New deep-water brachiopod resilient assemblage from the South-Iberian Palaeomargin (Western Tethys) and its significance for the brachiopod adaptive strategies around the Early Toarcian Mass Extinction Event


Authors: Baeza-Carratalá JF, Reolid M, García Joral F

Published in: Bulletin of Geosciences, volume 92, issue 2; pages: 233 - 256; Received 9 September 2016; Accepted in revised form 16 May 2017; Online 30 June 2017

Keywords: Jurassic, Brachiopods, Western Tethys, Early Toarcian Mass Extinction, adaptive strategies,

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The Pliensbachian-Toarcian transition was a period of changes in long-term environmental conditions leading up to the Early Toarcian Mass Extinction Event (ETMEE), which resulted in a noticeable extinction and turnover in the marine biota. The westernmost Tethyan basins, especially the peri-Iberian platforms, provide an exceptional brachiopod record to better understand the adaptive strategies and the severe ecological effects of these faunas within the marine ecosystems. This event marks a critical interval in the evolutionary history of the Phylum Brachiopoda as two orders, the Athyridida and Spiriferinida, became extinct. Evolutionary patterns displayed by several taxa from these groups and some rhynchonellids typifying deep-water habitats are analyzed across this biotic crisis spanning several Mediterranean and NW-European basins. New work performed in La Cerradura section, a deep pelagic trough from the South-Iberian palaeomargin, reveals two new taxa (Koninckodonta sumuntanensis and Atychorhynchia falsiorigo) herein described. This newly documented fauna supports pre-extinction dwarfing and resilience in deep refugia linked to the ETMEE, and an episode of speciation which is interpreted in terms of a pre-extinction radiation. In the ETMEE repopulation phase an opportunistic strategy occurs typified by Soaresirhynchia bouchardi, and a case of homoplasy involving post-extinction pioneers (Elvis taxon) is detected. Similar adaptive strategies occurred associated with other mass extinctions such as the Permian/Triassic and the Cretaceous/Paleocene events, supporting a possible standard pattern in the response of the brachiopod fauna to such biotic crises and shedding light on the ecological effects of the mass extinction events.


ABERHAN, M. 2001. Bivalve palaeobiogeography and the Hispanic Corridor: time of opening and effectiveness of a proto-Atlantic seaway. Palaeogeography, Palaeoclimatology, Palaeoecology 165, 375-394.View article

ABERHAN, M. & FÜRSICH, F.T. 1997. Diversity analysis of Lower Jurassic bivalves of the Andean Basin and the Pliensbachian-Toarcian mass extinction. Lethaia 29, 181-195.View article

AGER, D.V. 1959. The classification of the Mesozoic Rhynchonelloidea. Journal of Paleontology 33(2), 324-332. AGER, D.V. 1967. Some Mesozoic Brachiopods in the Tethys region, 135-151. In ADAMS, C.G. & AGER, D.V. (eds) Aspects of Tethyan Biogeography. Systematics Association Publication 7. AGER, D.V. 1990. British Liassic Terebratulida (Brachiopoda) Part I. Monograph of the Palaeontographical Society London 582, 143, 1-39.

ALMÉRAS, Y. & ELMI, S. 1993. Palaeogeography, physiography, palaeoenvironments and brachiopod communities. Example of the Liassic brachiopods in the Western Tethys. Palaeogeography, Palaeoclimatology, Palaeoecology 100, 95-108.View article

ALMÉRAS, Y. & FAURÉ, P. 2000. Les Brachiopodes liasiques des Pyrénées. Paléontologie, Biostratigraphie, Paléobiogéographie et Paléoenvironnements. Strata 36, 1-395.

ALMÉRAS, Y., ELMI, S. & FAURÉ, P. 2007. Les Brachiopodes Liasiques d’Algerie Occidentale. Documents des Laboratoires e Géologie Lyon 163, 3-241.

ALMÉRAS, Y., ELMI, S., MOUTERDE, R., RUGET, C. & ROCHA, R. 1988. Evolution Paléogéographique du Toarcien et influence sur les peuplements, 687-698. In ROCHA, R.B. & SOARES, A.F. (eds) International Symposium on Jurassic Stratigraphy. Lisboa.

