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The strophomenide brachiopod Ahtiella Öpik in the Ordovician of Gondwana and the early history of the plectambonitoids

  • Juan L. Benedetto (a1)

The Precordilleran species Ahtiella argentina Benedetto and Herrera, 1986 is redescribed and illustrated and Monorthis coloradoensis Benedetto, 1998b from northwestern Argentina is reassigned to the genus Ahtiella Öpik, 1932. Ahtiella famatiniana new species from volcaniclastic rocks of the Famatina range (western Argentina) and Ahtiella tunaensis new species from the Precordillera basin (Cuyania terrane) are proposed. Paleogeographic and stratigraphic evidence strongly suggests that Ahtiella originated in the Andean region of Gondwana to further migrate to Avalonia, Baltica, and Cuyania. Contrary to previous assumptions, the fossil record from the Famatina volcaniclastic succession suggests that the plectambonitoid Ahtiella famatiniana n. sp. evolved from the hesperonomiid orthoid Monorthis transversa Benedetto, 2003 that always occurs in the underlying strata. Phylogenetic analysis of Ahtiella species shows that A. famatiniana n. sp. and the Peruvian A. zarelae Villas in Gutiérrez-Marco and Villas, 2007 are not only the earliest species of the genus but also are morphologically intermediate between Monorthis Bates, 1968 and the later and more derived species of Ahtiella from Baltica and Cuyania. If, as empirical evidence presented here shows, Ahtiella originated from Monorthis through a series of minor transformations, then the impressive morphological gap between orthides and strophomenides was bridged through short-time cladogenesis events, suggesting that it might not have a definite discontinuity between the species level evolution and the origin of higher taxa (macroevolution).


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Aceñolaza, F.G., Miller, H., and Toselli, A.J., 2002, Proterozoic−early Paleozoic evolution in western South America—A discussion: Tectonophysics, v. 354, p. 121137. doi: 10.1016/s0040-1951(02)00295-0.
Albanesi, G.L., and Astini, R.A., 2000, New conodont fauna from Suri Formation (Early−Middle Ordovician), Famatina System, western Argentina: Reunión Anual de la Asociación Paleontológica Argentina, Mar del Plata, Abstracts, Ameghiniana, v. 37, no. 4, supplement, p. 68.
Albanesi, G.L., and Astini, R.A., 2002, Faunas de conodontes y Sacabambaspis janvieri (Vertebrata) en el Ordovícico Medio de la Cordillera Oriental Argentina: Implicancias estratigráficas y paleobiogeográficas: Congreso Argentino de Paleontología y Bioestratigrafía, 8th, Corrientes, Argentina, Abstracts, p. 17.
Albanesi, G.L., and Ortega, G., 2002, Advances on conodont-graptolite biostratigraphy of the Ordovician System of Argentina, in Aceñolaza, F.G., ed., Aspects of the Ordovician System in Argentina: INSUGEO Serie Correlación Geológica, v. 16, p. 143166.
Albanesi, G.L., and Vaccari, N.E., 1994, Conodontos del Arenig en la Formación Suri, Sistema de Famatina: Argentina, Revista Española de Micropaleontología, v. 26, p. 125146.
Astini, R.A., 1994, Interpretación estratigráfica de la Formación Sepulturas (Ordovícico Inferior) y unidades análogas del Noroeste Argentino: La Aloformación Sepulturas: 5th Reunión Argentina de Sedimentología, San Miguel de Tucumán, Argentina, Abstracts, p. 9–14.
Astini, R.A., 1998, Stratigraphical evidence supporting the rifting, drifting and collision of the Laurentian Precordillera terrane of western Argentina, in Pankhurst, R.J., and Rapela, C.W., eds., The Proto-Andean Margin of Gondwana: Geological Society London Special Publication, v. 142, p. 1133.
Astini, R.A., 1999, Sedimentary record, volcano-tectonic cyclicity and progressive emergence of an Early Ordovician perigondwanan volcanic arc: The Famatina System, in Kraft, P., and Fatka, O., eds., Quo vadis Ordovician?: Acta Universitatis Carolinae Geologica, v. 43, nos. 1/2 p. 115118.
Astini, R.A., 2003, The Ordovician proto-Andean basins, in Benedetto, J.L., ed., Ordovician Fossils of Argentina: Córdoba, Argentina, Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba, p. 174.
Astini, R.A., and Benedetto, J.L., 1996, Paleoenvironmental features and basin evolution of a complex island-arc region in the pre-Andean western Gondwana: The Famatina belt: 3rd International Symposium on Andean Geodynamics, St. Malo, France, Short Papers, p. 755–758. St. Malo.
Astini, R.A., and Dávila, D.F., 2002, El Grupo Cerro Morado (Ordovícico Medio) en el Famatina (28°−29°S), Andes centrales del oeste argentino: Revista Geológica de Chile, v. 29, p. 241254. doi: 10.4067/S0716-02082002000200006.
Astini, R.A., and Rapalini, A.E., 2003, Discussion. ‘Proterozoic-Early Paleozoic evolution in western South America - a discussion’ in: Tectonophysics, 354: 121–137 (2002): Tectonophysics, v. 366, p. 143–148.
