Late Cretaceous continental and marine vertebrate assemblages of the Laño Quarry (Basque-Cantabrian Region, Iberian Peninsula): an update

The vertebrate-bearing beds of the Laño quarry (Condado de Treviño) are among the most relevant sites from the Late Cretaceous of Europe. Geologically, Laño and the adjacent region are set on the southern limb of the South-Cantabrian Synclinorium (SE Basque-Cantabrian Region, northern Iberian Peninsula). The Laño sites were discovered in 1984; thousands of bones and teeth, including microfossils, have been collected during the prospection in the field and excavation campaigns. The vertebrate remains occur at two different stratigraphic horizons within a continental to shallow marine succession of Late Campanian-Maastrichtian age. The lower horizon contains the Laño 1 and Laño 2 sites, whereas the upper horizon contains the Albaina site. In the Laño sites, three fossiliferous beds (called L1A, L1B and L2) are known within an alluvial system composed mainly of fluvial sands and silts. The sedimentary structures are consistent with channel areas within an extensive braided river system. Based mainly on stratigraphic correlations, the fluvial beds of Laño are regarded as Late Campanian to Early Maastrichtian in age. These deposits have yielded a very diverse vertebrate assemblage, which consists of nearly 40 species, including actinopterygians, lissamphibians, lepidosaurs, turtles, crocodyliforms, dinosaurs, pterosaurs, and mammals. Seven genera and ten species have been erected to date in Laño. With reference to the marine vertebrate association of Albaina, it consists of at least 37 species, including sharks and rays, actinopterygians, mosasaurids, and plesiosaurs. Two genera and species of rhinobatoids (family indet.) and two new species of rhinobatids have been erected in Albaina. The fossil association indicates a Late (but not latest) Maastrichtian age. Recently, isolated turtle and dinosaur fossils have been discovered in the sublittoral beds of Albaina. The Laño quarry is one of the most noteworthy Campanian-Maastrichtian vertebrate localities of Europe by its taxonomic diversity, and provides useful information about the composition and affinities of both continental and marine vertebrate faunas from the latest Cretaceous of southwestern Europe.Los niveles con fósiles de vertebrados de la cantera de Laño (Condado de Treviño) se cuentan entre los más importantes del Cretácico Superior de Europa. Desde un punto de vista geológico, Laño y la región adyacente forman parte del flanco sur del Sinclinorio Subcantábrico (SE de la Región Vasco-Cantábrica). El descubrimiento de los niveles fosilíferos remonta a 1984; las prospecciones de campo y las campañas de excavación han proporcionado miles de huesos y dientes, incluyendo microfósiles. Los restos de vertebrados aparecen en dos horizontes estratigráficos diferentes formando parte de una sucesión continental a marina litoral de edad Campaniense superior a Maastrichtiense. El horizonte inferior contiene los yacimientos de Laño 1 y Laño 2, mientras que el superior contiene solo uno: Albaina. En los yacimientos de Laño, se reconocen tres niveles fosilíferos (llamados L1A, L1B y L2) formados en el seno de un sistema aluvial compuesto por arenas y limos fluviales. Las estructuras sedimentarias indican áreas de canal dentro de un sistema trenzado muy extendido. Según las correlaciones estratigráficas, los depósitos fluviales de Laño son de edad Campaniense superior a Maastrichtiense inferior. Estos depósitos han proporcionado una asociación muy diversa de vertebrados, que consiste en cerca de 40 especies, incluyendo actinopterigios, lisanfibios, lepidosaurios, tortugas, cocodrilos, dinosaurios, pterosaurios y mamíferos. En Laño se han definido hasta el momento siete géneros y diez especies. Por lo que respecta a la asociación de vertebrados marinos de Albaina, se han reconocido al menos 37 especies, que incluyen tiburones y rayas, actinopterigios, mosasaurios y plesiosaurios. Hasta la fecha se han definido en Albaina cuatro nuevos rinobatoideos: dos especies de rinobátidos y dos géneros y especies cuya familia es indeterminada. La asociación fósil es de edad Maastrichtiense superior no terminal. Recientemente se ha descrito el hallazgo de restos fósiles aislados de tortugas y dinosaurios en los niveles marinos de Albaina. La cantera de Laño es una de las localidades de vertebrados más destacadas del Campaniense-Maastrichtiense por su diversidad taxonómica, y proporciona información relevante sobre la composición y afinidades de las faunas de vertebrados continentales y marinos del Cretácico final del suroeste de Europa

Late cretaceous continental and marine vertebrate assemblages of the Laño Quarry (Basque-Cantabrian Region, Iberian Peninsula): An update