AL-SUWAIDI, A.H., ANGELOZZI, G.N., BAUDIN, F., DAMBORENEA, S.E., HESSELBO, S.P., JENKYNS, H.C., MANCENIDO, M.O. & RICCARDI, A.C. 2010. First record of the Early Toarcian Oceanic Anoxic Event from the Southern Hemisphere, Neuquén Basin, Argentina. Journal of the Geological Society of London 167, 633-636.View article

AL-SUWAIDI, A.H., HESSELBO, S.P., DAMBORENEA, S.E., MANCENIDO, M.O., JENKYNS, H.C., RICCARDI, A.C., ANGELOZZI, G.N. & BAUDIN, F. 2016. The Toarcian Oceanic Anoxic Event (Early Jurassic) in the Neuquén Basin, Argentina: A Reassessment of Age and Carbon Isotope Stratigraphy. The Journal of Geology 124, 171-193.View article

ANDRADE, B., DUARTE, L.V., GARCÍA JORAL, F., GOY, A. & HENRIQUES, M.H. 2016. Palaeobiogeographic patterns of the brachiopod assemblages of the Iberian Subplate during the Late Toarcian-Early Aalenian (Jurassic). Palaeogeography, Palaeoclimatology, Palaeoecology 442, 12-22.View article

ARIAS, C. 2009. Extinction pattern of marine Ostracoda across the Pliensbachian Toarcian boundary in the Cordillera Ibérica, NE Spain: causes and consequences. Geobios 42, 1-15.View article

ARIAS, C. 2013. The Early Toarcian (Early Jurassic) ostracod extinction events in the Iberian Range: The effect of temperature changes and prolonged exposure to low dissolved oxygen concentrations. Palaeogeography, Palaeoclimatology, Palaeoecology 387, 40-55.View article

BAEZA-CARRATALÁ, J.F. 2008. Patrimonio paleontológico en la colección Jiménez de Cisneros y su aplicación al estudio de los braquiópodos del Jurásico Inferior en la Cordillera Bética Oriental (provincias Alicante y Norte de Murcia). 906 pp. Ph.D. thesis, Universidad Alicante, Spain.

BAEZA-CARRATALÁ, J.F. 2013. Diversity patterns of Early Jurassic brachiopod assemblages from the westernmost Tethys (Eastern Subbetic). Palaeogeography, Palaeoclimatology, Palaeoecology 381-382, 76-91.View article

BAEZA-CARRATALÁ, J.F., GARCÍA JORAL, F., GIANNETTI, A. & TENT-MANCLÚS, J.E. 2015. Evolution of the last koninckinids (Athyridida, Koninckinidae), a precursor signal of the Early Toarcian mass extinction event in the Western Tethys. Palaeogeography, Palaeoclimatology, Palaeoecology 429, 41-56.View article

BAEZA-CARRATALÁ, J.F., GARCÍA JORAL, F. & TENT-MANCLÚS, J.E. 2011. Biostratigraphy and palaeobiogeographic affinities of the Jurassic brachiopod assemblages from Sierra Espuna (Maláguide Complex, Internal Betic Zones, Spain). Journal of Iberian Geology 37(2), 137-151.View article

BAILEY, T.R., ROSENTHAL, Y., MCARTHUR, J.M., VAN DE SCHOOTBRUGGE, B. & THIRLWALL, M.F. 2003. Paleoceanographic changes of the Late Pliensbachian-Early Toarcian interval: a possible link to the genesis of an Oceanic Anoxic Event. Earth and Planetary Science Letters 212, 307-320.View article

BASSOULLET, J.P., ELMI, S., POISSON, A., RICOU, L.E., CECCA, F., BELLIÓN, Y., GIRAUD, R. & BAUDIN, F., 1993. Middle Toarcian (184-182 Ma.). In DECOURT, J., RICOU, L. E. & VRIELYNCK, B. (eds) Atlas Tethys Palaeoenviromental Maps. Beicip-Franlab, Rueil-Malmaison.

BITTNER, A. 1893. Neue Koninckiniden des alpine Lias. Jahrbuch der kaiserlich-königlichen geologischen Reinchsalstat 43(1), 133-144.

BOUCOT, A.J., JOHNSON, J.G. & STATON, R.D. 1964. On some atrypoid, retzioid, and athyridoid Brachiopoda. Journal of Paleontology 38, 805-822. BRAGA, J.C. 1983. Ammonites del Domerense de la zona Subbética (Cordilleras Béticas, S. de Espana). 410 pp. Ph.D. thesis, Universidad Granada, Spain.

BRAGA, J.C., COMAS, M.C., DELGADO, F., GARCÍA-HERNÁNDEZ, M., JIMÉNEZ, A.P., LINARES, A., RIVAS, P. & VERA, J.A. 1981. The Liassic rosso ammonitico facies in the Subbetic Zone (Spain). Genetic consideration, 61-76. In FARINACCI, A. & ELMI, S. (eds) Rosso Ammonitico Symposium Proceedings. Tecnoscienza, Rome.

BUCEFALO PALLIANI, R., MATTIOLI, E. & RIDING, J.B. 2002. The response of marine phytoplankton and sedimentary organic matter to the Early Toarcian (Lower Jurassic) oceanic anoxic event in northern England. Marine Micropaleontology 46, 223-245.View article

CANIÇO, A., DUARTE, L.V., ROCHA, F., REOLID, M. & TERROSO, D. 2015. Palaeoenvironmental meaning of clay minerals assemblages across the late Pliensbachian-Early Toarcian (Early Jurassic) of Iberian Peninsula: Lusitanian, Algarve and Subbetic basins. Abstracts Euroclay 2015, p. 115.

