WR78
    |         |--H. reynaudii (Müller 1846) [=Comatula (Alecto) reynaudi] WR78
    |         `--H. savignyi L78
    `--Colobometridae [Pontiometridae] WR78
         |--Alisometra Clark 1947 WR78
         |--Pontiometra Clark 1907 WR78
         |    `--*P. andersoni (Carpenter 1889) [=Antedon andersoni; incl. P. insperatus Clark 1909] WR78
         |--Cotylometra Clark 1916 WR78
         |    `--*C. gracilicirra (Clark 1908) [=Oligometra gracilicirra] WR78
         |--Cyllometra Clark 1907 WR78
         |    `--*C. manca (Carpenter 1888) [=Antedon manca] WR78
         |--Colobometra Clark 1909 WR78
         |    `--*C. perspinosa (Carpenter 1881) [=Antedon perspinosa] WR78
         |--Austrometra Clark 1916 WR78
         |    `--*A. thetidis (Clark 1909) [=Oligometra thetidis] WR78
         |--Basilometra Clark 1936 [incl. Rhadinometra Gislén 1936] WR78
         |    `--*B. boschmai Clark 1936 [incl. *Rhadinometra dawydovi Gislén 1936] WR78
         |--Cenometra Clark 1909 WR78
         |    `--C. bella (Hartlaub 1890) (see below for synonymy) WR78
         |--Clarkometra Gislén 1922 WR78
         |    `--*C. elegans Gislén 1922 WR78
         |--Embryometra Gislén 1938 WR78
         |    `--*E. mortenseni Gislén 1938 WR78
         |--Epimetra Clark 1911 WR78
         |    `--*E. nympha Clark 1911 WR78
         |--Gislenometra Clark 1947 WR78
         |    `--*G. perplexa Clark 1947 WR78
         |--Oligometrides Clark 1913 WR78
         |    `--*O. adeonae (Lamarck 1816) [=Comatula adeonae; incl. Antedon bidens Bell 1884] WR78
         |--Petasometra Clark 1912 WR78
         |    `--*P. clarae (Hartlaub 1890) [=Antedon clarae] WR78
         |--Iconometra Clark 1929 WR78
         |    |--*I. speciosa Clark 1929 WR78
         |    `--I. anisa (Clark 1915) MG-H11
         |--Oligometra Clark 1908 WR78
         |    |--*O. serripinna (Carpenter 1881) (see below for synonymy) WR78
         |    `--O. carpenteri (Bell 1884) MG-H11
         |--Analcidometra Clark 1911 WR78
         |    |--*A. caribbea (Clark 1908) [=Oligometra caribbea] WR78
         |    `--A. armata MMR78
         `--Decametra Clark 1911 [incl. Prometra Clark 1912] WR78
              |--*D. moebius Clark 1911 WR78
              `--‘Colobometra’ chadwicki Clark 1911 [=*Prometra chadwicki] WR78

‘Alecto’ tesselata Müller 1841 [incl. Comatula jacquinoli Müller 1846, C. (Alecto) milberti Müller 1846, *Amphimetra milberti] WR78

‘Antedon’ protectus Lütken in Carpenter 1879 [incl. Himerometra acuta Clark 1908, A. flavomaculata Bell 1894, H. heliaster Clark 1908, A. marginata Carpenter 1888, A. monacantha Hartlaub 1890, *Stephanometra monacantha] WR78

Cenometra bella (Hartlaub 1890) [=Antedon bella; incl. A. abotti Clark 1907, Cenometra delicata Clark 1909, C. insueta Clark 1909, Himerometra unicornis Clark 1908, *C. unicornis] WR78

*Craspedometra acuticirra (Carpenter 1882) [=Antedon acuticirra; incl. A. australis Carpenter 1882, A. bipartipinna Carpenter 1882, A. ludovici Carpenter 1882] WR78

*Discometra rhodanica (Fontannes 1877) [=Eugeniacrinus rhodanicus; incl. Antedon meneghinianus Fontannes 1880] WR78

Eudiocrinus Carpenter 1882 [=Ophiocrinus Semper 1868 nec Salter 1856 nec Charlesworth 1865; Eudiocrinidae, Eudiocrininae] WR78

*Heterometra quinquedublicava (Carpenter 1888) [=Antedon quinquedublicava; incl. A. anceps Carpenter 1888, A. clemens Carpenter 1888] WR78

Himerometra robustipinna (Carpenter 1881) [=Actinometra robustipinna WR78; incl. Antedon crassipinna Hartlaub 1890 WR78, *Himerometra crassipinna WR78]

*Oligometra serripinna (Carpenter 1881) [=Antedon serripinna; incl. O. concinna Clark 1912, O. pulchella Clark 1908] WR78

*Zygometra microdiscus (Bell 1884) WR78 [=Antedon microdiscus WR78; incl. *Hyponome sarsi Lovén 1869 WR78, C50, WR78]

*Type species of generic name indicated

REFERENCES

[BL78] Breimer, A., & N. G. Lane. 1978. Ecology and paleoecology. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T316–T347. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[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.

[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.

[MMR78] Macurda, D. B., Jr., D. L. Meyer & M. Roux. 1978. The crinoid stereom. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T217–T228. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[WR78] Wienberg Rasmussen, H. 1978. Articulata. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 3 pp. T813–T927. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).