Specimens of Schwagerina huecoensis, from here.

Belongs within: Loeblichiidae.

The Schwagerininae are a group of fusulinid Foraminifera known from the Upper Carboniferous and Permian with outer test walls comprised of two layers. The test is tightly coiled in most genera, but becomes irregularly and loosely uncoiled in gerontic stages of the Japanese Permian genus Nipponitella (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Shell large, fusiform to irregularly cylindrical, planispiral, involute in most, irregularly uncoiled in some; spirotheca thick, composed of tectum and alveolar keriotheca; septa fluted in end zones of primitive genera, fluted completely across shell and to tops of chambers of more advanced genera; tunnel singular in most forms or multiple in one genus, axial fillings absent to massive; chomata massive to slight.

<==Schwagerininae [Polydiexodininae, Pseudofusulininae]
    |--Acervoschwagerina Hanzawa 1949 LT64
    |    `--*A. endoi (Hanzawa 1949) [=Paraschwagerina (*Acervoschwagerina) endoi] LT64
    |--Biwaella Morikawa & Isomi 1960 LT64
    |    `--*B. omiensis Morikawa & Isomi 1960 LT64
    |--Kwantoella Sakagami & Omata 1957 LT64
    |    `--*K. fujimotoi Sajagami & Omata 1957 LT64
    |--Oketaella Thompson 1951 LT64
    |    `--*O. fryei Thompson 1951 LT64
    |--Orientoschwagerina Miklukho-Maklay 1955 LT64
    |    `--*O. abichi Miklukho-Maklay 1955 LT64
    |--Robustoschwagerina Miklukho-Maklay 1959 LT64
    |    `--*R. tumida (Likharev 1939) [=Pseudoschwagerina tumida] LT64
    |--Rugosofusulina Rauser-Chernousova 1937 [=Rugofusulina (l. c.)] LT64
    |    `--*R. prisca (Ehrenberg 1842) LT64 [=Alveolina prisca LT64, Schwagerina (*Rugofusulina) prisca C40]
    |--Zellia Kahler & Kahler 1937 LT64
    |    `--*Z. heritschi (Kahler & Kahler 1937) [=Pseudoschwagerina (*Zellia) heritschi] LT64
    |--Nipponitella Hanzawa 1938 LT64
    |    |--*N. explicata Hanzawa 1938 LT64
    |    `--N. auricula LT64
    |--Paraschwagerina Dunbar & Skinner 1936
    |    |--*P. gigantea (White 1932) [=Schwagerina gigantea] LT64
    |    `--P. yabei C40
    |--Nagatoella Thompson 1936 LT64
    |    |--*N. orientis (Ozawa 1925) [=Fusulina (Schellwienia) ellipsoidalis var. orientis] LT64
    |    `--N. kobayashii LT64
    |--Rugososchwagerina Miklukho-Maklay 1959 LT64
    |    |--*R. yabei (Staff 1909) [=Schwagerina yabei] LT64
    |    `--R. fosteri LT64
    |--Polydiexodina Dunbar & Skinner 1931
    |    |--*P. capitanensis Dunbar & Skinner 1931
    |    |--P. afghanensis LT64
    |    `--P. shumardi C40
    |--Dunbarinella Thompson 1942 LT64
    |    |--*D. ervinensis Thompson 1942 LT64
    |    |--D. eoextenta LT64
    |    `--D. tumida LT64
    |--Kansanella Thompson 1957 LT64
    |    |--*K. (Kansanella) joensis Thompson 1957 LT64
    |    `--K. (Iowanella Thompson 1957) LT64
    |         `--K. (*I.) winterensis (Thompson, Verville & Lokke 1956) [=Triticites winterensis] LT64
    |--Pseudoschwagerina Dunbar & Skinner 1936 (see below for synonymy) LT64
    |    |--*P. uddeni (Beede & Kniker 1924) [=Schwagerina uddeni] LT64
    |    |--P. fusulinoides (Schellwien 1898) (see below for synonymy) LT64
    |    |--P. morikawai RR79
    |    |--P. muongthensis (Deprat 1915) [=Fusulina muongthensis, *Parazellia muongthensis] LT64
    |    |--P. robusta RR79
    |    `--P. roeseleri LT64
    |--Monodiexodina Sosnina in Kiparisova et al. 1956 LT64
    |    |--*M. sutschanica (Dutkevitch in Likharev et al. 1939) (see below for synonymy) LT64
    |    |    |--M. s. sutschanica DX84
    |    |    `--M. s. baotegensis Xia in Ding, Xia et al. 1984 DX84
    |    |--M. caracorumensis DX84
    |    |--M. neimonggolensis DX84
    |    |--M. shiptoni DX84
    |    |--M. wanneri DX84
    |    `--M. yongwangcunensis Xia in Ding, Xia et al. 1984 DX84
    |--Triticites Girty 1904 (see below for synonymy) LT64
    |    |--*T. secalicus (Say in James 1823) [=Miliolites secalicus] LT64
    |    |--T. beedei LT64
    |    |--T. ferganensis Miklukho-Maklay 1950 [=*Ferganites ferganensis] LT64
    |    |--T. globulus (Putrya 1948) [=*Protriticites globulus] LT64
    |    |--T. irregularis LT64
    |    |--T. milleri LT64
    |    |--T. montiparus (Ehrenberg 1854) [=Alveolina montipara, T. (*Montiparus) montiparus] LT64
