Thecostraca

The internal parasite Dendrogaster dichotomus, found in starfishes, from here.

Belongs within: Pancrustacea.
Contains: Thoracica.

The Thecostraca is a crustacean clade containing barnacles (Thoracica) and their close relatives. Within the clade, Ascothoracida and Rhizocephala are parasitic lineages, while Acrothoracica are sessile filter-feeders that burrow into the surface of an animal host. Hansenocaris is known only from juvenile specimens; adults are unknown but are perhaps also parasitic. Despite their divergent appearance as adults, members of the Thecostraca are united by characters of the larvae. The Acrothoracica, Rhizocephala and Thoracica are further united in a clade Cirripedia by features including modification of the antennules in the cypris larva for use in exploratory walking while seeking a settlement site, and in permanent cementation using glandular secretions. In contrast, cypris larvae of the Ascothoracida attach to their intended host by means of direct grasping with the antennules.

See also: The secret of Y-larvae; Forcing out the secret.

Synapomorphies (from Grygier 1987): Nauplii lacking post-maxillular limb buds; compound eyes with less than four crystalline cones per ommatidium; strongly developed frontal filaments associated with eyes.

<==Thecostraca
    |  i. s.: Rhamphoverritor LSE13
    |--Hansenocaris [Facetotecta] G87
    |    |--H. acutifrons G87
    |    |--H. hanseni [=Proteolepas hanseni] G87
    |    |--H. pacifica G87
    |    `--H. rostrata G87
    |--Ascothoracida G87
    |    |  i. s.: Ulophysema oeresundense Brattström 1936 BK77, H75
    |    |         Waginella sandersi G87
    |    |         Parascothorax G87
    |    |         Gorgonolaureus muzikae G87
    |    |         Endosacculus Voigt 1959 H75
    |    |           `--*E. moltkiae Voigt 1959 H75
    |    |--Laurida MD01
    |    |    |--Lauridae MD01
    |    |    |--Petrarcidae MD01
    |    |    `--Synagoga Norman 1888 LS02 [Synagogidae MD01]
    |    `--Dendrogastrida MD01
    |         |--Ctenosculum Heath 1910 BR05 [Ctenosculidae MD01]
    |         |--Ascothorax BK77 [Ascothoracidae MD01]
    |         |    `--A. ophioctenis BK77
    |         `--Dendrogaster BK77 [Dendrogastridae MD01]
    |              |--D. astericola BK77
    |              |--D. dichotomus BK77
    |              |--D. dogieli BK77
    |              |--D. murmanensis [=Myriocladus murmanensis] BK77
    |              `--D. rimskykorsakowi BK77
    `--Cirripedia G87
         |  i. s.: Mitella spinosa O81
         |         Epopella plicata HS01
         |         Notobalanus vestitus HS01
         |         Notomegabalanus decorus HS01
         |         Lepadites Blumenbach 1803 TMNZ64
         |         Pentalasmis vitrea H04
         |         ‘Amalthea’ coxi Sowerby 1908 H08
         |--Thoracica LS02
         |--Acrothoracica LS02
         |    |--Trypetesa Norman 1903 H75 [Apygophora, Trypetesidae MD01]
         |    `--Pygophora MD01
         |         |--Cryptophialidae MD01
         |         `--Lithoglyptidae MD01
         `--Rhizocephala LS02
              |  i. s.: Boschmaella japonica GLT03
              |--Akentrogonida GLT03
              |    |--Clistosaccidae [Sylidae, Sylonidae] MD01
              |    |--Duplorbidae MD01
              |    |--Mycetomorphidae MD01
              |    |--Polysaccidae MD01
              |    |--Chthamalophilidae MD01
              |    `--Thompsoniidae MD01
              `--Kentrogonida GLT03
                   |  i. s.: Peltogasterella GLT03
                   |--Peltogastridae MD01
                   |--Lernaeodiscus G87 [Lernaeodiscidae MD01]
                   |    `--L. porcellanae G87
                   `--Sacculinidae GLT03
                        |--+--Loxothylacus GLT03
                        |  |    |--L. panopaei GLT03
                        |  |    `--L. texanus RS10
                        |  `--+--Heterosaccus GLT03
                        |     |    |--H. californicus GLT03
                        |     |    `--H. dollfusi GLT03
                        |     `--Sacculina GLT03
                        |          |--*S. carcini GLT03
                        |          |--S. anceps B02
                        |          `--S. pinnotherae GLT03
                        `--+--+--‘Sacculina’ sinensis GLT03
                           |  `--+--‘Sacculina’ leptodiae GLT03
                           |     `--+--‘Sacculina’ confragosa GLT03
                           |        `--‘Sacculina’ oblonga GLT03
                           `--Polyascus Glenner, Lützen & Takahashi 2003 GLT03
                                |  i. s.: ‘Sacculina’ bourdoni GLT03
                                |         ‘Sacculina’ gibba GLT03
                                |         ‘Sacculina’ imberbis GLT03
                                |         ‘Sacculina’ insueta GLT03
                                |         ‘Sacculina’ nigra GLT03
                                |         ‘Sacculina’ robusta GLT03
                                |         ‘Sacculina’ schmitti GLT03
                                |--P. gregaria [=Sacculina gregaria] GLT03
                                `--+--*P. polygenea (Lützen & Takahashi 1997) [=Sacculina polygenea] GLT03
                                   `--P. plana [=Sacculina plana] GLT03