Astini, R.A., Benedetto, J.L., and Vaccari, N.E., 1995, The early Paleozoic evolution of the Argentine Precordillera as a Laurentian rifted, drifted and collided terrane: Geological Society of America Bulletin, v. 107, p. 253273.
Astini, R.A., Toro, B.A., Waisfeld, B.G., and Benedetto, J.L., 2004a, Revisión estratigráfica de la región de Los Colorados: Un perfil condensado del Paleozoico inferior en el borde occidental del Cordillera Oriental, Provincia de Jujuy: Revista de la Asociación Geológica Argentina, v. 59, p. 243260.
Astini, R.A., Thomas, W.A., and Yochelson, E.L., 2004b, Salterella in the Argentine Precordillera: An Early Cambrian paleobiogeographic indicator of Laurentian affinity: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 213, p. 125132. doi: 10.1016/j.palaeo.2004.07.008.
Baldo, E.G., Fanning, C.M., Rapela, C.W., Pankhurst, R.J., Casquet, C., and Galindo, C., 2003, U-Pd SHRIMP dating of rhyolite volcanism in the Famatina belt and K-bentonites in the Precordillera, in Albanesi, G.L., Beresi, M.S., and Peralta, S H., eds., Ordovician from the Andes: Proceedings of the 9th International Symposium on the Ordovician System, INSUGEO Serie Corelación Geológica, v. 17, p. 285–189.
Bancroft, B.B., 1949, Welsh Valentian brachiopods, in Lamont, A., ed., Welsh Valentian Brachiopods and the Strophomena antiquata Group of Fossil Brachiopods: Preprint from The Quarry Managers’ Journal, Mexborough, Yorkshire, UK, Times Printing Company, p. 2–10, pls. 1–2.
Bassett, M.G., 1984, Life strategies in Silurian brachiopods, in Bassett, M.G., and Lawson, J.D., eds., Autoecology of Silurian Ornanisms: The Paleontological Society, Special Paper in Palaeontology, v. 32, p. 237264.
Bassett, M.G., and Popov, L.E., 2017, Earliest ontogeny of the Silurian orthotetide brachiopod Coolinia and its significance for interpreting strophomenate phylogeny: Lethaia, v. 50, p. 504510. doi: 10.1111/let.12204.
Bassett, M.G., Popov, L.E., and Holmer, L.E., 2001, Functional morphology of articulatory structures and implications for patterns of musculature in Cambrian rhynchonelliform brachiopods, in Brunton, C.H.C., Cocks, L.R.M., and Long, S.L., eds., Brachiopods Past and Present: London, Taylor & Francis, p. 163176.
Bassett, M.G., Popov, L.E., and Holmer, L.E., 2002, Brachiopods: Cambrian-Tremadocian precursors to Ordovician radiation events, in Crame, J.A., and Owen, A.W., eds., Palaeobiogeography and Biodiversity Change: The Ordovician and Mesozoic-Cenozoic Radiations: Geological Society London Special Publication, v. 194, p. 1323.
Bassett, M.G., Popov, L.E., and Egerquist, E., 2008, Early ontogeny of some Ordovician-Silurian strophomenate brachiopods: Significance for interpreting evolutionary relationships within early Rhynchonelliformea: Fossils and Strata, v. 54, p. 1320.
Bates, D.E.B., 1968, The lower Paleozoic brachiopod and trilobites faunas of Anglesey: Bulletin of the British Museum (Natural History) Geology, v. 16, p. 125199.
Bates, D.E.B., 1969, Some early Arenig brachiopods and trilobites from Wales: Bulletin of the British Museum (Natural History) Geology, v. 18, p. 128.
Benedetto, J.L., 1994, Braquiópodos ordovícicos (Arenigíano) de la Formación Suri en la región de Chaschuil, Sistema de Famatina, Argentina: Ameghiniana, v. 31, p. 221238.
Benedetto, J.L., 1998a, Early Palaeozoic brachiopods and associated shelly faunas from western Gondwana: Its bearing on the geodynamic history of the pre-Andean margin, in Pankhurst, R.J., and Rapela, C.W., eds., The Proto-Andean Margin of Gondwana: Geological Society London Special Publication, v. 142, p. 5783.
Benedetto, J.L., 1998b, Early Ordovician (Arenig) brachiopods from the Acoite and Sepulturas formations: Cordillera Oriental, northwestern Argentina: Geologica et Palaeontologica, v. 32, p. 727.
Benedetto, J.L., 2001, Silicified Early Ordovician (Arenig) brachiopods from the San Juan Limestone, Argentine Precordillera: Geologica et Palaeontologica, v. 35, p. 129.
Benedetto, J.L., 2002, The Ordovician brachiopod faunas of Argentina: Chronology and biostratigraphic succession, in Aceñolaza, F.G, ed., Aspects of the Ordovician System in Argentina: 9th International Symposium on the Ordovician System, San Miguel de Tucumán, Argentina, Proceedings, INSUGEO, Serie Correlación Geológica, v. 16, p. 87–106.
Benedetto, J.L., 2003a, Brachiopods, in Benedetto, J.L., ed., Ordovician Fossils of Argentina: Córdoba, Argentina. Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba, p. 187272.
