Middle Ordovician harknessellid brachiopods (Dalmanellidina) from the Mediterranean margin of Gondwana


Authors: Reyes-Abril J, Villas E, Gutiérrez-Marco JC, Jiménez-Sánchez A, Colmenar J

Published in: Bulletin of Geosciences, volume 88, issue 4; pages: 813 - 828; Received 18 March 2013; Accepted in revised form 2 July 2013; Online 12 September 2013

Keywords: Ordovician, brachiopods, Orthida, Iberian Peninsula, Spain, Gondwana,

full text (PDF, 5.21 MB)

Export to RIS



The family Harknessellidae Bancroft, 1928 (Orthida, Dalmanellidina) was designed to embrace an assemblage of species referred previously to Harknessella Reed, 1917, and included five genera known mainly from the Middle and Upper Ordovician of England. Herein, we suggest reassigning to this family the genus Cacemia Mitchell, 1974, widespread in the middle Darriwilian (upper Middle Ordovician) of the Iberian and Armorican massifs. Since its designation, Cacemia was placed among the dalmanellidin heterorthids, in spite of its strongly mucronate hinge line, which is totally unknown within this Mediterranean family. A new harknessellid has been identified from the upper Darriwilian beds of the Central Iberian Zone (Central Spain): Isabelella fascicostellata Reyes-Abril   Villas gen. et sp. nov. It is similar to Horderleyella Bancroft, 1928 for its coarsely fascicostellate radial ornamentation and obtuse cardinal angles, although its convexoplane to convexoconcave profile allows discrimination from the typically dorsibiconvex Horderleyella. A phylogenetic analysis of the family places both Cacemia and Isabelella in basal positions of their clades, which fits with their early stratigraphic record. Based on our study, the family Harknessellidae appears to have originated in the high latitude Mediterranean margins of Gondwana during pre-Darriwilian times, before the detachment of Avalonia from Gondwana. The family reached its highest diversification in Avalonia throughout the Late Ordovician, keeping connections with the Mediterranean and Proto-Andean margins of Gondwana, as well as with the mid-latitude palaeocontinents of Baltica and South China.


Baarli, B.G. 1998. The Llandovery enteletacean brachiopods of the central Oslo region, Norway. Palaeontology 31, 1101-1129.

Babin, C. & Gutiérrez-Marco, J.C. 1991. Middle Ordovician bivalves from Spain and their phyletic and palaeogeographic significance. Palaeontology 34, 109-147.

Bancroft, B.B. 1928. The Harknessellinae. Memoirs and Proceedings of the Manchester Literary and Philosophical Society 72, 173-196.

Bancroft, B.B. 1945. The brachiopod zonal indices of the stages Costonian to Onnian in Britain. Journal of Paleontology 19, 181-252.

Benedetto, J.L. 2003. Brachiopods, 187-272. In Benedetto, J.L. (ed.) Ordovician Fossils of Argentina. Secretaría de Ciencia y Tecnología, Universidad Nacional de Córdoba.

Bergström, S.M., Chen, X., Gutiérrez-Marco, J.C. & Dronov, A.V. 2009. The new chronostratigraphic classification of the Ordovician System and its relations to major regional series and stages and ?13C chemostratigraphy. Lethaia 42, 97-107.View article

Bogoyavlenskaya, V.M. 1991. Brachiopods, 74-105. In Puchkov, V.N. (ed.) Ordovician of the Polar Urals. Palaeontology. Uralskoye otdeleniye Akademii nauk SSSR, Sverdlovsk.

Born, A. 1918. Die Calymene tristani-Stufe (mittleres Untersilur) bei Almaden, ihre Fauna, Gliederung und Verbreitung. Abhandlungen der senckenbergischen naturforschenden Gesellschaft 36 (for 1916), 309-358.

Cooper, A.H. 1980. The stratigraphy and palaeontology of the Ordovician to Devonian rocks of the area north of Dornes (near Figueiró dos Vinhos), central Portugal. 430 pp. Unpublished PhD thesis, University of Sheffield.

Cooper, G.A. 1956. Chazyan and related brachiopods. Smithsonian Miscellaneous Collections 127(1-2), 1-1245.

Cooper, G.A. & Muir-Wood, H.M. 1951. Brachiopod homonyms. Journal of the Washington Academy of Sciences 41, 195-196.

Delgado, J.F.N. 1897. Fauna Silúrica de Portugal. Novas observaçoes ácerca de Lichas (Uralichas) Ribeiroi. Memórias da Direcçao dos Trabalhos Geológicos de Portugal, 1-34.