CARIOU, E. & HANTZPERGUE, P. 1997. Groupe Français d’Études du Jurassique, biostratigraphie du Jurassique ouest-européen et méditerranéen. Zonations paralleles et distribution des invertébrés et microfossiles. Bulletin des Centres de Recherches Exploration-Production d’Elf-Aquitaine 17, 1-400.

CARUTHERS, A.H., GROCKE, D.R. & SMITH, P.L. 2011. The significance of an Early Jurassic (Toarcian) carbon-isotope excursion in Haida Gwaii (Quenn Charlotte Islands), British Columbia, Canada. Earth and Planetary Science Letters 307, 19-26.View article

CARUTHERS, A.H., SMITH, P.L. & GRÖCKE, D.R. 2013. The Pliensbachian-Toarcian (Early Jurassic) extinction, a global multi-phased event. Palaeogeography, Palaeoclimatology, Palaeoecology 386, 104-118.View article

CASELLATO, C.E. & ERBA, E. 2015. Calcareous nannofossil biostratigraphy and paleoceanography or the Toarcian Oceanic Anoxic Event at Colle di Sogno (Southern Alps, Northern Italy). Rivista Italiana di Paleontologia e Stratigrafia 121, 297-327.

CECCA, F. & MACCHIONI, F. 2004. The two Early Toarcian (Early Jurassic) extinction events in ammonoids. Lethaia 37, 35-56.View article

CHOFFAT, P. 1947. Description de la faune jurassique du Portugal. Brachiopodes. Mémoires du Service Géologique du Portugal, 1-46.

COMAS-RENGIFO, M.J., DUARTE, L.V., FÉLIX, F.F., GARCÍA JORAL, F., GOY, A. & ROCHA, R.B. 2015. Latest Pliensbachian-Early Toarcian brachiopod assemblages from the Peniche section (Portugal) and their correlation. Episodes 38, 2-8.

COMAS-RENGIFO, M.J., DUARTE, L.V., GARCÍA JORAL, F. & GOY, A. 2013. Los braquiópodos del Toarciense Inferior (Jurásico) en el área de Rabaçal-Condeixa (Portugal): distribución estratigráfica y paleobiogeografía. Comunicaçoes Geológicas 100(1), 37-42.

COMAS-RENGIFO, M.J., GARCÍA JORAL, F. & GOY, A. 2006. Spiriferinida (Brachiopoda) del Jurásico Inferior del NE y N de Espana: distribución y extinción durante el evento anóxico oceánico del Toarciense Inferior. Boletín de la Real Sociedad Espanola de Historia Natural, Sección Geológica 101, 147-157.

DAVIDSON, T. 1853. British Fossil Brachiopoda, vol. I. Introduction. 136 pp., 9 pl. Palaeontographical Society Monograph, London.

DAVIDSON, T. 1876-1878. A monograph of the British fossil Brachiopoda 4 (2), Triassic-Jurassic Supplement. Palaeontographical Society, London, 73-242.

DERA, G. & DONNADIEU, Y. 2012. Modeling evidences for global warming, Arctic seawater freshening, and sluggish oceanic circulation during the Early Toarcian anoxic event. Paleoceanography 27, PA2211.View article

DERA, G., NEIGE, P., DOMMERGUES, J.L., FARA, E., LAFFONT, R. & PELLENARD, P. 2010. High resolution dynamics of Early Jurassic marine extinctions: the case of Pliensbachian-Toarcian ammonites (Cephalopoda). Journal of the Geological Society of London 167, 21-33.View article

DESLONGCHAMPS, E.E. 1853. Mémoire sur les genres Loptaena et Thecidea des terrains Jurassiques du Calvados. Mémoires de la Société Linnéenne de Normandie 9, 213-250.

DULAI, A. & PÁLFY, J. 2003. The terebratulid brachiopod Lobothyris? subgregaria as an Early Jurassic Elvis species from Hungary. 3rd Workshop of the IGCP Project 458: “Triassic/Jurassic Boundary Changes”, Stará Lesná, Slovakia, 45-46.

DULAI, A., BITNER, M.A. & MÜLLER, P.Á.L. 2008. A monospecific assemblage of a new rhynchonellide brachiopod from the Paleocene of Austria. Fossils and Strata 54, 193-201.

ELMI, S., MAROK, A., SEBANE, A. & ALMÉRAS, Y. 2006. Importance of the Mellala section (Traras Mountains, NW Algeria) for the correlations of the Pliensbachian/Toarcian boundary. Volumina Jurassica 4, 158-160.

