Malacostraca

Paranebalia sp., from the Yale Peabody Museum of Natural History.


Belongs within: Euarthropoda.
Contains: Archaeostraca, Eumalacostraca, Nebaliidae.

The Malacostraca are a clade of crustaceans characterised by the possession of fourteen or fifteen trunk segments with the trunk divided into two sections bearing functionally distinct appendages. The anterior section of eight segments bears leaf-like phyllopods or rod-shaped stenopods whereas the following six segments bear swimming pleopods (there may be a terminal limbless segment). Those taxa with phyllopods have historically been recognised as the Phyllocarida but recent analysis have indicated this group is paraphyletic to the stenopod-bearing Eumalacostraca.

<==Malacostraca [Phyllocarida]
    |--+--Archaeostraca AC17
    |  `--+--Cinerocaris magnifica AC17, SC17
    |     `--Eumalacostraca AC17
    `--Leptostraca [Nebaliacea] AC17
         |  i. s.: Rhabdouraea Malzahn 1962 [Rhabdouraeidae] W-SP01
         |           `--*R. bentzi (Malzahn 1958) [=Nebalia bentzi) SM84
         |         Douglasocaris [Douglasocaridae] G88
         |--Nebaliopsis Sars 1887 [Nebaliopsidae, Nebaliopsididae] W-SP01
         |    `--*N. typica Sars 1887 W-SP01
         `--+--Nebaliidae W-SP01
            `--Paranebaliidae W-SP01
                 |--Paranebalia Claus 1880 W-SP01
                 |    |--*P. longipes (Willemöes-Suhm 1875) [=Nebalia longipes] W-SP01
                 |    `--P. belizensis Modlin 1991 W-SP01
                 `--Levinebalia Walker-Smith 2000 W-SP01
                      |--*L. maria Walker-Smith 2000 W-SP01
                      `--L. fortunata (Wakabara 1976) W-SP01

Malacostraca incertae sedis:
  Litopenaeus vannamei TL03
  Shinkaia crosnieri WS10
  Callizoe Barrande 1872 (n. d.) M61
  Orozoe Barrande 1872 (n. d.) M61
  Hymenocaris Salter 1853 B95 [Hymenocarididae, Hymenostraca FGS04]
    |--H. oelandica O68
    `--H. vermicauda Salter 1853 BWW93
  Sairocarididae [Hoplostraca] G88
    |--Sairocaris elongata (Peach 1882) BWW93
    `--Kellibrooksia macrogaster Schram 1973 BWW93
  Mytocaris Chlupáč 1970 BT04
    |--M. klouceki Chlupáč 1970 BT04
    `--‘Buffalopterus’ verrucosus Kjellesvig-Waering & Heubusch 1962 BT04
  Galenocaris Wells 1944 BT04
    `--G. campbelli Wells 1944 BT04
  Lebesconteia Jones & Woodward 1899 BT04
  Nothozoe Barrande 1872 (n. d.) BT04
    |--*N. pollens Barrande 1872 BB61
    `--N. barrandei Chlupáč 1970 BT04
  Saccocaris Salter 1873 BT04
    |--S. major Salter 1873 BT04
    |--S. minor Jones & Woodward 1891 BT04
    `--S. tetragona Chapman 1903 BT04
  Trigonocarys Barrois 1891 BT04

*Type species of generic name indicated

REFERENCES

[AC17] Aria, C., & J.-B. Caron. 2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature 545: 89–92.

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[B95] Bousfield, E. L. 1995. A contribution to the natural classification of Lower and Middle Cambrian arthropods: food-gathering and feeding mechanisms. Amphipacifica 2: 3–34.

[BT04] Braddy, S. J., V. P. Tollerton Jr, P. R. Racheboeuf & R. Schallreuter. 2004. Eurypterids, phyllocarids, and ostracodes. In: Webby, B. D., F. Paris, M. L. Droser & I. G. Percival (eds) The Great Ordovician Biodiversification Event pp. 255–265. Columbia University Press.

[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.

[FGS04] Feldmann, R. M., A. A. Garassino & C. E. Schweitzer. 2004. The presumed decapod, Palaeopemphix Gemmellaro, 1890, is a unique member of the Phyllocarida (Palaeopemphicida: Palaeopemphidae). Journal of Palaeontology 78: 340–348.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[M61] Moore, R. C. 1961. Editorial note. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q429. Geological Society of America and University of Kansas Press.

[O68] Öpik, A. A. 1968. Ordian (Cambrian) Crustacea Bradoriida of Australia. Commonwealth of Australia, Bureau of National Development, Bureau of Mineral Resources, Geology and Geophysics, Bulletin 103: 1–45.

[SM84] Schram, F. R., & E. Malzahn. 1984. The fossil leptostracan Rhabdouraea bentzi (Malzahn, 1958). Transactions of the San Diego Society of Natural History 20 (6): 95–98.

[SC17] Schwentner, M., D. J. Combosch, J. P. Nelson & G. Giribet. 2017. A phylogenomic solution to the origin of insects by resolving crustacean-hexapod relationships. Current Biology 27: 1818–1824.

[TL03] Thompson, F. L., Y. Li, B. Gomez-Gil, C. C. Thompson, B. Hoste, K. Vandemeulebroecke, G. S. Rupp, A. Pereira, M. M. De Bem, P. Sorgeloos & J. Swings. 2003. Vibrio neptunius sp. nov., Vibrio brasiliensis sp. nov. and Vibrio xuii sp. nov., isolated from the marine aquaculture environment (bivalves, fish, rotifers and shrimps). International Journal of Systematic and Evolutionary Microbiology 53: 245–252.

[W-SP01] Walker-Smith, G. K., & G. C. B. Poore. 2001. A phylogeny of the Leptostraca (Crustacea) with keys to families and genera. Memoirs of Museum Victoria 58 (2): 383–410.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

Euarthropoda

Reconstruction of Mollisonia plenovenatrix by J. Liang, from Aria & Caron (2019).


Belongs within: Arthropoda.
Contains: Pycnogonida, Euchelicerata, Artiopoda, Marrellomorpha, Myriapoda, Fuxianhuiida, Hymenocarina, Oligostraca, Allotriocarida, Tantulocarida, Thecostraca, Copepoda, Malacostraca.

