Camarotoechiidae

Various views of Leiorhynchus specimen, from the Peabody Museum of Natural History.

Belongs within: Rhynchonellida.

The Camarotoechiidae are a group of medium-sized to large brachiopods known from the Silurian to the Permian (Ager et al. 1965).

Characters (from Ager et al. 1965): Medium-sized to large, round or elliptical in outline, sides and front never truncated; costae generally rounded. Brachial valve with high median septum or ridge; hinge plate entire, or divided only in its most anterior portion, or divided by small shallow open septalium; cardinal process usually absent.

<==Camarotoechiidae AG65
    |--Septalariinae [Pseudopugnaxinae, Septalariidae] AG65
    |    |--Septalaria Leidhold 1928 AG65
    |    |    `--S. subtetragona (Schnur 1851) (see below for synonymy) AG65
    |    |--Amissopecten Havlíček 1960 AG65
    |    |    `--*A. velox (Barrande 1847) [=Terebratula velox] AG65
    |    |--Pseudocamarophoria Wedekind 1925 AG65
    |    |    `--*P. microrhyncha (Roemer 1844) [=Terebratula microrhyncha] AG65
    |    `--Pseudopugnax Likharev 1956 AG65
    |         `--*P. planissimus Likharev 1956 AG65
    `--Camarotoechiinae [Leiorhynchinae, Nudirostrinae] MM02
         |--Astutorhyncha Havlíček 1961 AG65
         |    `--*A. proserpina (Barrande 1847) [=Rhynchonella proserpina] AG65
         |--Calvinaria Stainbrook 1945 AG65
         |    `--*C. ambigua (Calvin 1878) [=Rhynchonella ambigua] AG65
         |--Felinotoechia Havlíček 1961 AG65
         |    `--*F. felina (Barrande 1879) [=Atrypa astuta var. felina] AG65
         |--Fenestrirostra Cooper 1955 AG65
         |    `--*F. glacialis (Billings 1862) [=Rhynchonella glacialis] AG65
         |--Hyborhynchella Cooper 1955 AG65
         |    `--*H. bransoni Cooper 1955 AG65
         |--Paranorella Cloud 1944 AG65
         |    `--*P. imperialis Cloud 1944 AG65
         |--Sanjuania Amos 1958 AG65
         |    `--*S. dorsisulcata Amos 1958 AG65
         |--Rhynchotretina Khalfin 1939 AG65
         |    `--*R. aequivalvis Khalfin 1939 AG65
         |--Rugaltarostrum Sartenaer 1961 AG65
         |    `--*R. madisonense (Haynes 1916) [=Leiorhynchus madisonense] AG65
         |--Camarotoechia Hall & Clarke 1893 AG65
         |    |--*C. congregata (Conrad 1841) [=Atrypa congregata] AG65
         |    `--C. allegania N79
         |--Caryorhynchus Crickmay 1952 AG65
         |    |--*C. carya (Crickmay 1952) [=Leiorhynchus carya] N79
         |    `--C. globiliforme N79
         |--Leiorhynchoidea Cloud 1944 AG65
         |    |--*L. schucherti Cloud 1944 AG65
         |    `--L. carboniferum (Girty 1911) [=Leiorhynchus carboniferum] AG65
         |--Paurorhyncha Cooper 1942 AG65
         |    |--*P. endlichi (Meek 1875) [=Rhynchonella endlichi] AG65
         |    `--P. cooperi AG65
         |--Plagiorhyncha McLearn 1918 MM02
         |    |--P. depressa (Sowerby 1839) (see below for synonymy) MM02
         |    |--P. niobe (Barrande 1847) MM02
         |    `--P. thisbe (Barrande 1847) MM02
         |--Hircinisca Havlíček 1960 AG65
         |    |--*H. sapho (Barrande 1879) [=Atrypa hircina var. sapho] AG65
         |    |--H. hebes AG65
         |    `--H. hircina AG65
         `--Leiorhynchus Hall 1860 (see below for synonymy) AG65
              |--*L. quadracostatus (Vanuxem 1842) [=Orthis quadracostata, *Nudirostra quadracostata] AG65
              |--L. castanea N79
              |--L. hippocastanea N79
              |--L. mesacostale N79
              |--L. miram N79
              |    |--L. m. miram N79
              |    `--L. m. alpha N79
              |--L. sinuatum N79
              `--L. (Zigania) ursus H79

Leiorhynchus Hall 1860 [=Liorhynchus Oehlert 1887 non Rudolphi 1801, Nudirostra Cooper & Muir-Wood 1951] AG65

Plagiorhyncha depressa (Sowerby 1839) [=Atrypa depressa; incl. A. analoga Wenjukow 1899, Plagiorhyncha analoga, Spirigerina cordata Lindström 1861, P. cordata, Rhynchonella glassii Davidson 1883, *P. glassii, Septatrypa subanaloga Rybnikova 1967, P. subanaloga] MM02

Septalaria subtetragona (Schnur 1851) [=Terebratula subtetragona; incl. T. ascendens Steininger 1853, *Septalaria ascendens] AG65

*Type species of generic name indicated

REFERENCES

[AG65] Ager, D. V., R. E. Grant, D. J. McLaren & H. Schmidt. 1965. Rhynchonellida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H552–H632. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[H79] House, M. R. 1979. Devonian in the Eastern Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A183–A217. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[MM02] Musteikis, P., & T. L. Modzalevskaya. 2002. Some Silurian brachiopods from Lithuania and their palaeobiogeographical significance. Palaeontology 45 (3): 595–626.

[N79] Norris, A. W. 1979. Devonian in the Western Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A218–A253. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

Delthyrididae

Howellella elegans, from Primitive Worlds.

Belongs within: Spiriferida.

The Delthyrididae are a group of spiriferid brachiopods known from the Lower Silurian to the Middle Devonian (Boucot et al. 1965).

Characters (from Boucot et al. 1965): Biconvex, weakly to strongly transverse; lateral slopes plicate or costate; fold and sulcus smooth or with median rib in sulcus and median groove on fold; micro-ornament typically consisting of distinct growth lamellae upon which are superposed capillae that become fimbriate at anterior margin of each growth lamella, more rarely consisting of capillae only or of teardrop-shaped granules; interior of pedicle valve generally with well-developed dental plates, some with distinct median septum but none with delthyrial plate; brachial-valve interior with area of diductor attachment deeply striate longitudinally, resulting in comblike cardinal process; crural plates absent or short; shell substance impunctate.