ELMI, S., ROULLEAU, L., GABILLY, J. & MOUTERDE, R. 1997. Toarcien. Biostratigraphie du Jurassique ouest-européen et méditerranéen: zonations paralleles et distribution des invertébrés et microfossiles. In CARIOU, E. & HANTZPERGUE, P. (eds) Bulletin des Centres de Recherches Exploration-Production d’Elf-Aquitaine 17, 25-36.

ERWIN, D. & DROSER, M. 1993. Elvis Taxa. Palaios 8, 623-624.View article

GAHR, M.E. 2002. Palökologie des Makrobenthos aus dem Unter-Toarc SW-Europas. Beringeria 31, 3-204. GAHR, M. 2005. Response of Lower Toarcian (Lower Jurassic) macrobenthos of the Iberian Peninsula to sea level changes and mass extinction. Journal of Iberian Geology 31, 197-215.

GARCÍA JORAL, F. & GOY, A. 2000. Stratigraphic distribution of Toarcian brachiopods from the Iberian Range and its relation to depositional sequences. Georesearch Forum 6, 381-386.

GARCÍA JORAL, F., GÓMEZ, J.J. & GOY, A. 2011. Mass extinction and recovery of the Early Toarcian (Early Jurassic) brachiopods linked to climate change in northern and central Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 302, 367-380.View article

GARCÍA-HERNÁNDEZ, M., LÓPEZ-GARRIDO, A.C., MARTÍNALGARRA, A., MOLINA, J.M., RUIZ-ORTIZ, P.A. & VERA, J.A. 1989. Las discontinuidades mayores del Jurásico de las Zonas Externas de las Cordilleras Béticas: Análisis e interpretación de los ciclos sedimentarios. Cuadernos de Geología Ibérica 13, 35-52.

GARCÍA-HERNÁNDEZ, M., LUPIANI, E. & VERA, J.A. 1987. La sedimentación liásica den el sector central del Subbético medio: registro de la evolución de un rift intracontinental. Acta Geológica Hispánica 21-22, 329-337.

GÓMEZ, J.J. & GOY, A. 2011. Warming-driven mass extinction in the Early Toarcian (Early Jurassic) of northern and central Spain. Correlation with other time-equivalent European sections. Palaeogeography, Palaeoclimatology, Palaeoecology 306, 176-195.View article

GÓMEZ, J.J., COMAS-RENGIFO, M.J. & GOY, A. 2016. Palaeoclimatic oscillations in the Pliensbachian (Early Jurassic) of the Asturian Basin (Northern Spain). Climate of the Past 12, 1199-1214.View article

GÓMEZ, J.J., GOY, A. & CANALES, M.L. 2008. Seawater temperature and carbon isotope variations in belemnites linked to mass extinction during the Toarcian (Early Jurassic) in Central and Northern Spain. Comparison with other European sections. Palaeogeography, Palaeoclimatology, Palaeoecology 258, 28-58.View article

GUEX, J. 2016. Retrograde evolution during major extinction crises. 77 pp. Springer, Cham.View article

HALLAM, A. 1986. The Pliensbachian and Tithonian extinction events. Nature 319, 765-768.View article

HALLAM, A. 1987. Radiations and extinctions in relation to environmental change in the marine Lower Jurassic of northwest Europe. Paleobiology 13, 152-168.View article

HALLAM, A. 1990. The end-Triassic mass extinction event. Geological Society of America Special Paper 247, 577-583.View article

HALLAM, A. 1996. Recovery of the marine fauna in Europe after the end-Triassic and early Toarcian mass extinctions, 231-236. In HART, M.B. (ed.) Biotic Recovery from Mass Extinction Events. Geological Society of London, Special Publications 102.

HAMMER, O., HARPER, D.A.T. & RYAN, P.D. 2001. Paleontological statistics software package for education and data analysis: Palaeontologia Electronica 4, 1-9.

HARPER, D.A.T. 1993. Suborders Athyrididina, Retziidina, Dayiidina, Koninckinidina, Thecideidina, 447-450. In BENTON, M.J. (ed.) The fossil record 2. Brachiopoda. Chapman & Hall, London.