Delgado, J.F.N. 1908. Systeme Silurique du Portugal. Étude de stratigraphie paléontologique. Memórias e Comunicaçoes dos Serviços Geológicos de Portugal, 1-245.

Gutiérrez-Marco, J.C., Aramburu, C., Arbizu, M., Bernárdez, E., Hacar Rodríguez, M.P., Méndez-Bedia, I., Montesinos López, R., Rábano, I., Truyols, J. & Villas, E. 1999. Revisión bioestratigráfica de las pizarras del Ordovícico Medio en el noroeste de Espana (Zonas Cantábrica, Asturoccidental-leonesa y Centroibérica septentrional). Acta Geologica Hispanica 34, 3-87.

Gutiérrez-Marco, J.C., Rábano, I., Prieto, M. & Martín, J. 1984. Estudio bioestratigráfico del Llanvirn y Llandeilo (Dobrotiviense) en la parte meridional de la zona Centroibérica (Espana). Cuadernos de Geología Ibérica 9, 287-319.

Gutiérrez-Marco, J.C., Robardet, M., Rábano, I., Sarmiento, G.N., San José Lancha, M.A., Herranz Araújo, P. & Pieren Pidal, A.P. 2002. Ordovician, 31-49. In Gibbons, W. & Moreno, T. (eds) The Geology of Spain. The Geological Society, London.

Hammann, W., Robardet, M. & Romano, M. 1982. The Ordovician System in southwestern Europe (France, Spain and Portugal). Correlation Chart and Explanatory Notes. International Union of Geological Sciences, Publication 11, 1-47.

Harper, D.A.T. 2000. Dalmanellidina, H782-H844. In Kaesler, R.L. (ed.) Treatise on Invertebrate Paleontology, Part H, Brachiopoda, Revised Volume 3. Geological Society of America & University of Kansas, Boulder & Lawrence.

Harper, D.A.T. 2007. Orthida, H2684-H2698. In Selden, R.L. (ed.) Treatise on Invertebrate Paleontology, Part H, Brachiopoda, Revised Volume 6. Geological Society of America &University of Kansas, Boulder & Lawrence.

Havlíček, V. 1967. Brachiopoda of the Suborder Strophomenidina in Czechoslovakia. Rozpravy Ústředního ústavu geologického 33, 334-235.

Havlíček, V. 1970. Heterorthidae (Brachiopoda) in the Mediterranean Province. Sborník geologických věd, Paleontologie 33, 7-39.

Havlíček, V. 1971. Brachiopodes de l’Ordovicien du Maroc. Notes et Mémoires du Service Géologique 230, 1-135.

Havlíček, V. 1977. Brachiopods of the order Orthida in Czechoslovakia. Rozpravy Ústředního ústavu geologického 44, 1-293.

Havlíček, V. & Branisa, L. 1980. Ordovician brachiopods of Bolivia (Succession of assemblages, climate control, affinity to Anglo-French and Bohemian provinces). Rozpravy Československé akademie věd 90, 1-54.

Havlíček, V., Kříž, J. & Serpagli, E. 1987. Upper Ordovician brachiopod assemblages of the Carnic Alps, middle Carinthia and Sardinia. Bollettino della Societa Paleontologica Italiana 25, 277-311.

Hints, L. & Harper, D.A.T. 2003. Review of the Ordovician rhynchonelliformean Brachiopoda of the East Baltic: Their distribution and biofacies. Bulletin of the Geological Society of Denmark 50, 29-43.

Maddison,W. & Maddison, D. 2003. MacClade, v. 4.06. Sinauer Associates, Sunderland, Massachusetts.

Mélou, M. 1976. Orthida (Brachiopoda) de la Formation de Postolonnec (Ordovicien), Finistere, France. Geobios 9, 693-717.View article

Mélou, M., Oulesbir, L. & Paris, F. 1999. Brachiopodes et chitinozoaires ordoviciens dans le NE du Sahara algérien: implications stratigraphiques et paléogeographiques. Geobios 32, 823-839.View article

Mélou,M. & Pillet, J. 1998. Orthida (Brachiopoda) des Schistes d’Angers (Llanvirn) du flanc nord du bassin de Saint-Julien de Voivantes. Bulletin de la Société d’Etudes scientifiques de l’Anjou 16, 89-97.

Mitchell, W.I. 1974. An outline of the stratigraphy and palaeontology of the Ordovician rocks of Central Portugal. Geological Magazine 111, 385-396.View article

Moore, R.C. 1952. Brachiopoda, 197-267. In Moore, R.C., Lalicker, C.G. & Fischer, A.G. Invertebrate Fossils. McGraw-Hill, New York.

