Scuticoxifera

Dorsal and lateral figures of female Tainisopus fontinalis, from here.

Belongs within: Eumalacostraca.
Contains: Oniscidea, Valvifera, Sphaeromatidea, Limnoriidea, Cymothoida.

The Scuticoxifera are a major clade within the isopods, uniting the oniscideans, valviferans, anthuroideans, and the polyphyletic 'flabelliferans'.

Synapomorphies (from Brandt & Poore 2003): Coxae I-VII transformed into plates in contact with entire length of tergite; penes medial on pereonite VII; anterior filter channels of stomach curving laterally or in transverse position (except Anthuroidea).

<==Scuticoxifera [Cymothoada, Cymothoidea, Flabellifera]
    |--+--Oniscidea BP03
    |  `--+--Valvifera BP03
    |     `--Sphaeromatidea BP03
    `--+--Tainisopidae [Tainisopidea] BP03
       |    |--Pygolabis Wilson 2003 BP03
       |    `--Tainisopus Wilson & Ponder 1992 BP03
       |         `--T. fontinalis Wilson & Ponder 1992 BP03
       `--+--Phoratopus [Phoratopidea, Phoratopodidae] BP03
          |    `--P. remex Hale 1925 BP03
          `--+--Limnoriidea BP03
             `--Cymothoida BP03

*Type species of generic name indicated

REFERENCES

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893-923.

Thermosbaenacea

Thermosbaena mirabilis, from Michigan Science Art.

Belongs within: Eumalacostraca.

The Thermosbaenacea are a group of small crustaceans that are only known from cave or marine interstitial habitats. Many species are found in waters with raised salinities.

Characters (from Stock 1986): Small (≤4 mm); strictly stygobiont. Carapace fused with first thoracomere; dorsal brood pouch present in female, internally connected with branchial chamber; oostegites absent. Free larval stages absent. Adults with five (Thermosbaena) or seven pereiopods. Penis present on 8th thoracomere of male.

<==Thermosbaenacea [Pancarida]
    |--Tulumellidae MD01
    |--Halosbaena Stock 1976 S86 [Halosbaenidae MD01]
    |    `--H. acanthura Stock 1976 S86
    `--+--Thermosbaenidae MD01
       |    |--Theobaena cambodjana Cals & Boutin 1985 S86
       |    |--Thermosbaena Monod 1924 S86
       |    |    `--T. mirabilis Monod 1924 S86
       |    `--Limnosbaena Stock 1976 S86
       |         `--L. finki (Meštrov & Lattinger-Penko 1969) S86
       `--Monodella Ruffo 1949 S86 [Monodellidae MD01]
            |--M. argentarii Stella 1951 [incl. M. halophila Karaman 1953] S86
            |--M. atlantomaroccana Cals 1985 S86
            |--M. relicta Por 1962 S86
            |--M. sanctaecrucis Stock 1976 S86
            |--M. somala Chelazzi & Messana 1982 S86
            |--M. stygicola Ruffo 1949 S86
            `--M. texana Maguire 1965 S86

*Type species of generic name indicated

REFERENCES

[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.

[S86] Stock, J. H. 1986. Thermosbaenacea. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 585-588. E. J. Brill / Dr. W. Backhuys: Leiden.

Aegidae

Specimen of Rocinela belliceps attached to a spotted ratfish Hydrolagus colliei, photographed by Upupa4me.

Belongs within: Cymothoida.

The Aegidae are a family of marine isopods, members of which are external parasites of fish. The first three pairs of pereopods are modified for clinging to their host. Members of the genus Aega have the maxillipedal palp reduced, so that it has only two segments, whereas five palpal segments are present in Rocinella (Menzies & Kruczynski 1983).

Characters (from Menzies & Kruczynski 1983): Peduncles and flagella of antennae well defined. First three pairs of pereopods prehensile, last four pairs ambulatory. Pleopods with plumose marginal setae. Maxillipedal palp bi- or pentarticulate, last two articles with series of stout recurved setae at apex. Maxilla I with single lobe; maxilla II with two dissimilar lobes armed with recurved teeth. Pleon of five subequal somites, followed by pleotelson bearing plumose marginal setae at apex.

<==Aegidae
    |--Barybrotes Schioedte & Meinert 1979 BP03
    |--Aega Leach 1815 MK83
    |    |--*A. psora (Linnaeus 1761) [=Oniscus psora; incl. A. emarginata Leach 1815] MK83
    |    |--A. antartica Hodgson 1910 BP03
    |    |--A. antillensis Schiödte & Meinert 1880 [incl. A. excisa Richardson 1910] MK83
    |    |--A. dentata Schiödte & Meinert 1880 MK83
    |    |--A. ecarinata Richardson 1898 MK83
    |    |--A. gracilipes Hansen 1895 MK83
    |    |--A. incisa Schiödte & Meinert 1880 MK83
    |    |--A. komai Bruce 1996 BP03
    |    |--A. ornata Richardson 1911 MK83
    |    |--A. plebeia Hansen 1897 BP03
    |    |--A. symmetrica H47
    |    |--A. tenuipes Schiödte & Meinert 1880 MK83
    |    `--A. webbii (Guérin 1836) MK83
    `--Rocinela Leach 1818 T08
         |--*R. danmonensis Leach 1818 MK83
         |--R. angustata H47
         |--R. belliceps H47
         |    |--R. b. belliceps H47
         |    `--R. b. pugettensis Hatch 1947 H47
         |--R. cubensis Richardson 1898 MK83
         |--R. dumerili (Lucas 1849) T08
         |--R. insularis Schiödte & Meinert 1880 MK83
         |--R. laticauda H47
         |--R. oculata Harger 1883 MK83
         |--R. orientalis Schioedte & Meinert 1879 T08
         |--R. propodialis H47
         |--R. quadrata BP03
         |--R. signata Schiödte & Meinert 1880 [incl. R. aries Schiödte & Meinert 1880] MK83
         `--R. tridens Hatch 1947 H47

*Type species of generic name indicated

REFERENCES

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893–923.

[H47] Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. University of Washington Publications in Biology 10 (5): 155–274.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1–126.

[T08] Trilles, J.-P. 2008. Some marine isopods from the Senckenberg Research Institute (Frankfurt am Main, Germany) (Crustacea, Isopoda: Cymothoidae, Aegidae, Corallanidae, Cirolanidae). Senckenbergiana Biologica 88 (1): 21–28.

Corallanidae

Dorsal and ventral view of specimen of Excorallana sp., photographed by Artour A.

Belongs within: Cymothoida.

The Corallanidae are a family of marine isopods. Most are parasites of fish, but some parasitise crustaceans (Brandt & Poore 2003).

Characters (from Menzies & Kruczynski 1983): Flagella and peduncles of antennae I and II well defined. Exite of maxilla with setae or single recurved hook or tooth. Last four pairs of pereopods ambulatory. Pleon with four or five somites in dorsal view, plus pleotelson; apex of pleotelson fringed with plumose marginal setae and stout two-pointed setae. Pleopods natatory with plumose setae marginally.

<==Corallanidae [Excorallanidae]
    |--Lanocira Hansen 1890 MK83
    |--Tachaea caridophaga (Riek 1953) BP03
    |--Alcirona Hansen 1890 [Alcironidae] T08
    |    `--*A. krebsii Hansen 1890 (see below for synonymy) MK83
    |--Nalicora Moore 1902 MK83
    |    `--*N. rapax Moore 1902 [=Excorallana rapax] MK83
    |--Argathona Stebbing 1905 [Argathonidae] T08
    |    |--A. macronema (Bleeker 1857) [incl. Brotherus longicornis Budde-Lund 1908] T08
    |    |--A. rhinoceros (Bleeker 1857) T08
    |    `--A. rostrata BP03
    |--Corallana Dana 1853 T08
    |    |--C. brevipes Schioedte & Meinert 1879 T08
    |    |--C. collaris Schioedte & Meinert 1879 T08
    |    |--C. hirsuta Schioedte & Meinert 1879 T08
    |    |--C. hirticauda Dana 1853 T08
    |    `--C. nodosa Schioedte & Meinert 1879 T08
    `--Excorallana Stebbing 1904 T08
         |--E. antillensis (Hansen 1890) [=Corallana antillensis, Exocorallana (l. c.) antillensis] MK83
         |--E. fissicauda (Hansen 1890) [=Corallana fissicauda] MK83
         |--E. mexicana Richardson 1905 MK83
         |--E. oculata (Hansen 1890) [=Corallana oculata] MK83
         |--E. quadricornis (Hansen 1890) [=Corallana quadricornis] MK83
         |--E. sexticornis (Richardson 1901) [=Corallana sexticornis] MK83
         |--E. subtilis (Hansen 1890) [=Corallana subtilis] MK83
         |--E. tricornis (Hansen 1890) [=Corallana tricornis, Exocorallana (l. c.) tricornis] MK83
         |    |--E. t. tricornis BP03
         |    `--E. t. occidentalis BP03
         `--E. warmingii (Hansen 1890) [=Corallana warmingii] MK83

*Alcirona krebsii Hansen 1890 [incl. A. hirsuta Moore 1902, A. insularis Hansen 1890, A. maldivensis Stebbing 1904] MK83

*Type species of generic name indicated

REFERENCES

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893-923.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1-126.