HARRIES, P.J. & KNORR, P.O. 2009. What does the ’Lilliput Effect’ mean? Palaeogeography, Palaeoclimatology, Palaeoecology 284, 4-10.View article

HARRIES, P.J. & LITTLE, C.T.S. 1999. The Early Toarcian (Early Jurassic) and the Cenomanian-Turonian (Late Cretaceous) mass extinctions: similarities and contrasts. Palaeogeography, Palaeoclimatology, Palaeoecology 154, 39-66.View article

HARRIES, P.J., KAUFFMAN, E.G. & HANSEN, T.A. 1996. Models for Biotic Survival Following Mass Extinction. Biotic recovery from mass extinction events, 41-60. In HART, M.B. (ed.) Geological Society of London, Special Publication 102.View article

HE, W., SHI, G.R., FENG, Q., CAMPI, M.J., GU, S., BU, J., PENG, Y. & MENG, Y. 2007. Brachiopod miniaturization and its possible causes during the Permian-Triassic crisis in deep water environments, South China. Palaeogeography, Palaeoclimatology, Palaeoecology 252, 132-144.View article

HERMOSO, M., MINOLETTI, F., LE CALLONNEC, L., JENKYNS, H.C., HESSELBO, S.P., RICKABY, R.E.M., RENARD, M., RAFAELIS, M. DE & EMMANUEL, L. 2009. Global and local forcing of Early Toarcian seawater chemistry: A comparative study of different paleoceanographic settings (Paris and Lusitanian basins). Paleoceanography 24, PA4208.View article

HESSELBO, S.P., JENKYNS, H.C., DUARTE, L.V. & OLIVEIRA, L.C.V. 2007. Carbon-isotope record of the Early Jurassic (Toarcian) oceanic anoxic event from fossil wood and marine carbonate (Lusitanian Basin, Portugal). Earth and Planetary Science Letters 253, 455-470.View article

JABLONSKI, D. 2002. Survival without recovery after mass extinctions. Proceedings of the National Academy of Sciences U.S.A. 99, 8139-8144.View article

JENKYNS, H.C. 1988. The Early Toarcian (Jurassic) anoxic event: stratigraphy, sedimentary and geochemical evidence. American Journal of Science 288, 101-151.View article

JENKYNS, H.C. & CLAYTON, C.J. 1997. Lower Jurassic epicontinental carbonates andmudstones from England and Wales: chemostratigraphic signals and the Early Toarcian anoxic event. Sedimentology 44, 687-706.View article

JENKYNS, H.C., GRÖCKE, D.R. & HESSELBO, S.P. 2001. Nitrogen isotope evidence for water mass denitrification during the early Toarcian (Jurassic) oceanic anoxic event. Paleoceanography 16, 593-603.View article

JIMÉNEZ, A.P., 1986. Estudio paleontológico de los ammonites del Toarciense inferior y medio de las Cordilleras Béticas (Dactylioceratidae e Hildoceratidae). 252 pp. Ph.D. thesis, Universidad de Granada, Spain.

JIMÉNEZ, A.P. & RIVAS, P. 1979. El Toarciense en la zona Subbética. Cuadernos de Geología de la Universidad de Granada 10, 397-411.

JIMÉNEZ, A.P., JIMÉNEZ DE CISNEROS, C., RIVAS, P. & VERA, J.A. 1996. The Early Toarcian Anoxic Event in the Westernmost Tethys (Subbetic): Paleogeographic and Paleobiogeographic significance. Journal of Geology 104, 399-416.View article

KAESLER, R.L. & SELDEN, P.A. (eds) 1997-2007. Treatise on Invertebrate Paleontology. Part H, Brachiopoda, Revised, Vols 1-6. Geological Society of America & Paleontological Institute, Boulder, Colorado & Lawrence.

KAUFFMAN, E.G., ERWIN, D.H. 1995. Surviving mass extinctions. Geotimes 14, 14-17.

KAUFFMAN, E.G. & HARRIES, P.J. 1996. The importance of crisis progenitors in recovery from mass extinction, 15-39. In HART, M.B. (ed.) Biotic Recovery from Mass Extinction Events, Geological Society of London, Special Publications 102. KUHN, O. 1949. Lehrbuch der Paläozoologie. 326 pp. E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart.

LATHUILIERE, B. & MARCHAL, D. 2009. Extinction, survival and recovery of corals from the Triassic to Middle Jurassic time. Terra Nova 21, 57-66.View article

LAURIN, B. & GARCÍA JORAL, F. 1990. Miniaturization and heterochrony in Homoeorhynchia meridionalis and H. cynocephala (Brachiopoda, Rhynchonelidae) from the Jurassic of the Iberian Range, Spain. Paleobiology 16, 62-76.View article

LEVINTON, J.S. 1970. The paleoecological significance of opportunistic species. Lethaia 3, 69-78.View article

LINDNER, A., CAIRNS, S.D. & CUNNINGHAM, C.W. 2008. From offshore to onshore: multiple origins of shallow water corals from deep-sea ancestors. PLoS One 3, e2429.View article

LITTLE, C.T.S. 1996. The Pliensbachian-Toarcian (Lower Jurassic) extinction event. Special Paper of the Geological Society of America 307, 505-512.View article

LITTLE, C.T.S. & BENTON, M.J. 1995. Early Jurassic mass extinction: A global long-term event. Geology 23, 495-498.View article