    |    |--T. moorei LT64
    |    |--‘Fusulina’ obsoleta Schellwien 1908 [=*Obsoletes obsoletus] LT64
    |    |--T. ohioensis LT64
    |    `--T. ventricosus (Meek & Hayden 1859) [=Fusulina cylindrica var. ventricosa, *Girtyina ventricosa] LT64
    |--Pseudofusulina Dunbar & Skinner 1931 [incl. Grabauina Lee 1924] LT64
    |    |--*P. huecoensis Dunbar & Skinner 1931 LT64
    |    |--P. ambigua RR79
    |    |--P. callosa RR79
    |    |--P. disca (Lee 1924) [=*Grabauina disca] LT64
    |    |--P. fecunda RR79
    |    |--P. firma RR79
    |    |--P. globosa RR79
    |    |--P. intermedia RR79
    |    |--P. japonica RR79
    |    |--P. moelleri RR79
    |    |--P. (Laxifusulina) neimonggolensis DX84
    |    |--P. nelsoni LT64
    |    |--P. plicatissima RR79
    |    |--P. schellwieni RR79
    |    |--P. urdalensis RR79
    |    |--P. verneuili RR79
    |    `--P. vulgaris RR79
    |--Leeina Galloway 1933 LT64
    |    |--*L. fusiformis (Schellwien in Dyhrenfurth 1909) (see below for synonymy) LT64
    |    `--L. kraffti VH01 [=Pseudofusulina kraffti DX84]
    |--Parafusulina Dunbar & Skinner 1931 [incl. Eoparafusulina Coogan 1960, Skinnerella Coogan 1960] LT64
    |    |--*P. wordensis Dunbar & Skinner 1931
    |    |--P. akasakensis (Deprat 1914) ZL04
    |    |--P. alaskensis LT64
    |    |--P. allisonensis RR79
    |    |--P. dongvanensis LT64
    |    |--P. durhami RR79
    |    |--P. gigantea LT64
    |    |--P. gracilis (Meek 1862) [=Fusulina gracilis, P. (*Eoparafusulina) gracilis] LT64
    |    |--P. gruperaensis ZL04
    |    |    |--P. g. gruperaensis ZL04
    |    |    `--P. g. brevica Sheng 1963 ZL04
    |    |--P. japonica VH01
    |    |--P. kaerimizensis RR79
    |    |--P. kattaensis C40
    |    |--‘Eoparafusulina’ linearis RR79
    |    |--P. lineata RR79
    |    |--P. lutugini RR79
    |    |--P. maleyi RR79
    |    |--P. matsubaishi RR79
    |    |--P. nosonensis LT64
    |    |--P. padangensis LT64
    |    |--P. parumvoluta LT64
    |    |--P. quasigruperaensis Sheng 1963 ZL04
    |    |--P. rothi DX84
    |    |--P. schucherti Dunbar & Skinner 1937 [=P. (*Skinnerella) schucherti] LT64
    |    |--P. spissisepta RR79
    |    |--P. subtilis Leven 1992 ZL04
    |    |--P. sullivanensis RR79
    |    |--P. wanneri LT64
    |    `--P. yunnanica DX84
    `--Schwagerina von Möller 1877 (see below for synonymy) LT64
         |--*S. princeps (Ehrenberg 1842) [=Borelis princeps] LT64
         |--S. arctica Xia in Ding, Xia et al. 1984 DX84
         |--S. bulegensis Xia in Ding, Xia et al. 1984 DX84
         |--S. campensis LT64
         |--S. conicocylindrica DX84
         |--S. crassitectoria RR79
         |--S. cushmani CB86
         |--S. douvellei DX84
         |--S. dugoutensis RR79
         |--S. fax LT64
         |--S. furoni LT64
         |--S. granumavenae DX84
         |--S. gruperaensis LT64
         |--S. guembeli RR79
         |--S. hawkinsi RR79
         |--S. huecoensis C40
         |--S. hupehensis DX84
         |--S. ishanensis (Hsu 1942) [=*Chusenella ishanensis] LT64
         |--S. jigulensis (Rauzer-Chernousova 1938) [=Triticites jigulensis, T. (*Jigulites) jigulensis] LT64
         |--‘Fusulina’ krotowi RR79
         |--S. kwangchiensis DX84
         |--S. longa DX84
         |--S. longipertica DX84
         |--S. mandulaensis Xia in Ding, Xia et al. 1984 DX84
         |--S. neimonggolensis Xia in Ding, Xia et al. 1984 DX84
         |--S. normalis Xia in Ding, Xia et al. 1984 DX84
         |--S. pactiruga DX84
         |--S. pseudoregularis DX84
         |--S. puerilis Xia in Ding, Xia et al. 1984 DX84
         |--S. pulchella Xia in Ding, Xia et al. 1984 DX84
         |--S. quasiregularis DX84
         |--S. rutschi LT64
         |--S. ruzhencevi (Rozovskaya 1949) [=*Daixina ruzhencevi] LT64
         |--S. setum RR79
         |--S. stuckenbergi (Rauzer-Chernousova 1938) [=*Triticites (*Rauserites) stuckenbergi] LT64
         |--S. thuanae (Viên 1959) [=*Codonoschwagerina thuanae] LT64
         |--S. trivialis Xia in Ding, Xia et al. 1984 DX84
         |--S. tschernyschewi CB86
         |--S. ulanqabensis Xia in Ding, Xia et al. 1984 DX84
         `--S. zhesiensis Xia in Ding, Xia et al. 1984 DX84