[T08] Trilles, J.-P. 2008. Some marine isopods from the Senckenberg Research Institute (Frankfurt am Main, Germany) (Crustacea, Isopoda: Cymothoidae, Aegidae, Corallanidae, Cirolanidae). Senckenbergiana Biologica 88 (1): 21-28.

Asellota

Female Munna minuta, from here.

Belongs within: Eumalacostraca.
Contains: Asellidae, Stenasellidae, Protojaniridae, Microparasellidae, Janiridae.

The Asellota are a group of aquatic isopods. The members of the Aselloidea are freshwater, while the members of the other superfamilies are marine. The Janiroidea are marked by a tendency to progressive fusion of the free pleonites into a single pleotelson (Henry et al. 1986).

Characters (from Menzies & Kruczynski 1983): First and second pairs of pleopods modified, unlike succeeding biramours pairs. Uropods usually biramous, tubular, leaf-like or absent. Pleon composed of one to three somites including pleotelson. Antenna II with or without antennal scale.

<==Asellota
    |  i. s.: Antias Richardson 1906 [Antiasidae] MK83
    |           |--*A. charcoti Richardson 1906 MK83
    |           `--A. milleri Menzies & Glynn 1968 MK83
    |         Munnoides caribea Carvacho 1977 MK83
    |         Notasellus N29
    |           `--‘Stenetrium’ inerme N29
    |--Protallocoxidae [Protallocoxoidea] MD01
    |--Aselloidea HLM86
    |    |--Asellidae HLM86
    |    `--Stenasellidae HLM86
    |--Gnathostenetroidoidea [Parastenetroidea] HLM86
    |    |--Protojaniridae HLM86
    |    |--Vermectias BP03 [Vermectiadidae MD01]
    |    |    `--V. nelladanae Just & Poore 1992 BP03
    |    `--Gnathostenetroididae HLM86
    |         |--Neostenetroides Carpenter & Magniez 1982 HLM86
    |         |    `--N. stocki Carpenter & Magniez 1982 HLM86
    |         `--Caecostenetroides Fresi & Schiecke 1968 HLM86
    |              |--C. ischitanum Fresi & Schiecke 1968 HLM86
    |              `--C. nipponicum Nunomura 1975 HLM86
    |--Stenetrioidea HLM86
    |    |--Pseudojaniridae [Pseudojaniroidea] MD01
    |    `--Stenetrium Haswell 1881 [Stenetriidae] HLM86
    |         |--*S. armatum Haswell 1881 MK83
    |         |--S. chiltoni N29
    |         |--S. crassimanus N29
    |         |--S. dagama N29
    |         |--S. fractum N29
    |         |--S. glauerti Nicholls 1929 N29
    |         |--S. haswelli N29
    |         |--S. longicorne N29
    |         |--S. macrochirium Nicholls 1929 N29
    |         |--S. minocule Menzies & Glynn 1968 MK83
    |         |--S. serratum Hansen 1905 MK83
    |         |--S. siamense N29
    |         |--S. spinirostrum Nicholls 1929 N29
    |         |--S. stebbingi Richardson 1902 (see below for synonymy) MK83
    |         |--S. suldhana N29
    |         `--S. truncatum Nicholls 1929 N29
    `--Janiroidea HLM86
         |  i. s.: Protocharon Chappuis & Delamare 1956 C86
         |           |--P. antarctica Chappuis 1958 C86
         |           `--P. arenicola Chappuis & Delamare 1956 C86
         |--Microparasellidae C86
         |--Janiridae C86
         |--Paramunnidae [Abyssianiridae, Pleurogoniidae] MD01
         |--Desmosomatidae [Pseudomesidae] MD01
         |--Acanthaspididae MD01
         |--Dendrotiidae MD01
         |--Echinothambematidae MD01
         |--Haplomunnidae MD01
         |--Ischnomesidae MD01
         |--Janirellidae MD01
         |--Joeropsididae MD01
         |--Katianiridae MD01
         |--Mesosignidae MD01
         |--Mictosomatidae MD01
         |--Nannoniscidae MD01
         |--Santiidae MD01
         |--Thambematidae MD01
         |--Munnopsididae [Ilyarachnidae] MD01
         |    |--Ilyarachna antarctica [incl. I. bicornis] BH88
         |    `--Bathybadistes spinosissima [=Ilyarachna spinosissima] BH88
         |--Haploniscus B88 [Haploniscidae MD01]
         |    `--H. tangaroae B88
         |--Munna H47 [Munnidae MD01]
         |    |--M. boecki BK77
         |    |--M. kroyeri H47
         |    `--M. minuta H47
         |--Jaeropsis Koehler 1885 H47, MK83 [Jaeropsidae]
         |    |--*J. curvicornis (Nicolet 1849) [=Jaera curvicornis] MK83
         |    |--J. brevicornis BBB-S95
         |    |--J. coralicola Schultz & McCloskey 1967 MK83
         |    |--J. lobata H47
         |    `--J. rathbunae Richardson 1902 MK83
         `--Macrostylis Sars 1864 M03 [Macrostylidae MD01]
              |--M. amplinexa Mezhov 1989 M03
              |--M. capito Mezhov 1989 M03
              |--M. confinis Mezhov 2003 M03
              |--M. medioxima Mezhov 2003 M03
              |--M. minuscularia Mezhov 2003 M03
              |--M. prolixa Mezhov 2003 M03
              |--M. sensitiva Birstein 1970 M03
              `--M. urceolata Mezhov 1989 M03

Stenetrium stebbingi Richardson 1902 [incl. S. antillense Hansen 1905, S. gilbertense Nordenstam 1946, S. medipacificum Miller 1941, S. occidentale Hansen 1905] MK83

*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.

[B88] Boxshall, G. A. 1988. A new genus of tantulocaridan (Crustacea: Tantulocarida) parasitic on a harpacticoid copepod from Tasmania. Bulletin of the British Museum (Natural History), Zoology series 54 (6): 271–274.

[BH88] Boxshall, G. A., & K. Harrison. 1988. New nicothoid copepods (Copepoda: Siphonostomatoida) from an amphipod and from deep-sea isopods. Bulletin of the British Museum (Natural History), Zoology series 54 (6): 285–299.

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893–923.

[C86] Coineau, N. 1986. Isopoda: Asellota: Janiroidea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 465–472. E. J. Brill/Dr. W. Backhuys: Leiden.

[H47] Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. University of Washington Publications in Biology 10 (5): 155–274.

[HLM86] Henry, J.-P., J. J. Lewis & G. Magniez. 1986. Isopoda: Asellota: Aselloidea, Gnathostenetroidoidea, Stenetrioidea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 434–464. E. J. Brill/Dr. W. Backhuys: Leiden.

[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.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1–126.

[M03] Mezhov, B. V. 2003. New abyssal species of the genus Macrostylis G. O. Sars, 1864 (Crustacea: Isopoda: Macrostylidae) from the northwestern part of the Indian Ocean. Arthropoda Selecta 12: 1–8.

[N29] Nicholls, G. E. 1929. Some new species of Stenetrium from Western Australia. Proceedings of the Linnean Society of New South Wales 54 (4): 361–374.

Anthuroidea

Diagnostic features of Paranthura floridensis, from Menzies & Kruczynski (1983).

Belongs within: Cymothoida.
Contains: Cyathura.