<==Delthyrididae [Delthyridae] M03
    |  i. s.: Eohowellella Lopushinskaya 1976 M03
    |           |--E. minimus Lopushinskaya 1976 M03
    |           `--E. yadrenkinae Lopushinskaya 1976 M03
    |--Cyrtinopsis Scupin 1896 [Cyrtinopsidae, Cyrtinopsinae] BJ65
    |    `--*C. undosa (Schnur 1853) [=Spirifer undosus] BJ65
    |         |--C. u. undosa TC71
    |         `--C. u. maiderensis TC71
    |--Kozlowskiellininae [Kozlowskiellinae] BJ65
    |    |--Hedeina Boucot 1957 BJ65
    |    |    `--*H. crispa (Linnaeus 1758) [=Anomia crispa, Delthyris crispa] BJ65
    |    `--Kozlowskiellina Boucot 1958 (see below for synonymy) BJ65
    |         |--*K. strawi (Boucot 1957) [=*Kozlowskiella strawi] BJ65
    |         |--‘Spirifer’ perlamellosus Hall 1857 [=*Megakozlowskiella perlamellosa] BJ65
    |         `--K. raricosta BJ65
    |--Delthyridinae BJ65
    |    |--Howittia Talent 1956 BJ65
    |    |    `--*H. howitti (Chapman 1905) [=Spirifer howitti] BJ65
    |    |--Ivanothyris Havlíček 1957 BJ65
    |    |    `--*I. gibbosa (Barrande 1879) [=Spirifer gibbosus] BJ65
    |    |--Neodelthyris Hou 1963 BJ65
    |    |    `--*N. sinensis Hou 1963 BJ65
    |    |--Uralospirifer Havlíček 1959 BJ65
    |    |    `--*U. mansy (Khodalevich 1951) [=Spirifer (Delthyris) mansy] BJ65
    |    `--Delthyris Dalman 1828 [incl. Quadrifarius Fuchs 1929] BJ65
    |         |--*D. elevata Dalman 1828 BJ65
    |         |--D. dumontianus H79
    |         `--D. loculata Fuchs 1923 [=*Quadrifarius loculatus] BJ65
    `--Acrospiriferinae BJ65
         |--Antispirifer Williams 1916 BJ65
         |    `--*A. harroldi Williams 1916 BJ65
         |--Australospirifer Caster 1939 BJ65
         |    `--*A. kayserianus (Clarke 1913) [=Spirifer kayserianus] BJ65
         |--Eospiriferina Grabau 1931 BJ65
         |    `--*E. lachrymosa (Grabau 1931) [=Spiriferina (*Eospiriferina) lachrymosa] BJ65
         |--Mauispirifer Allan 1947 BJ65
         |    `--*M. hectori Allan 1947 BJ65
         |--Pinguispirifer Havlíček 1957 BJ65
         |    `--*P. infirmus (Barrande 1879) [=Spirifer infirmus] BJ65
         |--Spinella Talent 1956 BJ65
         |    |--*S. buchanensis Talent 1956 BJ65
         |    `--S. yassensis [=Spirifer yassensis] TC71
         |--Acrospirifer Helmbrecht & Wedekind 1923 XY86
         |    |--*A. primaevus (Steininger 1853) [=Spirifer primaeva] BJ65
         |    |--A. coxi C68
         |    |--A. hercyniae H79
         |    |--A. kobehana N79
         |    |--A. lilydalensis H79
         |    |--A. macrothyris N79
         |    |--A. murchisoni N79
         |    |--A. peltica H79
         |    `--A. tonkinensis (Mansuy 1908) WL04
         `--Howellella Kozlowski 1843 BJ65, XY86 [=Crispella Kozlowski 1929 non Gray 1870 BJ65]
              |--*H. elegans (Muir-Wood 1925) (see below for synonymy) BJ65
              |--H. angustiplicata (Kozlowski 1829) [=Spirifer (Crispella) angustiplicatus] XY86
              |--H. cycloptera XY86
              |--H. fecunda Chen 1983 XY86
              |--H. minora XY86
              |--H. pauciplicata Waite 1956 M03
              |--H. prima Alekseeva 1967 M03
              |--H. subbragensis (Kulkov 1963) XY86
              |--H. vanuxemi N79
              `--H. yukiangensis XY86

*Howellella elegans (Muir-Wood 1925) [=Delthyris elegans; incl. Terebratula crispus Hisinger 1826, D. crispa non Anomia crispa Linné 1758] BJ65

Kozlowskiellina Boucot 1958 [=Kozlowskiella Boucot 1957 non Přibyl 1953; incl. Megakozlowskiella Boucot 1957, Megakozlowskiellina Amsden & Ventress 1963] BJ65

*Type species of generic name indicated

REFERENCES

[BJ65] Boucot, A. J., J. G. Johnson, C. W. Pitrat & R. D. Staton. 1965. Spiriferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H632–H728. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[C68] Cooper, R. A. 1968. Lower and Middle Paleozoic fossil localities of north-west Nelson. Transactions of the Royal Society of New Zealand, Geology 6 (7): 75–89.

[H79] House, M. R. 1979. Devonian in the Eastern Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A183–A217. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[M03] Modzalevskaya, T. L. 2003. Silurian and Devonian brachiopods from Severnaya Zemlya (Russian Arctic). Geodiversitas 25 (1): 73–107.

[N79] Norris, A. W. 1979. Devonian in the Western Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A218–A253. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[TC71] Thomson, K. S., & K. S. W. Campbell. 1971. The structure and relationships of the primitive Devonian lungfish—Dipnorhynchus sussmilchi (Etheridge). Yale University Peabody Museum of Natural History Bulletin 38: 1–109.

[WL04] Wang S.-Q. & R. F. Lundin. 2004. Sinoleperditiini (Ostracoda) from the Lower Emsian Shanglun Formation at the Shanglun village, central Guangxi, China. Journal of Paleontology 78 (2): 349–358.

[XY86] Xu H.-K. & Yao Z.-G. 1986. The Lower Devonian and its brachiopods from Lingshan, Guangxi. Acta Palaeontologica Sinica 25 (2): 169–179.

Cyrtospiriferoidea

Partial brachial valve of Cyrtiopsis animasensis, from here.

Belongs within: Spiriferida.
Contains: Cyrtospirifer.

The Cyrtospiriferoidea are a group of Devonian spiriferids characterised by the presence of a delthyrial plate.