Benedetto, J.L., 2003b, Early Ordovician (Arenig) brachiopods from volcaniclastic rocks of the Famatina Range, northwest Argentina: Journal of Paleontology, v. 77, p. 212242. doi: 10.1666/0022-3360(2003)077 0212:EOABFV 2.0.CO;2.
Benedetto, J.L., 2004, The allochthony of the Precordillera ten years later (1993–2003): A new paleobiogeographic test of the microcontinental model: Gondwana Research, v. 7, p. 10271039. doi: 10.1016/S1342-937X(05)71082-0.
Benedetto, J.L., 2007, Brachiopod succession in the Lower-Middle Ordovician carbonate platform of the Precordillera terrane, western Argentina: An example of interplay between environmental, biogeographic, and evolutionary processes: Acta Palaeontologica Sinica, v. 46, p. 2836.
Benedetto, J.L., 2008, Spatial and stratigraphic distribution of the rhynchonelliformean brachiopod Productorthis Kozlowski: Fast migrations or parallel evolution?, in Harper, D.A.T., Long, S.L., and Nielsen, C., eds., Brachiopoda: Fossil and Recent: Fossils and Strata, v. 54, p. 2129.
Benedetto, J.L., 2009, Chaniella, a new lower Tremadocian (Ordovician) brachiopod from northwestern Argentina and its phylogenetic relationships with basal rhynchonelliforms: Palaeontologische Zeitschrift, v. 83, p. 393405.
Benedetto, J.L., 2013, Presence of punctae in the ‘plectorthoidean’ brachiopod Famatinorthis turneri (Middle Ordovician) from western Argentina: Implications for early diversification of punctate orthides: Lethaia, v. 42, p. 170179. doi: 10.1111/j.1502-3931.2012.00330.x.
Benedetto, J.L., and Carrasco, P., 2002, Tremadoc (earliest Ordovician) brachiopods from the Purmamarca region and the Sierra de Mojotoro, Cordillera Oriental of northwestern Argentina: Geobios, v. 35, p. 647661. doi: 10.1016/S0016-6995(02)00079-7.
Benedetto, J.L., and Foglia, R.D., 2012, Lower and Middle Cambrian rhynchonelliformean brachiopods from the Precordillera terrane of Argentina: Journal of Paleontology, v. 86, p. 273281. doi: 10.2307/41480192.
Benedetto, J.L., and Herrera, Z.A., 1986, Braquiópodos del Suborden Strophomenidina de la Formación San Juan (Ordovícico temprano), Precordillera de San Juan, Argentina: Actas 4th Congreso Argentino Paleontología y Bioestratigrafía, Mendoza, Argentina, v. 1, p. 113123.
Benedetto, J.L., and Herrera, Z.A., 1987, Sanjuanella, un nuevo género de la subfamilia Ahtiellinae (Brachiopoda, Plectambonitacea) del Ordovícico de la Precordillera Argentina: Actas 4th Congreso Latinoamericano de Paleontología, Santa Cruz de la Sierra, Bolivia, v. 1, p. 97109.
Benedetto, J.L., and Muñoz, D.F., 2017, Plectorthoid brachiopods from the Lower Ordovician of NW Argentina: Phylogenetic relationships with Tarfaya Havlíček and the origin of heterorthids: Journal of Systematic Palaeontology, v. 15, p. 4367. doi: 10.1080/14772019.2016.1144086.
Benedetto, J.L., and Sánchez, T.M., 2003, The Ordovician radiation from a Gondwanan perspective: The early diversification of brachiopods and bivalves on mid- to high- latitude siliciclastic platforms, in Albanesi, G.L., Beresi, M.S., and Peralta, S.H., eds., Ordovician from the Andes, 9th International Symposium on the Ordovician System, San Miguel de Tucumán, Argentina, Proceedings, INSUGEO Serie Corelacion Geologica, v. 17, p. 245–250.
Benedetto, J.L., Dávila, D.F., and Astini, R.A., 2003, La presencia de Ahtiella Öpik y otros braquiópodos en la sucesión volcaniclástica ordovícica de la Sierra de Famatina: Connotaciones bioestratigráficas y paleogeográficas: Ameghiniana, v. 40, p. 80.
Benedetto, J.L., Sorrentino, L., Cech, N., and Sánchez, T.M., 2008, Functional morphology of the Ordovician plectambonitoid brachiopod Inversella (Reinversella) arancibiai Herrera & Benedetto from the San Juan Formation, Argentine Precordillera: Alcheringa, v. 32, p. 5363. doi: 10.1080/03115510701757373.
Benedetto, J.L., Vaccari, N.E., Waisfeld, B.G., Sánchez, T.M., and Foglia, R.D., 2009, Cambrian and Ordovician paleobiogeography of the South American margin of Gondwana and accreted terranes, in Bassett, M.G., ed., Early Palaeozoic Peri-Gondwanan Terranes: New Insights from Tectonics and Biogeography: The Geological Society London Special Publication, v. 325, p. 199230.
Benton, M.J., and Pearson, P.N., 2001, Speciation in the fossil record: Trends in Ecology and Evolution, v. 16, p. 405411. doi: 10.1016/S0169-5347(01)02149-8.