The Anthuroidea are a group of elongate, predatory marine isopods. The scientific name refers the 'flower-like' appearance of the lateral uropods.

See also: Flower-tails.

Characters (from Menzies & Kruczynski 1983): Cylindrical, body over six times as long as wide. Pleon consisting of six somites plus telson (though some pleonites may be fused). Uropods lateral, often folding over telson. Mouthparts piercing and sucking, or cutting or biting.

<==Anthuroidea [Aberrantia, Anthuridea]
    |  i. s.: Belura pilara Poore & Lew Ton 1988 BP03
    |         Kupellonura biriwa Poore & Lew Ton 1988 BP03
    |--Oscanthura [Hyssuridae] B86
    |--Expanathuridae MD01
    |--Leptanthura B86 [Leptanthuridae MD01]
    |--Anthelura MK83 [Antheluridae MD01]
    |    `--A. affinis Richardson 1902 MK83
    |--Paranthuridae B86
    |    |--Curassanthura Kensley 1981 B86
    |    |    `--C. halma Kensley 1981 B86
    |    |--Cruregens Chilton 1881 B86
    |    |    `--C. fontanus Chilton 1881 B86
    |    |--Accalathura Barnard 1925 MK83
    |    |    |--*A. crenulata (Richardson 1901) [=Calathura crenulata] MK83
    |    |    `--A. crassa Barnard 1925 MK83
    |    |--Colanthura B86
    |    |    |--C. coeca B86
    |    |    |--C. ornata Carvacho 1977 MK83
    |    |    `--C. tenuis Richardson 1902 MK83
    |    `--Paranthura Bate & Westwood 1868 MK83
    |         |--*P. costana Bate & Westwood 1868 MK83
    |         |--P. antillensis Barnard 1925 MK83
    |         |--P. barnardi Paul & Menzies 1971 MK83
    |         |--P. floridensis Menzies & Kruczynski 1983 MK83
    |         `--P. infundibulata Richardson 1902 [incl. P. verrillii Richardson 1902] MK83
    `--Anthuridae B86
         |--Cyathura B86
         |--Xenanthura brevitelson Barnard 1925 MK83
         |--Pendanthura tanaiformis Menzies & Glynn 1968 MK83
         |--Ptilanthura tricarina Menzies & Frankenberg 1966 MK83
         |--Apanthuroides millae Menzies & Glynn 1968 MK83
         |--Horoloanthura irpex Menzies & Frankenberg 1966 MK83
         |--Eisothistos atlanticus Vanhöffen 1914 MK83
         |--Panathura formosa Menzies & Frankenberg 1966 MK83
         |--Apanthuretta B86
         |    |--A. lathridia Wägele 1982 B86
         |    `--A. pori B86
         |--Caenanthura B86
         |    |--C. indica B86
         |    `--C. siamensis B86
         |--Skuphonura Barnard 1925 MK83
         |    |--*S. laticeps Barnard 1925 MK83
         |    `--S. lindae Menzies & Kruczynski 1983 MK83
         |--Malacanthura MK83
         |    |--M. caribbica Paul & Menzies 1971 MK83
         |    `--M. cumanensis Paul & Menzies 1971 MK83
         |--Haliophasma MK83
         |    |--H. curri Paul & Menzies 1971 MK83
         |    |--H. irmae Paul & Menzies 1971 MK83
         |    `--H. valeriae Paul & Menzies 1971 MK83
         |--Apanthura MK83
         |    |--A. magnifica Menzies & Frankenberg 1966 MK83
         |    |--A. signata Menzies & Glynn 1968 MK83
         |    `--A. significa Paul & Menzies 1971 MK83
         `--Mesanthura Barnard 1914 MK83
              |--*M. catenula (Stimpson 1855) [=Anthura catenula] MK83
              |--M. decorata Menzies & Glynn 1968 MK83
              |--M. floridensis Menzies & Kruczynski 1983 MK93
              |--M. looensis Kensley & Schotte 1987 BP03
              |--M. paucidens Menzies & Glynn 1968 MK83
              `--M. pulchra Barnard 1925 MK83

Nomen nudum: Curassanthura canariensis Wägele in Botosaneanu 1986 B86

*Type species of generic name indicated

REFERENCES

[B86] Botosaneanu, L. 1986. Isopoda: Anthuridea. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 428-433. E. J. Brill / Dr. W. Backhuys: Leiden.

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893-923.

[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.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1-126.

Synocheta

Trichoniscus pusillus, photographed by Krister Hall.

Belongs within: Oniscidea.

The Synocheta are a clade of small slaters restricted to damp habitats. Some species have become aquatic cave-dwellers. Males of this group have the genital ducts fusing within the penis, so that they penis bears only a single opening.

Synapomorphies (from Schmidt 2008): Male genital ducts distally fused; eyes with at most 3 ommatidia; remotor muscles of pleopod protopodites M9, 11 and 13 composed of 2 strands each.

<==Synocheta MD01
    |--Styloniscoidea MD01
    |    |--Schoebliidae MD01
    |    |--Titaniidae MD01
    |    |--Tunanoniscidae MD01
    |    `--Styloniscidae [Patagoniscidae] MD01
    |         |--Notoniscus [Notoniscinae] Va73
    |         |    `--N. tasmanicus (Chilton 1915) Va73
    |         `--Styloniscus Dana 1852 [Styloniscinae] Va73
    |              |--S. (Styloniscus) Va73
    |              |    |--S. (S.) australiensis Vandel 1973 Va73
    |              |    |    |--S. a. australiensis Va73
    |              |    |    `--S. a. cavernicolus Vandel 1973 Va73
    |              |    |--S. (S.) otakensis (Chilton 1901) [=Trichoniscus otakensis] Va73
    |              |    |--S. (S.) squarrosus Green 1961 Va73
    |              |    |--S. (S.) thomsoni (Chilton 1901) [=Trichoniscus thomsoni] Va73
    |              |    `--S. (S.) verrucosus (Budde-Lund 1906) [=Trichoniscus verrucosus] Va73
    |              `--S. (Euroniscus Vandel 1973) Va73
    |                   `--S. (*E.) commensalis (Chilton 1910) Va73
    `--Trichoniscoidea [Endophora] MD01
         |--Buddelundiellidae MD01
         `--Trichoniscidae MD01
              |  i. s.: Cantabroniscus Vandel 1965 S86
              |           `--C. primitivus Vandel 1965 S86
              |         Scotoniscus Racovitza 1908 S86
              |           `--S. macromelos Racovitza 1908 S86
              |         Bureschia Verhoeff 1926 S86
              |           `--B. bulgarica Verhoeff 1926 S86
              |         Typhlotricholigioides Rioja 1953 S86
              |           `--T. aquaticus Rioja 1953 S86
              |         Brackenridgia Eigenmann & Ulrich 1902 [incl. Protrichoniscus] S86
              |           |--B. bridgesi (van Name 1942) S86
              |           |--B. cavernarum M90
              |           |--B. heroldi M90
              |           `--B. reddelli M90
              |         Titanethes Schioedte 1849 S86
              |           |--T. (Titanethes) S86
              |           |    |--T. (T.) albus Schioedte 1849 S86
              |           |    |--T. (T.) biseriatus Verhoeff 1900 S86
              |           |    `--T. (T.) dahli Verhoeff 1926 S86
              |           `--T. (Cyphonethes Verhoeff 1926) S86
              |                `--T. (C.) hercegovinensis Verhoeff 1900 S86
              |         Alpioniscus Racovitza 1908 S86
              |           |  i. s.: A. feneriensis Vu73
              |           |         A. kratochvili Frankenberger 1939 UO05
              |           |--A. (Illyrionethes Verhoeff 1927) S86
              |           |    |--A. (I.) balthasari PN02
              |           |    |--A. (I.) bosniensis Frankenberger 1939 S86
              |           |    `--A. (I.) heroldii Verhoeff 1931 S86
              |           `--A. (Macedonethes Buturović 1955) S86
              |                `--A. (M.) skopjensis Buturović 1955 S86
              |         Hyloniscus riparius M90
              |         Haplophthalmus danicus M90
              |         Trichoniscoides Sars 1899 M90, S86
              |           |--T. mixtus Racovitza 1908 S86
              |           `--T. sarsi M90
              |         Amerigoniscus paynei M90
              |         Titana mirabilis N55
              `--Trichoniscinae H47
                   |--Cordioniscus stebbingi H47
                   |--Oregoniscus Hatch 1947 H47
                   |    `--*O. nearcticus [=Trichoniscus nearcticus] H47
                   |--Miktoniscus H47
                   |    |--M. halophilus M90
                   |    `--M. medcofi H47
                   |--Androniscus H47
                   |    |--A. dentiger H47
                   |    `--A. subterraneus Vu73
                   `--Trichoniscus D37
                        |--T. elisabethae D37
                        |--T. flavescens Lucas 1846 E12
                        |--T. provisorius M90
                        |--T. pusillus [incl. T. demivirgo] M90
                        `--T. pygmaeus H47

*Type species of generic name indicated

REFERENCES

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[H47] Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. University of Washington Publications in Biology 10 (5): 155–274.