LITTLER, K., HESSELBO, S.P. & JENKYNS, H.C. 2010. A carbon-isotope perturbation at the Pliensbachian-Toarcian boundary: evidence from the Lias Group, NE England. Geological Magazine 147, 181-192.View article

MACCHIONI, F. & CECCA, F. 2002. Biodiversity and biogeography of middle-late liassic ammonoids: implications for the Early Toarcian mass extinction. Geobios 35, 165-175.View article

MANCENIDO, M.O. 2000. A systematic summary of the stratigraphic distribution of Jurassic rhynchonellide genera (brachiopoda). Advances in Jurassic Research 2000, 387-396. In HALL, R.L., SMITH, P.L. (eds) Proceedings of the Fifth International Symposium on the Jurassic System, GeoResearch Forum 6.

MANCENIDO, M.O. 2002. Paleobiogeography of Mesozoic brachiopod faunas from Andean-Patagonian areas in a global context. Geobios 35, 176-192.View article

MARSHALL, L.G. & CORRUCCINI, R.S. 1978. Variability, evolutionary rates, and allometry in dwarfing lineages. Paleobiology 4, 101-118.View article

MATTIOLI, E. & ERBA, E. 1999. Synthesis of calcareous nannofossil events in the Tethyan Lower and Middle Jurassic. Rivista Italiana di Paleontologia e Stratigrafia 105, 343-376.

MATTIOLI, E., PITTET, B., BUCEFALO PALLIANI, R., RÖHL, H.J., SCHMID-RÖHL, A. & MORETTINI, E. 2004. Phytoplankton evidence for the timing and correlation of palaeoceanographical changes during the Early Toarcian oceanic anoxic event (Early Jurassic). Journal of the Geological Society of London 161, 685-693.View article

MATTIOLI, E., PITTET, B., PETITPIERRE, L. & MAILLIOT, S. 2009. Dramatic decrease of the pelagic carbonate production by nannoplankton across the Early Toarcian Anoxic Event (TOAE). Global and Planetary Change 65, 134-145.View article

MATTIOLI, E., PITTET, B., SUAN, G. & MAILLIOT, S. 2008. Calcareous nannoplankton across the Early Toarcian Anoxic Event: implications for paleoceanography within the western Tethys. Paleoceanography 23, PA3208.View article

MATTIOLI, E., PLANCQ, J., BOUSSAHA, M., DUARTE, L.V. & PITTET, B. 2013. Calcareous nannofossil biostratigraphy: new data from the Lower Jurassic of the Lusitanian Basin. Comunicaçoes Geológicas 100, Especial I, 69-76.

MCARTHUR, J.M., ALGEO, T.J., VAN DE SCHOOTBRUGGE, B., LI, Q. & HOWARTH, R.J. 2008. Basinal restriction, black shales, Re-Os dating, and the Early Toarcian (Jurassic) oceanic anoxic event. Paleoceanography 23, PA4217.View article

MCARTHUR, J.M., DONOVAN, D.T., THIRVALL, M.F., FOUKE, B.W. & MATTEY, D. 2000. Strontium isotope of the Early Toarcian (Jurassic) oceanic anoxic event, the duration of ammonite biozones and belemnite palaeotemperatures. Earth and Planetary Science Letters 179, 269-285.View article

MORTEN, S.D. & TWITCHETT, R.J. 2009. Fluctuations in the body size of marine invertebrates through the Pliensbachian-Toarcian extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 284, 29-38.View article

MOUTERDE, R., BUSNARDO, R. & LINARES, A. 1971. Le Domérien dans le Subbetique Central (Andalusie). Données préliminaires. Cuadernos de Geología Ibérica 2, 237-254.

NIKITENKO, B.L. & MICKEY, M.B. 2004. Foraminifera and ostracodes across the Pliensbachian-Toarcian boundary in the Arctic Realm. Geological Society of London, Special Publication 230, 137-173.View article

POZZA, G.C. & BAGAGLIA, A. 2001. A Koninckella fauna recorded from the s.s. type-locality of the Marne di M. Serrone Formation (Umbria, Central Italy, Northern Apennine). 1 Part. Revue de Paléobiologie 20, 19-29.

REOLID, M., CHAKIRI, S. & BEJJAJI, Z. 2013a. Adaptative strategies of the Toarcian benthic foraminiferal assemblages from the Middle Atlas (Morocco): palaeoecological implications. Journal of African Earth Sciences 84, 1-12.View article

REOLID, M., NIETO, L.M. & SÁNCHEZ-ALMAZO, I.M. 2013b. Caracterización geoquímica de facies pobremente oxigenadas en el Toarciense inferior (Jurásico inferior) del Subbético Externo. Revista de la Sociedad Geológica de Espana 26, 69-84.