The name Euarthropoda refers to the most exclusive clade formed by crown-group arthropods. Recent analyses agree on a division of living arthropods between three major clades, the Chelicerata, Myriapoda and Pancrustacea, with the latter two usually being united to form a clade of arthropods possessing mandibles. Fossil euarthropod taxa not belonging to these clades are best known from the Cambrian period with the exception of the Artiopoda (the clade including the trilobites). The clade Pancrustacea includes the modern crustaceans and insects with the latter a derived subgroup of the former (Regier et al. 2010). This last grouping is primarily supported by molecular data; potential synapomorphies of the Pancrustacea have all become modified in various subclades. Relationships within the pancrustaceans remain subject to analysis. A relationship between the Tantulocarida and Thecostraca has been suggested on the basis of gonopore position in the two groups; tantulocarids and basal thecostracans are also both sessile parasites.

<==Euarthropoda (see below for synonymy)
    |  i. s.: Dala peilertae Müller 1983 ZS07
    |         Walossekia quinquespinosa Müller 1983 ZS07
    |--Panchelicerata AC19
    |    |--+--Sanctacaris Briggs & Collins 1988 AC19, CB04 [Sanctacarida, Sanctacaridea]
    |    |  |    |--‘Utahcaris’ orion Conway Morris & Robison 1988 B95
    |    |  |    `--Sanctacaris uncata Briggs & Collins 1988 WD02
    |    |  `--Habeliida B95
    |    |       |--Habelia Walcott 1912 AC19, CB04
    |    |       |    `--H. optata AC19
    |    |       `--Ecnomocaris LSE13
    |    `--Chelicerata AC19
    |         |  i. s.: Wisangocaris barbarahardyae Jago, García-Bellido & Gehling 2016 BS20
    |         |--Mollisoniidae [Mollisoniida] AC19
    |         |    |--Mollisonia Walcott 1912 AC19, CB04 [incl. Houghtonites Raymond 1931 AC19]
    |         |    |    |--*M. symmetrica Walcott 1912 [incl. M. rara Walcott 1912] AC19
    |         |    |    |--M. gracilis Walcott 1912 [=Houghtonites gracilis] AC19
    |         |    |    |--M. plenovenatrix Aria & Caron 2019 AC19
    |         |    |    `--M. sinica Zhang et al. 2002 AC19
    |         |    |--Thelxiope Simonetta & Delle Cave 1975 AC19
    |         |    |--Corcorania Jell 1980 AC19
    |         |    |    `--C. trispinosa AC19
    |         |    `--Urokodia Hou, Chen & Lu 1989 AC19
    |         |--Pycnogonida RS10
    |         `--Euchelicerata RS10
    `--+--Retifaciida LSE13
       |    |--Retifacies Hou, Chen & Lu 1989 LSE13, CB04
       |    |    `--R. abnormalis Hou et al. 1989 CB04
       |    `--+--Pygmaclypeatus LSE13
       |       `--+--+--Kiisortoqia AC19
       |          |  `--Siriocars LSE13
       |          `--Squamacula LSE13
       |               |--S. buckorum LSE13
       |               `--S. clypeatus LSE13
       `--+--Artiopoda AC19
          `--+--Marrellomorpha AC19
             `--+--Aquilonifer Briggs, Siveter et al. 2016 AC19, BS16
                |    `--*A. spinosus Briggs, Siveter et al. 2016 BS16
                `--Mandibulata (see below for synonymy) AC17
                     |  i. s.: Ercaia [Promandibulata] C12
                     |           `--E. minuscule C12
                     |         Musacaris LSE13
                     |         Cambropachycopidae LSE13
                     |           |--Cambropachycope clarksoni Walossek & Müller 1990 BWW93
                     |           `--Goticaris longispinosa Walossek & Müller 1990 BWW93
                     |--Myriapoda AC19
                     `--+--+--Fuxianhuiida AC19
                        |  `--Hymenocarina AC19
                        `--Pancrustacea (see below for synonymy) AC17
                             |  i. s.: Nippotantulus heteroxenus Huys, Ohtsuka & Boxshall 1994 MA01
                             |         Neomysis M02
                             |           |--N. americana M02
                             |           `--N. vulgaris CS77
                             |         Perosquilla armata K-MC02
                             |           |--P. a. armata K-MC02
                             |           `--P. a. capensis K-MC02
                             |         Mesosoma Otto 1821 C92
                             |         Mixodiaptomus laciniatus D51
                             |         Pleuroncodes planipes CH97
                             |         Phalangites priscus Münster 1836 D07
                             |         Cryptophthalmus Rafinesque 1814 BR05
                             |         Cymodocea Rafinesque 1814 BR05
                             |         ‘Cymodocea’ Leach 1818 nec Rafinesque 1814 nec Lamouroux 1816 BR05
                             |         ‘Dactylopus’ Claus 1862 non Gill 1859 BR05
                             |         Donovania Leach 1814 BR05
                             |         Orientalina Kolosnitsyna 1973 BR05
                             |         Pagodina van Beneden 1853 BR05
                             |         ‘Trichia’ de Haan 1839 non Haller 1768 BR05
                             |         ‘Typhis’ Risso 1816 non Montfort 1810 [Typhidae] BR05
                             |         Fenneropenaeus chinensis DH08
                             |         Limnadella kitei Girard 1854 DS89
                             |         Anatanais robusto BBB-S95
                             |         Anthonema Walther 1904 H75
                             |           `--*A. problematica Walther 1904 H75
                             |         ‘Acherusia’ Lucas 1846 non Laporte & Gory 1837 E12
                             |           `--A. dumerilii Lucas 1846 E12
                             |         Naesea edwardsii Lucas 1846 E12
                             |         Ovalia Latreille in Griffith & Pidgeon 1833 KA-Z11
                             |         Cycloidea FGS04
                             |           |--Cyclus de Koninck 1841 [incl. Paraprosopon Gemmellaro 1890] FGS04
                             |           `--Oonocarcinus Gemmellaro 1890 FGS04
                             |         Heymonsicambria EL11
                             |         Macleayia Haswell 1830 HW92
                             |         Pseudalibrotus littoralis CS77
                             |         Hymenodora glacialis CS77
                             |         Ozolus Latreille 1802 L02a
                             |           `--*O. gasterostei Latreille 1802 L02b (see below for synonymy)
                             |         Amymone L02b
                             |           |--A. baccha [=Monoculus bacchus] L02b
                             |           |--A. faunus [=A. fauna, Monoculus faunus] L02b
                             |           |--A. moenas L02b
                             |           |--A. satyrus [=A. satyra, Monoculus satyrus] L02b
                             |           |--A. silene [=A. silenus, Monoculus silenus] L02b
                             |           `--A. thyas L02b
                             |         Nauplius L02b
                             |           |--N. bracteatus [=Monoculus bracteatus] L02b
                             |           `--N. saltatorius [=Monoculus saltatorius] L02b
                             |         Microdentopus gryllotalpa PP64
                             |         Aristeomorpha foliacea PP64
                             |         Ligur ensiferus PP64
                             |         Paromola cuvieri PP64
                             |         Anamathia rissoana PP64
                             |         Ergasticus clouei PP64
                             |         Dorynchus thompsoni PP64
                             |         Cymopolia caroni PP64
                             |         Halopsyche de Saussure 1857 BR17
                             |         Palpipes Roth 1851 K18
                             |         Berndtia purpurea CG97
                             |         Trypetasa lampas CG97
                             |         Montagua monoculoides M01
                             |         Amblyops crozetii O01
                             |         Harpactocarcinus tumidus H79
                             |         Tuzoiidae O68
                             |           |--Tuzoia O68
                             |           `--*Dioxycaris argenta (Walcott 1888) [=Leperditia argenta, Isoxys argenta] O68
                             |         Corynuropsis Scott 1894 E00
                             |--Oligostraca AC19
                             `--Altocrustacea [Ostraca, Phyllopodomorpha, Thoracopoda] RS10
                                  |--Allotriocarida SC17
                                  `--Multicrustacea [Communostraca, Vericrustacea] SC17
                                       |  i. s.: Arenosicaris inflata SC17
                                       |--+--Tantulocarida LS02
                                       |  `--Thecostraca SC17
                                       `--+--Copepoda SC17
                                          `--Malacostraca SC17