<==Cyrtospiriferoidea RG04
    |--Spinocyrtiidae [Guerichellinae, Spinocyrtiinae] RG04
    |    |--Acutatheca Stainbrook 1945 BJ65
    |    |    `--*A. propria Stainbrook 1945 BJ65
    |    |--Alatiformia Struve 1963 M65
    |    |    `--*A. alatiformis (Drevermann 1907) [=Spirifer alatiformis] M65
    |    |--Allanella Crickmay 1953 [=Allanaria Crickmay 1953] BJ65
    |    |    `--*A. allani (Warren 1944) [=Spirifer allani, *Allanaria allani] BJ65
    |    |--Chimaerothyris Paulus, Struve & Wolfart 1963 M65
    |    |    `--*C. hotzi Paulus, Struve & Wolfart 1963 M65
    |    |--Eosyringothyris Stainbrook 1943 BJ65
    |    |    `--*E. aspera (Hall 1858) [=Spirifera aspera] BJ65
    |    |--Subcuspidella Mittmeyer 1965 [incl. Tenuicostella] RG04
    |    |    |--*S. subcuspidata (Schnur 1851) [=Spirifer subcuspidatus] RG04
    |    |    `--S. incerta (Fuchs 1909) RG04
    |    |--Mediospirifer Bublichenko 1956 BJ65
    |    |    |--M. audacula (Conrad 1842) (see below for synonymy) BJ65
    |    |    `--M. mediotextus H79
    |    |--Spinocyrtia Frederiks 1916 [incl. Platyrachella Fenton & Fenton 1924] BJ65
    |    |    |--*S. granulosa (Conrad 1839) [=Delthyris granulosa] BJ65
    |    |    |--S. macbridei (Calvin 1883) [=Spirifera macbridei, *Platyrachella macbridei] BJ65
    |    |    |--S. marcyi N79
    |    |    `--‘Platyrachella’ mesastralis N79
    |    `--Adolfia Gürich 1909 (see below for synonymy) BJ65
    |         |--*A. deflexa (Roemer 1843) [=Spirifer deflexus, *Guerichella deflexa] BJ65
    |         |--‘Guerichella’ angustistellata BJ65
    |         |--A. bifida BJ65
    |         |--‘Spirifer (*Plectospirifer)’ heimi Grabau 1931 BJ65
    |         `--A. talasica H79
    `--Cyrtospiriferidae [Cyrtospiriferinae] BJ65
         |--Cyrtospirifer RG04
         |--Platyspirifer Grabau 1931 BJ65
         |    `--*P. paronai (Martelli 1902) [=Schizophoria paronai, Theodossia (*Platyspirifer) paronai] BJ65
         |--Tenticospirifer Tien 1938 BJ65
         |    `--*T. tenticulum (de Verneuil 1845) [=Spirifer tenticulum] BJ65
         |--Arctospirifer Stainbrook 1950 BJ65
         |    `--*A. constrictus Stainbrook 1950 BJ65
         |--Austrospirifer Glenister 1955 BJ65
         |    `--*A. variabilis Glenister 1955 BJ65
         |--Dmitria Sidyachenko 1961 BJ65
         |    `--*D. romanowskii (Nalivkin 1930) [=Spirifer (Cyrtospirifer) romanowskii] BJ65
         |--Prospira Maxwell 1954 BJ65
         |    `--*P. typa Maxwell 1954 BJ65
         |--Schizospirifer Grabau 1931 BJ65
         |    `--*S. latistriatus (Frech 1911) [=Spirifer aperturatus var. latistriatus] BJ65
         |--Sphenospira Cooper 1954 BJ65
         |    `--*S. alta (Hall 1866) [=Spirifera alta] BJ65
         |--Sulcatospirifer Maxwell 1954 BJ65
         |    `--*S. primus Maxwell 1954 BJ65
         |--Syringospira Kindle 1909 BJ65
         |    `--*S. prima Kindle 1909 BJ65
         |--Indospirifer Grabau 1931 BJ65
         |    `--*I. padaukpinensis (Reed 1908) [=Spirifer padaukpinensis] BJ65
         |         |--I. p. padaukpinensis BJ65
         |         `--I. p. maoerhchuanensis BJ65
         `--Cyrtiopsis Grabau 1923 (see below for synonymy) BJ65
              |--*C. davidsoni Grabau 1923 BJ65
              |--C. animasensis N79
              |--C. graciosa Grabau 1923 [=C. (*Grabauicyrtiopsis) graciosa] BJ65
              |--C. intermedia BJ65
              |--‘Uchtospirifer’ murchisonianus H79
              |--‘*Uchtospirifer’ nalivkini Lyashenko 1957 BJ65
              `--C. transversa Grabau 1931 [=C. (*Sinocyrtiopsis) transversa] BJ65

Adolfia Gürich 1909 [=Guerichella Paeckelmann 1931, Guerichia (l. c.); incl. Plectospirifer Grabau 1931] BJ65

Cyrtiopsis Grabau 1923 [incl. Alphacyrtiopsis Gatinaud 1949 (nom. vet.), Betacyrtiopsis Gatinaud 1949 (nom. vet.), Grabauicyrtiopsis Gatinaud 1949, Paracyrtiopsis Gatinaud 1949 (nom. vet.), Sinocyrtiopsis Gatinaud 1949, Uchtospirifer Lyashenko 1957] BJ65

Mediospirifer audacula (Conrad 1842) [=Delthyris audacula; incl. D. medialis Hall 1843, *Mediospirifer medialis] BJ65

*Type species of generic name indicated

REFERENCES

[BJ65] Boucot, A. J., J. G. Johnson, C. W. Pitrat & R. D. Staton. 1965. Spiriferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H632–H728. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[H79] House, M. R. 1979. Devonian in the Eastern Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A183–A217. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[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).

[N79] Norris, A. W. 1979. Devonian in the Western Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A218–A253. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[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.

Pentamerida

Porambonites (Porambonites) schmidti, from here.

Belongs within: Rhynchonellata.
Contains: Stricklandioidea, Gypidulinae, Pentameridae, Virgianidae, Huenellidae, Clarkellidae, Camerellidae, Parastrophinidae.

The Pentamerida are a group of brachiopods known from the Middle Cambrian to the Upper Devonian, with an impunctate shell and an unrestricted pedicle opening. In the Middle Cambrian to Lower Devonian Porambonitacea, the brachiophores are united by supporting plates of variable length. In the Middle Ordovician to Upper Devonian Pentameridina, the inner such plates are long and curved outward to meet the lateral walls of the valve (Amsden & Biernat 1965).

Characters (from Amsden & Biernat 1965): Generally biconvex shells with pedicle spondylium; delthyrium may be unmodified or at least partly closed by deltidium. Brachial valve with brachial processes, commonly braced at their posterior end by supporting plates and usually terminating blindly (ending in a loop in Enantiosphenidae). Shell impunctate.