Boring: Trypetesa caveata Tomlinson 1963 H75

*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.

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

[B02] Boyko, C. B. 2002. A worldwide revision of the recent and fossil sand crabs of the Albuneidae Stimpson and Blepharipodidae, new family (Crustacea: Decapoda: Anomura: Hippoidea). Bulletin of the American Museum of Natural History 272: 1–396.

[GLT03] Glenner, H., J. Lützen & T. Takahashi. 2003. Molecular and morphological evidence for a monophyletic clade of asexually reproducing Rhizocephala: Polyascus, new genus (Cirripedia). Journal of Crustacean Biology 23: 548–557.

[G87] Grygier, M. J. 1987. New records, external and internal anatomy, and systematic position of Hansen’s y-larvae (Crustacea: Maxillopoda: Facetotecta). Sarsia 72: 261–278.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

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

[H08] Hedley, C. 1908. Studies on Australian Mollusca. Part X. Proceedings of the Linnean Society of New South Wales 33: 456–489, pls 7–10.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

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

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4 (2485): 1–7.

[MD01] Martin, J. W., & G. E. Davis. 2001. An updated classification of the Recent Crustacea. Natural History Museum Los Angeles County, Science Series 39: 1–124.

[O81] O'Brien, C. O. 1981. A. A. Book of New Zealand Wildlife: A guide to the native and introduced animals of New Zealand. Lansdowne Press: Auckland.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[TMNZ64] Teichert, C., R. C. Moore & D. E. Nodine Zeller. 1964. Rhyncholites. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt K. Mollusca 3. Cephalopoda—General Features—Endoceratoidea—Actinoceratoidea—Nautiloidea—Bactritoidea pp. K467–K484. The Geological Society of America and the University of Kansas Press.

Thoracica

Individuals of Calantica spinosa, photographed by Peter E. Smith.

Belongs within: Thecostraca.
Contains: Balanomorpha.

The Thoracica are the barnacles, sessile crustaceans in which the legs have been modified into a filter-feeding fan, while the body is armoured with calcareous plates. The Thoracica have been divided into the Pedunculata (goose barnacles, etc.) in which the body is supported by a fleshy peduncle, and Sessilia (acorn barnacles, etc.) in which there is no peduncle between the body and the substrate; however, it seems likely that the Pedunculata is paraphyletic (Martin & Davis 2001).