[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.

[M90] Muchmore, W. B. 1990. Terrestrial Isopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 805–817. John Wiley & Sones: New York.

[N55] Noirot, C. 1955. Termites du centre et du sud-ouest de l'Angola récoltés par A. de Barros Machado. Diamang Publicações Culturais 27: 139–150.

[PN02] Poggi, R., G. Nonveiller, A. Colla, D. Pavićević & T. Rađa. 2001–2002. Thaumastocephalini, a new tribe of Pselaphinae for Thaumastocephalus folliculipalpus n. gen., n. sp., an interesting new troglobious species from central Dalmatia (Croatia) (Coleoptera, Staphylinidae, Pselaphinae). Annali del Museo Civico di Storia Naturale “Giacomo Doria” 94: 1–20.

Schmidt, C. 2008. Phylogeny of the terrestrial Isopoda (Oniscidea): a review. Arthropod Systematics and Phylogeny 66 (2): 191–226.

[S86] Sket, B. 1986. Isopoda: Oniscidea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 482–485. E. J. Brill / Dr. W. Backhuys: Leiden.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[Va73] Vandel, A. 1973. Les isopodes terrestres de l'Australie. Étude systématique et biogéographique. Mémoires du Muséum National d'Histoire Naturelle, Série A, Zoologie 82: 1–171.

[Vu73] Vuilleumier, F. 1973. Insular biogeography in continental regions. II. Cave faunas from Tessin, southern Switzerland. Systematic Zoology 22 (1): 64–76.

Oniscidea

Sea slater Ligia oceanica, photographed by Mark Robinson.

Belongs within: Scuticoxifera.
Contains: Synocheta, Crinocheta.

The Oniscidea contain the terrestrial isopods, the familiar woodlice. Characters supporting this group include a minute antenna I, and pseudotracheal pleopods (Brandt & Poore 2003). Some members of the group are amphibious or littoral, but most are fully terrestrial. Tylos are supra-littoral isopods distinguished by the ability to roll into a ball, and the presence of a well-developed process on the front of the head. The Ligiidae, rock slaters, are found on rocky shores and are characterised by a long second antenna with ten or more flagellar articles, and very long and slender uropod rami (Muchmore 1990).

<==Oniscidea [Cloportides, Oniscides, Oniscodea, Oniscoida]
    |--Tylos Latreille 1826 BP03 [Tylinae, Tylidae, Tylomorpha MD01]
    |    |--*T. latreillei B94
    |    |--T. australis Lewis & Bishop 1990 BP03
    |    |--T. europaeus PP64
    |    |--T. neozelanicus V73
    |    |--T. niveus M90
    |    |--T. nudulus V73
    |    |--T. punctatus M90
    |    `--T. sardous PP64
    `--Ligiamorpha MD01
         |--+--Synocheta MD01
         |  `--Crinocheta MD01
         `--Diplocheta MD01
              |--Mesoniscus M78 [Mesoniscidae MD01]
              |    `--M. alpicola M78
              `--Ligiidae MD01
                   |--Euphiloscia elrodii H47
                   |--Ligioides Wahrberg 1922 V73
                   |    `--L. intermedius Wahrberg 1922 V73
                   |--Ligidium M90
                   |    |--L. elrodii M90
                   |    |--L. (Ligidium) gracile H47
                   |    |--L. longicaudatum H47
                   |    `--L. splendidum Strouhal 1940 BWW93
                   `--Ligia Fabricius 1798 JP13 [incl. Ligyda MK83]
                        |--L. australiensis Dana 1853 [=L. gaudichaudii var. australiensis] V73
                        |--L. exotica Roux 1828 V73
                        |--L. italica PP64
                        |--L. novaezelandiae HS01
                        |--L. oceanica H47
                        `--L. (Ligia) pallasii H47

Oniscidea incertae sedis:
  Mexiconiscus Schultz 1964 [incl. Xilitloniscus Bowman 1965] S86
    `--M. laevis (Rioja 1956) S86
  Lucasius pallidus [=Porcellio pallidus] S77
  Trichorhina [Squamiferidae] M90
    |--T. australiensis Wahrberg 1922 V73
    |--T. heterophthalma M90
    |--T. tomentosa M90
    `--T. triocis M90
  Alloniscus Dana 1854 [Alloniscidae] JP13
    |--A. nicobaricus Budde-Lund 1885 V73
    |--A. oahuensis Budde-Lund 1885 V73
    |--A. pallidulus Budde-Lund 1885 V73
    `--A. perconvexus H47

*Type species of generic name indicated

REFERENCES

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BP03] Brandt, A., & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893–923.

[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.

[H47] Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. University of Washington Publications in Biology 10 (5): 155–274.

[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.

[JP13] Judd, S., & G. Perina. 2013. An illustrated key to the morphospecies of terrestrial isopods (Crustacea: Oniscidea) of Barrow Island, Western Australia. Records of the Western Australian Museum Supplement 83: 185–207.

[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.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1–126.

[M90] Muchmore, W. B. 1990. Terrestrial Isopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 805–817. John Wiley & Sones: New York.

[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.

[S77] Schmalfuss, H. 1977. Eine neue Nagurus-Art aus der Süd-Agäis (Crustacea: Isopoda: Oniscoidea: Trachelipidae). Senckenbergiana Biologica 57 (4–6): 359–365.

[S86] Sket, B. 1986. Isopoda: Oniscidea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 482–485. E. J. Brill/Dr W. Backhuys: Leiden.

[V73] Vandel, A. 1973. Les isopodes terrestres de l'Australie. Étude systématique et biogéographique. Mémoires du Muséum National d'Histoire Naturelle, Série A, Zoologie 82: 1–171.

Last updated: 21 May 2019.

Valvifera

Male and female of Arcturella dilatata, from here.

Belongs within: Scuticoxifera.
Contains: Mesidoteinae, Idothea.

The Valvifera are a group of aquatic isopods found in both marine and freshwater habitats. They are characterised by having the uropods rotated ventrally to form a branchial chamber enclosing the pleopods (Brandt & Poore 2003).

Characters (from Brandt & Poore 2003): Antenna 1 without scale. Mandibular lacinia mobilis present; mandibular molar a cylindrical process with triturative flat end, occasionally obsolete; mandibular palps absent. Dorsal coxal plate 7 similar to others (rarely narrower). Pleonite 6, usually more pleonites, fused to pleotelson, pleonite 1 similar to others (rarely reduced). Pleotelson underside vaulted, branchial chamber defined by ridges along mesial margin of lateral edge. Pleopod 3 endopod roughly triangular (or at least with inner angle sharper than outer). Uropods lateral to margin of pleotelson, peduncle and short rami enclosing pleopods in branchial chamber.