Euarthropoda [Arachnata, Arachnomorpha, Aspidonia, Aspidota, Cormogonida, Eotrilobitacea, Myriochelata, Paradoxopoda, Polymeria, Skaracarida]

Mandibulata [Antennata, Atelocerata, Biantennata, Labiophora, Labrophora, Opisthogoneata, Tracheata] AC17

*Ozolus gasterostei Latreille 1802 L02b [incl. Argulus delphini L02b, Monoculus delphinus L02a, M. gyrini L02b]

Pancrustacea [Archoptera, Copepodoida, Copepodomorpha, Crustacea, Crustaceanomorpha, Entomostraca, Eucrustacea, Gymnota, Maxillopoda, Pneumonura, Progonomorpha, Pseudopoda, Tetraconata, Thecostracomorpha]

*Type species of generic name indicated

REFERENCES

[AC17] Aria, C., & J.-B. Caron. 2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature 545: 89–92.

[AC19] Aria, C., & J.-B. Caron. 2019. A middle Cambrian arthropod with chelicerae and proto-book gills. Nature 573: 586–589.

[BS20] Baker, C. M., K. Sheridan, S. Derkarabetian, A. Pérez-González, S. Vélez & G. Giribet. 2020. Molecular phylogeny and biogeography of the temperate Gondwanan family Triaenonychidae (Opiliones: Laniatores) reveals pre-Gondwanan regionalisation, common vicariance, and rare dispersal. Invertebrate Systematics 34: 637–660.

[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.

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

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[B95] Bousfield, E. L. 1995. A contribution to the natural classification of Lower and Middle Cambrian arthropods: food-gathering and feeding mechanisms. Amphipacifica 2: 3–34.

[BS16] Briggs, D. E. G., D. J. Siveter, D. J. Siveter, M. D. Sutton & D. Legg. 2016. Tiny individuals attached to a new Silurian arthropod suggest a unique mode of brood care. Proceedings of the National Academy of Sciences of the USA 13 (16): 4410–4415.

[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.

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology 2nd ed. WCB McGraw-Hill: Boston.

[C12] Chen, J.-Y. 2012. Evolutionary scenario of the early history of the animal kingdom: evidence from Precambrian (Ediacaran) Weng’an and Early Cambrian Maotianshan biotas, China. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 239–379. Springer.

[CG97] Cohen, B. L., & A. B. Gawthrop. 1997. The brachiopod genome. In: Kaesler, R. L. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda, Revised vol. 1. Introduction pp. 189–211. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[CB04] Cotton, T. J., & S. J. Braddy. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 169–193.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species 3rd ed. Columbia University Press: New York.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[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.

[E00] Engel, M. S. 2000. Classification of the bee tribe Augochlorini (Hymenoptera: Halictidae). Bulletin of the American Museum of Natural History 250: 1–89.

[EL11] Erwin, D. H., M. Laflamme, S. M. Tweedt, E. A. Sperling, D. Pisani & K. J. Peterson. 2011. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334: 1091–1097.

[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.

[FGS04] Feldmann, R. M., A. A. Garassino & C. E. Schweitzer. 2004. The presumed decapod, Palaeopemphix Gemmellaro, 1890, is a unique member of the Phyllocarida (Palaeopemphicida: Palaeopemphidae). Journal of Palaeontology 78: 340–348.

[H79] Haast, J. von. 1879. Geology of the Provinces of Canterbury and Westland, New Zealand. A report comprising the results of official explorations. "Times" Office: Christchurch.

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

[HW92] Houston, W. W. K., & T. A. Weir. 1992. Melolonthinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 174–358. AGPS Press: Canberra.

[K-MC02] Klein-MacPhee, G., & B. B. Collette. 2002. Scorpionfishes. Family Scorpaenidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 331–338. Smithsonian Institute Press: Washington.

[K18] Kury, A. B. 2018. Familial nomina in harvestmen (Arachnida, Opiliones). Bionomina 13: 1–27.

[KA-Z11] Kury, A. B., & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the "Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)". Zootaxa 3034: 47–68.

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

[L02a] Latreille, P. A. 1802a. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L02b] Latreille, P. A. 1802b. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 4. Familles naturelles des genres. F. Dufart: Paris.

[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.

[MA01] Martínez Arbizu, P. 2001. Psammocyclopinidae fam. n., a new monophyletic group of marine Cyclopoida (Copepoda, Crustacea), with the description of Psammocyclopina georgei sp. n. from the Magellan region. Revista Bras. Zool. 18 (4): 1325–1339.

[M01] M'Intosh, W. C. 1901. The coloration of marine animals. Annals and Magazine of Natural History, series 7, 7: 221–240.

[M02] Munroe, T. A. 2002. Herrings. Family Clupeidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 111–160. Smithsonian Institute Press: Washington.

[O01] Ohlin, A. 1901. On a new "bipolar" schizopod. Annals and Magazine of Natural History, series 7, 7: 371–374.