Bergström, S.M., Chen, X., Gutiérrez-Marco, J.C., and Dronov, A., 2009, The new chronostratigraphic classification of the Ordovician System and its relations to major regional series and stages and to δ13C chemostratigraphy: Lethaia, v. 42, p. 97107. doi: 10.1111/j.1502-3931.2008.00136.x.
Billings, E., 1859, On some new genera and species of Brachiopoda from the Silurian and Devonian rocks of Canada: Canadian Naturalist and Geologist, v. 4, p. 1131.
Bruton, D.L., and Harper, D.A.T., 1981, Brachiopods and trilobites of the Early Ordovician serpentine Otta Conglomerate, south central Norway: Norks Geologisk Tidsskrift, v. 61, p. 153181.
Butts, C., 1926, The Paleozoic rocks, in Adams, G.I., Butts, C., Stephenson, L.W., and Cooke, C.W., eds., Geology of Alabama: Alabama Geological Survey Special Report 14, p. 41–230.
Cañas, F.L., 1999, Facies sequences of late Cambrian to Early Ordovician carbonates of the Argentine Precordillera: A physical stratigraphic comparison with Laurentian platforms, in Keppie, D., and Ramos, V.A., eds., Laurentia-Gondwana Connections Before Pangea: Geological Society of America Special Paper 336, p. 4362.
Carlorosi, J., Heredia, S., and Aceñolaza, G., 2017, Conodontes de la Formación Suri en la región de Chaschuil, Sistema de Famatina, Argentina: Congreso Geológico Argentino, 20th, San Miguel de Tucumán, Argentina, Actas, p. 2123.
Carlson, S.J., 2016, The evolution of Brachiopoda: Annual Review of Earth and Planetary Sciences, v. 44, p. 409438. doi: 10.1146/annurev-earth-060115-012348.
Carrera, M.G., 1997, Significado paleoambiental de los poríferos y briozoos de la Formación San Juan (Ordovícico), Precordillera argentina: Ameghiniana, v. 34, p. 179199.
Carrera, M.G., 2003, Sponges and Bryozoans, in Benedetto, J.L., ed., Ordovician Fossils of Argentina: Córdoba, Argentina, Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba, p. 155185.
Carrera, M.G., and Astini, R.A., 1998, Valoración de las restricciones ambientales durante la transición Arenigiano-Llanvirniano: Ordovícico de la Precordillera Argentina: Revista de la Asociación Geológica Argentina, v. 53, p. 4156.
Carrera, M.G., and Ernst, A., 2010, Darriwilian bryozoans from the San Juan Formation (Ordovician), Argentine Precordillera: Ameghiniana, v. 47, p. 343354. doi: 10.5710/AMGH.v47i3.3.
Carrera, M.G., and Rigby, J.K., 1999, Biogeography of the Ordovician sponges: Journal of Paleontology, v. 73, p. 637.
Cisterna, C.E., 2001, Volcanismo subácueo en el Eopaleozoico del Sistema de Famatina, noroeste de Argentina: Revista de la Asociación Geológica Argentina, v. 56, p. 1624.
Cocks, L.R.M., and Rong, J.Y., 1989, Classification and review of the brachiopod superfamily Plectambonitacea: Bulletin of the British Museum of Natural History (Geology), v. 45, p. 77163.
Cocks, L.R.M., and Rong, J.Y., 2000, Strophomenida, in Kaesler, R.L., ed., Treatise on Invertebrate Paleontology, Part H (Revised), Brachiopoda 2: Boulder, Colorado, and Lawrence, Kansas, Geological Society of America (and University of Kansas Press), p. H216H349.
Cocks, L.R.M., and Torsvik, T.H., 2002, Earth geography from 500 to 400 million years ago: A faunal and palaeomagnetic review: Journal of the Geological Society London, v. 159, p. 631644. doi: 10.1144/0016-764901-118.
Congreve, C.R., Krug, A.Z., and Patzkowsky, M.E., 2015, Phylogenetic revision of the Strophomenida, a diverse and ecologically important Palaeozoic brachiopod order: Palaeontology, v. 58, p. 745758. doi: 10.1111/pala.12177.
Cooper, G.A., 1956, Chazyan and related brachiopods: Smithsonian Miscellaneous Collections, v. 127, p. 11245.
Dahlquist, J.A., Pankhurst, R.J., Rapela, C.W., Galindo, C., Casquet, C., Fanning, C.M., Saavedra, J., Baldo, E., and González-Casado, J., 2005, New SHRIMP ages in the Sierra de Famatina, NW Argentina: Implications for the Famatinian Orogen, in Pankhurst, R.J., and Veiga, G.D., eds., Gondwana 12, Geological and Biological Heritage of Gondwana, Mendoza, Argentina, Abstracts, p. 123.
Davidson, T., 1868, On the earliest forms of Brachiopoda hitherto discovered in the British Palaeozoic rocks: Geological Magazine, v. 5, p. 303316.
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, v. 78, p. 275286. doi: 10.1666/0022-3360(2004)078 0275:SSAIBO 2.0.C);2.
