Strophomenida

Rafinesquina sp., from here.

Belongs within: Rhynchonelliformea.
Contains: Lyttoniidae, Stropheodontidae, Strophomenidae, Leptaenidae.

The Strophomenida are a lineage of strophomenate brachiopods known from the Ordovician to the Triassic. The Upper Carboniferous to Triassic Lyttoniacea are among the most unusual brachiopod groups, with an irregularly shaped shell in which the primary brachial valve is reduced to a small remnant. In its place is a leaf-like, lobate secondary shell that does not extend as broadly as the pedicle shell, meaning that in life at least part of the animal was permanently exposed. The Ordovician to Carboniferous Strophomenacea had more standard concavo-convex to resupinate shells, with the apical foramen sealed in some later lineages so that the adult shell was unattached to the substrate. Within the Strophomenacea, the Rafinesquinidae are known from the Middle and Upper Ordovician, and possess a usually concavo-convex shell with a small apical foramen and vestigial pseudodeltidium (Muir-Wood & Williams 1965).

Characters (from Dewing 2004): Shell structure laminar, pseudopunctae small, smooth-sided, randomly distributed, cored by small taleolae, without associated muscle-attachment pits. Cardinal process bilobed. Socket plates thin, upwardly concave sheets, forming long high-socket ridges extending beyond posterior platform.

