Belongs within: Rhynchonellata.
Contains: Atriboniidae, Stenoscismatidae, Rhynchotrematidae, Trigonirhynchidae, Eatoniidae, Uncinulidae, Wellerellinae, Lacunosellinae, Cirpinae, Camarotoechiinae, Rhynchonellininae, Rhynchonellidae, Oligorhynchiidae, Yunnanellidae, Basiliolidae, Frieleiidae, Leptocoeliidae, Allorhynchiidae.
The Rhynchonellida are a lineage of brachiopods that are first known from the Middle Ordovician, surviving into the present day. The classification of the order is complicated by a historical difference in study methods applied to Palaeozoic and post-Palaeozoic taxa (Ager et al. 1965).
Palaeozoic taxa include the Middle Devonian to Upper Permian Stenoscismatoidea which possess an elongate trough-shaped structure, called the camarophorium, on a high median septum duplex in the brachial valve, and typically also possess a spondylium in the pedicle valve. The Mesozoic Rhynchonelloidea lack a camarophorium, and commonly also lack a spondylium. Both are also absent in the Carboniferous to Permian genus Rhynchopora, which possesses a punctate shell in contrast to the impunctate shell of Rhynchonelloidea (Ager et al. 1965).
The Lower Devonian to Lower Carboniferous Pugnacidae had shells with relatively few costae, sometimes none, and often with fine radial striae (Ager et al. 1965). The Lower Ordovician to Lower Carboniferous Rhynchotrematoidea had a subpentagonal outline and contained long, often raduliform crura. The Lower Silurian to Upper Devonian Uncinuloidea had a high tongue on the front of the ventral valve and usually bore anterior and lateral margin spines. The Pennsylvanian Cardiarinidae bore an elongate beak with strong inner beak ridges and an elaborate parathyridium (Ager et al. 1965). The Lower Carboniferous to Upper Cretaceous Wellerellidae were uniplicate with entire hinge plates and no septalium or cardinal processes (Ager et al. 1965). The Lower Carboniferous to Lower Permian Rhynchotetradidae were wedge-shaped posteriorly with an acute apical angle and large concave interareas. The Silurian to Permian Camarotoechiidae were medium-sized to large brachiopods with a round or elliptical outline and generally rounded costae (Ager et al. 1965). The Mesozoic Dimerellidae are usually sulcate and very small, with reduced deltidial plates and very long crura (Ager et al. 1965).
Characters (Ager et al. 1965): Articulate brachiopods, usually with rostrate shell, functional pedicle developed, delthyrium partially closed by deltidial plates. Mantle canals much branched, with one pair of main trunks in each mantle. Median septum commonly supporting septalium or hinge plates in brachial valve; dental plates usually present; spondylia normally absent. Recent representatives mostly with two pairs of metanephridia, lophophore spirolophous, with ventrodorsally directed cones supported by crura. Shell substance normally impunctate, rarely with inner fibrous layer punctate.