<=-=Thoracica
    |  i. s.: Conopea C79
    |         Octolasmis C79
    |         Alepas minuta BK77
    |         Trilasmis kaempferi [incl. Poecilasma aurantium Richard 1903] BK77
    |         Euraphia withersi C03
    |         Tetraclitella karandei Ross 1971 C03
    |         Chamaesipho Y03
    |           |--C. brunnea Y03
    |           |--C. columna Y03
    |           `--C. tasmanica Y03
    |         Octomeris sulcata Y03
    |         Catomerus Y03
    |         Cryptolepas rachianecti W81
    |         Conchoderma auritum W81
    |         Xenobalanus W81
    |           |--X. globicipitis W81
    |           `--X. strictus W81
    |         Austrominius modestus HS01
    |         Striatobalanus amaryllis (Darwin 1854) WMJ09
    |         Austromegabalanus nigrescens WMJ09
    |--Sessilia MD01
    |    |--Neobrachylepadidae [Brachylepadomorpha] MD01
    |    |--Balanomorpha MD01
    |    `--Verrucomorpha MD01
    |         |--Neoverrucidae MD01
    |         `--Verrucidae MD01
    `--Pedunculata MD01
         |--Ibla [Iblidae, Iblomorpha] MD01
         |--Lepadomorpha MD01
         |    |--Oxynaspididae MD01
         |    |--Poecilasmatidae MD01
         |    `--Lepas R85 [Lepadidae MD01]
         |         |--L. anserifera Linnaeus 1767 WMJ09
         |         `--L. australis R85
         |--Heteralepadomorpha MD01
         |    |--Anelasmatidae MD01
         |    |--Heteralepadidae MD01
         |    |--Koleolepadidae MD01
         |    |--Malacolepadidae MD01
         |    |--Microlepadidae MD01
         |    `--Rhizolepadidae MD01
         `--Scalpellomorpha MD01 [Scalpelliformes]
              |--Calanticidae [Calanticinae] JH09
              |    |--Calantica Gray 1825 [incl. Protomitella Broch 1922] JH09
              |    |    |--C. villosa (Leach 1824) [incl. *C. homii Gray 1825] JH09
              |    |    |--C. darwini Jones & Hosie 2009 JH09
              |    |    |--C. moskalebi Zevina & Galkin 1989 JH09
              |    |    |--C. pusilla Utinomi 1970 JH09
              |    |    `--C. spinosa (Quoy & Gaimard 1834) JH09
              |    |--Scillaelepas Seguenza 1876 JH09
              |    |    `--S. ginginensis [=Calantica (Scillaelepas) ginginensis] W26
              |    |--Pisiscalpellum Utinomi 1958 JH09
              |    |--Euscalpellum Hoek 1907 JH09
              |    |--Paracalantica Utinomi 1949 JH09
              |    |--Smilium Gray 1825 JH09
              |    |    |--S. horridum Pilsbry 1912 JH09
              |    |    `--S. hypocrites (Barnard 1924) JH09
              |    |--Capitulum Oken 1815 JH09
              |    |    `--C. mitella C03
              |    |--Zeugmatolepas Withers 1913 JH09
              |    |--Titanolepas Withers 1913 JH09
              |    |--Aurivillialepas Newman 1980 JH09
              |    |--Gruvelialepas Newman 1980 JH09
              |    |--Newmanilepas Zevina & Yakhontov 1987 JH09
              |    |--Cretiscalpellum Withers 1922 JH09
              |    |--Pachyscalpellum Buckeridge 1991 JH09
              |    |--Virgiscalpellum Withers 1935 JH09
              |    |--Zeascalpellum Buckeridge 1983 JH09
              |    `--Crosnieriella Jones 1998 JH09
              |         `--C. acanthosubcarinae Jones 1998 JH09
              |--Pollicipes Leach 1817 JH09 [Pollicipedidae MD01, Pollicipinae]
              |    |--P. acuminatus Darwin 1851 W26
              |    |--P. cornucopia BBB-S95
              |    `--P. spinosus B79
              `--Scalpellum Leach 1817 W26 [Scalpellidae MD01]
                   |--*S. (Scalpellum) scalpellum BK77
                   `--S. (Neoscalpellum Pilsbry 1907) W26
                        |--S. (*N.) dicheloplax W26
                        `--S. (N.) glauerti Withers 1926 W26

*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.

[BBB-S95] Boubezari, K., G. Bitar & D. Bellan-Santini. 1995. Structure et organisation de trois moulières (Mytilus galloprovincialis et Perna perna) de la région d'Alger. Mésogée 54: 63-72.

[B79] Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: Why colonial animals have more hiding places. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds) pp. 459-497. Academic Press: London.