[O68] Öpik, A. A. 1968. Ordian (Cambrian) Crustacea Bradoriida of Australia. Commonwealth of Australia, Bureau of National Development, Bureau of Mineral Resources, Geology and Geophysics, Bulletin 103: 1–45.

[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.

[SC17] Schwentner, M., D. J. Combosch, J. P. Nelson & G. Giribet. 2017. A phylogenomic solution to the origin of insects by resolving crustacean-hexapod relationships. Current Biology 27: 1818–1824.

[WD02] Waloszek, D., & J. A. Dunlop. 2002. A larval sea spider (Arthropoda: Pycnogonida) from the Upper Cambrian ‘Orsten’ of Sweden, and the phylogenetic position of pycnogonids. Palaeontology 45 (3): 421–446.

[ZS07] Zhang, X.-G., D. J. Siveter, D. Waloszek & A. Maas. 2007. An epipodite-bearing crown-group crustacean from the Lower Cambrian. Nature 449: 595–598.

Echinocaridina

Fossil specimen of Paraechinocaris punctata, from the Digital Atlas of Ancient Life.


Belongs within: Archaeostraca.

The Echinocaridina are a group of phyllocarid crustaceans known from the Silurian (or possibly the Ordovician) to the early Carboniferous (Briggs et al. 1993).

Characters (from Rode & Lieberman 2002): Carapace subovate to subquadrate; anterior horn rounded or absent; posterior margin sinuous or convex. Rostrum free or absent, lacking longitudinal ridge. Carapace divided into anterior and posterior regions by distinct groove; one to five raised nodes present in anterior region. Marginal groove present on carapace and demarcating marginal rim.

Echinocaridina RL02
    |--Aristozoe Barrande 1872 FGS04 [Aristozoidae RL02]
    |    |--A. memoranda Barrande 1872 RL02 [incl. *A. amica Barrande 1868 BB61]
    |    |--A. parabolica O68
    |    `--A. porcula BWW93
    |--Ptychocarididae RL02
    |    |--Kerfornecaris Racheboeuf & Rolfe 1990 RL02
    |    |    `--K. roscanvelensis Racheboeuf & Rolfe 1990 RL02
    |    `--Ptychocaris Novák 1885 RL02
    |         `--P. novaki Copeland 1960 RL02
    `--Echinocarididae RL02
         |--Montecaridinae RL02
         |    |--Eleutherocaris whitfieldi (Clarke 1885) RL02
         |    `--Montecaris RL02
         |         |--M. antecedens Chlupáč 1960 BWW93
         |         |--M. lehmannii Jux 1960 RL02
         |         `--M. strunensis Jux 1960 RL02
         `--Echinocaridinae RL02
              |--Paraechinocaris Rode & Lieberman 2002 RL02
              |    |--*P. punctata (Hall 1863) (see below for synonymy) RL02
              |    |--P. asiatica (Reed 1908) [=Echinocaris asiatica] RL02
              |    `--P. spiniger (Racheboeuf 1998) [=Echinocaris spiniger] RL02
              `--Echinocaris Whitfield 1880 FGS04
                   |--*E. sublaevis Whitfield 1880 (see below for synonymy) RL02
                   |--E. auricula Eller 1935 RL02
                   |--E. beecheri Copeland 1960 RL02
                   |--E. castorensis Copeland 1960 RL02
                   |--E. clarkii Beecher 1902 RL02
                   |--E. condylepis Hall & Clarke 1988 RL02
                   |--E. consanguina Eller 1935 RL02
                   |--E. crosbyensis Eller 1937 RL02
                   |--E. hunanensis Han & Zhou 1993 RL02
                   |--E. multinodosa Whitfield 1880 RL02
                   |--E. multispinosis Feldmann & McKenzie 1981 RL02
                   |--E. ohioensis Sturgeon, Hlavin & Kesling 1964 RL02
                   |--E. randallii Beecher 1902 RL02
                   |--E. sloliensis Partridge 1902 RL02
                   |--E. socialis Beecher 1884 RL02
                   |--E. turgida Eller 1935 RL02
                   `--E. whidbornei (Jones & Woodward 1889) RL02

Echinocaris sublaevis Whitfield 1880 [incl. E. pulcra Sturgeon, Hlavin & Kesling 1964, E. pustulosa Whitfield 1880] RL02

Paraechinocaris punctata (Hall 1863) [=Ceratiocaris punctatus, Echinocaris punctata, Echinocaris punctatus; incl. Ceratiocaris armatus Hall 1863] RL02

*Type species of generic name indicated

REFERENCES

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[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.

[FGS04] Feldmann, R. M., A. A. Garassino & C. E. Schweitzer. 2004. The presumed decapod, Palaeopemphix Gemmellaro, 1890, is a unique member of the Phyllocarida (Palaeopemphicida: Palaeopemphidae). Journal of Palaeontology 78: 340–348.

[O68] Öpik, A. A. 1968. Ordian (Cambrian) Crustacea Bradoriida of Australia. Commonwealth of Australia, Bureau of National Development, Bureau of Mineral Resources, Geology and Geophysics, Bulletin 103: 1–45.

[RL02] Rode, A. L., & B. S. Lieberman. 2002. Phylogenetic and biogeographic analysis of Devonian phyllocarid crustaceans. Journal of Paleontology 76 (2): 271–286.

Cypridinidae

Left view of female Vargula norvegica, from Benson et al. (1961).


Belongs within: Myodocopida.

The Cypridinidae are a group of relatively large planktic ostracods, often over two millimetres in length.

Characters (from Benson et al. 1961, as Cypridininae): Carapace more or less strongly calcified, usually smooth; dorsal border arched, anterior margin of rostrum evenly curved or sinuous. Mostly rather large forms (more than 2 mm); sexual dimorphism weak or absent.