<==Valvifera [Idoteides, Idoteoidea, Idotheoidea]
    |  i. s.: Ronalea Menzies & Bowman 1956 MK83
    |         Eusymmerus Richardson 1899 MK83
    |--Antarcturus MK83 [Antarcturidae BP03]
    |--Arcturides MK83 [Arcturididae BP03]
    |--Austracturellidae BP03
    |--Holidoteidae BP03
    |--Pseudidotheidae BP03
    |--Rectarcturidae BP03
    |--Xenarcturellidae BP03
    |--Saduria [Chaetiliidae] BP03
    |    `--S. entomon BP03
    |--Holognathus [Holognathidae] BP03
    |    `--H. karamea BP03
    |--Arcturidae [Amesopodidae] BP03
    |    |--Arcturus MK83
    |    |    |--A. deshayesii Lucas 1846 E12
    |    |    `--A. floridanus Richardson 1900 MK83
    |    |--Thermoarcturus venezuelensis Paul & Menzies 1971 MK883
    |    |--Dolichisus MK83
    |    |--Neoarcturus MK83
    |    |--Pseudarcturella MK83
    |    |--Microarcturus MK83
    |    |--Edwinjoycea Menzies & Kruczynski 1983 MK83
    |    |    `--*E. horologium Menzies & Kruczynski 1983 MK83
    |    |--Arcturella Sars 1899 MK83
    |    |    |--*A. dilatata (Sars 1882) [=Astacilla dilatata] MK83
    |    |    |--A. bispinata Menzies & Kruczynski 1983 MK83
    |    |    `--A. spinata Menzies & Kruczynski 1983 MK83
    |    `--Astacilla MK83
    |         |--A. cymodocea Menzies & Glynn 1968 MK83
    |         |--A. lasallae Paul & Menzies 1971 MK83
    |         `--A. lauffi Menzies & Frankenberg 1966 MK83
    `--Idoteidae N37
         |  i. s.: Cleantis MK83
         |           |--C. planicauda Benedict in Richardson 1899 MK83
         |           `--C. tubicola HS01
         |         Edotea triloba (Say 1818) MK83
         |         Tropedotea Menzies & Kruczynski 1983 MK83
         |           `--*T. lyonsi Menzies & Kruczynski 1983 MK83
         |         Chiridotea N37
         |           |--C. arenicola Wigley 1960 MK83
         |           |--C. coeca (Say 1818) MK83
         |           |--C. excavata Harper 1974 MK83
         |           `--C. stenops Menzies & Frankenberg 1966 MK83
         |         Erichsonella MK83
         |           |--E. attenuata (Harger 1873) MK83
         |           |--E. filiformis (Say 1818) MK83
         |           |    |--E. f. filiformis MK83
         |           |    |--E. f. isabelensis Menzies 1951 MK83
         |           |    `--E. f. tropicalis Menzies & Glynn 1968 MK83
         |           `--E. floridana Benedict in Richardson 1901 MK83
         |         Synidotea N37
         |           |--S. angulata H47
         |           |--S. bicuspida [=Edotea bicuspida, Edotia bicuspida] H47
         |           |--S. consolidata H47
         |           |--S. hirtipes N37
         |           |--S. nebulosa H47
         |           |--S. nodulosa [=Edotea nodulosa] H47
         |           |--S. pettiboneae Hatch 1947 H47
         |           `--S. ritteri H47
         |         Symmius N37
         |         Paridotea N37
         |         Pentidotea N37
         |           |--P. resecata N37 [=Idotaea resecata H47, Idotea resecata H47]
         |           |--P. rotunda N37
         |           |--P. stenops H47
         |           |--P. whitei N37 [=Idothea whitei H47, Idotea whitei H47]
         |           `--P. wosnesenskii N37 (see below for synonymy)
         |         Synisoma F05
         |           |--S. albertoi Castellanos & Junoy 2005 F05
         |           `--S. capito BBB-S95
         |--Mesidoteinae N37
         |--Glyptonotus [Glyptonotinae] N37
         |    |--G. acutus N37
         |    `--G. antarcticus Eights 1852 FZA07
         `--Idothea N37

Pentidotea wosnesenskii N37 [=Idotaea wosnesenskii H47, Idotea wossnessenskii H47; incl. Pentidotea wosnesenskii var. exlineae Hatch 1947 H47]

*Type species of generic name indicated

REFERENCES

[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.

[BP03] Brandt, A. & G. C. B. Poore. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships. Invertebrate Systematics 17: 893–923.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[FZA07] Fiege, D., H. Zibrowius & P. M. Arnaud. 2007. New deep-water records of cocoons of undescribed species of Fecampiidae from Antarctica to the Bay of Biscay (Platyhelminthes, Turbellaria, Rhabdocoela). Senckenbergiana Biologica 87 (1): 1–6.

[H47] Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. University of Washington Publications in Biology 10 (5): 155–274.

[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.

[MK83] Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1–126.

[N37] Nicholls, G. E. 1937. On the freshwater Idoteidae of New Zealand (Crustacea, Isopoda). Annals and Magazine of Natural History, series 10, 19: 113–136.

Epicaridea

Generalised male of Entoniscidae, from S. J. Keable et al..

Belongs within: Cymothoida.
Contains: Bopyridae.

The Epicaridea are a group of isopods parasitic on other crustaceans. The females have slightly to greatly asymmetrical bodies, and both pairs of antennae are rudimentary (Menzies & Kruczynski 1983).

<==Epicaridea [Bopyrida, Epicarida] MD01
    |  i. s.: Microniscus [Microniscidae] MD01
    |--Bopyroidea MD01
    |    |--Bopyridae MD01
    |    |--Dajidae MD01
    |    `--Entoniscidae MD01
    `--Cryptoniscoidea MD01
         |--Asconiscidae MD01
         |--Cabiropidae [Cabiropsidae] MD01
         |--Crinoniscidae MD01
         |--Cryptoniscidae MD01
         |--Cyproniscidae MD01
         |--Hemioniscidae MD01
         `--Podasconidae MD01

*Type species of generic name indicated

REFERENCES

[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.

Menzies, R. J., & W. L. Kruczynski. 1983. Isopod Crustacea (exclusive of Epicaridea). Memoirs of the Hourglass Cruises 6 (1): 1-126.

Squillidae

The mantis shrimp Squilla empusa, photographed by Richard King.

Belongs within: Hoplocarida.

The Squillidae is a family of mantis shrimps whose members have spearing claws, adapted for catching soft prey (Manning 1980).

Characters (from Manning 1980): Propodi of posterior three maxillipeds slender, not beaded or ribbed ventrally. Telson with distinct median carina. At most submedian teeth of telson with movable apices. Four or more intermediate denticles present on telson.

<==Squillidae [Harpiosquillidae, Squilloidea] MD01
    |--Meiosquilla C73
    |    |--M. quadridens (Bigelow 1893) [=Squilla quadridens] C73
    |    `--M. schmitti (Lemos de Castro 1955) [=Squilla schmitti] C73
    `--Squilla Fabricius 1787 B02
         |--S. chydaea Manning 1962 C73
         |--S. deceptrix Manning 1969 C73
         |--S. discors Manning 1962 C73
         |--S. edentata (Lunz 1937) C73
         |--S. empusa Say 1818 C73
         |--S. granti G75
         |--S. grenadensis Manning 1969 C73
         |--S. heptacantha C73
         |--S. interrupta G75
         |--S. investigatoris H15
         |--S. massawensis P71
         |--S. neglecta Gibbes 1850 [=Chloridella neglecta] C73
         |--S. prasinolineata Dana 1852 C73
         |--S. raphidea G75
         `--S. rugosa Bigelow 1893 C73
              |--S. r. rugosa C73
              `--S. r. pinensis (Lunz 1937) [=Chloridella rugosa var. pinensis] C73

*Type species of generic name indicated

REFERENCES

[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.

[C73] Camp, D. K. 1973. Stomatopod Crustacea. Memoirs of the Hourglass Cruises 3 (2): 1-100.

[G75] Grant, E. M. 1975. Guide to Fishes. The Co-ordinator-General's Department: Brisbane (Australia).

[H15] Harmer, S. F. 1915. The Polyzoa of the Siboga Expedition – Part I. Entoprocta, Ctenostomata and Cyclostomata. E. J. Brill: Leyden.

Manning, R. B. 1980. The superfamilies, families, and genera of recent stomatopod Crustacea, with diagnoses of six new families. Proceedings of the Biological Society of Washington 93 (2): 362-372.

[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.

[P71] Por, F. D. 1971. One hundred years of Suez Canal—a century of Lessepsian migration: retrospect and viewpoints. Systematic Zoology 20 (2): 138-159.

Bivalvia

Outer shell (above) and inner mould of Fordilla troyensis, from Walcott (1886).

Belongs within: Mollusca.
Contains: Daonella, Halobia, Polidevciidae, Sanguinolariidae, Ferganoconcha, Astartila, Myonia, Perrierinidae, Palaeomutelidae, Pleurophorus, Opponobrachia, Anatinidae, Leptonidae, Palaeoheterodonta, Anomalodesmata, Pteriomorphia, Heterodonta.