Eldredge, N., and Gould, S.J., 1972, Punctuated equilibria: An alternative to phyletic gradualism, in Schopf, T.J.M., ed., Models in Paleobiology: San Francisco, California, W.H. Freeman & Company, p. 82115.
Erwin, D.H., 2007, Disparity: Morphological pattern and developmental context: Palaeontology, v. 50, p. 5773. doi: 10.1111/j.1475-4983.2006.00614.x.
Espisúa, E., 1968, El Paleozoico inferior del Río de Las Chacritas, Dpto. de Jáchal, provincia de San Juan, con especial referencia al Silúrico: Revista de la Asociación Geológica Argentina, v. 23, p. 297311.
Fang, Z.J., and Sánchez, T.M., 2012, Origin and early evolution of the Bivalvia, in Seldon, P.A., ed., Treatise [on Invertebrate Paleontology] Online, Part N (Revised), Mollusca 6, Bivalvia 1: Lawrence, Kansas, The Paleontological Institute, 21 p. doi: 10.17161/to.v0i0.4275.
Finney, S.C., 2007, The parautochthonous Gondwanan origin of the Cuyania (greater Precordillera) terrane of Argentina: A reevaluation of evidence used to support an allochthonous Laurentian origin: Geologica Acta, v. 5, p. 127158.
Goloboff, P.A., and Catalano, S.A., 2016, TNT version 1.5, including a full implementation of phylogenetic morphometrics: Cladistics, v. 32, p. 221238. doi: 10.1111/cla.12160.
Gómez, F.J., and Astini, R.A., 2015, Sedimentology and sequence stratigraphy from a mixed (carbonate-siliciclastic) rift to passive margin transition: The Early to Middle Cambrian of the Argentine Precordillera: Sedimentary Geology, v. 316, p. 3961. doi: 10.1016/j.sedgeo.2014.11.003.
Gould, S.J., and Eldredge, N., 1977, Punctuated equilibria: The tempo and mode of evolution reconsidered: Paleobiology, v. 3, p. 115151.
Gutiérrez-Marco, J.C., and Villas, E., 2007, Brachiopods from the uppermost Lower Ordovician of Peru and their palaeogeographical significance: Acta Palaeontologica Polonica, v. 52, p. 547562.
Gutiérrez-Marco, J.C., Albanesi, G.L., Sarmiento, G.N., and Carlotto, V., 2008, An Early Ordovician (Floian) conodont fauna from the eastern Cordillera of Peru (Central Andean Basin): Geologica Acta, v. 6, p. 147160.
Hall, J., and Clarke, J.M., 1892, An introduction to the study of the Brachiopoda: New York Geological Survey, v. 9, p. 1367.
Harper, D.A.T., MacNiocaill, C., and Williams, S.H., 1996, The palaeogeography of Early Ordovician Iapetus terranes: An integration of faunal and palaeomagnetic constraint: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 121, p. 297312.
Harper, D.A.T., Cocks, L.R.M., Popov, L.E., Sheehan, P.M., Bassett, M.G., Copper, P., Holmer, L.E., Jin, J., and Rong, J.Y., 2004, Brachiopods, in Webby, B.D., Droser, M., Paris, F., and Percival, I.G., eds., The Great Ordovician Biodiversification Event: New York, Columbia University Press, p. 157178.
Harper, D.A.T., Owen, A.W., and Bruton, D.L., 2009, Ordovician life around Celtic fringes: Diversifications, extintions and migrations of brachiopod and trilobite faunas at middle latitudes, in Bassett, M.G., ed., Early Palaeozoic Peri-Gondwanan Terranes: New Insights from Tectonics and Biogeography: Geological Society of London Special Publication, v. 325, p. 157170.
Harper, D.A.T., Rasmussen, C.M.Ø., Liljeroth, M., Blodgett, R.B., Candela, Y., Jin, J., Percival, I.G., Rong, J.Y., Villas, E., and Zhan, R., 2013, Biodiversity, biogeography and phylogeography of Ordovician rhynchonelliform brachiopods: The Geological Society London Memoirs, v. 38, p. 127144. doi: 10.1144/M38.11.
Harper, D.A.T., Popov, L.E., and Holmer, L.E., 2017, Brachiopods: Origin and early history: Palaeontology, v. 60, p. 609631. doi: 10.1111/pala.12307.
Harrington, H.J., and Leanza, A.F., 1957, Ordovician trilobites of Argentina: University of Kansas Department of Geology Special Publication, v. 1, 276 p.
Havlíček, V., and Branisa, L., 1980, Ordovician brachiopods of Bolivia: Rozpravy Ceskolovenské Akademie Věd, v. 90, p. 153.
Havliček, V., and Josopait, V., 1972, Articulate brachiopods from the Iberian Chais, northeast Spain (middle Cambrian-upper Cambrian-Tremadoc): Neues Jahrbuch für Geologie und Palaöntologie, v. 140, p. 328353.
Heredia, S.E., Beresi, M., and Peralta, S., 2011, Estratigrafía y bioestratigrafía del Ordovícico Medio del río Las Chacritas, Precordillera Central de San Juan: INSUGEO Serie Correlación Geológica, v. 27, p. 1826.