<==Pentamerida
    |  i. s.: Tcherskidium Nikolaev & Sapelnikov 1969 RZJ04
    |         Kirkidium Amsden, Boucot & Johnson 1967 RZJ04
    |         Galeatellina Sapelnikov & Rukavishnikova 1976 RZJ04
    |         Eoconchidium Rozman 1967 RZJ04
    |         Vosmiverstum Breivel & Breivel 1970 RZJ04
    |         Prostricklandia Rukavishnikova & Sapelnikov 1973 RZJ04
    |         Proconchidium Rukavishnikova & Sapelnikov 1973 RZJ04
    |--Pentameridina [Pentameracea] RZJ04
    |    |  i. s.: Parallelelasmatidae AB65
    |    |           |--Parallelelasma Cooper 1956 AB65
    |    |           |    `--*P. pentagonum Cooper 1956 AB65
    |    |           |--Didymelasma Cooper 1956 AB65
    |    |           |    `--*D. longicrurum Cooper 1956 AB65
    |    |           |--Metacamerella Reed 1917 AB65
    |    |           |    `--*M. balcletchiensis (Davidson 1883) [=Stricklandinia balcletchiensis] AB65
    |    |           `--Salonia Cooper & Whitcomb 1933 AB65
    |    |                `--*S. magnaplicata Cooper & Whitcomb 1933 AB65
    |    |         Enantiosphen Whidborne 1893 [Enantiosphenidae] AB65
    |    |           `--*E. vicaryi (Davidson 1882) [=Meganteris vicaryi] AB65
    |    |--Stricklandioidea RZJ04
    |    |--Gypidulinae RZJ04
    |    |--Clorindinae AB65 [Clorindoidea RZJ04]
    |    |    |--Clorinda Barrande 1879 RZJ04
    |    |    |    `--*C. armata Barrand 1879 AB65
    |    |    |--Clorindella Amsden 1964 AB65
    |    |    |    `--*C. areyi (Hall & Clarke 1894) [=Barrandella areyi] AB65
    |    |    |--Antirhynchonella Oehlert in Fischer 1887 [=Barrandella Hall & Clarke 1894 (nom. rej.)] AB65
    |    |    |    `--*A. linguifera (Sowerby in Murchison 1839) [=Atrypa linguifera, *Barrandella linguifera] AB65
    |    |    `--Clorindina Khodalevich 1939 AB65
    |    |         `--*C. uralica Khodalevich 1939 AB65
    |    `--Pentameroidea RZJ04
    |         |  i. s.: Eokirkidium Khodalevich & Sapelnikov 1970 RZJ04
    |         |         Vagranella Sapelnikov 1960 RZJ04
    |         |           `--V. diversoplicata Sapelnikov 1960 RZJ04
    |         |         Subriana Sapelnikov 1960 RZJ04
    |         |         Mariannaella Sapelnikov & Rukavishnikova 1975 RZJ04
    |         |         Virgianella Nikiforova & Sapelnikov 1971 RZJ04
    |         |           |--V. glabella (Rong & Yang 1981) RZJ04
    |         |           `--V. sogdianica Nikiforova & Sapelnikov 1973 [=Holorhynchus (Nondia) sogdianica] RZJ04
    |         |         Pseudoconchidium Nikiforova & Sapelnikov 1971 RZJ04
    |         |         Lissidium Lenz 1989 RZJ04
    |         |--Pentameridae RZJ04
    |         `--Virgianidae RZJ04
    `--Porambonitacea [Syntrophiidina, Syntrophioidea] AB65
         |--Huenellidae AB65
         |--Clarkellidae AB65
         |--Camerellidae AB65
         |--Parastrophinidae AB65
         |--Cambrotrophia Ulrich & Cooper 1937 [=Eostrophia Ulrich & Cooper 1936 non Dall 1890; Eostrophiidae] AB65
         |    `--*C. cambria (Walcott 1908) [=Syntrophia cambrica, *Eostrophia cambria] AB65
         |--Lycophoria Lahusen 1886 [Lycophoriidae] AB65
         |    `--*L. nucella (Dalman 1828) [=Atrypa nucella] AB65
         |--Brevicamera Cooper 1956 [Brevicameridae] AB65
         |    `--*B. camerata Cooper 1956 AB65
         |--Alimbellidae AB65
         |    |--Alimbella Andreeva 1960 AB65
         |    |    `--*A. armata Andreeva 1960 AB65
         |    `--Medesia Andreeva 1960 AB65
         |         `--*M. uralica Andreeva 1960 AB65
         |--Syntrophiidae AB65
         |    |--Syntrophia Hall & Clarke 1893 AB65
         |    |    |--*S. lateralis (Whitfield 1886) [=Triplecia lateralis] AB65
         |    |    `--S. torynifera AB65
         |    `--Xenelasma Ulrich & Cooper 1936 [Xenelasmatinae, Xenelasminae] aB65
         |         `--*X. syntrophioides Ulrich & Cooper 1936 AB65
         |--Syntrophopsidae AB65
         |    |--Syntrophopsis Ulrich & Cooper 1936 AB65
         |    |    `--*S. magna Ulrich & Cooper 1936 AB65
         |    |--Hesperotrophia Ulrich & Cooper 1936 AB65
         |    |    `--*H. obscura Ulrich & Cooper 1936 AB65
         |    `--Rhysostrophia Ulrich & Cooper 1936 AB65
         |         `--*R. nevadensis Ulrich & Cooper 1936 AB65
         |--Tetralobulidae AB65
         |    |--Tetralobula Ulrich & Cooper 1936 AB65
         |    |    `--*T. delicatula Ulrich & Cooper 1936 AB65
         |    |--Doloresella Sando 1957 AB65
         |    |    `--*D. concentrica Sando 1957 AB65
         |    |--Imbricatia Cooper 1952 AB65
         |    |    `--*I. lamellata Cooper 1952 AB65
         |    `--Punctolira Ulrich & Cooper 1936 AB65
         |         `--*P. punctolira Ulrich & Cooper 1936 AB65
         `--Porambonites Pander 1830 [Porambonitidae] AB65
              |--P. (Porambonites) AB65
              |    |--*P. (P.) intermedia Pander 1830 AB65
              |    |--P. (P.) reticulatus Pander 1830 AB65
              |    `--P. (P.) schmidti AB65
              |--P. (Equirostra Cooper & Muir-Wood 1951) [=Isorhynchus King 1850 non Schoenherr 1833] AB65
              |    `--P. (*E.) aequirostris (von Schlotheim 1820) (see below for synonymy) AB65
              `--P. (Noetlingia Hall & Clarke 1894) AB65
                   `--P. (*N.) tscheffkini (de Verneuil 1840) [=Spirifer tscheffkini] AB65

Porambonites (*Equirostra) aequirostris (von Schlotheim 1820) [=Terebratulites aequirostris, P. (*Isorhynchus) aequirostris] AB65

*Type species of generic name indicated

REFERENCES

[AB65] Amsden, T. W., & G. Biernat. 1965. Pentamerida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H523–H552. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[RZJ04] Rong, J.-Y., R.-B. Zhan & J. Jin. 2004. The Late Odovician and Early Silurian pentameride brachiopod Holorhynchus Kiaer, 1902 from north China. Journal of Paleontology 78 (2): 287–299.

Atrypida

Anoplotheca venusta, from here.

Belongs within: Rhynchonellata.
Contains: Atrypidae, Lissatrypidae.

The Atrypida are a group of spiralia-bearing brachiopods known from the Middle Ordovician to Upper Devonian. Members of this group may be divided between the Atrypacea, in which the spiralia are directed medially or dorsomedially, and the Dayiacea, in which the spiralia are directed ventrally or laterally, or are planispiral parallel to the median plane. Within the Dayiacea, members of the Dayiidae have smooth shells whereas members of the other families are variously ornamented (Boucot et al. 1965).

Characters (from Boucot et al. 1965, as Atrypidina): Impunctate, mostly biconvex spire-bearing brachiopods, commonly with narrow cardinal margin. Interarea low, obsolescent, or absent. Pedicle-valve beak may be truncated by foramen or foramen may reside within delthyrium. Spiralia directed medially, dorsomedially, or laterally; crura projecting to join primary lamellae of spiralia at posteromedial position, or deflected laterally more or less parallel to plane of commissure to join primary lamellae in posterolateral position; simple jugum commonly present connecting primary lamellae.