<==Cypridinidae [Cypridinadae, Cypridininae]
    |--Codonocera Brady 1902 BB61
    |    `--*C. cruenta Brady 1902 BB61
    |--Gigantocypris Müller 1895 BB61
    |    `--*G. agassizi Müller 1895 BB61
    |--Eocypridina Kesling & Ploch 1960 LH61
    |    `--*E. campbelli Kesling & Ploch 1960 LH61
    |--Cyprosis Jones 1881 M61, BB61
    |    `--*C. haswelli Jones 1881 BB61
    |--Azygocypridina Sylvester-Bradley 1950 [=Crossophorus Brady 1880 non Hemprich & Ehrenberg 1828] BB61
    |    |--*A. imperator (Brady 1880) [=*Crossophorus imperator] BB61
    |    `--A. gibber BB61
    |--Heterodesmus Brady 1865 [incl. Siphonostra Skogsberg 1920] BB61
    |    |--*H. adamsi Brady 1866 BB61
    |    `--H. spinifer BB61
    |--Cypridina Milne Edwards 1840 (see below for synonymy) BB61
    |    |--C. inermis [incl. *C. reynaudi Milne Edwards 1840] BB61
    |    |--C. globosa BK77
    |    `--C. homoedwardsiana S61
    `--Vargula Skogsberg 1920 BB61
         |  i. s.: V. bullae M83
         |         V. hilgendorfi (Müller 1890) HM03
         |--*V. (Vargula) norvegica (Baird 1860) [=Cypridina norvegica] BB61
         `--V. (Doloria Skogsberg 1920) BB61
              `--V. (*D.) levis (Skogsberg 1920) [=Cypridina (*D.) levis] BB61

Cypridina Milne Edwards 1840 [incl. Daphnia M’Coy 1844 non Müller 1776, Eupathistoma Brady 1898, Pyrocypris Müller 1890] BB61

*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.

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[HM03] Hanamura, Y., & M. Matsuoka. 2003. Feeding habits of the sand shrimp, Crangon uritai Hayashi & Kim, 1999, in the central Seto Inland Sea, Japan. Crustaceana 76 (8): 1017–1024.

[LH61] Levinson, S. A., & H. V. Howe. 1961. Addendum. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q421–Q422. Geological Society of America and University of Kansas Press.

[M61] Moore, R. C. 1961. Supplement—USSR treatise on Ostracoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q422–Q429. Geological Society of America and University of Kansas Press.

[M83] Morin, J. G. 1983. Coastal bioluminescence: patterns and functions. Bulletin of Marine Science 33 (4): 787–817.

[S61] Scott, H. W. 1961. Shell morphology of Ostracoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q21–Q37. Geological Society of America and University of Kansas Press.

Graviidae

Left view of Ceratocypris longispina, from Benson et al. (1961).


Belongs within: Ostracoda.

The Graviidae are a group of Palaeozoic ostracods characterised by a carapace with epibolic ventral margins.

<==Graviidae M61
    |--Gravia Polenova 1952 (n. d.) BB61
    |    `--*G. aculata Polenova 1952 BB61
    |--Pribylina Ivanova 1959 LH61
    |    `--*P. levis Ivanova 1959 LH61
    |--Boucekites Přibyl 1951 M61, BB61
    |    `--*B. devonicus Přibyl 1951 BB61
    |--Selebratina Polenova 1952 (n. d.) M61, BB61
    |    `--*S. dentata (Polenova 1952) [=Gravia (*Selebratina) dentata] BB61
    |--Pinnatulites Hessland 1949 M61, BB61
    |    `--*P. procera (Kummerow 1924) [=Primitiella procera] BB61
    `--Ceratocypris Poulsen 1934 M61, BB61
         |--*C. symmetrica Poulsen 1934 BB61
         `--C. longispina BB61

*Type species of generic name indicated

REFERENCES

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[LH61] Levinson, S. A., & H. V. Howe. 1961. Addendum. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q421–Q422. Geological Society of America and University of Kansas Press.

[M61] Moore, R. C. 1961. Supplement—USSR treatise on Ostracoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q422–Q429. Geological Society of America and University of Kansas Press.

Drepanellacea

Left valve of Kirkbyella delicata, from Hoare & Merrill (2004).


Belongs within: Ostracoda.

The Drepanellacea are a group of non-dimorphic ostracods known from the Ordovician to the Permian (Whatley et al. 1993). Members of the group vary greatly in development of nodes and features of the marginal rim but are united by the existence of intermediate forms (Benson et al. 1961).

Characters (from Benson et al. 1961): Carapace subquadrate to subrectangular, with or without marginal rim; surface smooth, pitted, or reticulate; one or more nodes present, sometimes terminated by spines; nodes rarely lobate; S2 poorly to well defined; hingement by tongue and groove; dimorphism unknown.