Herrera, Z.A., and Benedetto, J.L., 1987, El género Reinversella (Brachiopoda) en el Ordovícico temprano de la Precordillera Argentina: Actas 10th Congreso Geológico Argentino, San Miguel de Tucumán, Argentina, v. 3, p. 7780.
Herrera, Z.A., and Benedetto., J.L., 1991, Early Ordovician brachiopod faunas from the Precordillera basin, western Argentina: Biostratigraphy and paleobiogeographical affinities, in Mackinnon, D.I., Lee, D.E., and Campbell, J.D, eds., Brachiopods through Time: Rotterdam, The Netherlands, A.A. Balkema, p. 283301.
Hessland, I., 1949, Investigations of the Lower Ordovician of the Siljan District, Sweden: Notes on Swedish Ahtiella species: Bulletin of Geology, v. 33, p. 511527.
Jones, O.T., 1928, Plectambonites and some allied genera: Memoirs of the Geological Survey of Great Britain, Palaeontology, v. 1, p. 367527.
Keller, M., 1999, Argentine Precordillera: Sedimentary and plate tectonic history of a Laurentian crustal fragment in South America: Geological Society of America Special Paper 341, p. 1131.
Keller, M., 2012, The Argentine Precordillera: A little American carbonate bank, in Derby, J.R., Fritz, R.D., Longacre, S.A., Morgan, W.A., and Sternbach, C.A., eds., The Great American Carbonate Bank: The geology and economic resources of the Cambrian–Ordovician sauk megasequence of Laurentia: AAPG Memoir, v. 98, p. 9851000.
Kozlowski, R., 1927, Sur certains Orthides ordoviciens des environs de St. Petersburg: Bibliotheca Universitatis Libirae Polonae, v. 17, p. 321.
Lamansky, V.V., 1905, Die ältesten silurischen Schichten Russlands: Comité Géologique Mémoires St. Petersburg, n. ser, v. 20, p. 1203.
Laurie, J.R., 1991, Articulate brachiopods from the Ordovician and lower Silurian of Tasmania, in Jell, P.A., ed., Australian Ordovician Brachiopod Studies: Association of Australasian Paleontologists Memoir, v. 11, p. 1106.
Lavié, F.J., and Benedetto, J.L., 2016, Middle Ordovician (Darriwilian) linguliform and craniiform brachiopods from the Precordillera (Cuyania terrane) of west-central Argentina: Journal of Paleontology, v. 90, p. 10681080. doi: 10.1017/jpa.2016.111.
Lehnert, O.T., Bergström, S.M., and Vaccari, N.E., 1997, Arenig conodonts from the Famatina Range, northwestern Argentina: Faunal affinities and paleogeographic implications: 18th IAS Regional European Meeting of Sedimentology, Heidelberg, Germany, Gaea Heidelbergensis, Abstracts, v. 3, p. 216217.
MacArthur, R.H., and Wilson, E.O., 1967, The Theory of Island Biogeography: New York, Princeton University Press, 203 p.
Mángano, M.G., and Buatois, L.A., 1996, Shallow marine event sedimentation in a volcanic arc-related setting: The Ordovician Suri Formation, Famatina Range, northwest Argentina: Sedimentary Geology, v. 105, p. 6390.
Mángano, M.G., and Buatois, L.A., 1997, Slope apron deposition in an arc-related setting: The Vuelta de las Tolas Member (Suri Formation), Famatina Basin, northwest Argentina: Sedimentary Geology, v. 109, p. 155180.
Mángano, M.G., Astini, R.A., Buatois, L.A., and Dávila, D.F., 2002, The Ordovician System in the Famatina Belt: Depositional and tectonic evolution, in Aceñolaza, F.G., ed., Aspects of the Ordovician System in Argentina: INSUGEO, Serie Correlación Geológica, v. 16, p. 295312.
Mestre, A., and Heredia, S.E., 2012, Darriwilian species of Histiodella (Conodonta) in the Argentine Precordillera: Alcheringa, v. 36, p. 141150. doi: 10.1080/03115518.2011.593334.
Mestre, A., and Heredia, S.E., 2013, La Zona de Yangtzeplacognathus crassus (Conodonta), Darriwiliano de la Precordillera Central, San Juan, Argentina: Ameghiniana, v. 50, p. 407417. doi: 10.5710/AMGH.15.05.2013.558.
Muñoz, D.F., and Benedetto, J.L., 2016, The eoorthid brachiopod Apheoorthina in the Lower Ordovician of NW Argentina and the dispersal pathways along western Gondwana: Acta Palaeontologica Polonica, v. 61, p. 633644. doi: 10.4202/app.00241.2016.
Neuman, R.B., 1971, An early Middle Ordovician brachiopod assemblage from Maine, New Brunswick, and northern Newfoundland, in Dutro, J.T., Jr., ed., Paleozoic Perspectives: A Paleontological Tribute to G. Arthur Cooper: Smithsonian Contributions to Paleobiology, v. 3, p. 113124.
Neuman, R.B., 1976, Early Ordovician (late Arenig) brachiopods from Virgin Arm, New World Island, Newfoundland: Geological Survey of Canada Bulletin, v. 261, p. 1161.
Neuman, R.B., 1984, Geology and paleobiology of islands in Iapetus Ocean: Review and implications: Geological Society of America Bulletin, v. 95, p. 1181201.