<==Atrypida [Atrypidina, Atrypoidea]
    |--Atrypacea BJ65
    |    |  i. s.: Loilemia Reed 1936 BJ65
    |    |           `--*L. proxima Reed 1936 BJ65
    |    |--Atrypidae M03
    |    `--Lissatrypidae BJ65
    `--Dayiacea BJ65
         |--Kayseria Davidson 1882 [Kayseriidae] BJ65
         |--Dayiidae BJ65
         |    |--Aulidospira Williams 1962 [Aulidospiridae, Aulidospirinae] BJ65
         |    |    `--*A. trippi Williams 1962 BJ65
         |    |--Cyclospira Hall & Clarke 1893 [incl. Triplecella Wilson 1932; Cyclospiridae, Cyclospirinae] BJ65
         |    |    |--*C. bisulcata (Emmons 1842) [=Orthis bisulcata] BJ65
         |    |    `--‘*Triplecella’ duplicata Wilson 1932 BJ65
         |    `--Dayiinae [Dayinae, Protozeugidae] BJ65
         |         |--Dayia Davidson 1881 [=Daya (l. c.) non Bleeker 1877] BJ65
         |         |    `--*D. navicula (Sowerby in Murchison 1839) [=Terebratula navicula] BJ65
         |         `--Protozeuga Twenhofel 1913 BJ65
         |              `--*P. mawei (Davidson 1881) [=Waldheimia mawei] BJ65
         |--Leptocoeliidae BJ65
         |    |--Leptocoelia Hall 1857 BJ65
         |    |    `--L. flabellites (Conrad 1841) [=Atrypa flabellites; incl. *Leptocoelia propria Hall 1857] BJ65
         |    |--Anabaia Clarke 1893 BJ65
         |    |    `--*A. paraia Clarke 1893 BJ65
         |    |--Australocoelia Boucot & Gill 1956 BJ65
         |    |    `--*A. tourteloti Boucot & Gill 1956 BJ65
         |    `--Eocoelia Nikiforova 1961 BJ65
         |         |--*E. hemisphaerica [=Atrypa hemisphaerica] BJ65
         |         |--E. curtisi EB01
         |         `--E. intermedia EB01
         `--Anoplothecidae [Coelospiridae] BJ65
              |--Coelospira Hall 1863 [Coelospirinae] BJ65
              |    |--*C. concava (Hall 1857) [=Leptocoelia concava] BJ65
              |    `--C. virginia BJ65
              `--Anoplothecinae BJ65
                   |--Bifida Davidson 1882 BJ65
                   |    `--*B. lepida (d’Archiac & de Verneuil 1842) [=Terebratula lepida] BJ65
                   |--Coelospirina Havlíček 1956 BJ65
                   |    `--*C. modica Havlíček 1956 BJ65
                   `--Anoplotheca Sandberger 1855 [=Hoplotheca Bigsby 1878] BJ65
                        |--*A. venusta (Schnur 1853) (see below for synonymy) BJ65
                        `--A. australis Dun 1907 F71

*Anoplotheca venusta (Schnur 1853) [=Terebratula venusta, *Hoplotheca venusta; incl. Productus lamellosus Sandberger 1850-1856] BJ65

*Type species of generic name indicated

REFERENCES

[BJ65] Boucot, A. J., J. G. Johnson, C. W. Pitrat & R. D. Staton. 1965. Spiriferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H632–H728. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[EB01] Eckert, J. D., & C. E. Brett. 2001. Early Silurian (Llandovery) crinoids from the Lower Clinton Group, western New York State. Bulletins of American Paleontology 360: 1–88.

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

[M03] Modzalevskaya, T. L. 2003. Silurian and Devonian brachiopods from Severnaya Zemlya (Russian Arctic). Geodiversitas 25 (1): 73–107.

Brachiopoda

Mickwitzia monilifera, from here.

Belongs within: Brachiozoa.
Contains: Linguliformea, Craniiformea, Rhynchonelliformea.

The Brachiopoda are a phylum of animals superficially similar to bivalve molluscs, but with a very different anatomical structure. The valves of the shell are dorsal and ventral in position, with the ventral larger than the dorsal, with a posterior opening between the valves through with emerges the fleshy pedicle in most species (members of the Craniiformea lack a pedicle). The shell contains a large mantle cavity, which is occupied by a feathery feeding structure, the lophophore. Brachiopods were ecologically predominant during the Palaeozoic, but are a relatively minor part of the recent fauna (Prothero 1998).

The Lower Cambrian Mickwitzia possessed an oval to subcircular, ventri-biconvex shell with an elevated submarginal or posteriorly recurved apex on the pedicle valve, and a low submarginal apex on the brachial valve (Rowell 1965). This genus has been identified as a stem-group brachiopod, retaining setigerous tubes passing through the pseudointerarea in the rear of the pedicle valve similar to those found in the tommotiids, a group of likely fossil brachiopod relatives. Such setigerous tubes have been lost in the crown group (Williams & Homer 2002). Brachiopods are also represented in the Lower Cambrian by the genus Salanygolina which shares some features with members of the crown subgroup Rhynchonelliformea (Erwin et al. 2011). Living brachiopods are divided between the Linguliformea and Calciata, with the latter characterised by calcareous shells in contrast to the phosphatic shells of the former (Holmer et al. 1995).

Ptychopeltis incola is an arched, saddle-shaped shell from the Middle Ordovician, originally described as a monoplacophoran mollusc but since re-interpreted as a brachiopod (Rowell 1965).

See also: Taxon of the Week: The lamp (shell) post.

Brachiopoda (see below for synonymy)
    |--Mickwitzia Schmidt 1888 WH02, R65 [incl. Causea Wiman 1902 R65]
    |    |--*M. monilifera (Linnarsson 1869) [=Lingula monilifera] R65
    |    `--‘*Causea’ formosa Wiman 1902 R65
    |--Salanygolina EL11
    `--+--Linguliformea WH02
       `--Calciata HP95
            |--Craniiformea WH02
            `--Rhynchonelliformea WH02