<==Drepanellacea [Aechminacea, Binodicopa] BB61
    |  i. s.: Saffordellina Bassler & Kellett 1934 [=Saffordella Ulrich & Bassler 1923 non Dunbar 1920] BB61
    |           `--*S. muralis (Ulrich & Bassler 1923) [=*Saffordella muralis] BB61
    |--Spinigeritidae WSB93
    |    |--Susus sus Schallreuter 1991 WSB93
    |    `--Eocytherella Bonnema 1933 WSB93, BB61
    |         |--E. nioni Vannier 1986 WSB93
    |         |--E. smithi (Jones 1887) [=Cytherella smithi] BB61
    |         `--E. troedssoni (Bonnema 1933) [=Cytherella troedssoni, Primitia tenera Troedsson 1918 (preoc.)] BB61
    |--Kirkbyellidae BB61
    |    |--Psilokirkbyella prominens Becker & Wang 1992 WSB93
    |    `--Kirkbyella Coryell & Booth 1933 HM04
    |         |--*K. (Kirkbyella) typa Coryell & Booth 1933 BB61
    |         `--K. (Berdanella Sohn 1961) HM04
    |              |--K. (B.) delicata Hoare & Merrill 2004 HM04
    |              |--K. (B.) gutkei (Croneis & Bristol 1939) HM04
    |              `--K. (B.) quadrata (Croneis & Gutke 1939) HM04
    |--Richinidae [Circulinidae] BB61
    |    |--Rivillina schallreuteri Vannier 1983 WSB93
    |    |--Crescentilla Barrande 1872 BB61
    |    |    `--*C. pugnax Barrande 1872 BB61
    |    |--Pseudulrichia Schmidt 1941 BB61
    |    |    |--*P. bivertex (Ulrich 1879) [=Leperditia bivertex] BB61
    |    |    `--P. simplex BB61
    |    |--Richina Coryell & Malkin 1936 BB61
    |    |    |--*R. truncata Coryell & Malkin 1936 BB61
    |    |    |--R. selenicristata Stover 1956 WSB93
    |    |    `--R. zygalis BB61
    |    `--Parulrichia Schmidt 1941 BB61
    |         `--*P. diversa (Jones & Holl 1886) [=Primitia diversa] BB61
    |              |--P. d. diversa BB61
    |              `--P. d. antiqua BB61
    |--Aechminellidae BB61
    |    |--Aechminella Harlton 1933 [incl. Balantoides Morey 1935, Boursella Turner 1939] BB61
    |    |    `--*A. trispinosa Harlton 1933 BB61
    |    |--Bicornella Coryell & Cuskley 1934 BB61
    |    |    `--*B. tricornis Coryell & Cuskley 1934 BB61
    |    |--Cornigerella Warthin 1930 BB61
    |    |    `--‘Beyrichia’ tuberculospinosa Jones & Kirkby 1886 [incl. *Cornigerella minuta Warthin 1930] BB61
    |    |--Evlanovia Egorov 1950 BB61
    |    |    `--*E. tichonovitchi Egorov 1950 BB61
    |    |--Mauryella Ulrich & Bassler 1923 [incl. Verrucolsella Neave 1950, Verrucosella Croneis & Gale 1938] BB61
    |    |    `--*M. mammilata Ulrich & Bassler 1923 BB61
    |    |--Milanovskya Egorov 1950 BB61
    |    |    `--*M. bicornis Egorov 1950 BB61
    |    `--Beyrichiana Kellett 1933 BB61
    |         `--*B. permiana Kellett 1933 BB61
    |--Aechminidae [Aechmininae] BB61
    |    |--Lindsayella Coryell & Williamson 1936 BB61
    |    |    `--*L. rugosa Coryell & Williamson 1936 BB61
    |    |--Sigynus Kesling 1953 BB61
    |    |    `--*S. dictyotus Kesling 1953 BB61
    |    |--Waldronites Coryell & Williamson 1942 [=Cornulina Coryell & Williamson 1936 non Conrad 1853] BB61
    |    |    `--*W. bispinosa (Coryell & Williamson 1936) [=*Cornulina bispinosa] BB61
    |    |--Mammoides Bradfield 1935 WSB93, BB61
    |    |    `--*M. mammilata Bradfield 1935 BB61, WSB93 [=Beyrichiana mammilata BB61]
    |    |--Aechmina Jones & Holl 1869 BB61
    |    |    |--*A. cuspidata Jones & Holl 1869 BB61
    |    |    |--A. bovina Jones 1887 [incl. A. bovina var. punctata Krause 1892] C01
    |    |    `--A. ventadorni Vannier 1986 WSB93
    |    |--Aechminaria Coryell & Williamson 1936 BB61
    |    |    |--*A. nodosa Coryell & Williamson 1936 BB61
    |    |    `--A. robusta BB61
    |    `--Paraechmina Ulrich & Bassler 1923 BB61
    |         |--*P. spinosa (Hall 1852) [=Cytherina spinosa, Beyrichia spinosa] BB61
    |         `--P. postica BB61
    |--Drepanellidae [Depranellidae, Drepanellinae, Neodrepanellinae] BB61
    |    |--Pilla sinensis (Hou 1953) WSB93
    |    |--Duplexibollia duplex (Kraus 1892) WSB93
    |    |--Drepanella Ulrich 1890 [=Depranella Ulrich 1890] BB61
    |    |    `--*D. crassinoda Ulrich 1890 BB61
    |    |--Acantonodella Zaspelova 1952 [incl. Acanthodella Howe 1955] BB61
    |    |    `--*A. terciocornuta Zaspelova 1952 BB61
    |    |--Kinnekullea Henningsmoen 1948 BB61
    |    |    `--*K. waerni Henningsmoen 1948 BB61
    |    |--Neodrepanella Zaspelova 1952 [incl. Tetracornella Zaspelova 1952] BB61
    |    |    `--*N. tricornis (Batalina 1941) [=Drepanella tricornis] BB61
    |    |--Scofieldia Ulrich & Bassler 1908 BB61
    |    |    `--*S. bilateralis (Ulrich 1894) [=Drepanella bilateralis] BB61
    |    |--Planusella Ivanova 1959 LH61
    |    |    `--*P. bicornis Ivanova 1959 LH61
    |    |--Pseudonodella Zaspelova 1952 M61, BB61
    |    |    `--*P. plana Zaspelova 1952 BB61
    |    `--Pseudonodellina Polenova 1955 M61, BB61
    |         |--*P. parvula (Polenova 1953) [=Nodella parvula] BB61
    |         |--P. rotundata BB61
    |         `--P. strelniensis BB61
    |--Bolliidae [Bolliinae, Ulrichiinae] BB61
    |    |--Klimphores kuemperi Schallreuter 1992 WSB93
    |    |--Neoulrichia pulchra Kozur 1983 WSB93
    |    |--Bichilina Sarv 1959 BB61
    |    |    `--*B. prima Sarv 1959 BB61
    |    |--Jonesites Coryell 1930 [=Placentula Jones & Holl 1886 non Lamarck 1822; incl. Placentella Wilson 1935] BB61
    |    |    `--*J. excavata (Jones & Holl 1869) [=Primitia excavata, *Placentula excavata] BB61
    |    |--Maratia Kay 1940 BB61
    |    |    `--*M. mara Kay 1940 BB61
    |    |--Tetrastorthynx Kesling 1953 BB61
    |    |    `--*T. diabolicus Kesling 1953 BB61
    |    |--Tmemolophus Kesling 1953 BB61
    |    |    `--*T. margarotus Kesling 1953 BB61
    |    |--Jonesella Ulrich 1890 M61, BB61 [incl. Melanella Wade 1911 non Bowdich 1822 BB61, Vogdesella Baker 1924 BB61]
    |    |      `--*J. crepidiformis (Ulrich 1879) [=Leperditia crepidiformis] BB61
    |    |--Kayatia Öpik 1953 M61, BB61
    |    |    `--*K. prima Öpik 1953 BB61
    |    |--Signetopsis Henningsmoen 1954 M61, BB61
    |    |    `--*S. quadrilobata Henningsmoen 1954 BB61
    |    |--Parajonesites Ivanova 1955 (n. d.) M61, BB61
    |    |    `--*P. notabilis Ivanova 1955 BB61
    |    |--Parenthatia Kay 1940 BB61
    |    |    |--*P. punctata (Ulrich 1894) [=Moorea punctata] BB61
    |    |    `--P. camerata BB61
    |    |--Ulrichia Jones 1890 [incl. Limbatula Zaspelova 1952, Warthinia Spivey 1939] BB61
    |    |    |--*U. conradi Jones 1890 BB61
    |    |    `--U. nodosa [=*Warthinia nodosa] BB61
    |    |--Xystinotus Kesling 1953 BB61
    |    |    |--*X. wrighrotum Kesling 1953 BB61
    |    |    `--X. subnodatus BB61
    |    `--Bollia Jones & Holl 1886 BB61
    |         |--*B. uniflexa Jones & Holl 1886 BB61
    |         |--B. auricularis Jones 1887 C01
    |         |--B. subaequata BB61
    |         `--B. ungula BB61
    `--Quadrijugatoridae [Ceratopsinae, Quadrijugatidae] WSB93
         |--Harpabollia harparum (Troedsson 1918) WSB93
         |--Hesslandella Henningsmoen 1953 BB61 BB61
         |    `--*H. macroreticulata (Hessland 1949) [=Ctenentoma macroreticulata] BB61
         |--Protallinnella Jaanusson 1957 BB61
         |    `--*P. grewingki (Bock 1867) [=Beyrichia grewingki] BB61
         |--Pseudorakverella Sarv 1959 BB61
         |    `--*P. optata Sarv 1959 BB61
         |--Rigidella Öpik 1937 BB61
         |    `--*R. mitis (Öpik 1935) [=Steusloffia mitis] BB61
         |--Tallinnopsis Sarv 1959 BB61
         |    `--*T. calkeri (Bonnema 1909) [=Tetradella calkeri] BB61
         |--Tetrada Neckaja 1958 BB61
         |    `--*T. memorabilis (Neckaja 1953) [=Tetradella memorabilis] BB61
         |--Zygobolboides Spivey 1939 BB61
         |    `--*Z. grafensis Spivey 1939 BB61
         |--Ivaria Neckaja 1960 M61 [=Glossopsis Neckaja 1953 nec Bush 1904 nec Hessland 1949 BB61]
         |    `--*I. robusta (Hessland 1949) [=*Glossopsis robusta] M61
         |--Piretopsis Henningsmoen 1953 M61, BB61
         |    `--*P. donsi Henningsmoen 1953 BB61
         |--Quadrijugator Kesling & Hussey 1953 BB61
         |    |--*Q. permarginatus (Foerste 1917) [=Bollia permarginata] BB61
         |    `--Q. marcoi Vannier 1986 WSB93
         |--Tallinnella Öpik 1937 BB61
         |    |--*T. dimorpha Öpik 1937 BB61
         |    `--T. marchica (Krause 1889) TM02
         |--Ceratopsis Ulrich 1894 (see below for synonymy) BB61
         |    |--*C. chambersi (Miller 1875) [=Beyrichia chambersi] BB61
         |    `--C. oculifera BB61
         |--Kiesowia Ulrich & Bassler 1908 BB61
         |    |--*K. dissecta (Krause 1892) [=Beyrichia dissecta] BB61
         |    `--K. radians BB61
         |--Quadrilobella Ivanova 1955 [incl. Tetradellina Harris 1957] BB61
         |    |--*Q. recta Ivanova 1955 BB61
         |    `--Q. arpilobata BB61
         `--Ullerella Henningsmoen 1950 M61, BB61 [=Ullia Henningsmoen 1949 non Roewer 1943 BB61]
              |--*U. ulli (Dons 1949) [=*Ullia ulli] BB61
              `--U. ventroplicata BB61