Öpik, A.A. 1933. Über einige Dalmanellacea aus Estland. Universitatis Tartuensis (Dorpatensis) Acta et Commentationes A 25, 1-25.

Rábano, I. 1989. Trilobites del Ordovícico Medio del sector meridional de la zona Centroibérica espanola. Parte I. Yacimientos, bioestratigrafía y aspectos paleobiogeográficos. Boletín Geológico y Minero 100, 307-338.

Reed, F.R.C. 1917. The Ordovician and Silurian brachiopods of the Girvan district. Transactions of the Royal Society of Edinburgh 51, 795-998.View article

Reyes-Abril, J. 2009. Braquiópodos del Ordovícico Medio de la Zona Centroibérica Meridional (Espana). 374 pp. Unpublished PhD thesis, University of Zaragoza.

Reyes-Abril, J., Gutiérrez-Marco, J.C. & Villas, E. 2011. Biostratigraphy of the Middle Ordovician brachiopods from Central Spain, 463-472. In Gutiérrez-Marco, J.C., Rábano, I. & García-Bellido, D. (eds) Ordovician of the World. Serie Cuadernos del Museo Geominero 14, Instituto Geológico y Minero de Espana, Madrid.

Reyes-Abril, J., Villas, E. & Gutiérrez-Marco, J.C. 2010. Orthid brachiopods from the Middle Ordovician of the Central Iberian Zone, Spain. Acta Palaeontologica Polonica 55, 285-308.View article

Ribeiro, C., Sharpe, D., Salter, J.W., Jones, T.R. & Bunbury, C.J.F. 1853. On the Carboniferous and Silurian formations of the neighbourhood of Bussaco in Portugal. Quarterly Journal of the Geological Society of London 9, 135-160.View article

Romao, J.M.C. 2000. Estudo tectono-estratigráfico de um segmento do bordo SW da Zona Centro-Ibérica, e as suas relaçoes com a Zona Ossa-Morena. 323 pp. Unpublished PhD thesis, Universidade de Lisboa.

Schuchert, C. 1913. Class Brachiopoda. The Lower Devonian deposits of Maryland. Maryland Geological Survey, Baltimore, 290-449.

Schuchert, C. & Cooper, G.A. 1931. Synopsis of the brachiopod genera of the suborders Orthoidea and Pentameroidea, with notes on the Telotremata. American Journal of Science 222, 241-251.View article

Schuchert, C. & Cooper, G.A. 1932. Brachiopod genera of the suborders Orthoidea and Pentameroidea. Memoirs of the Peabody Museum of Natural History 4, 270 pp.

Swofford, D.L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts.

Temple, J.T. 1965. Upper Ordovician brachiopods from Poland and Britain. Acta Palaeontologica Polonica 10, 379-427.

Tromelin, G.L. de & Lebesconte, P. 1876. Essai d’un catalogue raisonné des fossiles siluriens des départements de Maine-et-Loire, de la Loire-Inférieure et du Morbihan, avec des observations sur les terrains Paléozoiques de l’Ouest de la France. Comptes Rendus du 4eme Session de la Association Française pour l’Avancement des Sciences, Nantes, 601-661.

Verneuil, E. DE & Barrande, J. 1855. Description des fossiles trouvés dans les terrains Silurien et Dévonien d’Almadén, d’une partie de la Sierra Morena et des Montagnes de Tolede. Bulletin de la Société Géologique de France (2eme série) 12, 964-1025.

Villas, E. 1992. New Caradoc brachiopods from the Iberian Chains (Northeastern Spain) and their stratigraphic significance. Journal of Paleontology 66, 772-793.

Williams,A. 1949. New Lower Ordovician brachiopods from the Llandeilo-Llangadock District. Geological Magazine 86, 161-174.View article

Williams, A. & Harper, D.A.T. 2000. Orthida, H714-H782. In Kaesler, R.L. (ed.) Treatise on Invertebrate Paleontology, Part H, Brachiopoda; Revised Volume 3. Geological Society of America & University of Kansas, Boulder, Colorado & Lawrence, Kansas.

Zhan, R.-B. & Jin, J. 2005. Brachiopods from the Middle Ordovician Shihtzupu Formation of Yunnan Province, China. Acta Palaeontologica Polonica 50, 365-393.

Zhu, C.-Y. 1985. Late Ordovician Brachiopoda from Wulongtun district of eastern Da Hinggan Ling region, northeast China. Bulletin of the Shengyang Institute of Geology and Mineral Resources, Chinese Academy of Sciences 12, 24-54. [in Chinese, English abstract]