Belongs within: Athyridida.
Contains: Athyris, Composita.
The Athyrididae were a family of brachiopods that lived from the Upper Silurian to the Triassic. In members of the subfamilies Athyridinae and Cleiothyridinae, a jugal saddle is present with bifurcations extending at least partway into the spiralia; this saddle is absent in the Protathyridinae. Protathyridinae are known from the Upper Silurian to the Middle Devonian and had the cardinal plate pierced apically. More advanced athyridids appeared in the Lower Devonian (Boucot et al. 1965).
The Athyrididae: spiralia and lamellae
Published 28 February 2012
The Athyrididae were a family of brachiopods that lived from the Silurian to the Permian, or until the end of the Triassic if the Diplospirellidae and Retzioidea are derived from the athyridids (Alvarez et al. 1998). Many athyridids possessed concentric lamellae on the outside of the shell; in the Upper Devonian to Permian Cleiothyridina, these lamellae were developed into a dense forest of flat spines. Specimens of the Devonian species Athyris vittata with preserved colour patterns indicate that the presence of radial stripes (Blodgett et al. 1988).
The Athyrididae are members of the order Athyridida, one of a number of brachiopod groups to possess a calcified spiral support (called, funnily enough, a spiralium) for the lophophore (see the link above for an explanation of the brachiopod lophophore). In the 1965 Treatise on Invertebrate Paleontology volume for brachiopods (Moore 1965), all the spiralia-possessing brachiopods were combined as the Spiriferida; however, other features of the shell are not consistent with a single origin for the spiralium, and they are now divided between the Spiriferida, Atrypida and Athyridida.
Many spiralium-bearing species appear to have lived on soft sediments, and it is possible that the spiralium was developed primarily as an adaptation for such habitats (Alvarez & Brunton 1990). For filter-feeders living in such habitats, the greatest challenge for feeding is not so much taking in food particles, but keeping the filter (in this case, the lophophore) from becoming clogged by indigestible particles such as sand. Reversing the direction of beat of the lophophore cilia moves such particles back towards the shell opening, where rapidly closing the valves will give the final impetus to ‘spit out’ the offending particles. In those species with lamellae, the slow-down of water currents as they hit the lamellae before the water enters between the valves may have also reduced the amount of particulate matter getting in. The lamellae may have also helped to support the shell opening above the sediment surface, though more significant in this regard would have been the pedicle, the fleshy stalk emerging from an opening in the back of the shell that anchored the athyridid in the sediment.
Systematics of Athyrididae
Characters (Boucot et al. 1965): Smooth or pauciplicate shells commonly with cardinal plate pierced apically, not depressed to form septalium; jugal saddle may occur and lamellar expansions may be developed at growth lines.
<==Athyrididae [Athyridae, Didymothyrididae]
|--DidymothyridinaeMM02
| |--DidymothyrisB75
| |--Corallothyris Modzalevskaya 1970MM02
| | `--*C. canaliculata (Wenjukow 1899) [=Meristella canaliculata]MM02
| `--Pseudoprotathyris Modzalevskaya 1979MM02
| `--*P. infantilis (Kozłowski 1929) [=Protathyris infantile]MM02
