ALLAGECRINOIDEA AND CODIACRINOIDEA: MICROCRINOIDS (ECHINODERMATA) FROM THE UPPER KUNGURIAN (LOWER PERMIAN) OF OMAN
DOI:
https://doi.org/10.54103/2039-4942/30643Keywords:
Permian; Crinoidea; disparids; eucladids; Russia; West Timor.Abstract
Upper Kungurian (Lower Permian) crinoids of the Allagecrinoidea and Codiacrinoidea are described from the Ash Sharqiyah South region of the Sultanate of Oman. These two taxonomic groups include both microcrinoids and small macrocrinoids that were isolated from washing bulk samples of the Qarari Unit from two sample sites near Wadi Khawr Al Jaramah, Batain, Oman. Eight genera, one new, and fourteen species, twelve new, are described. The Allagecrinoidea and Codiacrinoidea described herein complete our present understanding of crinoids from the Permian of Oman and help to better understand Permian echinoderm paleocommunities in the Neo-Tethys. The generic composition of this Oman microcrinoid fauna is intermediate in composition between those of Russia and West Timor.
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References
Angiolini L., Balini M., Garzanti E., Nicora A., Tintori A., Crasquin S. & Muttoni G. (2003) - Permian climatic and paleogeographic changes in Northern Gondwana: the Khuff Formation of Interior Oman. Palaeogeography, Palaeoclimatology, Palaeoecology, 191: 269-300.
Arendt Y.A. (1970) - Morskie Lilii Gipokrinidy [Crinoids Hypocrinidae]. Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSSR, 128: 220 p.
Arendt Y.A. & Hecker R.T. (1964) – Klass Crinoidea. Morski lilii sistematitcheckaia tchast. In: Orlov Y.A. (Ed) - Osnovi Paleontologii, Iglokozhi, Gemikhordovye, Pogonofory, i Shchetinkochelyustnye. Moscow, Izdatelstvo Nedra: 76-105, 214-231.
Ausich W.I. & Donovan S.K. (2023) - Chapter 7: Glossary of morphological terms. In: Ausich W. I. (Ed) - Treatise on Invertebrate Paleontology, Part T, Revised, Volume 1, University of Kansas Press, Lawrence, Kansas, 26 p.
Ausich W.I., Wright D.F., Cole S.R. & Sevastopulo G.D. (2020) - Homology of posterior interray plates in crinoids: A review and new perspectives from phylogenetics, the fossil record, and development. Palaeontology, 63: 525-545. http://doi.org/10.1111/pala.12475
Bassler R.S. & Moodey M.W. (1943) - Bibliographic and faunal index of Paleozoic pelmatozoan echinoderms. Geological Society of America Special Paper, 45: 734 pp.
Bather F.A. (1890) - British fossil crinoids. II The classification of the Inadunata Fistulata (cont’d.). Annals and Magazine of Natural History, ser. 6, 5, 373-388, 485-486.
Bather F.A. (1899) - A phylogenetic classification of the Pelmatozoa. British Association for the Advancement of Science, 1898, 916-923.
Branson C.C. (1948) - Bibliographic index of Permian invertebrates. Geological Society of America Memoir, 22: 163-164.
Carpenter P.H. & Etheridge R. Jr. (1881) - Contributions to the study of the British Paleozoic crinoids. - No. 1. On Allagecrinus, the representative of the Carboniferous limestone series. Annals and Magazine of Natural History, ser. 5, 7: 281-298.
Charlton T.R., Barber A.J., Harris R.A., Barkham S.T., Bird P.R., Archbold N.W., Morris N.J., Nicoll R.S., Owen H.G., Owens R.M., Sorauf J.E., Taylor P.D., Webster G.D. & Whittaker J.E. (2002) - The Permian of Timor: stratigraphy, palaeontology and palaeogeography. Journal of Asian Earth Sciences. 20(6): 719-774.
Eagle M.K., Hoskin L. & Hayward B.W. (2018) -The first macrofossil (Crinoidea: Cladida) from the Caples Terrane, Northland, North Island, New Zealand. New Zealand Journal of Geology and Geophysics, 61(4): 498-507.
Erwin D.H. (2006) - Extinction. Princeton University Press; Princeton, New Jersey, 296 pp.
Fortey R.A. & Heward A.P. (2015) - A new morphologically diverse Permian trilobite fauna from Oman. Acta Palaeontologica Polonica, 60(1): 201-216.
Girty G.H. (1908) - The Guadalupian fauna. U. S. Geological Survey, Professional Paper, 58: 512 pp.