<==Rhynchonellida
| i. s.: Oligoptycherhynchus Sartenaer 1970RG04
| |--*O. daleidensis (Roemer 1844)RG04
| `--O. hexatomaRG04
| |--O. h. hexatomaRG04
| `--O. h. wetteldorfensis (Schmidt 1941)RG04
| Uncinuloides King 1931AG65
| `--*U. guadelupae (Shumard 1858) [=Rhynchonella guadelupae, U. guadelupensis King 1931]AG65
| Disphenia [Dispheniidae]HB93
| `--D. myiodes Grant 1988HB93
|--Stenoscismatoidea [Camerophoriacea, Stenoscismacea, Stenoscismatacea]HB93
| |--AtriboniidaeAG65
| `--StenoscismatidaeAG65
|--RhynchotrematoideaRG04
| |--RhynchotrematidaeM03
| `--TrigonirhynchidaeRG04
|--UncinuloideaRG04
| |--EatoniidaeRG04
| `--UncinulidaeAG65
|--Cardiarinidae [Lambdarinoidea]WCB00
| |--Loborina lobata Balinski 1982HB93
| |--Lambdarina iota Grant 1988HB93
| `--Cardiarina Cooper 1956AG65
| `--*C. cordata Cooper 1956AG65
|--Wellerellidae [Wellerelloidea]WCB00
| | i. s.: Yakukijaella dubatolovi Baranov 1977HB93
| | Tetratomia Schmidt 1941HB93, AG65
| | |--*T. tetratoma (Schnur 1851) [=Terebratula tetratoma]AG65
| | `--T. amanshauseri (Dahmer 1942)HB93
| |--WellerellinaeAG65
| |--LacunosellinaeAG65
| `--CirpinaeAG65
|--Rhynchotetradidae [Rhynchotetradoidea, Rhynchotetraidae, Rhynchotetridae]WCB00
| |--Axiodeaneia Clark 1917AG65
| | `--*A. platypleura Clark 1917AG65
| |--Rhynchotetra Weller 1910AG65
| | |--*R. caputtestudinis (White 1862) [=Rhynchonella caputtestudinis]AG65
| | `--R. missouriensisAG65
| `--Goniophoria Yanishevskiy 1910AG65
| |--*G. monstrosa Yanishevskiy 1910AG65
| `--G. carinataAG65
|--Ancistrorhynchidae [Ancistrorhynchinae, Ancistrorhynchoidea]WCB00
| |--Kritorhynchia seclusa Rong & Yang 1981HB93
| |--Drepanorhyncha Cooper 1956AG65
| | `--*D. ottawaensis (Billings 1862) [=Porambonites ottawaensis]AG65
| `--Ancistrorhyncha Ulrich & Cooper 1942AG65
| |--*A. costata Ulrich & Cooper 1942AG65
| |--A. modesta Popov 1981HB93
| |--A. perplexa Cooper 1956HB93
| |--A. prisca Popov 1981HB93
| `--A. vacua Cooper 1956HB93
|--Camarotoechiidae [Camarotoechioidea]WCB00
| | i. s.: ParacamarotoechiaJB12
| |--CamarotoechiinaeMM02
| `--Septalariinae [Pseudopugnaxinae, Septalariidae]AG65
| |--Septalaria Leidhold 1928AG65
| | `--S. subtetragona (Schnur 1851) (see below for synonymy)AG65
| |--Amissopecten Havlíček 1960AG65
| | `--*A. velox (Barrande 1847) [=Terebratula velox]AG65
| |--Pseudocamarophoria Wedekind 1925AG65
| | `--*P. microrhyncha (Roemer 1844) [=Terebratula microrhyncha]AG65
| `--Pseudopugnax Likharev 1956AG65
| `--*P. planissimus Likharev 1956AG65
|--Dimerellidae [Dimerelloidea]AG65
| |--RhynchonellininaeAG65
| |--Dimerella Zittel 1870 [Dimerellinae]AG65
| | `--*D. guembeli Zittel 1870AG65
| |--Peregrinella Oehlert 1887AG65
| | |--P. multicarinata (Lamarck 1819) (see below for synonymy)AG65
| | `--P. whitneyiAG65
| `--HalorellinaeAG65
| |--Halorelloidea Ager 1960AG65
| | `--*H. rectifrons (Bittner 1884) [=Halorella rectifrons]AG65
| `--Halorella Bittner 1884 [incl. Barzellinia deGregorio 1930]AG65