Brachiopoda incertae sedis:
  Araratella WM03
  Spinocarinifera arcuata CSZ03
  Echinoconchella elegans CSZ03
  Latiproductus latissima CSZ03
  *Bruntonites mellingeni CSZ03
  Loriolithyris russillensis BO02
  Malurostrophia TC71
  Parachonetes macrostriatus TC71
  Altaethyrella Severgina 1978 RZJ04
  Ovalospira Fu 1982 RZJ04
  Pectenospira Popov, Nikitin & Sokiran 1999 RZJ04
  Jonesea grayi MM02
  Tecnocyrtina missouriensis P98
  Muhuathyris circularis Sun, Ma et al. 2004 SB04
  Aucklandrhynchia E01
    |--A. lavis E01
    `--A. opuatiaensis E01
  Pleuropugnoides panderi ZP86
  Vitiliproductus groberi ZP86
  Rhombospirifer zhesiensis Duan & Li in Ding, Xia et al. 1984 DX84
  Gypospirifer volatilis Duan & Li in Ding, Xia et al. 1984 DX84
  Kwangsirhynchus luwuensis XY86
  Hoskingia nobilis F71
  Sulciplica F71
    |--S. tasmaniensis (Morris in Strzelecki 1845) [=Spirifer tasmaniensis] F71
    `--S. transversa [incl. Spirifer vespertilio Morris in Strzelecki 1845] F71
  Harttites Howell & Knight 1936 [=Harttia Walcott 1884 non Steindachner 1877] F62
    `--*H. matthewi (Walcott 1884) [=*Harttia matthewi] F62
  Cariorhynchus tumida Hou79
  Spinacytia ostiolatus Hou79
  Eurekaspirifer pinyonensis N79
  Cortezorthis N79
    |--C. bathurstensis N79
    `--C. cortezensis N79
  Carinapyga loweryi N79
  Toquimaella kayi N79
  Septoliphoria astieriana Hol79
  Linguella viridens L95
  Camarophora globulina Phillips 1834 W77
  Amblotrema Rafinesque 1831 [=Amblytrema Agassiz 1847] M65
  Arctitreta Whitfield 1908 M65
    `--*A. pearyi Whitfield 1908 M65
  Biarea Torbakova 1959 M65
  Brachiopus Rafinesque 1831 M65
  Branconia Gagel 1890 M65
    `--*B. borussica Gagel 1890 M65
  Bufocephalus Linné 1779 M65
  Bursula Herrmannsen 1846 M65
  Comelicania Frech 1901 M65
    `--*C. megalotis (Stache 1878) [=Athyris megalotis] M65
  Cornwallia Wilson 1932 M65
    `--*C. minuta Wilson 1932 M65
  Diclipsites Rafinesque 1831 M65
  Diclisma Rafinesque 1820 M65
  Didymospira Salomon 1895 M65
  Diphyites Herrmannsen 1846 [=Diphytes Schröter 1779 (non-binomial)] M65
  Dirinus M’Coy 1844 M65
    `--*D. bucklandi M’Coy 1844 M65
  Euorthisina Havlíček 1950 M65
  Gamdaella Miloradovich 1947 M65
  Gasconsia Northrop 1939 M65
    `--*G. schucherti Northrop 1939 M65
  Gaspesia Clarke 1907 M65
    `--*G. aurelia (Billings 1874) [=Orthis aurelia] M65
  Goniclis Rafinesque 1818 M65
  Hemisterias Rafinesque 1832 M65
  Ivanovia Ivanova 1949 non Khvorova 1946 (ICBN) M65
  Lampas Humphrey 1797 [=Lampus (l. c.)] M65
  Liocoelia Schuchert & Cooper 1931 M65
    `--*L. proxima (Barrande 1879) [=Pentamerus proximus] M65
  Martinigisis Lebedev 1926 M65
  Megarites Rafinesque 1820 M65
  Mesotreta Kutorga 1848 M65
    `--*M. tentorium (Kutorga 1848) [=Siphonotreta tentorium] M65
  Neogypidula Likharev 1934 M65
  Peridiolithus Hupsch 1768 M65
  Platilites Rafinesque 1820 M65
  Pleuranisis Rafinesque 1820 M65
  Plicoprothyris Dahmer 1940 M65
  Pomatospirella Bittner 1892 M65
    `--*P. thecidium (Bittner 1892) [=Spirigera (*Pomatospirella) thecidium] M65
  Reflexa Rotai 1931 M65
    `--*R. reflexa Rotai 1931 M65
  ‘Rhynchoferella’ Spriestersbach 1942 non Strand 1915 M65
  Rictia de Gregorio 1930 M65
    `--*R. simplex de Gregorio 1930 M65
  Spondylobolus M’Coy 1851 [=Spondilobolus (l. c.), Spondylobus (l. c.)] M65
    `--*S. craniolaris M’Coy 1851 M65
  Swantonia Walcott 1905 M65
    `--*S. antiquata (Billings 1861) [=Camerella antiquata] M65
  Syntrophoides Schuchert & Cooper 1931 M65
    `--*S. harlanensis (Walcott 1905) [=Billingsella harlanensis] M65
  Telistrophis Rafinesque 1832 M65
  Thecospirella Bittner 1900 M65
    `--*T. loczyi Bittner 1900 M65
  Venezuelia Weisbord 1926 M65
  Virbium deGregorio 1930 M65
  Ptychopeltis Perner 1903 [Ptychopeltidae] R65
    `--*P. incola Perner 1903 R65
  Zhanatella utahensis LM05
  Quadrisonia lavadamensis LM05
  Eurytreta sublata LM05

Brachiopoda [Brachionacephala, Brachionopoda, Ecardines, Gastrocaulia, Inarticulata, Lyopomata, Palliobraches, Palliobrachiata, Pleuropygia, Spirobrachiophora, Spirobranchia, Tretenterata]

*Type species of generic name indicated

REFERENCES

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707–737.

[CSZ03] Chen, Z.-Q., G. R. Shi & L.-P. Zhan. 2003. Early Carboniferous athyridid brachiopods from the Qaidam Basin, northwest China. Journal of Paleontology 77: 844–862.

[DX84] Ding Y., Xia G., Duan C., Li W., Liu X. & Liang Z. 1984. Study on the early Permian stratigraphy and fauna in Zhesi district, Nei Mongol Zizhiqu (Inner Mongolia). Bulletin Tianjin Institure Geol. Min. Res. 10.

[E01] Eagle, M. K. 2001. A new species of Cottreauaster (Asteroidea: Echinodermata) from the Middle Jurassic of New Zealand. Records of the Auckland Museum 37: 93–100.

[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.

[F62] Fisher, D. W. 1962. Small conoidal shells of uncertain affinities. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W98–W143. Geological Society of America, and University of Kansas Press.

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

[Hol79] Hölder, H. 1979. Jurassic. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A390–A417. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[HP95] Holmer, L. E., L. E. Popov, M. G. Bassett & J. Laurie. 1995. Phylogenetic analysis and ordinal classification of the Brachiopoda. Palaeontology 38: 713–741.

[Hou79] House, M. R. 1979. Devonian in the Eastern Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A183–A217. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[L95] Landing, E. 1995. Upper Placentian–Branchian series of mainland Nova Scotia (middle–upper Lower Cambrian): faunas, paleoenvironments, and stratigraphic revision. Journal of Paleontology 69 (3): 475–495.

[LM05] Lehnert, O., J. F. Miller, S. A. Leslie, J. E. Repetski & R. L. Ethington. 2005. Cambro-Ordovician sea-level fluctuations and sequence boundaries: the missing record and the evolution of new taxa. Special Papers in Palaeontology 73: 117–134.

[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.

[N79] Norris, A. W. 1979. Devonian in the Western Hemisphere. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A218–A253. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

[RZJ04] Rong, J.-Y., R.-B. Zhan & J. Jin. 2004. The Late Odovician and Early Silurian pentameride brachiopod Holorhynchus Kiaer, 1902 from north China. Journal of Paleontology 78 (2): 287–299.

[R65] Rowell, A. J. 1965. Inarticulata. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H260–H296. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[SB04] Sun, Y., A. Baliński, X. Ma & Y. Zhang. 2004. New bizarre micro-spiriferid brachiopod from the Early Carboniferous of China. Acta Palaeontologica Polonica 49 (2): 267–274.

[TC71] Thomson, K. S., & K. S. W. Campbell. 1971. The structure and relationships of the primitive Devonian lungfish—Dipnorhynchus sussmilchi (Etheridge). Yale University Peabody Museum of Natural History Bulletin 38: 1–109.

[WM03] Webster, G. D., C. G. Maples, R. Mawson & M. Dastanpour. 2003. A cladid-dominated Early Mississippian crinoid and conodont fauna from Kerman Province, Iran and revision of the glossocrinids and rhenocrinids. Journal of Paleontology 77 (Suppl. 3): 1–35.

[W77] White, C. A. 1877. Report upon the invertebrate fossils collected in portions of Nevada, Utah, Colorado, New Mexico, and Arizona, by parties of the expeditions of 1871, 1872, 1873, and 1874. U.S. Geographical Surveys West of the One Hundredth Meridian 4 (1): 1–219, pls 1–21.

[WH02] Williams, A., & L. E. Holmer. 2002. Shell structure and inferred growth, functions and affinities of the sclerites of the problematic Micrina. Palaeontology 45 (5): 845–873.