Golonka J. (2007) - Phanerozoic paleoenvironmental and paleolithofaces maps: Late Paleozoic. Geologia, 33(2): 145-209.
Immenhauser A., Schreurs G., Peters T., Matter A., Hauser M. & Dumitrica P. (1998) - Stratigraphy, sedimentology, and depositional frameworks of the Permian to uppermost Cretaceous Batain Group, eastern-Oman. Eclogae Geologicae Helvetiae, 91: 217-235.
Immenhauser A., Schreurs G., Gnos E., Oterdoom H.W. & Hartmann B. (2000) - Late Palaeozoic to Neogene geodynamic evolution of northeastern Oman margin. Geological Magazine, 137(1): 1-18.
Jaekel O. (1894) - Über die Morphogenie und Phylogenic der Crinoiden. Sitzungsberichten der Gesellschaft Naturforschender Freunde, Jahrgang 1894, 4: 101-121.
Jell P.A. & Willink R.J. (1993) – Early Permian cladid crinoids in the Gharif Formation of Oman. Jell P.A. (Ed) – Paleontological studies in honour of Ken Campbell. Memoir Association of Australasian Palaeontologists, 15: 305-312.
Kammer T.W., Sumrall C.D., Zamora S., Ausich W.I. & Deline B. (2013) - Oral region homologies in Paleozoic crinoids and other plesiomorphic pentaradial echinoderms. PLoS ONE 8(11): e77989 [doi:10.1371/journal.pone.0077989].
Kirk E. (1914) - Notes on the fossil crinoid genus Homocrinus Hall. Proceedings of the U.S. National Museum, 46: 473-483.
Lane N.G. (1967) - Revision of suborder Cyathocrinina (Class Crinoidea). University of Kansas Paleontological Contributions, Paper, 24: 1-13.
Lane N.G. & Sevastopulo G.D. (1981) - Functional of a microcrinoid: Kallimorphocrinus punctatus n. sp. Journal of Paleontology, 55: 13-28.
Lane N.G. & Sevastopulo G.D. (1982) - Growth and systematic revision of Kallimorphocrinus astrus, a Pennsylvanian microcrinoid. Journal of Paleontology, 56: 244-259.
Lane N.G., Sevastopulo G.D. & Strimple H.L. (1985) - Amphipsalidocrinus: a monocyclic camerate microcrinoid. Journal of Paleontology, 59: 78-84.
Leven E. & Heward A.P. (2013) - Fusulinids from isolated Qarari outcrops (Permian) occurring among Jurassic–Cretaceous Batain Group (Baitain Plain, Eastern Oman). Rivista Italiana di Paleontologia e Stratigrafia, 119(2): 153-162.
Marez Oyens F.A.H.W. de. (1940) - Neue Permische Krinoiden von Timor, mit Bemerkungen über deren Vorkommen in Basleogebiet. Geological Expedition to the Lesser Sunda Island under leadership of H. A. Brouwer, 1: 285-348.
Miller J.S. (1821) - A natural history of the Crinoidea, or lily-shaped animals; with observations on the genera, Asteria, Euryale, Comatula and Marsupites. Bryan & Co., Bristol, England, 150 pp.
Moore R.C. & Ewers J.D. (1942) - A new species of Synbathocrinus from the Mississippian rocks of Texas, with description of ontogeny. Journal of the Scientific Laboratories of Denison University, 42(7): 92-106.
Moore R.C. & Laudon L.R. (1943) - Evolution and classification of Paleozoic crinoids. Geological Society of America, Special Paper, 46: 151 pp.
Moore R.C. & Teichert C. (Eds). (1978) - Treatise on Invertebrate Paleontology, Part T, Echinodermata 2, Crinoidea. 3 vols., Geological Society of America and University of Kansas; Boulder, Colorado and Lawrence, Kansas, 1027 pp.
Peck R.E. (1936) - Lower Mississippian microcrinoids from the Kinderhook and Osage groups of Missouri. Journal of Paleontology, 10: 282-293.
Peters T., Al-Battashy M., Bläsi H., Hauser M., Immenhauser A., Moser L. & Al-Rajhi A. (2001) - Geological map of Sur and Al Ashkharah, Sheet NF 40-8F and Sheet NF 40-12C, Scale 1:100,000, Explanatory Notes. Sultanate of Oman Ministry of Commerce and Industry, Directorate General of Minerals, 9 pp.