Neuman, R.B., and Bates, D.E.B., 1978, Reassessment of Arenig and Llanvirn age (Early Ordovician) brachiopods from Anglesey, north-west Wales: Palaeontology, v. 21, p. 571613.
Neuman, R.B., and Harper, D.A.T., 1992, Paleogeographic significance of Arenig-Llanvirn Toquima−Table Head and Celtic brachiopod assemblage, in Webby, B.D., and Laurie, J.R., eds., Global Perspectives on Ordovician Geology: Rotterdam, The Netherlands, Balkema, p. 241254.
Öpik, A.A., 1930, Brachiopoda Protremata der Estländischen Ordovizischen Kukruse-Stufe: Acta et Commentationes Universitatis Tartuensis, v. A17, p. 1262.
Öpik, A., 1932, Über die Plectellinen: Acta et Commentationes Universitatis Tartuensis, v. A23, p. 185.
Öpik, A., 1933, Über Plectamboniten: Acta et Commentationes Universitatis Tartuensis, v. A24, p. 179.
Öpik, A., 1934, Über Klitamboniten: Acta et Commentationes Universitatis Tartuensis, v. A26, no. 5, p. 1239.
Ortega, G., Albanesi, G.L., and Frigerio, S., 2007, Graptolite-conodont biostratigraphy and biofacies of the Middle Ordovician Cerro Viejo succession, San Juan, Precordillera, Argentina: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 245, p. 245264. doi: 10.1016/j.palaeo.2006.02.023.
Pander, C.H., 1830, Beiträge zur Geognosie des russichen Reiches: St. Petersburg, Gedrukt bei K. Kray, 165 p.
Pankhurst, R.J., Rapela, C.W., and Fanning, C.M., 2001, Age and origin of coeval TTG, I- and S-type granites in the Famatinian belt of NW Argentina: Transactions of the Royal Society of Edinburgh Earth Sciences, v. 91, p. 151168. doi: 10.1017/S0263593300007343.
Peralta, S., Heredia, S.E., and Beresi, M., 1999, Upper Arenig–Lower Llanvirn sequence of the Las Chacritas River, Central Precordillera, San Juan Province, Argentina, in Kraft, P., and Fatka, O., eds., Quo vadis Ordovician?: Acta Universitatis Carolinae Geologica, v. 43, 1/2, p. 123126.
Popov, L.E., Ghobadi Pour, M., Bassett, M.G., and Kebriaee-Zadeh, M., 2009, Billingsellidae and orthidae brachiopods: New insights into earliest Ordovician evolution and biogeography from northern Iran: Palaeontology, v. 52, p. 3552. doi: 10.1111/j.1475-4983.2008.00833.x.
Ramos, V.A., 2004, Cuyania, an exotic block to Gondwana: Review of a historical success and the present problems: Gondwana Research, v. 7, p. 10091026. doi: 10.1016/S1342-937X(05)71081-9.
Sánchez, T.M., 1997, Additional Mollusca (Bivalvia and Rostroconchia) from the Suri Formation, Early Ordovician (Arenig), western Argentina: Journal of Paleontology, v. 71, p. 10461054.
Sánchez, T.M., and Babin, C., 1993, Un insolite mollusque bivalve, Catamarcaia n.g. de l’Arenig (Ordovicien inférieur) d’Argentine: Comptes Rendus de l’Academie des Sciences, v. 316, p. 265271.
Schuchert, C., and Cooper, G.A., 1931, Synopsis of the brachiopod genera of the suborders Orthoidea and Pentameroidea, with notes on the Telotremata: American Journal of Science, ser. 5, v. 22, p. 241255.
Serra, F., Albanesi, G.L., Ortega, G., and Bergström, S.M., 2015, Biostratigraphy and palaeoecology of Middle-Late Ordovician conodont and graptolite faunas of the Las Chacritas River section, Precordillera of San Juan, Argentina: Geological Magazine, v. 153, p. 813829. doi: 10.1017/S0016756814000752.
Severgina, L.G., 1967, [New species and genera of the Ordovician brachiopods from the Sayano-Altai Hill district]: Uchenye Zapinski Tomskogo Gosudartvennogo Universiteta, v. 63, p. 120140. (in Russian).
Sorrentino, L., Benedetto, J.L., and Carrera, M.G., 2009, Diversidad taxonómica y distribución de morfotipos de braquiópodos en la Zona de Ahtiella argentina (Ordovícico Medio), Formación San Juan, Precordillera Argentina: Ameghiniana, v. 46, p. 241253.
Spjeldnaes, N., 1957, The Middle Ordovician of the Oslo region, Norway 8, Brachiopoda of the suborder Strophomenoidea: Norsk Geologisk Tidsskrift, v. 37, p. 1214.
Suyarkova, A.A., and Koren, T.N., 2009, Advances in the Ordovician graptolite biostratigraphy of the St. Petersburg area, Russia: Estonian Journal of Earth Sciences, v. 58, p. 39. doi: 10.3176/earth.2009.1.01.
Thomas, W.A., and Astini, R.A., 1996, The Argentine Precordillera: A traveller from the Ouachita Embayment of North American Laurentia: Science, v. 273, p. 752757.