|--LochengiinaeCSZ03
| |--Galeatathyris Jin 1983CSZ03
| |--Titanothyris Jin & Hu in Wang et al. 1982CSZ03
| `--Lochengia Yoh 1929CSZ03 [=Cryptospirifer Grabau 1931BJ65; incl. Flexathyris Grunt 1980CSZ03]
| |--*L. lochengensis (Grabau 1931)CSZ03 [=*Cryptospirifer lochengensisBJ65]
| |--L. anormalisG31
| |--L. aplicataG31
| |--L. enteletiformisCSZ03
| |--L. prokofievi [=Flexathyris prokofievi]CSZ03
| |--L. qinghaiensis Chen, Shi & Zhan 2003CSZ03
| `--L. variabilis (Möller 1862) [=Athyris (Actinoconcha) variabilis, Flexathyris variabilis]CSZ03
|--CleiothyridininaeCSZ03
| |--Leiothyrcridina Grunt 1980CSZ03
| |--Bruntonathyris Chen, Shi & Zhan 2003CSZ03
| | |--*B. amunikeensis Chen, Shi & Zhan 2003CSZ03
| | |--B. heijianshanensis Chen, Shi & Zhan 2003CSZ03
| | `--B. patula (Brunton 1980) [=Actinoconchus expansus patulus]CSZ03
| `--Cleiothyridina Buckman 1906CSZ03 (see below for synonymy)
| |--*C. deroissii (Léveillé 1835)BJ65 (see below for synonymy)
| |--C. capillata (Waagen 1883)HB93
| |--C. mediaZP86
| |--C. pectiniferaBJ65
| `--C. seriataWBM97
|--ProtathyridinaeBJ65
| |--Buchanathyris Talent 1956BJ65
| | `--*B. westoni Talent 1956BJ65
| |--Glassina Hall & Clarke 1893BJ65
| | |--*G. laeviuscula (Sowerby 1839)BJ65, HB93 [=Terebratula laeviusculaBJ65]
| | `--G. usitata Modzalevskaya 1979HB93
| |--Protathyris Kozlowski 1929XY86
| | |--*P. praecursor Kozlowski 1929BJ65
| | |--P. didymaBJ65
| | |--P. ovalis Tong 1982XY86
| | `--P. sibiricaTY12
| `--Greenfieldia Grabau 1910BJ65
| |--*G. whitfieldi Grabau 1910BJ65
| |--G. orbiculata Modzalevskaya 2003M03
| `--G. uberisM03
| |--G. u. uberisM03
| `--G. u. tanta Modzalevskaya 1980M03
`--Athyridinae [Athyrinae]CSZ03
| i. s.: Anathyrella Khalfin 1961BJ65
| `--*A. ussovi (Khalfin 1933) [=Anathyris ussovi]BJ65
| Actinoconchus McCoy 1844BJ65, CSZ03
| |--*A. planosulcata (Phillips 1836)BJ65 (see below for synonymy)
| `--A. expansaCSZ03 [=Athyris expansaCSZ03; incl. At. expansa var. kwangsiensisG31]
| Lamellosathyris Jin & Fang 1983CSZ03
| |--*L. lamellosa (Léveillé 1835) (see below for synonymy)CSZ03
| |--L. bradyensis (Carter 1967) [=Athyris bradyensis]CSZ03
| |--L. forschi (Janischewsky 1954) [=Athyris forschi]CSZ03
| |--L. hannibalensis [=Athyris hannibalensis]CSZ03
| `--L. qaidamensis Chen,Shi & Zhan 2003CSZ03
|--Leptathyris Siehl 1962B75, BJ65
| `--*L. gryphis Siehl 1962BJ65
`--+--+--Triathyris Comte 1938B75, BJ65
| | `--*T. mucronata (de Verneuil 1850) [=Terebratula mucronata]BJ65
| `--Anathyris von Peetz 1901B75, BJ65 [incl. Plicathyris Khalfin 1946BJ65]
| |--*A. phalaena (Phillips 1841)BJ65 [=Spirifera phalaenaBJ65, Licharewia phalaenaF71, Spirifer phalaenaF71]
| |--A. ezquerraiBJ65
| `--‘Athyris’ sibirica Khalfin 1940 [=*Plicathyris sibirica]BJ65
`--+--AthyrisB75
|--Septathyris Boucot, Johnson & Staton 1964B75, BJ65
| |--*S. aliena (Drevermann 1904) [=Athyris aliena]BJ65
| `--S. davoustiBM05
`--Spirigerellinae [Spirigerellidae]HB93
|--CompositaCSZ03
|--Spirigerella Waagen 1883 [incl. Athyrella Renz 1913 (n. n.)]BJ65
| `--*S. derbyi Waagen 1883BJ65
`--Meristospira Grabau 1910B75, BJ65
`--*M. michiganense Grabau 1910BJ65
Athyrididae incertae sedis:
Atrythyris Struve 1965CSZ03
Anomactinella Bittner 1890BJ65
`--*A. ‘flexuosa’ (Münster 1841) [=Terebratula flexuosa non King 1835]BJ65
Dioristella Bittner 1890BJ65
`--*D. indistincta (von Beyrich 1862) [=Terebratula indistincta]BJ65
Janiceps Frech 1901BJ65