The Bivalvia are one of the main subgroups of the molluscs, readily distinguished from other classes by the division of the shell into two hinged lateral valves. Bivalves also lack a developed head and mostly feed by filtering organic particles from the water (with some notable exceptions). The valves of the shell are connected at the hinge by a ligament of keratinous proteins that may also be partially calcified. The hinge may also include a series of interlocking teeth on each valve. In members of the clade Autobranchia, the gills are enlarged and folded within the mantle cavity to function in filter-feeding. More basal bivalve lineages have the gills as simple non-reflected lamellae that function for respiration only, with the labial palps as the main food-collecting organs. Bivalves outside the clade Amarsipobranchia have at least the first few anterior filaments of the gills inserted unfused into a distal oral groove. In most members of the Amarsipobranchia, however, these filaments are fused or not inserted (Plazzi et al. 2011).

The earliest fossils that have been assigned to the bivalves come from the Early and Middle Cambrian and include the genera Fordilla, Pojetaia and Arhouriella (Budd & Jensen 2000). These are small forms, only a few millimetres in length, that may have had a burrowing lifestyle similar to that of the modern Nuculidae (Morton 2012). Fordilla and Pojetaia both had laterally compressed, subequally valved and suboval shells with few teeth in the hinge. Fordilla has a small, anteriorly placed umbo whereas that of Pojetaia is central to subcentral in position. Arhouriella had a veneriform shell with two teeth overhanging the hinge plate on the right valve (the left valve is unknown).

See also: Pig's toes and water nymphs.

Characters (from Giribet & Wheeler 2002): Body laterally compressed with an external bivalved shell that is hinged dorsally. Valves connected by a partially calcified elastic ligament and held together by 1 or 2 adductor muscles. Buccal or radular apparatus absent, mantle lobes either joined or free ventrally. Mantle cavity spacious, extending upwards on each side of visceral mass and containing a pair of laterally suspended ctenidia. Ctenidia may be enlarged, lamellate and plicate. Mouth and anus located at opposite ends of body, gut typically convoluted. Pair of ciliated labial palps connecting ctenidia to the mouth, directing food particles into it. Foot extensible, either elongated or laterally compressed.

<==Bivalvia [Acephala, Arcacea, Aviculae, Cryptodonta, Nuculiformii, Nuculoida, Palaeotaxodonta, Pelecypoda, Protobranchia]
    |--Fordillidae [Fordilloida] C96
    |    |--Neofordilla elegans Krasilova 1977 SB93
    |    |--Pojetaia Jell 1980 C96, M12
    |    |    `--P. runnegari Jell 1980 SB93
    |    `--Fordilla BJ00
    |         |--F. sibirica P08
    |         `--F. troyensis Barrande 1881 M12
    `--+--Opponobranchia PC11
       `--Autobranchia (see below for synonymy) PC11
            |  i. s.: Anatinidae W77
            |         Mactromyopsis [Mactromyidae] SB93
            |           `--M. (Mactromyella) inflata (Thévenin 1909) SB93
            |         Montanaria devonicus (Beushausen 1884) SB93
            |         Plesiocyprinella carinata Holdhaus 1918 SB93
            |         Leptonidae SB93
            |         Vokesella inopinata Chavan 1952 SB93
            |         Neoleptonidae SB93
            |           |--Goodalliopsis terminalis (Deshayes 1858) SB93
            |           |--Neolepton Monterosato 1875 F26
            |           |    `--N. citrinum F26
            |           `--Puyseguria Powell 1927 SB93, P61
            |                |--*P. cuneata Powell 1927 P61
            |                |--P. crenulifera Maxwell 1969 SB93
            |                |--P. prognata Powell 1927 P61
            |                `--P. tani Powell 1939 P61
            |         Cardiolarioidea C04
            |           |  i. s.: Cardiolaria beirensis C96
            |           |         Eritropis C04
            |           `--Praenuculidae SB93
            |                |--Ledopsis JB12
            |                |--Nuculodonta JB12
            |                `--Deceptrix C04
            |                     |--D. carinata Fuchs 1919 SB93
            |                     `--D. levata C96
            |--Palaeoheterodonta PC11
            `--Amarsipobranchia PC11
                 |--Anomalodesmata PC11
                 `--+--Pteriomorphia PC11
                    `--Heterodonta PC11