<==Strophomenida
    |  i. s.: Australostrophia Caster 1939 M65
    |           `--*A. mesembria (Clarke 1913) [=Leptostrophia mesembria] M65
    |         Socraticum de Gregorio 1930 M65
    |           `--*S. firmum de Gregorio 1930 M65
    |--Lyttoniacea [Oldhaminidina, Oldhaminoidea] M-WW65
    |    |--Lyttoniidae M-WW65
    |    |--Bactrynium Emmrich 1855 [=Pterophloios Emmrich 1855; Bactryniidae] M-WW65
    |    |    `--*B. bicarinatum Emmrich 1855 [=*Pterophloios bicarinata] M-WW65
    |    |--Spinolyttonia Sarycheva 1964 [Spinolyttoniidae] M-WW65
    |    |    `--*S. arakeljani Sarycheva 1964 M-WW65
    |    `--Poikilosakidae M-WW65
    |         |--Poikilosakos Watson 1917 M-WW65
    |         |    `--*P. petaloides Watson 1917 M-WW65
    |         |--Cardinocrania Waagen 1885 M-WW65
    |         |    `--*C. indica Waagen 1885 M-WW65
    |         |--Pseudoleptodus Steli 1956 M-WW65
    |         |    `--*P. getawayensis Stehli 1956 M-WW65
    |         `--Adriana de Gregorio 1930 [incl. Stita de Gregorio 1930] M-WW65
    |              |--*A. osiensis de Gregorio 1930 M-WW65
    |              |--A. guadelupensis M-WW65
    |              `--‘Lyttonia (*Stita)’ paupera de Gregorio 1930 M-WW65
    `--Strophomenacea [Strophomenidina, Strophomenoidea] M-WW65
         |  i. s.: Leptostrophiidae [Leptostrophiinae, Leptostrophiini] D04
         |           |--Gamphalosia Stainbrook 1946 M-WW65
         |           |    `--*G. tenuissima Stainbrook 1946 M-WW65
         |           |--Protoleptostrophia Caster 1939 M-WW65
         |           |    `--*P. blainvillii (Billings 1874) [=Strophomena blainvillii] M-WW65
         |           |--Maoristrophia Allan 1947 M-WW65
         |           |    |--*M. neozelanica Allan 1947 M-WW65
         |           |    `--M. banksi H79
         |           |--Leptostrophia Hall & Clarke 1892 [incl. Rhytistrophia Caster 1939] M-WW65
         |           |    |--*L. magnifica (Hall 1857) [=Stropheodonta magnifica] M-WW65
         |           |    |--‘Stropheodonta’ beckii Hall 1859 [=*Rhytistrophia beckii] M-WW65
         |           |    `--L. quadrata (Mitchell 1923) [=Stropheodonta (Leptostrophia) quadrata] F71
         |           `--Nervostrophia Caster 1939 (see below for synonymy) M-WW65
         |                |--*N. nervosa (Hall 1843) [=Strophomena nervosa] M-WW65
         |                |--‘Leptostrophia’ camerata Fenton & Fenton 1924 [=*Sulcatostrophia camerata] M-WW65
         |                |--‘*Pseudodouvillina’ euglyphea Stainbrook 1945 M-WW65
         |                `--N. rockfordensis M-WW65
         |         Lamanskya Moberg & Segerberg 1906 M65
         |           `--*L. splendens Moberg & Segerberg 1906 M65
         |         Foliomena Havlíček 1952 [Foliomenidae] M-WW65
         |           `--*F. folium (Barrande 1879) [=Strophomena folium] M-WW65
         |         Christiania Williams 1953 [Christianiidae] M-WW65
         |           `--*C. subquadrata (Hall 1883) [=Leptaena subquadrata] M-WW65
         |         Stropheodontidae M-WW65
         |--+--Strophomenidae D04
         |  `--Glyptomenidae [Glyptomeninae] D04
         |       |--Pionomena Cooper 1956 M-WW65
         |       |    `--*P. neumani Cooper 1956 M-WW65
         |       |--Platymena Cooper 1956 M-WW65
         |       |    `--*P. plana Cooper 1956 M-WW65
         |       |--Glyptomena Cooper 1956 [incl. Oslomena Spjeldnaes 1957] M-WW65
         |       |    |--*G. sculpturata Cooper 1956 M-WW65
         |       |    `--‘*Oslomena’ osloensis Spjeldnaes 1957 M-WW65
         |       `--Mjoesina Spjeldnaes 1957 D04, M-WW65
         |            |--*M. mjoesensis (Holtedahl 1916) [=Rafinesquina mjoesensis] M-WW65
         |            |--M. ceres (Billings 1860) D04
         |            `--M. rugata M-WW65
         `--+--+--Archeochonetes primigenius (Twenhofel 1914) D04
            |  `--+--+--Aphanomena planoconvexa Dewing 1999 D04
            |     |  `--Leptaenidae D04
            |     `--Megamyonia Wang 1949 D04, M-WW65
            |          |--*M. knighti Wang 1949 M-WW65
            |          |--M. arethusa D04
            |          `--M. nitens (Billings 1860) D04
            `--Rafinesquinidae [Rafinesquininae] D04
                 |--Kjerulfina Bancroft 1929 M-WW65
                 |    `--*K. trigonalis Bancroft 1929 M-WW65
                 |--Kjaerina Bancroft 1929 M-WW65
                 |    |--*K. typa Bancroft 1929 M-WW65
                 |    `--K. hartae Jin, Caldwell & Norford 1995 D04
                 |--Hedstroemina Bancroft 1929 M-WW65
                 |    |--*H. fragilis Bancroft 1929 [=Kjaerina (*Hedstroemina) fragilis] M-WW65
                 |    `--H. reticulata D04
                 `--Rafinesquina Hall & Clarke 1892 D04, M03 [incl. Playfairia Reed 1917 M-WW65]
                      |--*R. alternata (Conrad 1838) [=Leptaena alternata] M-WW65
                      |--R. crispa Andreeva 1961 M03
                      |--‘Strophomena’ deltoidea Conrad 1839 [=*Playfairia deltoidea] M-WW65
                      |--R. nasuta WR65
                      |--R. planulata M-WW65
                      |--R. stropheodontoides (Savage 1913) (see below for synonymy) M03
                      `--R. trentonensis M-WW65

Nervostrophia Caster 1939 [incl. Pseudodouvillina Stainbrook 1945, Sulcatostrophia Caster 1939] M-WW65

Rafinesquina stropheodontoides (Savage 1913) [=Brachyprion stropheodontoides; incl. R. inaequicostata Lopushinskaya 1967] M03

*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.

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

[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.

[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).

[M-WW65] Muir-Wood, H., & A. Williams. 1965. Strophomenida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H361–H521. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).

[WR65] Williams, A., & A. J. Rowell. 1965. Morphology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H57–H138. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).