Ceratopsis Ulrich 1894 [incl. Ceratella Ulrich 1890 (n. n.) nec Hook. f. 1844 (ICBN) nec Gray 1869 nec Pat. 1887 (ICBN) nec (Quél.) Bigeard & Guill. 1913 (ICBN)] BB61

*Type species of generic name indicated

REFERENCES

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[C01] Chapman, F. 1901. On some fossils of Wenlock age from Mulde, near Klinteberg, Gotland; with notes by Prof. T. Rupert Jones and Dr. F. A. Bather. Annals and Magazine of Natural History, series 7, 7: 141–160, pl. 3.

[HM04] Hoare, R. D., & G. K. Merrill. 2004. A Pennsylvanian (Morrowan) ostracode fauna from Texas. Journal of Paleontology 78 (1): 185–204.

[LH61] Levinson, S. A., & H. V. Howe. 1961. Addendum. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q421–Q422. Geological Society of America and University of Kansas Press.

[M61] Moore, R. C. 1961. Supplement—USSR treatise on Ostracoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q422–Q429. Geological Society of America and University of Kansas Press.

[TM02] Tinn, O., & T. Meidla. 2002. An enigmatic early ?palaeocope ostracode from the Arenig of NW Russia. Acta Palaeontologica Polonica 47: 685–690.

[WSB93] Whatley, R. C., D. J. Siveter & I. D. Boomer. 1993. Arthropoda (Crustacea: Ostracoda). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 343–356. Chapman & Hall: London.

Beyrichiacea

Male (above) and female Kloedenia normalis, from Benson et al. (1961).


Belongs within: Ostracoda.
Contains: Beyrichia.

The Beyrichiacea are a group of ostracods known from the late Ordovician to the early Permian (Whatley et al. 1993) characterised by the dimorphic development of a ventral to anteroventral lobe or pouch.

Characters (from Benson et al. 1961): Straight-hinged ostracodes with subequal ends or forward swing, mostly with well-developed lobes and sulci and showing tendency to have carinal, velate, and marginal structures; dimorphism usually well defined.