REOLID, M., MAROK, A. & SEBANE, A. 2014b. Foraminiferal assemblages and geochemistry for interpreting the incidence of Early Toarcian environmental changes in North Gondwana palaeomargin (Traras Mountains, Algeria). Journal of African Earth Sciences 95, 105-122.View article

REOLID, M., MATTIOLI, E., NIETO, L.M. & RODRÍGUEZ-TOVAR, F.J. 2014a. The Early Toarcian Ocanic Anoxic Event in the External Subbetic (Southiberian Palaeomargin, Westernmost Tethys): geochemistry, nannofossils and ichnology. Palaeogeography, Palaeoclimatology, Palaeoecology 411, 79-94.View article

REOLID, M., RIVAS, P. & RODRÍGUEZ-TOVAR, F.J. 2015. Toarcian ammonitico rosso facies from the South Iberian Paleomargin (Betic Cordillera, southern Spain): paleoenvironmental reconstruction. Facies 61, 22.View article

RITA, P., REOLID, M. & DUARTE, L.V. 2016. The incidence of the Late Pliensbachian -Early Toarcian biotic crisis from ecostratigraphy of benthic foraminiferal assemblages: new insights from the Peniche reference section, Portugal. Palaeogeography, Palaeoclimatology, Palaeoecology 454, 267-281.View article

RODRÍGUEZ-TOVAR, F.J. & REOLID, M. 2013. Environmental conditions during the Toarcian Oceanic Anoxic Event (T-OAE) in the westernmost Tethys: influence of the regional context on a global phenomenon. Bulletin of Geosciences 88, 697-712.View article

RODRÍGUEZ-TOVAR, F.J. & UCHMAN, A. 2010. Ichnofabric evi- dence for the lack of bottom anoxia during the Lower Toarcian Oceanic Anoxic Event (T-OAE) in the Fuente de la Vidriera section, Betic Cordillera, Spain. Palaios 25, 576-587.View article

ROLDÁN, F.J., LUPIANI, E., VILLALOBOS, M. & JEREZ-MIR, L. 1991. Mapa y Memoria explicativa de la Hoja 947, Jaén. MAGNA. Instituto Geológico y Minero de Espana.

RUBAN, D.A. 2004. Diversity dynamics of Early-Middle Jurassic brachiopods of Caucasus, and the Pliensbachian-Toarcian mass extinction. Acta Palaeontologica Polonica 49, 275-282.

RUBAN, D.A. 2009. Brachiopod decline preceded the Early Toarcian mass extinction in the Northern Caucasus (northern Neo-Tethys Ocean): A palaeogeographical context. Revue de Paléobiologie 28, 85-92.

RUBAN, D.A., RADULOVIC, B.V., & RADULOVIC, V.J. 2015. Diversity dynamics of Early and Middle Jurassic brachiopods in the Getic and Danubian tectonic units of eastern Serbia. Palaeogeography, Palaeoclimatology, Palaeoecology 425, 97-108.View article

SANDOVAL, J., BILL, M., AGUADO, R., O’DOGHERTY, L., RIVAS, P., MORARD, A. & GUEX, J. 2012. The Toarcian in the Subbetic basin (southern Spain): Bioevents (ammonite and calcareous nannofossils) and carbon-isotope stratigraphy. Palaeogeography, Palaeoclimatology, Palaeoecology 342-343, 40-63.View article

SAVAGE, N.M., MANCENIDO, M.O. & OWEN, E.F. 2002. Order Rhynchonellida. In WILLIAMS, A., BRUNTON, C.H.C., CARLSON, S.J. et al. (eds) Treatise on Invertebrate Palaeontology. Part H, Brachiopoda (Revised), Volume 4, Rhynchonelliformea (part). 807 pp. Geological Society of America and University of Kansas, Boulder, Colorado and Lawrence.

SUAN, G., MATTIOLI, E., PITTET, B., LÉCUYER, C., SUCHÉRASMARX, B., DUARTE, L.V., PHILIPPE, M., REGGIANI, L. & MARTINEAU, F. 2010. Secular environmental precursors to Early Toarcian (Jurassic) extreme climate changes. Earth and Planetary Science Letters 290, 448-458.View article

SUAN, G., PITTET, B., BOUR, I., MATTIOLI, E., DUARTE, L.V. & MAILLIOT, S. 2008. Duration of the Early Toarcian carbon isotope excursion deduced from spectral analysis: consequence for its possible causes. Earth and Planetary Science Letters 267, 666-679.View article

THUY, B., KIEL, S., DULAI, A., GALE, A.S., KROH, A., LORD, A.R., NUMBERGER-THUY, L.D., STÖHR, S. & WISSHAK, M. 2014. First glimpse into Lower Jurassic deep-sea biodiversity: in situ diversification and resilience against extinction. Proceedings of the Royal Society B 281, 20132624.View article