Reich M. (2007) – Linguaserra spandeli sp. nov. (Echinodermata: Ophiocistioidea) from the Late Permian (Zechstein) of Thuringia, Germany/ Annales de Paléontologie, 93: 317-330.
Sepkoski J.J., Jr. (1981) - A factor analytical description of the Phanerozoic marine fossil record. Palaeobiology, 7: 36-53.
Sevastopulo G.D. (2005) - The early ontogeny of blastoids. Geological Journal, 40: 351-362.
Shellnut J.G., Bhat G.M., Brookfield M.E. & Jahn B.-M. (2011) - No link between the Panjal Traps (Kashmir) and the Late Permian mass extinctions. Geophysical Research Letters, 38: L19308, doi:10.1029/2011GL049032, 2011.
Strimple H.L. (1966) - Some notes concerning the Allagecrinidae. Oklahoma Geology Notes, 26: 99-111.
Strimple H.L. & Koenig J.W. (1956) - Mississippian microcrinoids from Oklahoma and New Mexico. Journal of Paleontology, 30: 1225-1247.
Strimple H.L. & Sevastopulo G.D. (1982) - A Permian microcrinoid from Sicily. Journal of Paleontology, 56: 1451-1452.
Strimple H.L., Allison R.C. & Kline G.L. (1971) - Pennsylvanian crinoids from Alaska, Part 2. In Fossil Crinoid Studies. University of Kansas Paleontological Contributions, Paper, 56(2): 9-15.
Ubaghs G. (1978) - General morphology. In: Moore R.C. & Teichert C. (Eds) - Treatise on Invertebrate Paleontology. Pt. T. Echinodermata 2 Geological Society of America and University of Kansas Press, Boulder, Colorado and Lawrence, Kansas, T58-T216.
Viaretti M., Heward A.P., Gementi A. & Angiolini L. (2022) - Upper Cisuralian-lower Guadalupian brachiopods from the Qarari Unit, Batain Plain, northeastern Oman: systematics, palaeoecology and correlations. Rivista Italiana di Paleontologia e Stratigrafia, 128(3): 643-694.
Wachsmuth C. & Springer F. (1880–1886) - Revision of the Palaeocrinoidea. Proceedings of the Academy of Natural Sciences of Philadelphia Pt. I. The families Ichthyocrinidae and Cyathocrinidae (1880), 226-378 (separate repaged 1-153. Pt. II. Family Sphaeroidocrinidae, with the sub-families Platycrinidae, Rhodocrinidae, and Actinocrinidae (1881), 177-411 (separate repaged, 1-237). Pt. III, Sec. 1. Discussion of the classification and relations of the brachiate crinoids, and conclusion of the generic descriptions (1885), 225-364, (separate repaged, 1-138). Pt. III, Sec. 2. Discussion of the classification and relations of the brachiate crinoids, and conclusion of the generic descriptions (1886), 64-226 (separate repaged to continue with section 1, 139-302).
Wanner J. (1916) - Die Permischen echinodermen von Timor, I. Teil. Palaontologie von Timor, 11: 329 pp.
Wanner J. (1920) - Über Armlose Krinoiden aus dem jüngeren Palaeozoikum. Verhandelingen Geologie Mijnbouw Genootoch, Geologic Series, 5: 21-35.
Wanner J. (1924) - Die permischen Krinoiden von Timor. Jaarbook van net Mijnwezen Nederlandes Oost-Indie, Verhandlungen (1921), Gedeelte, 3: 348 pp.
Wanner J. (1929) - Neue Beitrage zur Kenntnis der Permischen Echinodermen von Timor. I. Allagecrinus. II. Hypocrinites. Dienst van den Mijnbouw in Nederlandsch-Indie, Wetenschappelijke Mededeelingen, 11: 1-116.
Wanner J. (1930) - Neue Beitrage zur Kenntnis der Permischen Echinoderm von Timor, III. Hypocrinae, Paracatillocrinus und Allagecrinus dux. Dienst van den Mijnbouw in Nederlandsch-Indie, Wetenschappelijke Mededeelingen, 13: 31 pp.
Webster G. D. (1973) - Bibliography and index of Paleozoic crinoids, 1942-1968. Geological Society of America, Memoir 137: 341 pp.
Webster G.D. (1974) - Crinoid pluricolumnal noditaxis patterns. Journal of Paleontology, 48: 1283-1288.
Webster G.D. (1987) - Permian crinoids from the type-section of the Callytharra Formation, Callytharra Springs, Western Australia. Alcheringa, 11: 95-135.