`--*J. peracuta (Stache 1878) [=Athyris peracuta]BJ65
Pentactinella Bittner 1890BJ65
`--*P. quinquecostata (Münster 1841) [=Terebratula quinquecostata]BJ65
Pradoia Comte 1938BJ65
|--*P. torenoi (de Verneuil & d’Archiac 1845) [=Terebratula torenoi]BJ65
`--P. colletteiBJ65
Stolzenburgiella Bittner 1903BJ65
`--*S. bukowskii Bittner 1903BJ65
Tetractinella Bittner 1890BJ65
`--*T. trigonella (von Schlotheim 1820) [=Terebratulites trigonellus]BJ65
Apheathyris Fu 1982HB83, HC04
`--A. guyuanensis Fu 1982HB93
DichozygopleuraB75
Nomen nudum: Lochengia holoensis Grabau 1931BJ65
*Actinoconchus planosulcata (Phillips 1836)BJ65 [=Spirifera planosulcataBJ65, Athyris (Actinoconchus) planosulcatusF71, Spirifer planosulcataW77, Terebratula planosulcataW77; incl. Actinoconchus paradoxus M’Coy 1844BJ65]
Cleiothyridina Buckman 1906CSZ03 [=Cleiothyris King 1850 non Phillips 1841BJ65, Cliothyris Hall & Clarke 1893 non Agassiz 1846BJ65]
*Cleiothyridina deroissii (Léveillé 1835)BJ65 [=Spirifer deroissyiBJ65, Athyris royssii Davidson 1860BJ65, *Cleiothyris deroissyiBJ65, Cliothyris royssiiG31, Spirigera royssiiG31; incl. Athyris royssii var. orientalisG31]
*Lamellosathyris lamellosa (Léveillé 1835) [=Spirifer lamellosus, Actinoconchus lamellosus, Athyris lamellosa, A. lamellosus (l. c.), Cleiothyridina lamellosa, Lamellosathyris lamellosus (l. c.); incl. Spirifer squamosa Phillips 1836]CSZ03
*Type species of generic name indicated
References
Alvarez, F., & C. H. C. Brunton. 1990. The shell-structure, growth and functional morphology of some Lower Devonian athyrids from northwest Spain. Lethaia 23: 117–131.
Alvarez, F., Rong J.-Y. & A. J. Boucot. 1998. The classification of athyridid brachiopods. Journal of Paleontology 72 (5): 827–855.
Blodgett, R. B., A. J. Boucot & W. F. Koch II. 1988. New occurrences of color patterns in Devonian articulate brachiopods. Journal of Paleontology 62 (1): 46–51.
[BM05] Botquelen, A., & E. Mayoral. 2005. Early Devonian bioerosion in the Rade de Brest, Armorican Massif, France. Palaeontology 48 (5): 1057–1064.
[B75] Boucot, A. J. 1975. Evolution and Extinction Rate Controls. Elsevier Scientific Publishing Company.
[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).
[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.
[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.
[HC04] Harper, D. A. T., L. R. M. Cocks, L. E. Popov, P. M. Sheehan, M. G. Bassett, P. Copper, L. E. Holmer, J. Jin & Rong J. 2004. Brachiopods. In: Webby, B. D., F. Paris, M. L. Droser & I. G. Percival (eds) The Great Ordovician Biodiversification Event pp. 157–178. Columbia University Press.
[M03] Modzalevskaya, T. L. 2003. Silurian and Devonian brachiopods from Severnaya Zemlya (Russian Arctic). Geodiversitas 25 (1): 73–107.
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.
[TY12] Talent, J. A., Y. A. Yokin, V. N, Yolkina, T. P. Kipriyanova & R. T. Gratsianova. 2012. Changes in the pattern of brachiopod biogeography in northern Asia through early and middle Devonian times. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 821–829. Springer.
[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.
[WBM97] Williams, A., C. H. C. Brunton & D. I. MacKinnon. 1997. Morphology. In: Kaesler, R. L. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda, Revised vol. 1. Introduction pp. 321–422. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).
[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.