Bivalvia incertae sedis:
  Crytodonta saffordi PR79
  Pyrenomoeus EB01
  Batolites Montfort 1808 D-DP03
    `--B. organisans Montfort 1808 D-DP03
  Lapeirousiinae D-DP03
    |--Lapeirousia Bayle 1878 D-DP03
    |--Praelapeirousia D-DP03
    `--Lapeirousella D-DP03
  Plicatomytilus AV03
  Cataceramus HS03
    |--C. barabini (Morton 1834) HS03
    `--C. subcircularis (Meek 1876) HS03
  Horiopleura lamberti HS02
  Polyconites verneuili HS02
  Pseudotoucasia santanderensis HS02
  Sphaerulites HS02
  Chondrodonta HS02
  Nodipecten nodosus TL03
  Callithaca adamsii L10
  Swiftopecten swifti GAS03
  Liocyma XZ10
    |--L. chinensis XZ10
    `--L. fluctuosa GAS03
  Alveinus ojianus GAS03
  Panomya arctica KBC03
  Sertifer keenae KBC03
  Gouldia M62
    |--G. cerina WE63
    |--G. minima M62 [=Circe minima C64]
    `--G. pacifica C64
  Carolia placunoides G92
  Cubitostrea G92
  Nicaisolopha G92
  Devonia perrieri (Malard 1903) (see below for synonymy) BK77
  Halobiidae [Halobioidea] S04
    |--Daonella S04
    `--Halobia KI70
  Natasia boliviensis SV03
    |--N. b. boliviensis SV03
    `--N. b. fragilis SV03
  Arisaigia postornata P98
  Potamocorbula XZ10
    |--P. amurensis (Schrenck 1867) XZ10
    |--P. laevis (Hinds 1843) XZ10
    `--P. ustulata (Reeve 1865) XZ10
  Polidevciidae SB93
  Pseudophytina rugifera GW02
  Mactromeris polynyma GR98
  Sarepta Adams 1860 [Sareptidae] DK08
    |--S. obolella (Tate 1886) [=Leda obolella) F26
    `--S. planiuscula (Tate 1886) [=Leda planiuscula, Ovaleda planiuscula] DK08
  Saxicava [Saxicavidae] F26
    |--S. antarctica F26
    |--S. australis F26
    |--S. norvegica N79
    |--S. pholadis Middendorff 1851 N79 (see below for synonymy)
    |--S. rugosa [incl. S. rugosa var. praecisa] U79
    `--S. subalata F26
  Barcoona Finlay 1927 [=Pachydomella Etheridge 1907 non Ulrich 1891] F27
    `--*B. chutus (Etheridge 1907) [=*Pachydomella chutus] F27
  Modiolarca impacta HS01
  Pseudoarcopagia disculus HS01
  Varinucula gallinacea HS01
  Melina cumingii (Reeve 1858) [=Perna cumingii] H09
  Crenatula H09
    |--C. flammea Reeve 1858 H09
    `--C. picta (Gmelin 1791) [=Ostrea picta] H09
  Meleagrina H09
    |--M. anomioides (Reeve 1857) [=Avicula anomioides] H09
    |--M. fimbriata [incl. M. mazatlanica] C64
    |--M. flexuosa (Reeve 1857) [=Avicula flexuosa] H09
    |--M. lucunata (Reeve 1857) [=Avicula lucunata] H09
    |--M. margaritifera (Linnaeus 1758) [=Mytilus margaritiferus] H09
    |--M. maxima (Jameson 1901) [=Margaritifera maxima] H09
    |--M. sugillata (Reeve 1857) [=Avicula sugillata] H09
    `--M. tegulata (Reeve 1857) [=Avicula tegulata] H09
  Aenigma aenigmatica [=Tellina aenigmatica] H09
  Placenta H09
    |--P. lobata Soerby 1871 H09
    `--P. placenta (Linnaeus 1758) [=Anomia placenta] H09
  Myrina coppingeri Smith 1885 H09
  Coelodon elongatus Carpenter 1846 H09
  Crassatellites H09
    |--C. janus Hedley 1906 H09
    |--C. rhomboides (Smith 1885) [=Crassatella rhomboides] H09
    |--C. torresi (Smith 1885) [=Crassatella torresi] H09
    `--C. ziczac [=Crassatella ziczac] H09
  Carditella H09
    |--C. infans Smith 1885 H09
    `--C. torresi Smith 1885 H09
  Opisocardium subretusum (Sowerby 1841) [=Cardium subretusum] H09
  Glaucomya H09
    |--G. desmoyersii H09
    `--G. rugosa Hanley 1828 [=Glauconomya rugosa] H09
  Metis H09
    |--M. meyeri (Philippi 1846) [=Tellina meyeri] H09
    `--M. spectabilis (Hanley 1844) [=Tellina spectabilis] H09
  Phylloda foliacea (Linnaeus 1758) [=Tellina foliacea] H09
  Sanguinolariidae P61
  Psammosolen H09
    |--P. australis (Dunker 1861) [=Macha australis] H09
    |--P. coarctatus (Gmelin 1791) [=Solen coarctatus] H09
    |--P. deshayesii (Dunker 1861) [=Macha deshayesii] H09
    |--P. minutus (Dunker 1861) [=Azor minutus] H09
    `--P. sulcatus (Dunker 1861) [=Macha sulcatus] H09
  Labiosa grayi (Adams 1872) [=Raeta grayi] H09
  Davila plana (Hanley 1843) [=Mesodesma plana] H09
  Roccellaria pupina (Deshayes 1855) [=Gastrochaena pupina] H09
  Nausitoria H09
    |--N. manni Wright 1865 H09
    `--N. thoracites (Gould 1856) [=Calobates thoracites] H09
  Margarita Leach 1814 BR05
    |--M. maculata [incl. M. elegantula Jeffreys 1864 (n. n.)] J64
    `--M. minutissima Mighels 1843 J49
  Peloris Poli 1791 BR05
  ‘Venilia’ Morton 1833 non Rafinesque 1815 BR05
  Fasciculiconcha guangxiensis ZP86
  Aloidis [Aloididae] M54
    `--A. iredalei M54
  Conradilla caelata USDI77
  Cyrtonaias tampicoensis USDI77
  Plethobasis USDI77
    |--P. cicatricosus USDI77
    |--P. cooperianus USDI77
    `--P. cypha B66
  Eotrigonia semiundulata Q72
  Papyridea BD86
    |--P. australe (Sowerby 1834) BD86
    `--P. papyraceum (Gmelin 1791) BD86
  Bathymodiolus Kenk & Wilson 1985 AJH07
    `--B. brevior R01
  Mesosaccella Chavan 1946 SY85
  Danlengiconcha Lu 1978 K84
  Volsellina zhesiensis Liang in Ding, Xia et al. 1984 DX84
  Ferganoconcha ZZ88
  Nakamuranaia ZZ88
    |--N. chingshanensis ZZ88
    |--N. subrotunda ZZ88
    `--N. zhejiangensis ZZ88
  Sibireconcha ZZ88
    |--S. anodontoides ZZ88
    `--S. jenissejensis ZZ88
  Solenaia menyinensis ZZ88
  Pseudocardiniidae SB93
    |--Tutuella chachlovi Ragozin 1938 SB93
    |--Utschamiella tungussica Ragozin 1938 SB93
    `--Kija SB93
         |--K. kibetenensis Lebedev 1958 SB93
         `--K. tjazhinensis Lebedev 1958 SB93
  Actinoceramus KK08
    |--A. concentricus KK08
    `--A. salomoni (d’Orbigny 1850) KK08
  Actinopterella F71
    |--A. lamellosa Sherrard 1959 F71
    `--A. minuta Sherrard 1959 F71
  Actinopteria F71 [incl. Clathrocoelia Hall 1879 F62]
    |--A. australis Dun 1896 F71
    `--A. decussata [incl. *Clathrocoelia eborica Hall 1879] F62
  Astartila F71
  Aucella F71
    |--A. hausmanni G31
    `--A. hughendensis F71
  Girtypecten tessellatus [=Aviculopecten tessellatus] F71
  Burrier dunii Laseron 1910 F71
  Stutchburia F71
    |--S. compressa F71
    |--S. costata [=Cardinia costata] F71
    |--S. farleyensis Etheridge 1900 F71
    |--S. obliqua Etheridge 1900 F71
    |--S. praerupta (Dana 1849) [=Cypricardia praerupta] F71
    |--S. recta (Dana 1849) [=Cardinia recta] F71
    |--S. simplex (Dana 1849) [=Cypricardia simplex] F71
    `--S. veneris [=Cypricardia veneris] F71
  Cleobis F71
    |--C. grandis (Dana 1849) (see below for synonymy) F71
    `--C. gryphoides [=Cardiomorpha gryphoides, Megadesmus (Megadesmus) gryphoides] F71
  Corimya primula F71
  Cypricardinia Hall 1859 SB93
    |--C. gregarius (Etheridge 1900) (see below for synonymy) F71
    |--C. sinensis G31
    `--C. subelegans G31
  Cytheria moorei F71
  Diaphragmella merismopteroides Etheridge 1918 F71
  Permopecten aviculatum [=Entolium aviculatum] F71
  Glyptodesma bundanoonensis Etheridge 1917 F71
  Glyptoleda F71
    |--G. glomerata Fletcher 1945 F71
    `--G. reidi Fletcher 1945 F71
  Grammatodon daintreei Etheridge 1907 F71
  Grammysioides declivis Sherrard 1959 F71
  Maccoyella F71
    |--M. barklyi F71
    |--M. corbiensis F71
    |--M. mariaeburnensis C16
    |--M. reflecta F71
    `--M. umbonalis F71
  Myonia F71
  Nucundata darwini [=Nuculana darwini, Phestia darwini, Tellinomya darwini] F71
  Pachydomus sacculus M’Coy 1847 F71
  Protovirgus F71
    |--P. dunstani (Etheridge 1888) [=Unio dunstani] F71
    `--P. flemingi McMichael 1956 F71
  Solenomya translineata Laseron 1910 F71
  Tatella aptiana F71
  Nototrigonia cinctuta (Etheridge 1902) [=Trigonia cinctuta] F71
  Undulomya rugulata Fletcher 1946 F71
  Unionella F71
    |--U. bowralensis Etheridge 1888 F71
    |--U. carnei Etheridge 1888 F71
    `--U. wianamattensis (Etheridge 1888) [=Unio wianamattensis] F71
  Mysidioptera Salomon 1895 [=Latemaria Wilckens 1909] W60
    |--*M. ornata Salomon 1895 (see below for synonymy) W60
    |--M. exima Diener 1913 W60
    |--M. globosa Broili 1930 [=M. (Latemaria) globosa] W60
    |--M. globularis (Trechmann 1918) [=Megalodon globularis] W60
    |--M. kittlii Bittner 1895 [=M. (*Latemaria) kittlii] W60
    |--M. obliqua Broili 1903 W60
    |--M. riceae Waterhouse 1960 W60
    |--M. vixcostata W60
    `--M. woehrmanni Salomon 1895 W60
  Hokonuia Trechmann 1918 W60
    `--H. limaeformis Trechmann 1918 (see below for synonymy) W60
  Cardiomorpha nuggetensis Trechmann 1918 W60
  Foramelina exempla MG-H11
  Fissilunula clarkei H75
  Apidopholas Fischer 1887 H75
  Karadjalia venusta H79
  Volviceramus K79
    |--V. exogyroides K79
    `--V. involutus K79
  Cremnoceramus K79
    |--C. inconstans K79
    |--C. koeneni K79
    `--C. wandereri K79
  Endocostea K79
    |--E. baltica K79
    `--E. typica K79
  Enantiostreon difforme (Schlotheim 1820) KL04
  Eurynia D56 [=Euryma (l. c.) B64]
    `--E. iris D56
  Camya Hinz-Schallreuter 1995 M12
  Arhouriella Geyer & Streng 1998 M12
  Mixtipecten amuriensis (Woods 1917) S98
  Pectenoproductus Likharev 1930 M65
    `--*P. proprius Likharev 1930 M65
  Halophiala Koken 1925 KC60
  Shaanxiconcha cliovata HH09
  Perrierinidae P61
  Obovaria V16
    |--O. levigata (Rafinesque 1820) [=Unio levigata, Elliptio levigata; incl. Obovaria lens Lea 1831] V16
    |--O. olivaria (Rafinesque 1820) (see below for synonymy) V16
    |--O. retusa Lam. 1819 [incl. O. torsa Rafinesque 1820, Unio torsa] V16
    `--O. subrotunda (Rafinesque 1820) (see below for synonymy) V16
  Symphynota viridis (Rafinesque 1820) [=Unio viridis, Elliptio viridis] V16
  Hiatula H86
    |--H. compacta C64
    |--H. diphos F66
    |--H. incerta H86
    `--H. nuttalli C64
  Mysia neozelanica H86
  Solenella H86
    |--S. australis [incl. Nucula ornata] H86
    `--S. funiculata H86
  Macrodon H86
    |--M. australis H86
    |--M. kingianum G31
    |--M. naliwkini G31
    |--M. striatum G31
    `--M. verneuilianum G31
  Mactrinula dolabrata (Reeve 1854) XZ10
  Raetellops pulchella (Adams & Reeve 1850) XZ10
  Coelomactra antiquata (Spengler 1802) XZ10
  Nitidotellina iridella (Mastensi 1868) XZ10
  Cyclina XZ10
    |--C. sinensis (Gmelin 1791) XZ10
    `--C. subquadrata C64
  Trigonothracia XZ10
    |--T. jinxingae Xu 1980 XZ10
    `--T. pusilla (Gould 1861) XZ10
  Avella sinica (Wang & Qi 1984) XZ10
  Talonostrea talonata Li & Qi 1994 XZ10
  Tugonia XZ10
    |--T. anatina PP64
    `--T. huanghaiensis Xu 1987 XZ10
  Sinolyonsia sinica Xu 1992 XZ10
  Porterius dalli (Smith 1885) XZ10
  Asthenothaerus XZ10
    |--A. huaughaensis Xu 1989 XZ10
    `--A. villosior C64
  Crassithracia ovata (Xu 1989) XZ10
  Bulluniella borealis P08
  Pseudomyonia Runnegar 1983 P08
  Gastrana fragilis PP64
  Alba tenuis PP64
  ‘Pleurodonta’ Conrath 1887 non Herrmanssen 1847 BR17
  Caesariella JB12
  Paleostraba JB12
  Chesapecten middlesexensis P12
  Mansfieldostrea geraldjohnsoni P12
  Dualina T08
    |--‘Nautilus’ desideratus Barrande 1865–1867 T08
    `--D. longiuscula (Barrande 1881) T08
  Algerina algena Kříž 2008 T08
  Sainshandia V-RR93
    |--S. bainshirensis V-RR93
    |--S. dushiulensis V-RR93
    |--S. mongolica V-RR93
    `--S. ongonulensis V-RR93
  Pseudohyria V-RR93
    |--P. cardiiformis V-RR93
    |--P. debilicostata V-RR93
    `--P. marginodentata V-RR93
  Plicatotrigonioides V-RR93
    |--P. gobiaensis V-RR93
    `--P. khamariensis V-RR93
  Gogiola multicostata V-RR93
  Buginella V-RR93
    |--B. buginica V-RR93
    `--B. pseudoanodonta V-RR93
  Plicatella transaltica V-RR93
  Carnalpia rostrata Křiž 1974 D94
  Thoralia [Thoraliidae] SB93
    `--T. languedociana (Thoral 1935) SB93
  Pristiglomidae SB93
  Siliculidae SB93
  Lametilidae SB93
  Tindaria [Tindariidae] SB93
    `--T. arata Bellardi 1875 SB93
  Pachymytilus Zittel 1881 SB93
  Pseudopachymytilus Krumbeck 1923 SB93
  Boiomytilus Růžička & Prantl 1961 SB93
  Glendella dickinsi Runnegar 1970 SB93
  Sichuania marwicki Waterhouse 1980 SB93
  Carditomantea spinata Quenstedt 1929 SB93
  Cypricardella Hall 1858 SB93
  Tellidorella Berry 1962 SB93
  Rhaetidia zittellii Bittner 1895 SB93
  Ceromyella miotaurina Sacco 1901 SB93
  Roxoa intrigans (Mendes 1944) SB93
  Prilukiellidae SB93
    |--Amnigeniella kumsassiana (Ragozin 1960) SB93
    `--Microdontella problematica Lebedev 1944 SB93
  Palaeomutelidae SB93
  Acharax Dall 1908 AJH07
    `--A. cretacea Kanie & Nishida 2000 AJH07
  Linga De Gregorio 1884 AJH07
  Thyella lamellosa [incl. Cumingia elegans] M01
  Ciliatocardium ciliatum HG98
  Serratina eugonia PG98
  Stagnestesta Tasch 1961 T61
    `--*S. solitaria Tasch 1961 T61
  Rostrotortus Tasch 1961 T61
    `--*R. dissimilis Tasch 1961 T61
  Productae Tasch 1961 T61
    `--*P. dunbaris Tasch 1961 T61
  Elliptoidea Tasch 1961 T61
    `--*E. vulgaris Tasch 1961 T61
  Harvella elegans C64
  Gratelupia hydeana C64
  Mactrella C64
    |--M. carinata [incl. M. alata] C64
    `--M. exoleta C64
  Pseudobakewellia krasnowidowoensis G31
  Netschajewia G31
    |--N. elongata G31
    `--N. globosa G31
  Pleurophorina simplex G31
  Pleurophorus G31
  Alula kutorgi G31
  Bakewellia G31
    |--B. antiqua G31
    |--B. bicarinata G31
    |--B. ceratophaga G31
    |--B. ladina G31
    |--B. nikitowkensis G31
    `--B. sulcata G31
  Cyrtodontarca bakewellioides G31
  Clinopristha radiata G31
  Angustella praeangusta [=Gervilleia praeangusta] G31
  Palaeostrea sinensis G31
  Cucullopsis quadrata G31
  Acanthopecten shansiensis G31
  Posidoniella G31
    |--P. elongata G31
    |--P. laevis G31
    |--P. pyriformis G31
    |--P. variabilis G31
    |--P. vetusta G31
    `--P. yuani G31
  Vola praecursor G31
  Posidonomya becheri G31
  Jousseaumiella concharum Knudsen 1944 H64
  Trigonicardia antillarum WE63
  Canthyria N65b
    |--C. collina N65b
    `--C. spinosa N65b
  Schistodesmus lampreyanus N65b
  Dozierella gouldii N65a
  Terraia altissima EB99