[XY86] Xu H.-K. & Yao Z.-G. 1986. The Lower Devonian and its brachiopods from Lingshan, Guangxi. Acta Palaeontologica Sinica 25 (2): 169–179.

[ZP86] Zhang, R., & J. Pojeta Jr. 1986. New bivalves from the Datang Stage, Lower Carboniferous, Guangdong Province, China. Journal of Paleontology 60 (3): 669–679.

Rhynchonelliformea

Bicia gemma, from Walcott (1886).

Belongs within: Brachiopoda.
Contains: Rhynchonellata, Orthotetida, Billingsellida, Strophomenida, Productida.

The Rhynchonelliformea is the clade containing the articulate brachiopods (Articulata) as well as their stem relatives. The clade is characterised by the possession of an organocalcitic shell with fibrous microstructure (Williams & Holmer 2002). Articulate brachiopods have the valves connected by an arrangement of hinge teeth and dental sockets, though this has been lost in a very few derived subgroups.

<==Rhynchonelliformea
    |--Obolellida HP95 [Obolellata P98]
    |    |  i. s.: Nochoriella EL11
    |    `--Obolellidae [Obolellacea] R65a
    |         |--Obolella Billings 1861 R65a
    |         |    `--*O. chromatica Billings 1861 R65a
    |         |--Alisina Rowell 1962 R65a
    |         |    `--*A. atlantica (Walcott 1890) [=Obolella atlantica] R65a
    |         |--Bicia Walcott 1901 R65a
    |         |    `--*B. gemma (Billings 1871) [=Obolella gemma] R65a
    |         |--Magnicanalis Rowell 1962 R65a
    |         |    `--*M. mobergi (Walcott 1901) [=Obolella mobergi] R65a
    |         `--Trematobolus Matthew 1893 (see below for synonymy) R65a
    |              |--*T. insignis Matthew 1893 R65a
    |              `--T. kempanum (Matthew 1897) [=*Protosiphon kempanum] R65a
    `--Articulata [Arthropomata, Clistenterata, Pygocaulia, Testicardines] P98
         |  i. s.: Eichwaldiacea [Dictyonellidina] R65c
         |           |--Eichwaldiidae R65c
         |           |    |--Eichwaldia Billings 1858 R65c
         |           |    |    `--*E. subtrigonalis Billings 1858 R65c
         |           |    `--Dictyonella Hall 1868 [=Dyctionella (l. c.)] R65c
         |           |         |--*D. coralifera (Hall 1852) [=Atrypa coralifera] R65c
         |           |         `--D. capewelli R65c
         |           `--Isogrammidae R65c
         |                |--Megapleuronia Cooper 1952 R65c
         |                |    |--*M. fabianii (Greco 1947) [=Productus fabianii] R65c
         |                |    `--M. grecoi R65c
         |                `--Isogramma Meek & Worthen 1870 (see below for synonymy) R65c
         |                     |--*I. millipunctatum (Meek & Worthen 1870) [=Chonetes millipunctatus] R65c
         |                     |--‘*Aulacorhynchus’ pachti Dittmar 1872 [=*Aulacorhyna pachti] R65c
         |                     |--I. renfrarum R65c
         |                     `--I. texanum R65c
         |--Rhynchonellata P98
         `--Strophomenata P98
              |  i. s.: Cyphomenoidea julia D04
              |         Biparetis paucirugosus D04
              |         Nasutimena fluctuosa (Billings 1860) D04
              |         Katastrophomena D04
              |--Orthotetida D04
              |--Billingsellida D04
              `--+--Strophomenida D04
                 `--Productida D04

Rhynchonelliformea incertae sedis:
  Nikolaevirhynchus Baranov 1988 M03
  Pseudocamarotoechia gljadensis Kul’kov 1974 M03
  Kotujella Andreeva 1962 EL11, WW65 [Chileata WH02]
    `--*K. calva Andreeva 1962 WW65
  Kutorginata [Kutorginacea, Kutorginida, Palaeotremata, Rustellacea] WH02
    |  i. s.: Khasagtina EL11
    |--Yorkia Walcott 1897 [incl. Quebecia Walcott 1905; Yorkiidae] R65b
    |    |--*Y. wanneri Walcott 1897 R65b
    |    `--‘Obolella’ circe Billings 1872 [=*Quebecia circe] R65b
    |--Kutorginidae [Rustellidae, Schuchertinidae] R65b
    |    |--Schuchertina Walcott 1905 R65b
    |    |    `--*S. cambria Walcott 1905 R65b
    |    `--Kutorgina Billings 1861 [=Koturgina; incl. Rustella Walcott 1905] R65b
    |         |--*K. cingulata (Billings 1861) [=Obolella cingulata] R65b
    |         |--‘*Rustella’ edsoni Walcott 1905 R65b
    |         `--K. reticulata R65b
    `--Nisusiidae [Nisusiinae] WW65
         |--Eoconcha Cooper 1951 WW65
         |    `--*E. austini Cooper 1951 WW65
         |--Matutella Cooper 1951 WW65
         |    `--*M. clarki Cooper 1951 WW65
         `--Nisusia Walcott 1905 WH02, WW65
              |--*N. festinata (Billings 1861) [=Orthisina festinata] WW65
              `--N. deissi WW65

Isogramma Meek & Worthen 1870 [incl. Aulacorhyna Strand 1928, Aulacorhynchus Dittmar 1872 non Gould 1834] R65c

Trematobolus Matthew 1893 [=Trematobolos (l. c.); incl. Protosiphon Matthew 1897, Protosyphon (l. c.)] R65a

*Type species of generic name indicated

REFERENCES

[D04] Dewing, K. 2004. Shell structure and its bearing on the phylogeny of Late Ordovician–Early Silurian strophomenoid brachiopods from Anticosti Island, Québec. Journal of Paleontology 78: 275–286.

[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.

[HP95] Holmer, L. E., L. E. Popov, M. G. Bassett & J. Laurie. 1995. Phylogenetic analysis and ordinal classification of the Brachiopoda. Palaeontology 38: 713–741.

[M03] Modzalevskaya, T. L. 2003. Silurian and Devonian brachiopods from Severnaya Zemlya (Russian Arctic). Geodiversitas 25 (1): 73–107.

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

[R65a] Rowell, A. J. 1965a. Inarticulata. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H260–H296. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[R65b] Rowell, A. J. 1965b. Class uncertain. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H296–H297. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[R65c] Rowell, A. J. 1965c. Order uncertain—Dictyonellidina. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H359–H361. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[WH02] Williams, A., & L. E. Holmer. 2002. Shell structure and inferred growth, functions and affinities of the sclerites of the problematic Micrina. Palaeontology 45 (5): 845–873.

[WW65] Williams, A., & A. D. Wright. 1965. Orthida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H299–H359. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

Pentameridae

Borealis cristiformis, from Modzalevskaya (2003).

Belongs within: Pentamerida.

The Pentameridae were a family of brachiopods that lived from the Upper Ordovician to the Lower Devonian (Amsden & Biernat 1965). In some genera, such as Conchidium and Lissocoelina, the pedicle valve is distinctly rostrate, arching over the brachial valve.