[C03] Chan, B. K. K. 2003. Studies on Tetraclita squamosa and Tetraclita japonica (Cirripedia: Thoracica) II: Larval morphology and development. Journal of Crustacean Biology 23 (3): 522-547.

[C79] Crisp, D. J. 1979. Dispersal and re-aggregation in sessile marine invertebrates, particularly barnacles. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds) pp. 319-327. Academic Press: London.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45-80.

[HJ08] Huisman, J. M., D. S. Jones, F. E. Wells & T. Burton. 2008. Introduced marine biota in Western Australian waters. Records of the Western Australian Museum 25 (1): 1-44.

[JH09] Jones, D. S. & A. M. Hosie. 2009. A new species of Calantica from Western Australian waters (Thoracica: Scalpellomorpha: Calanticidae). Records of the Western Australian Museum 25 (3): 239-246.

[MD01] Martin, J. W., & G. E. Davis. 2001. An updated classification of the Recent Crustacea. Natural History Museum Los Angeles County, Science Series 39: 1-124.

[R85] Robertson, C. J. R. (ed.) 1985. Reader’s Digest Complete Book of New Zealand Birds. Reader’s Digest: Sydney.

[W81] Watson, L. 1981. Sea Guide to Whales of the World. Hutchinson: London.

[WMJ09] Wells, F. E., M. Mulligan & D. S. Jones. 2009. Prevention of introduction of species brought into Geraldton Harbour, Western Australia, by the dredge Leonardo da Vinci. Records of the Western Australian Museum 25 (2): 153-158.

[W26] Withers, T. H. 1926. The crinoid Marsupites, and a new cirripede from the Upper Cretaceous of Western Australia. Journal of the Royal Society of Western Australia 12 (11): 97-104, pl. XI.

[Y03] Yan, Y. 2003. Larval development of the barnacle Chinochthmalus scutelliformis (Cirripedia: Chthmalidae) reared in the laboratory. Journal of Crustacean Biology 23 (3): 513-521.

Balanomorpha

Armatobalanus allium on horny coral, from Shahdadi et al. (2014).

Belongs within: Thoracica.
Contains: Tetraclita, Balanidae, Pyrgomatidae.

The Balanomorpha are the clade of barnacles containing the familiar acorn barnacles. Members of the clade are characterised by a symmetrical wall formed of 1, 4, 6 or 8 plates, and with the operculum usually formed from paired terga and scuta. At least the first two pairs of cirri are modified as mouth appendages with the third pair also being somewhat modified in Coronuloidea and more extensively modified in Balanoidea.

See also: Reference review: Barnacles among the coral.