Thomas, W.A., and Astini, R.A., 2003, Ordovician accretion of the Argentine Precordillera terrane to Gondwana: A review: Journal of South American Earth Sciences, v. 16, p. 6779. doi: 10.1016/S0895-9811(03)00019-1.
Thomas, W.A., Astini, R.A., Mueller, P.A., Gehrels, G.E., and Wooden, J.L., 2004, Transfer of the Argentine Precordillera terrane from Laurentia: Constraints from detrital-zircon geochronology: Geology, v. 32, p. 965968. doi: 10.1130/G20727.1.
Tolmacheva, T., Egerquist, E., Meidla, T., Vinn, O., and Holmer, L.E., 2003, Faunal composition and dynamics in the Middle Ordovician of the East Baltic: Geological Magazine, v. 140, p. 3144. doi: 10.1017/S001675680200701X.
Topper, T.P., Harper, D.A.T., and Brock, G.A., 2013, Ancestral billingsellides and the evolution and phylogenetic relationships of early rhynchonelliform brachiopods: Journal of Systematic Palaeontology, v. 11, p. 821833. doi: 10.1080/14772019.2012.728253.
Toro, B.A., and Brussa, E.D., 1997, Graptolitos de la Formación Suri (Arenig) en el Sistema de Famatina, Argentina: Revista Española de Paleontología, v. 12, p. 175188.
Ulrich, O., and Cooper, G.A., 1936, New genera and species of Ozarkian and Canadian brachiopods: Journal of Paleontology, v. 10, p. 616631.
Vaccari, N.E., 1995, Early Ordovician trilobite biogeography of Precordillera and Famatina, western Argentina: Preliminary results, in Cooper, J.D., Droser, M.L., and Finney, S.C., eds., Ordovician Odyssey: Short Papers for the Seventh International Symposium on the Ordovician System, Las Vegas, Nevada: Fullerton, California, Pacific Section Society for Sedimentary Geology, v. 77, p. 193–196.
Vaccari, N.E., and Waisfeld, B.G., 1994, Nuevos trilobites de la Formación Suri (Ordovícico temprano) en la región de Chaschuil, Provincia de Catamarca: Implicancias bioestratigráficas: Ameghiniana, v. 31, p. 7386.
Waisfeld, B.G., and Vaccari, N.E., 2003, Trilobites, in Benedetto, J.L., ed., Ordovician Fossils of Argentina: Córdoba, Spain, Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba, p. 295409.
Waisfeld, B.G., Vaccari, N.E., Chatterton, B.D.E., and Edgecombe, G.D., 2001, Systematics of Shumardiidae (Trilobita), with new species from the Ordovician of Argentina: Journal of Paleontology, v. 75, p. 827859. doi: 10.1666/0022-3360(2001)075 0827:SOSTWN 2.0.CO;2.
Waisfeld, B.G., Sánchez, T.M., Benedetto, J.L., and Carrera, M.G., 2003, Early Ordovician (Arenig) faunal assemblages from western Argentina: Biodiversification trends and paleogeographic constraints: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 196, p. 343373. doi: 10.1016/S0031-0182(03)00464-4.
Webby, B.D., 1992, Ordovician island biotas: New South Wales record and global implications: Journal and Proceedings of the Royal Society of New South Wales, v. 124, p. 5177.
Wiley, E O., 1979, Annotated Linnaean Hierarchy, with comments on natural taxa and competing systems: Systematic Zoology, v. 28, p. 308337.
Williams, A., and Curry, G.B., 1985, Lower Ordovician Brachiopoda from the Tourmakeady Limestone, Co. Mayo, Ireland: Bulletin of the British Museum (Natural History) Geology Series, v. 38, p. 183269.
Williams, A., and Hurst, J.M., 1977, Brachiopod evolution, in Hallam, A., ed., Patterns of Evolution: Amsterdam, Elsevier, p. 79121.
Williams, A., Carlson, S.J., Brunton, C.H.C., Holmer, L.E., and Popov, L.E., 1996, A supra-ordinal classification of the Brachiopoda: Philosophical Transactions of the Royal Society of London, ser. B, v. 351, p. 11711193.
Xu, H.K., and liu, d.y., 1984, Late Early Ordovician brachiopods of southwestern China: Bulletin Nanjing Institute of Geology and Palaeontology, Academia Sinica, v. 8, p. 217235.
Zalasiewicz, J.A., Taylor, L., Rushton, A.W.A., Loydell, D.K., Rickards, R.B., and Williams, M., 2009, Graptolites in British stratigraphy: Geological Magazine, v. 146, p. 785850. doi: 10.1017/S0016756809990434.
Zhan, R.B., Jin, J., and Rong, J.Y., 2006, β-diversity fluctuations in Early–Mid Ordovician brachiopod communities of South China: Geological Journal, v. 41, p. 271288. doi: 10.1002/gj.1040.
Zhan, R.B., Jin, J., Rong, J.Y, and Liang, Y., 2015, The earliest known strophomenoids (Brachiopoda) from early Middle Ordovician rocks of South China: Palaeontology, v. 56, p. 11211148. doi: 10.1111/pala.12039.
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