Characters (from Amsden & Biernat 1965, as Pentamerinae): Moderate to large size, with smooth, costate or costellate shells generally lacking well-developed interareas; fold and sulcus absent or weakly developed. Brachial processes long, rodlike; outer plates commonly discrete, but in a few genera uniting to make cruralium.

<==Pentameridae [Pentamerinae]
    |--Aliconchidium St. Joseph 1942 AB65
    |    `--*A. yassi St. Joseph 1942 AB65
    |--Brooksina Kirk 1922 AB65
    |    `--*B. alaskensis Kirk 1922 AB65
    |--Callipentamerus Boucot 1964 M65
    |    `--*C. corrugatus (Weller & Davidson 1896) [=Pentamerus corrugatus] M65
    |--Capelliniella Strand 1928 [=Capellinia Hall & Clarke 1894 non Trichese 1874] AB65
    |    `--*C. mira (Hall & Clarke 1894) [=*Capellinia mira] AB65
    |--Cymbidium Kirk 1926 AB65
    |    `--*C. actum Kirk 1926 AB65
    |--Jolvia Sapelnikov 1960 AB65
    |    `--*J. multiplexa Sapelnikov 1960 AB65
    |--Lissocoelina Schuchert & Cooper 1931 AB65
    |    `--*L. pergibbosa (Hall & Whitfield 1875) [=Pentamerus pergibbosus] AB65
    |--Pentamerifera Khodalevich 1939 AB65
    |    `--*P. taltiensis (Chernyshev 1893) [=Pentamerus taltiensis] AB65
    |--Pentameroides Schuchert & Cooper 1931 AB65
    |    `--*P. subrectus (Hall & Clarke 1894) [=Pentamerus oblongus subrectus] AB65
    |--Rhipidium Schuchert & Cooper 1931 AB65
    |    `--*R. knappi (Hall & Whitfield 1872) [=Pentamerus knappi] AB65
    |--Harpidium Kirk 1925 M03
    |    |--*H. insigne (Kirk 1925) [=H. insignis] AB65
    |    `--H. angustum Poulsen 1943 M03
    |--Pentamerus Sowerby 1813 (see below for synonymy) AB65
    |    |--*P. oblongus Sowerby in Murchison 1839 AB65 (see below for synonymy)
    |    |--P. costatus Etheridge 1892 F71
    |    `--P. hospes F71
    |--Borealis Boucot, Johnson & Staton 1971 M03
    |    |--*B. borealis (Eichwald 1842) M03
    |    |    |--B. b. borealis M03
    |    |    `--B. b. osloensis Mørk 1981 M03
    |    |--B. cristiformis (Modzalevskaya 1985) [=Pentamerus cristiformis] M03
    |    `--B. schmidti (Lebedev 1892) M03
    `--Conchidium Oehlert 1887 AB65
         |--*C. biloculare (Hisinger 1799) [=Anomia bilocularis] AB65
         |--C. alaskense AB65
         |--C. etheridgei (Dun 1907) [=Pentamerus (Conchidium) etheridgei] F71
         |--C. knightii [=Pentamerus knightii; incl. C. knightii var. stricta] F71
         |--C. munsteri St. Joseph 1937 AB65
         |--C. occidentale (Hall 1852) [=Pentamerus occidentalis] AB65
         `--C. vogulicum AB65

Nomen rejiciendum: Pentamerus laevis Sowerby 1813 AB65

Pentamerus Sowerby 1813 [incl. Grypidia Dalman 1828 (nom. rej.), Miopentamerus Alexander 1936 (nom. rej.), Trimurus Caldwell 1934 (nom. rej.)] AB65

*Pentamerus oblongus Sowerby in Murchison 1839 AB65 [incl. P. samojedicus Keyserling 1846 M03, Borealis samojedicus M03]

*Type species of generic name indicated

REFERENCES

[AB65] Amsden, T. W., & G. Biernat. 1965. Pentamerida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda, vol. 2, pp. H523-H552. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

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

[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).

Rhynchonellata

Ventral (left) and dorsal (right) views of Sphenorthis niagarensis, from Grubbs (1939).

Belongs within: Rhynchonelliformea.
Contains: Thecideida, Pentamerida, Rhynchonellida, Atrypida, Spiriferida, Terebratulida, Athyridida, Enteletacea, Orthacea.

The Rhynchonellata is the clade of brachiopods that includes all surviving articulate species. They are distinguished from other articulate brachiopods by the presence of a pedicle rudiment and projecting brachiophores (Williams et al. 1996). The pedicle opening is generally unrestricted in the Orthida and Pentamerida, but is restricted by deltidial plates in the more derived orders. These latter groups also developed a persistent non-fibrous primary layer in the shell. Members of the extinct orders Atrypida, Spiriferida and Athyridida are further characterised by the presence of spiral brachidia that would have supported the lophophore in life.

Characters (from Williams et al. 1996): Shells fibrous, endopunctate or impunctate, biconvex, strophic or astrophic, articulated by deltidiodont or cyrtomatodont teeth and sockets buttressed by brachiophores or supported by parallel socket or hinge plates that may converge to form septalium or cruralium; pedicle opening as delthyrium or rounded foramen; cardinal areas and notothyrium commonly vestigial or absent, wide in some later groups; dental plates less commonly converge to form spondylium; posteriomedial ventral adductor scars flanked or enclosed by ventral diductor and laterally placed adjustor scars; dorsal adductor scars petaloid or grouped and quadripartite; crura present in later groups and commonly extended as spiralia or loops; mantle canal systems variable as saccate, digitate, pinnate, or lemniscate impressions.

<==Rhynchonellata
    |  i. s.: Thecideida F05
    |         Wangyuia EL11
    |         Sphenorthis Grubbs 1939 M65
    |           `--*S. niagarensis Grubbs 1939 M65
    |--+--Pentamerida P98
    |  `--+--Rhynchonellida P98
    |     `--+--Atrypida P98
    |        `--+--Spiriferida P98
    |           `--+--Terebratulida P98
    |              `--Athyridida P98
    `--Orthida [Orthidina, Orthoidea] P98
         |  i. s.: Glyptoria HP95
         |         Israeleria HP95
         |         Leioria HP95
         |         Minororthis Ivanov 1950 M65
         |           `--*M. malivkini Ivanov 1950 M65
         |         Wynnia Walcott 1908 M65
         |           `--*W. warthi (Walcott 1908) [=Orthis warthi] M65
         |         Yeosinella Reed 1932 M65
         |           `--*Y. consignata Reed 1932 M65
         |--Enteletacea WW65
         `--Orthacea WW65

*Type species of generic name indicated

REFERENCES

[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.

[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.

Grubbs, D. M. 1939. Fauna of the Niagaran nodules of the Chicago area. Journal of Paleontology 13 (6): 543–560.

[HP95] Holmer, L. E., L. E. Popov, M. G. Bassett & J. Laurie. 1995. Phylogenetic analysis and ordinal classification of the Brachiopoda. Palaeontology 38: 713–741.

[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).

[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

Williams, A., S. J. Carlson, C. H. C. Brunton, L. E. Holmer & L. Popov. 1996. A supra-ordinal classification of the Brachiopoda. Philosophical Transactions: Biological Sciences 351 (1344): 1171-1193.

[WW65] Williams, A., & A. D. Wright. 1965. Orthida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H299–H359. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).