<==Beyrichiacea BB61
    |  i. s.: Gillatia Öpik 1953 BB61
    |           `--*G. trinacria Öpik 1953 BB61
    |--Treposellidae [Treposellinae] WSB93
    |    |--Loutriella jupiterensis Copeland 1974 WSB93
    |    |--Parabouckekius martinssoni Jones 1985 WSB93
    |    |--Katatona romei Jones 1985 WSB93
    |    `--Treposella Ulrich & Bassler 1908 BB61
    |         |--*T. lyoni (Ulrich 1891) [=Beyrichia lyoni] BB61
    |         `--T. stellata BB61
    |--Craspedobolbinidae WSB93
    |    |--Malnina spinosa Jones 1989 WSB93
    |    |--Armilla sibirica Bushmina 1975 WSB93
    |    |--Craspedobolbina Kummerow 1923 BB61
    |    |    `--*C. dietrichi Kummerow 1923 BB61
    |    |--Copelandella Bless & Jordan 1971 WSB93, G88
    |    |    `--C. novascotia (Jones & Kirby 1884) WSB93
    |    `--Bolbineossia Kesling, Heany et al. 1958 WSB93, BB61
    |         |--*B. didictyosa Kesling, Heany et al. 1958 BB61
    |         `--B. pineaulti Copeland 1974 WSB93
    |--Zygobolbidae [Zygobolbinae] BB61
    |    |--Bonnemaia Ulrich & Bassler 1923 BB61
    |    |    `--*B. celsa Ulrich & Bassler 1923 BB61
    |    |--Glymmatobolbina Harris 1957 BB61
    |    |    `--*G. quadrata Harrs 1957 BB61
    |    |--Mastigobolbina Ulrich & Bassler 1923 BB61
    |    |    `--*M. typa Ulrich & Bassler 1923 BB61
    |    |--Plethobolbina Ulrich & Bassler 1923 BB61
    |    |    `--*P. typicalis Ulrich & Bassler 1923 BB61
    |    `--Zygobolba Ulrich & Bassler 1923 BB61
    |         |--*Z. (Zygobolba) decora (Billings 1866) [=Beyrichia decora] BB61
    |         |--Z. (Zygobolbina Ulrich & Bassler 1923) BB61
    |         |    `--Z. (*Z.) conradi (Ulrich & Bassler 1923) [=*Zygobolbina conradi] BB61
    |         `--Z. (Zygosella Ulrich & Bassler 1923) BB61
    |              `--Z. (*Z.) vallata (Ulrich & Bassler 1923) [=*Zygosella vallata] BB61
    `--Beyrichiidae [Beyrichiinae, Beyrichinae, Kloedeniinae, Kloedeninae] BB61
         |--Fallaticella schaeferi Schallreuter 1984 WSB93
         |--Apatobolbina Ulrich & Bassler 1923 BB61
         |    `--*A. granifera Ulrich & Bassler 1923 BB61
         |--Bolbibollia Ulrich & Bassler 1923 BB61
         |    `--*B. labrosa Ulrich & Bassler 1923 BB61
         |--Bolbiprimitia Kay 1940 BB61
         |    `--*B. fissurella (Ulrich & Bassler 1923) [=Halliella fissurella] BB61
         |--Cornikloedenia Henningsmoen 1954 BB61
         |    `--*C. ventralis (Ulrich & Bassler 1923) [=Drepanellina ventralis] BB61
         |--Ctenobolbinella Kummerow 1953 BB61
         |    `--*C. carinata Kummerow 1953 BB61
         |--Dolichoscapha Kesling & Ehlers 1958 BB61
         |    `--*D. escharota Kesling & Ehlers 1958 BB61
         |--Drepanellina Ulrich & Bassler 1923 BB61
         |    `--*D. clarki Ulrich & Bassler 1923 BB61
         |--Halliella Ulrich 1891 BB61
         |    `--*H. retifera Ulrich 1891 BB61
         |--Hibbardia Kesling 1953 BB61
         |    `--*H. lacrimosa (Swartz & Oriel 1948) [=Amphissites lacrimosus] BB61
         |--Mesomphalus Ulrich & Bassler 1913 [=Mezomphalus Zaspelova 1952] BB61
         |    `--*M. hartleyi Ulrich & Bassler 1913 BB61
         |--Myomphalus Swartz & Whitmore 1956 BB61
         |    `--*M. dorsinodosus Swartz & Whitmore 1956 BB61
         |--Phlyctiscapha Kesling 1953 BB61
         |    `--*P. rockportensis Kesling 1953 BB61
         |--Pseudobeyrichia Swartz & Whitmore 1956 BB61
         |    `--*P. perornata Swartz & Whitmore 1956 BB61
         |--Saccarchites Swartz & Whitmore 1956 BB61
         |    `--*S. saccularis Swartz & Whitmore 1956 BB61
         |--Welleria Ulrich & Bassler 1923 BB61
         |    `--*W. obliqua Ulrich & Bassler 1923 BB61
         |--Bodenia Ivanova 1959 LH61
         |    `--*B. aspera Ivanova 1959 LH61
         |--Balticella Thorslund 1940 M61, BB61
         |    `--*B. oblonga Thorslund 1940 BB61
         |--Pseudoleperditia Schneider 1956 WSB93, BB61
         |    `--*P. tuberculifera Schneider 1956 BB61
         |--Dibolbina Ulrich & Bassler 1923 BB61
         |    |--*D. cristata Ulrich & Bassler 1923 BB61
         |    `--D. steusloffi BB61
         |--Lophokloedenia Swartz & Whitmore 1956 BB61
         |    |--*L. manliensis (Weller 1903) [=Beyrichia manliensis] BB61
         |    `--L. kummeli BB61
         |--Welleriopsis Swartz & Whitmore 1956 BB61
         |    |--*W. diplocystulis Swartz & Whitmore 1956 BB61
         |    `--W. jerseyensis BB61
         |--Zygobeyrichia Ulrich 1916 BB61
         |    |--*Z. apicalis Ulrich 1916 BB61
         |    `--Z. barretti BB61
         |--Kyamodes Jones 1888 [=Kyammodes Ulrich & Bassler 1908] BB61
         |    |--*K. whidbornei Jones 1888 BB61
         |    |--K. kiesowi BB61
         |    `--K. tricornis BB61
         |--Kloedenia Jones & Holl 1886 BB61
         |    |--*K. wilckensiana (Jones 1855) [=Beyrichia wilckensiana] BB61
         |    |--K. apiculata Jones 1888 C01
         |    |--K. australis Chapman 1920 F71
         |    |--K. fifieldensis Chapman 1920 F71
         |    |--K. gotlandica Chapman 1901 C01
         |    |--K. normalis BB61
         |    `--K. sussexensis BB61
         `--Beyrichia BB61

*Type species of generic name indicated

REFERENCES

[BB61] Benson, R. H., J. M. Berdan, W. A. van den Bold, T. Hanai, I. Hessland, H. V. Howe, R. V. Kesling, S. A. Levinson, R. A. Reyment, R. C. Moore, H. W. Scott, R. H. Shaver, I. G. Sohn, L. E. Stover, F. M. Swain & P. C. Sylvester-Bradley. 1961. Systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q99–Q421. Geological Society of America and University of Kansas Press.

[C01] Chapman, F. 1901. On some fossils of Wenlock age from Mulde, near Klinteberg, Gotland; with notes by Prof. T. Rupert Jones and Dr. F. A. Bather. Annals and Magazine of Natural History, series 7, 7: 141–160, pl. 3.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[LH61] Levinson, S. A., & H. V. Howe. 1961. Addendum. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q421–Q422. Geological Society of America and University of Kansas Press.

[M61] Moore, R. C. 1961. Supplement—USSR treatise on Ostracoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt Q. Arthropoda 3: Crustacea: Ostracoda pp. Q422–Q429. Geological Society of America and University of Kansas Press.

[WSB93] Whatley, R. C., D. J. Siveter & I. D. Boomer. 1993. Arthropoda (Crustacea: Ostracoda). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 343–356. Chapman & Hall: London.