Webster G.D. (1998) - Distortion of the stratigraphy and biostratigraphy of Timor, a historical review with an analysis of the crinoid and blastoid faunas. Proceedings of the Royal Society of Victoria, 110: 45-72.
Webster G.D. & Jell P. A. (1992) - Permian echinoderms from Western Australia. Memoirs of the Queensland Museum, 32(1): 311-373.
Webster G.D. & Jell P.A. (1999) - New Permian crinoids from eastern Australia. Memoirs of the Queensland Museum, 43(1): 279-340.
Webster G.D. & Sevastopulo G.D. (2007) - Paleogeographic significance of Early Permian crinoids and blastoids from Oman. Paläontologische Zeitschrift, 81(4): 399-405.
Webster G.D. & Webster D.W. (2014) - Bibliography and index of Paleozoic crinoids, coronates, and hemistreptocrinoids. http://crinoids.azurewebsites.net/ (accessed 2025-12-12).
Webster G.D., Angiolini L. & Tintori A. (2009) - Permian crinoids from the Saiwan and Khuff Formations, southeastern Oman. Rivista, Italiana, di Paleontologia e Stratigrafia, 115(1): 27-48.
Webster G.D., Ausich W.I., Waters J.A. & Heward A.P. (2025) - New Permian echinoderms from northeastern Oman have connections with Tethyan echinoderm faunas from West Timor, Western Australia, and Russia. Rivista, Italiana, di Paleontologia e Stratigrafia, 131(3): 631-676 https://doi.org/10.54103/2039-4942/28160
Webster G.D., Heward A.P. & Ausich W.I. (2022) - First crinoid crown from the Permian Khuff Formation of Oman. Proceedings of the Geologists’ Association, 133: 154-161. doi.org/10.1016/jpgeola2022.02005)
Webster G.D., Maples C.G., Yazdi M., Marcus S. & Waters J.A. (2011) - Early Pennsylvanian, Bashkirian, echinoderms from eastern Iran, a potential transitional fauna between Laurentia/Avalonia and the Paleotethys, and a Permian cromyocrinid from central Iran. Palaeobiodiversity and Palaeoenvironments, 91: 1-61.
Weller J. M. (1930) - A group of larviform crinoids from Lower Pennsylvanian strata of the eastern Interior Basin. Illinois Geological Survey, Report of Investigations, 21: 43 p.
Wright D.F. (2017a) - Bayesian estimation of fossil phylogenies and the evolution of early to middle Paleozoic crinoids (Echinodermata). Journal of Paleontology, 91: 799-814. https://doi.org/10.1017/jpa.2016.141
Wright D.F. (2017b) - Phenotypic innovation and adaptive constraints in the evolutionary radiation of Palaeozoic crinoids. Scientific Reports, 7: 13745. https://doi.org/10.1038/s41598-017-13979-9
Wright D.F., Ausich W.I., Cole S.R., Rhenberg E.C. & Peter M.E. (2017) - Phylogenetic taxonomy and classification of the Crinoidea (Echinodermata). Journal of Paleontology, 91: 829-846. https://doi.org/10.1017/jpa.2016.142
Wright J. (1933) - Two new crinoids from the Scottish Carbonifeous Limestones, with notes on the Allagecrinidae. Geological Magazine, 70: 193-208.
Yakovlev N.N. (1926) - Fauna iglokozhikh permokarbona iz Krasnoufimska na Urale. I [Echinoderm fauna from the Permocarboniferous of Krasnoufimsk in the Urals, I]. Izvestiy Geologicheskogo Komiteta, Leningrad, 45(2): 51-57.
Yakovlev N.N. (1927) - Fauna iglokozhikh permokarbona iz Krasnoufimska na Urale. II [Echinoderm fauna from the Permocarboniferous of Krasnoufimsk in the Urals, II]. Izvestiy Geologicheskogo Komiteta, Leningrad, 46(3): 181-192.
Yakovlev N.N. (1930) - Notes sur les Crinoïdes paléozoiques. Part 1. Contribution a la connaissance de Hexacrinus patereformis Schultze et du genre Hexacrinus, 907–908, Part 2. Crinoïdes du Permien de la Sicile, 908–910, Part 3. Un nouveau crinoide du bassin du Donetz. Izvestiya Akademii Nauk SSSR (Bulletin de l’Academie Sciences de l’URSS), 907-911.
Zittel K.A. von. (1880) - Handbuch der Palaeontologie, v. 1, Palaeozoologie. München, Leipzig, R. Oldenbourg (1879), 1:765 pp.
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