<==Balanomorpha
    |--Chionelasmatidae [Chionelasmatoidea] MD01
    |    |--Chionelasmus darwini BWW93
    |    `--Eochionelasmus BWW93
    |--Pachylasma BWW93 [Pachylasmatidae, Pachylasmatoidea MD01]
    |    `--P. veteranum Buckeridge 1983 BWW93
    |--Tetraclitoidea MD01
    |    |--Bathylasma JB12 [Bathylasmatidae MD01]
    |    |    `--B. rangatira Buckeridge 1983 BWW93
    |    `--Tetraclitidae C03
    |         |--Tetraclita C03
    |         |--Tesseropora rosea (Krauss 1848) HJ08
    |         |--Eopopella eosimplex Buckeridge 1983 BWW93
    |         `--Elminius C03
    |              |--E. modestus DL77
    |              `--E. plicatus Gray 1843 C03
    |--Coronuloidea MD01
    |    |--Platylepas C79 [Platylepadidae MD01]
    |    |--Chelonibia C79 [Chelonibiidae MD01]
    |    |    |--C. patula C79
    |    |    `--C. testudinaria A99
    |    `--Coronulidae MD01
    |         |--Emersonius cybosyrinx Ross 1967 BWW93
    |         `--Coronula W81
    |              |--C. balaenaris W81
    |              |--C. diadema W81
    |              `--C. reginae W81
    |--Balanoidea MD01
    |    |--Balanidae MD01
    |    `--+--Pyrgomatidae A92
    |       `--Archaeobalanidae A92
    |            |--Solidobalanus BWW93
    |            `--Armatobalanus Hoek 1913 [Archaeobalaninae] A92
    |                 |--A. allium (Darwin 1854) [=Balanus allium] A92
    |                 |--A. arcuatus (Hoek 1913) [=Balanus (Armatobalanus) arcuatus; incl. Cantellius tredecimus] A92
    |                 |--A. cepa (Darwin 1854) [=Balanus cepa] A92
    |                 |--A. circe (Kolosvary 1947) [=Balanus circe] A92
    |                 |--A. filigranus (Broxh 1916) [=Balanus filigranus] A92
    |                 |--A. funiculorum (Annandale 1906) [=Balanus funiculorum] A92
    |                 |--A. nefrens (Zullo 1963) [=Balanus (Armatobalanus) nefrens] A92
    |                 |--A. oryza (Broch 1931) [=Balanus (Armatobalanus) oryza] A92
    |                 |--A. palaoensis (Hiro 1937) [=Balanus (Armatobalanus) palaoensis] A92
    |                 |--A. quadrivittatus (Darwin 1854) [=Balanus quadrivittatus] A92
    |                 |--A. quinquevittatus (Hoek 1913) [=Balanus (Armatobalanus) quinquevittatus] A92
    |                 `--A. terebratus (Darwin 1854) [=Balanus terebratus] A92
    `--Chthamaloidea MD01
         |--Catophragmidae MD01
         |    |--Catophragmus polymerus H04
         |    `--Pachydiadema cretaceum BWW93
         `--Chthamalidae Y03
              |--Hexelasma hirsutum JB12, PP64
              |--Chinochthamalus Foster 1980 Y03
              |    `--*C. scutelliformis (Darwin 1854) [=Chamaesipho scutelliformis] Y03
              `--Chthamalus Y03
                   |--C. anisopoma R96
                   |--C. antennatus H04
                   |--C. challengeri Y03
                   |--C. dalli Y03
                   |--C. depressus PP64
                   |--C. fissus Y03
                   |--C. fragilis RS10
                   |--C. malayensis Y03
                   |--C. montagui Southward 1976 C03
                   `--C. stellatus Poli 1971 C03

*Type species of generic name indicated

REFERENCES

[A92] Anderson, D. T. 1992. Structure, function and phylogeny of coral-inhabiting barnacles (Cirripedia, Balanoidea). Zoological Journal of the Linnean Society 106: 277–339.

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (transl. 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[BWW93] Briggs, D. E. G., M. J. Weedon & M. A. Whyte. 1993. Arthropoda (Crustacea excluding Ostracoda). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 321–342. Chapman & Hall: London.

[C03] Chan, B. K. K. 2003. Studies on Tetraclita squamosa and Tetraclita japonica (Cirripedia: Thoracica) II: larval morphology and development. Journal of Crustacean Biology 23 (3): 522–547.

[C79] Crisp, D. J. 1979. Dispersal and re-aggregation in sessile marine invertebrates, particularly barnacles. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 319–327. Academic Press: London.

[DL77] Dinamani, P., & P. A. Lenz. 1977. Some aspects of spatfall of the New Zealand rock oyster during 1974. Veliger 20 (1): 17–26.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[HJ08] Huisman, J. M., D. S. Jones, F. E. Wells & T. Burton. 2008. Introduced marine biota in Western Australian waters. Records of the Western Australian Museum 25 (1): 1–44.

[JB12] Johnson, M. E., & B. G. Baarli. 2012. Development of intertidal biotas through Phanerozoic time. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 63–128. Springer.

[MD01] Martin, J. W., & G. E. Davis. 2001. An updated classification of the Recent Crustacea. Natural History Museum Los Angeles County, Science Series 39: 1–124.

[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.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[R96] Roff, D. A. 1996. The evolution of threshold traits in animals. Quarterly Review of Biology 71 (1): 3–35.

[W81] Watson, L. 1981. Sea Guide to Whales of the World. Hutchinson: London.

[Y03] Yan, Y. 2003. Larval development of the barnacle Chinochthmalus scutelliformis (Cirripedia: Chthmalidae) reared in the laboratory. Journal of Crustacean Biology 23 (3): 513–521.

Last updated: 7 March 2022.