| |--*H. amphitoma (Bronn 1832) [=Rhynchonella amphitoma]AG65
| `--‘*Barzellinia’ primogenita deGregorio 1930AG65
|--Rhynchoporoidea [Rhynchoporacea]HB93
| |--Aratella [Aratellidae]HB93
| | |--A. aratica (Abrahamians 1957)HB93
| | `--A. dichotomians (Abrahamians 1954)HB93
| `--Rhynchopora King 1865 [=Rhynchoporina Oehlert 1887; Rhynchoporidae]AG65
| |--*R. geinitziana (de Verneuil 1845) [=Terebratula geinitziana, *Rhynchoporina geinitziana]AG65
| |--R. basedowi Etheridge 1918F71
| |--R. beecheriB86
| |--R. depressaG31
| |--R. emeritaG31
| |--R. morini (Drot 1964)HB93
| |--R. nikitiniG31
| |--R. variabilisG31
| `--R. wynneiG31
|--Hemithyrididae [Hemithiridoidea, Hemithyrinae]WCB00
| |--Protegulorhynchia meridionalis Owen 1980HB93
| |--Notosaria Cooper 1959 [Notosariidae]AG65
| | `--*N. nigricans (Sowerby 1846) [=Terebratula nigricans]AG65
| |--Plicirhynchia Allan 1947AG65
| | `--*P. plicigera (von Ihering 1897) [=Rhynchonella plicigera]AG65
| |--Hemithiris d’Orbigny 1847 [=Hemithyris Bronn 1848]AG65
| | |--*H. psittacea (Gmelin 1790) [=Anomia psittacea]AG65
| | `--H. woodwardiAG65
| `--Tegulorhynchia Chapman & Crespin 1923AG65
| |--*T. squamosa (Hutton 1873) [=Rhynchonella squamosa]AG65
| |--T. boongeroodaensis McNamara 1983HB93
| |--T. doederleiniAG65
| `--T. nigricansN86
|--Norellidae [Norellinae, Norelloidea]WCB00
| |--Norella Bittner 1890AG65
| | `--*N. refractifrons (Bittner 1890) [=Rhynchonella refractifrons]AG65
| |--Apringia deGregorio 1886AG65
| | `--*A. giuppa deGregorio 1886AG65
| |--Austriellula Strand 1928 (see below for synonymy)AG65
| | `--*A. dilatata (Suess 1855) [=Rhynchonella dilatata, *Austriella dilatata, *Austriellina dilatata]AG65
| |--Pisirhynchia Buckman 1918AG65
| | `--*P. pisoides (Zittel 1869) [=Rhynchonella pisoides]AG65
| |--Rectirhynchia Buckman 1918AG65
| | `--*R. lopensis (Moore 1855) [=Rhynchonella lopensis]AG65
| |--Monticlarella Wisniewska 1932AG65
| | |--*M. czenstochowiensis (Roemer 1870) [=Rhynchonella czenstochowiensis]AG65
| | `--M. jefferiesi Owen 1968HB93
| `--Nannirhynchia Buckman 1918AG65
| |--*N. subpygmaea Buckman 1918AG65
| `--N. pygmaeaAG65
|--Pugnacidae [Pugnacoidea, Pugnaxinae]WCB00
| |--Donella Rotay 1931AG65
| | `--*D. minima Rotay 1931AG65
| |--Isopoma Torley 1934AG65
| | `--*I. brachyptyctum (Schnur 1853) [=Terebratula brachyptycta]AG65
| |--Parapugnax Schmidt 1964AG65
| | `--*P. brecciae (Schmidt 1941) [=Pugnax pugnus brecciae]AG65
| |--Shumardella Weller 1910AG65
| | `--*S. missouriensis (Shumard 1855) [=Rhynchonella missouriensis]AG65
| |--XeniopugnaxHB93
| | |--X. lynx (Barrande 1879)HB93
| | `--X. modicus (Barrande 1847)HB93
| |--Pugnoides Weller 1910HB93, AG65
| | |--*P. ottumwa (White 1862) [=Rhynchonella ottumwa]AG65
| | `--P. tardivensis Waterhouse 1982HB93
| `--Pugnax Hall & Carke 1893AG65
| |--*P. acuminatus (Sowerby 1822) [=Terebratula acuminata]AG65
| |--P. connivensG31
| |--P. granulumG31