TOMAŠOVÝCH, A. & SIBLÍK, M. 2007. Evaluating compositional turnover of brachiopod communities during the end-Triassic mass extinction (Northern Calcareous Alps): Removal of dominant groups, recovery and community reassembly. Palaeogeography, Palaeoclimatology, Palaeoecology, 244, 170-200.View article

TWITCHETT, R.J. 2001. Incompleteness of the Permian-Triassic fossil record: a consequence of productivity decline? Geological Journal 36, 341-353.View article

TWITCHETT, R.J. 2006. The palaeoclimatology, palaeoecology and palaeoenvironmental analysis of mass extinction events. Palaeogeography, Palaeoclimatology, Palaeoecology 232, 190-213.View article

TWITCHETT, R.J. 2007. The Lilliput effect in the aftermath of the end-Permian extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 252, 132-144.View article

URBANEK, A. 1993. Biotic crises in the history of Upper Silurian graptoloids: a palaeobiological model. Historical Biology 7, 29-50.View article

VERA, J.A. 2001. Evolution of the South Iberian Continental Margin. Mémoires du Muséum National d’Histoire Naturel de Paris 186, 109-143.

VERA, J.A., MARTÍN-ALGARRA, A., SÁNCHEZ-GÓMEZ, M., FORNÓS, J.J. & GELABERT, B. 2004. Cordillera Bética y Baleares, 345-464. In VERA, J.A. (ed.). Geología de Espana, SGE-IGME.

VÖRÖS, A. 1993. Jurassic microplate movements and brachiopod migrations in the western part of the Tethys. Palaeogeography, Palaeoclimatology, Palaeoecology 100, 125-145.View article

VÖRÖS, A. 2002. Victims of the Early Toarcian anoxic event: the radiation and extinction of Jurassic Koninckinidae (Brachiopoda). Lethaia 35, 345-357.View article

VÖRÖS, A. 2003. Early Jurassic Koninckinids (Athyridida, Brachiopoda) from Tivoli near Rome, Italy. Geologica Romana 36, 169-197.

VÖRÖS, A. 2005. The smooth brachiopods of the Mediterranean Jurassic: refugees or invaders? Palaeogeography, Palaeoclimatology, Palaeoecology 223, 222-242.View article

VÖRÖS, A. 2009. The Pliensbachian brachiopods of the Bakony Mountains (Hungary). Geologica Hungarica, series Paleontologica 58, 1-300.

VÖRÖS, A. 2016. Early Jurassic (Pliensbachian) brachiopod biogeography in the Western Tethys: The Euro-Boreal and Mediterranean faunal provinces revised. Palaeogeography, Palaeoclimatology, Palaeoecology 457, 170-185.View article

VÖRÖS, A., KOCSIS, Á.T. & PÁLFY, J. 2016. Demise of the last two spire-bearing brachiopod orders (Spiriferinida and Athyridida) at the Toarcian (Early Jurassic) extinction event. Palaeogeography, Palaeoclimatology, Palaeoecology 457, 233-241.View article

WADE, B.S. & OLSSON, R.K. 2009. Investigation of pre-extinction dwarfing in Cenozoic planktonic foraminifera. Palaeogeography, Palaeoclimatology, Palaeoecology 284, 39-46.View article

WESTERMANN, G.E.G. 2000. Marine faunal realms of the Mesozoic: review and revision under the new guidelines for biogeographic classification and nomenclature. Palaeogeography, Palaeoclimatology, Palaeoecology 163, 49-68.View article

WIGNALL, P.B. & BOND, D.P.G. 2008. The end-Triassic and Early Jurassic mass extinction records in the British Isles. Proceedings of the Geologists’ Association 119, 73-84.View article

WIGNALL, P.B., NEWTON, R.J. & LITTLE, C.T.S. 2005. The timing of paleoenvironmental change and cause-and-effect relationships during the Early Jurassic mass extinction in Europe. American Journal of Science 305, 1014-1032.View article

ZAKHAROV, V.A., SHURYGIN, B.N., IL’INA, V.I. & NIKITENKO, B.L. 2006. Pliensbachian-Toarcian biotic turnover in North Siberia and the Arctic Region. Stratigraphy and Geological Correlation 14, 399-417.View article

ZEZINA, O.N. 2008. Biogeography of the recent brachiopods. Paleontological Journal 42, 830-858.View article

ZHANG, Y., SHI, G.R., HE, W., WU, H., LEI, Y., ZHANG, K., DU, C., YANG, T., YUE, M. & XIAO, Y. 2016. Significant pre-mass extinction animal body-size changes: Evidences from the Permian-Triassic boundary brachiopod faunas of South China. Palaeogeography, Palaeoclimatology, Palaeoecology 448, 85-95.View article