Autobranchia [Anatinacea, Autolamellibranchiata, Dimyaria, Eulamellibranchia, Heteroconchia, Lamellibranchia, Lucinata, Lucinoida, Mesosyntheta, Metabranchia, Mytilacea, Mytiliformii, Prionodesmacea] PC11

Cleobis grandis (Dana 1849) [=Maeonia grandis, Megadesmus (Cleobis) grandis; incl. Pachydomus gigas M’Coy 1847, Maeonia gigas, Myonia gigas, Ma. gracilis Dana 1849, Megadesmus (Cleobis) gracilis, Myonia gracilis, Globicarina grossula Waterhouse 1965, Cleobis robusta Laseron 1910] F71

Cypricardinia gregarius (Etheridge 1900) [=Pleurophorus gregarius; incl. P. gregarius var. inflata Laseron 1910] F71

Devonia perrieri (Malard 1903) [=Synapticola perrieri, Entovalva perrieri; incl. Montacuta donacina Bateson 1923] BK77

Hokonuia limaeformis Trechmann 1918 [incl. Nuggetia morganiana Wilckens 1927, H. parki Wilckens 1927, H. rotundata Trechmann 1918] W60

*Mysidioptera ornata Salomon 1895 [incl. M. ornata var. laevigata Bittner 1895, M. ornata var. lombardica Bittner 1895] W60

Obovaria olivaria (Rafinesque 1820) [=Amblema olivaria, Unio olivaria; incl. Obovaria ellipsis Lea 1828] V16

Obovaria subrotunda (Rafinesque 1820) [=Obliquaria subrotunda, Unio subrotunda; incl. Obovaria circulus Lea 1829, O. striata Rafinesque 1820, Unio striata] V16

Saxicava pholadis Middendorff 1851 N79 [incl. S. clava C64, S. distorta C64, S. gastrochaenoidea C64, S. legumen C64]

*Type species of generic name indicated

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Last updated: 6 March 2022.