*Leeina fusiformis (Schellwien in Dyhrenfurth 1909) [=Fusulina vulgaris fusiformis, Pseudofusulina vulgaris fusiformis] LT64

*Monodiexodina sutschanica (Dutkevitch in Likharev et al. 1939) [=Schwagerina wanneri sutschanica, Parafusulina sutschanica] LT64

Pseudoschwagerina Dunbar & Skinner 1936 [incl. Occidentoschwagerina Miklukho-Maklay 1959, Parazellia Rauzer-Chernousova 1961] LT64

Pseudoschwagerina fusulinoides (Schellwien 1898) [=Schwagerina fusulinoides, *Occidentoschwagerina fusulinoides] LT64

Schwagerina von Möller 1877 [incl. Chusenella Hsu 1942, Codonoschwagerina Viên 1959, Daixina Rozovskaya 1949, Jigulites Rozovskaya 1948, Rauserites Rozovskaya 1950] LT64

Triticites Girty 1904 [incl. Ferganites Miklukho-Maklay 1959, Girtyina Staff 1909, Montiparus Rozovskaya 1948, Obsoletes Kireeva 1950, Protriticites Putrya 1948] LT64

*Type species of generic name indicated


[CB86] Chen D.-Q. & Bao H. 1986. Lower Permian ostracodes from the Chihsia Formation of Jurong and Longtan, Jiangsu Province. Acta Micropalaeontologica Sinica 3 (2): 107–132.

[C40] Cushman, J. A. 1940. Foraminifera: Their classification and economic use 3rd ed. Harvard University Press: Cambridge (Massachusetts).

[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.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[RR79] Ross, C. A., & J. R. P. Ross. 1979. Permian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A291–A350. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[VH01] Vachard, D., M. Hauser, R. Martini, L. Zaninetti, A. Matter & T. Peters. 2001. New algae and problematica of algal affinity from the Permian of the Aseelah Unit of the Batain Plain (East Oman). Geobios 34 (4): 375–404.

[ZL04] Zhou, Z., & M. Liengjarern. 2004. Lower Permian perrinitid ammonoid faunas from Thailand. Journal of Paleontology 78: 317–339.

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