| |--P. osagensisG31
| |--P. pugnusNU83
| |--P. swallowianaG31
| `--P. utaG31
`--Rhynchonelloidea [Rhynchonellacea]HB93
| i. s.: Diabolirhynchia Drot 1964M65
| `--*D. hollardi Drot 1964M65
| Dorsisinus Sanders 1958AG65
| `--*D. louisianensis (Weller 1914) [=Centronella louisianensis]AG65
| Ladogifornix Schmidt 1964AG65
| `--*L. fornicata (Schnur 1853) [=Terebratula fornix]AG65
| Laosia Mansuy 1913AG65
| `--*L. dussaulti Mansuy 1913AG65
| Leiorhynchoides Dovgal 1953AG65
| `--*L. gratianovae Dovgal 1953AG65
| Lissopleura Whitfield 1896AG65
| `--*L. aequivalvis (Hall 1857) [=Rhynchonella aequivalvis]AG65
| Monadotoechia Havlíček 1960AG65
| |--*M. monas (Barrande 1847) [=Terebratula monas]AG65
| `--M. monadinaAG65
| Nantanella Grabau 1936AG65
| `--*N. mapingensis Grabau 1936AG65
| Nemesa Schmidt 1941AG65
| `--*N. nemesana Schmidt 1941AG65
| Nyege Veevers 1959AG65
| `--*N. scopimus Veevers 1959AG65
| Payuella Grabau 1934AG65
| `--*P. obscura Grabau 1934AG65
| Phoenitoechia Havlíček 1960AG65
| `--*P. phoenix (Barrande 1847) [=Terebratula phoenix]AG65
| Pleiopleurina Schmidt 1964AG65
| `--*P. pleiopleura (Conrad 1841) [=Atrypa pleiopleura]AG65
| Praegnantenia Havlíček 1961AG65
| `--*P. praegnans (Barrande 1847) [=Terebratula praegnans]AG65
| Protorhyncha Hall & Clarke 1893AG65
| `--*P. dubia (Hall 1847) [=Atrypa dubia]AG65
| Straelenia Maillieux 1935 [incl. Dinapophysia Maillieux 1935]AG65
| `--*S. dunensis (Drevermann 1902) (see below for synonymy)AG65
| Zilimia Nalivkin 1947AG65
| |--*Z. polonica (Gürich 1896) [=Rhynchonella polonica]AG65
| `--Z. mugodjaricaAG65
| Basilicorhynchus Crickmay 1952AG65
| `--*B. basilicum (Crickmay 1952) [=Leiorhynchus basilicum]AG65
| Ladogioides McLaren 1961 [incl. Athabaschia Crickmay 1963]AG65
| |--*L. pax McLarren 1961AG65
| |--L. asmenista (Crickmay 1963)NU83
| |--L. kakwaensisNU83
| `--L. mollicomus Crickmay 1963NU83
| Linguopugnoides Havlíček 1960AG65
| `--*L. carens (Barrande 1879) [=Rhynchonella nympha carens]AG65
| Trifidorostellum Sartenaer 1961 [incl. Pseudoleiorhynchus Rozman 1962]AG65
| |--*T. dunbarense (Haynes 1916) [=Leiorhynchus dunbarense]AG65
| `--‘Leiorhynchus’ uralicus Nalivkin 1947 [=*Pseudoleiorhynchus uralicus]AG65
|--RhynchonellidaeAG65
|--OligorhynchiidaeAG65
|--YunnanellidaeAG65
|--BasiliolidaeAG65
|--FrieleiidaeAG65
|--LeptocoeliidaeHB93
|--AllorhynchiidaeHB93
|--Camerophorina Schmidt 1941 [Camarophorinidae, Camerophorinidae]AG65
| |--*C. pachyderma (Quenstedt 1871) [=Terebratula pachyderma]AG65
| `--C. bijugata (Schnur 1851)HB93
|--Erymnaria Cooper 1959 [Erymnariidae]AG65
| |--*E. polymorpha (Massalongo 1850) [=Terebratula polymorpha]AG65
| `--E. cubensis Cooper 1959HB93
|--Pygmaella [Pygmaellidae]HB93
| |--P. orbiculata Baranov 1977HB93
| `--P. pygmaea Baranov 1977HB93
|--PhoenicitoechiidaeHB93
| |--Phoenicitoechia phoenix (Barrande 1847)HB93
| `--Kotysex simulans (Barrande 1879)HB93
|--RozmanariidaeHB93
| |--Rackirhynchia lacerata (Barrande 1879)HB93
| `--Hadyrhyncha hadyensis Havlíček 1979HB93
|--AntistrixidaeHB93
| |--Antistrixia invicta Johnson 1972HB93
| `--Kindleina suemezensis Savage, Erberlein & Churkin 1978HB93
|--Amphipella [Amphidellidae]HB93
| |--A. arcaria Cooper & Grant 1976HB93
| `--A. attenuata Cooper & Grant 1976HB93
|--PetasmatheridaeHB93
| |--Iotina minuta Cooper & Grant 1976HB93
| `--Petasmatherus opulus Cooper & Grant 1976HB93
|--Septirhynchia Muir-Wood 1935 [Septirhynchiidae]AG65
| |--*S. azaisi (Cottreau 1924) [=Rhynchonella azaisi]AG65
| |--S. budulcaensis (Steffanini 1932)HB93
| `--S. hirschi Feldman 1987HB93
|--InnaechiidaeHB93
| |--Alekseevaella salagaensis (Rzonsnickaja 1967)HB93
| `--Lepidocycloides Nikiforova 1961HB93, AG65
| `--*L. baikiticus Nikiforova 1961AG65
|--PlectorhynchellidaeM03
| |--Sibiritoechia Alekseeva 1966M03
| | `--S. convexa Alekseeva 1966 [incl. S. lata Alekseeva 1966, S. oblonga Alekseeva 1966]M03
| `--Plectorhynchella Cooper & Muir-Wood 1951 [=Monticola Nalivkin 1930 non Boie 1822]AG65
| |--*P. collinensis (Frech 1902) [=Athyris collinensis, *Monticola collinensis]AG65
| `--P. markovskiiAG65
|--Cryptopora Jeffreys 1869 (see below for synonymy)AG65
| |--*C. gnomon (Jeffreys 1876) [=*Atretia gnomon, *Neatretia gnomon]AG65
| |--C. antiqua Bitner & Pisera 1979HB93
| |--C. brazieri (Crane 1886) [=Atretia brazieri]H09
| |--‘*Mannia’ nysti Davidson 1874AG65
| `--C. parvillima (Sacco 1902)HB93
|--DogdoidaeHB93
| |--Selennjachia abaimovae Baranov 1982HB93
| |--Tatjanica trigona Baranov 1982HB93
| `--Corvinopugnax Havlíček 1961HB93, AG65
| |--*C. corvinus (Barrande 1847) [=Rhynchonella corvina]AG65
| `--C. bimbax Havlíček 1990HB93
|--Tetracameridae [Tetracamerinae]AG65
| |--Tetracamera Weller 1910AG65
| | `--*T. subcuneata (Hall 1858) [=Rhynchonella subcuneata]AG65
| |--Rotaia Rzhonsnitskaya 1959 [=Welleria Rotay 1941 non Ulrich & Bassler 1923]AG65
| | `--*R. subtrigona (Meek & Worthen 1860) [=Rhynchonella subtrigona, *Welleria subtigona]AG65
| `--Yanishewskiella Likharev 1957AG65
| `--*Y. angulata (Linné 1767) [=Anomia angulata, Goniophoria angulata]AG65
|--PontisiidaeHB93
| |--Pontisia leonica (Martinez-Chacon 1979)HB93
| |--Lirellaria costellata Cooper & Grant 1976HB93
| |--AphaurosiaHB93
| | |--A. rotundata Cooper & Grant 1976HB93
| | `--A. scutata Cooper & Grant 1976HB93
| `--AnteridocusHB93
| |--A. bicostatus Cooper & Grant 1976HB93
| `--A. swallovianus (Shumard 1860)HB93
`--KatuniidaeHB93
|--Katunia Kulkov 1963AG65
| `--*K. subtrigonata Kulkov 1963AG65
|--Sulcatina Schmidt 1964HB93, AG65
| `--*S. sulcata (Cooper 1942) [=Trigonirhynchia sulcata]AG65
`--Terebratuloides Waagen 1883HB93, AG65
|--*T. davidsoni Waagen 1883AG65
|--T. elongataG31
|--T. senexG31
`--T. triplicataG31
Austriellula Strand 1928 [=Austriella Bittner 1890 non Tenison-Woods 1881, Austriellina Schuchert & LeVene 1929; incl. Jacobella Patte 1926 non Jeannet 1908]AG65
Cryptopora Jeffreys 1869 [=Atretia Jeffreys 1870, Neatretia Fischer & Oehlert 1891; incl. Mannia Davidson 1874 nec Opiz 1829 (ICBN) nec Trevis. 1857 (ICBN); Cryptoporidae]AG65
Peregrinella multicarinata (Lamarck 1819) [=Terebratula multicarinata; incl. T. peregrina von Buch 1835 non Schlotheim 1913, *Peregrinella peregrina]AG65
Septalaria subtetragona (Schnur 1851) [=Terebratula subtetragona; incl. T. ascendens Steininger 1853, *Septalaria ascendens]AG65
*Straelenia dunensis (Drevermann 1902) [=Rhynchonella dunensis; incl. R. dannenbergi mut. minor Drevermann 1902]AG65
*Type species of generic name indicated
References
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[B86] Brezinski, D. K. 1986. Trilobites from the Keokuk Limestone (Mississippian) of Missouri. Annals of Carnegie Museum 55 (8): 137–143.
[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.
[G31] Grabau, A. W. 1931. The Permian of Mongolia: A report on the Permian fauna of the Jisu Honguer limestone of Mongolia and its relations to the Permian of other parts of the world. American Museum of Natural History: New York.
[HB93] Harper, D. A. T., C. H. C. Brunton, L. R. M. Cocks, P. Copper, E. N. Doyle, A. L. Jeffrey, E. F. Owen, M. A. Parkes, L. E. Popov & C. D. Prosser. 1993. Brachiopoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 427–462. Chapman & Hall: London.
[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.
[JB12] Johnson, M. E., & B. G. Baarli. 2012. Development of intertidal biotas through Phanerozoic time. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 63–128. Springer.
[M03] Modzalevskaya, T. L. 2003. Silurian and Devonian brachiopods from Severnaya Zemlya (Russian Arctic). Geodiversitas 25 (1): 73–107.
[M65] Moore, R. C. (ed.) 1965. Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).
[MM02] Musteikis, P., & T. L. Modzalevskaya. 2002. Some Silurian brachiopods from Lithuania and their palaeobiogeographical significance. Palaeontology 45 (3): 595–626.
[N86] Nield, E. W. 1986. Liljevallia gotlandica: encrustation patterns in the earliest cemented articulate brachiopod, and their implications for its larval behaviour. Palaeogeography, Palaeoclimatology, Palaeoecology 56: 277–290.
[NU83] Norris, A. W., & T. T. Uyeno. 1983. Biostratigraphy and paleontology of middle-upper Devonian boundary beds, Gypsum Cliffs area, northeastern Alberta. Geological Survey of Canada Bulletin 313: 1–65.
[RG04] Racheboeuf, P. R., R. Gourvennec, M. Deynoux & D. Brice. 2004. The Devonian of the Hodh area (Islamic Republic of Mauritania): paleontology and stratigraphy. Journal of Paleontology 78 (1): 98–110.
[WCB00] Williams, A., S. J. Carlson & C. H. C. Brunton. 2000. Brachiopod classification. In: Kaesler, R. L. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda, Revised vol. 2. Linguliformea, Craniiformea, and Rhynchonelliformea (part) pp. 1–27. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).