The vertebrate-bearing beds of the Laño quarry (Condado de Treviño) are among the most relevant sites from the Late Cretaceous of Europe. Geologically, Laño and the adjacent region are set on the southern limb of the South-Cantabrian Synclinorium (SE Basque-Cantabrian Region, northern Iberian Peninsula). The Laño sites were discovered in 1984; thousands of bones and teeth, including microfossils, have been collected during the prospection in the field and excavation campaigns. The vertebrate remains occur at two different stratigraphic horizons within a continental to shallow marine succession of Late Campanian-Maastrichtian age. The lower horizon contains the Laño 1 and Laño 2 sites, whereas the upper horizon contains the Albaina site. In the Laño sites, three fossiliferous beds (called L1A, L1B and L2) are known within an alluvial system composed mainly of fluvial sands and silts. The sedimentary structures are consistent with channel areas within an extensive braided river system. Based mainly on stratigraphic correlations, the fluvial beds of Laño are regarded as Late Campanian to Early Maastrichtian in age. These deposits have yielded a very diverse vertebrate assemblage, which consists of nearly 40 species, including actinopterygians, lissamphibians, lepidosaurs, turtles, crocodyliforms, dinosaurs, pterosaurs, and mammals. Seven genera and ten species have been erected to date in Laño. With reference to the marine vertebrate association of Albaina, it consists of at least 37 species, including sharks and rays, actinopterygians, mosasaurids, and plesiosaurs. Two genera and species of rhinobatoids (family indet.) and two new species of rhinobatids have been erected in Albaina. The fossil association indicates a Late (but not latest) Maastrichtian age. Recently, isolated turtle and dinosaur fossils have been discovered in the sublittoral beds of Albaina. The Laño quarry is one of the most noteworthy Campanian-Maastrichtian vertebrate localities of Europe by its taxonomic diversity, and provides useful information about the composition and affinities of both continental and marine vertebrate faunas from the latest Cretaceous of southwestern EuropeLos niveles con fósiles de vertebrados de la cantera de Laño (Condado de Treviño) se cuentan entre los más importantes del Cretácico Superior de Europa. Desde un punto de vista geológico, Laño y la región adyacente forman parte del flanco sur del Sinclinorio Subcantábrico (SE de la Región Vasco-Cantábrica). El descubrimiento de los niveles fosilíferos remonta a 1984; las prospecciones de campo y las campañas de excavación han proporcionado miles de huesos y dientes, incluyendo microfósiles. Los restos de vertebrados aparecen en dos horizontes estratigráficos diferentes formando parte de una sucesión continental a marina litoral de edad Campaniense superior a Maastrichtiense. El horizonte inferior contiene los yacimientos de Laño 1 y Laño 2, mientras que el superior contiene solo uno: Albaina. En los yacimientos de Laño, se reconocen tres niveles fosilíferos (llamados L1A, L1B y L2) formados en el seno de un sistema aluvial compuesto por arenas y limos fluviales. Las estructuras sedimentarias indican áreas de canal dentro de un sistema trenzado muy extendido. Según las correlaciones estratigráficas, los depósitos fluviales de Laño son de edad Campaniense superior a Maastrichtiense inferior. Estos depósitos han proporcionado una asociación muy diversa de vertebrados, que consiste en cerca de 40 especies, incluyendo actinopterigios, lisanfibios, lepidosaurios, tortugas, cocodrilos, dinosaurios, pterosaurios y mamíferos. En Laño se han definido hasta el momento siete géneros y diez especies. Por lo que respecta a la asociación de vertebrados marinos de Albaina, se han reconocido al menos 37 especies, que incluyen tiburones y rayas, actinopterigios, mosasaurios y plesiosaurios. Hasta la fecha se han definido en Albaina cuatro nuevos rinobatoideos: dos especies de rinobátidos y dos géneros y especies cuya familia es indeterminada. La asociación fósil es de edad Maastrichtiense superior no terminal. Recientemente se ha descrito el hallazgo de restos fósiles aislados de tortugas y dinosaurios en los niveles marinos de Albaina. La cantera de Laño es una de las localidades de vertebrados más destacadas del Campaniense-Maastrichtiense por su diversidad taxonómica, y proporciona información relevante sobre la composición y afinidades de las faunas de vertebrados continentales y marinos del Cretácico final del suroeste de EuropaField research in the Laño quarry has been supported by financial aids of the Ministerio de Economía y Competitividad (MINECO projects CGL2010-18851/BTE and CGL2013-47521-P), Ministerio de Ciencia e Innovación (CGL2007-64061/BTE), Ministerio de Ciencia y Tecnología (CGL2004-02338/BTE, BTE2001-0185-C02-01, BOS2000-1369), Ministerio de Educación y Ciencia (Acción Integrada hispano-francesa 201-B), Diputación Foral de Álava/Arabako Foru Aldundia (93/23, 95/A23), Gobierno Vasco/Eusko Jaurlaritza (IT834-13, IT-320-10, IT-361-07, GV121.310-4/87), Universidad del País Vasco/Euskal Herriko Unibertsitatea (9/ UPV00121.310-15303/2003), Centre National de la Recherche Scientifique (CNRS, France), Institut National des Sciences de l’Univers (INSU), The Dinosaur Society (research grant 1997) and the National Geographic Research (grant #6597-99). This work is part of a palaeontological collaboration (Convenio específico de colaboración) between the Universidad del País Vasco/EHU, the Centre National de la Recherche Scientifique (CNRS, France) and the Muséum National d’Histoire Naturelle (MNHN, Paris). Contribution ISEM n° 2015-04

Hadrodus Leidy 1858

Genus Hadrodus Leidy, 1858 TYPE SPECIES. — By subsequent designation: Hadrodus priscus Leidy, 1858. Cretaceous from Colombus, Mississippi, USA. “The genus and species are founded upon the fragment of a bone with two teeth, apparently a Pycnodont fish allied to Placodus ” (Leidy 1858: 167). Holotype of Hadrodus priscus: isolated premaxilla figured in Leidy (1873: pl. 19, figs 17-20). OTHER SPECIES. — H. marshi Gregory, 1950, based on premaxilla, prearticular, and fragments of skull roof, lower Senonian from the Niobrara Chalk, Smoky Hill River, Kansas, USA; H. hewletti (Applegate, 1970), Mooreville Chalk (Campanian) in Greene County, Alabama. Bell (1986) gathered in a single pycnodontiform taxon the dentitions named Hadrodus and the cranial roof named Hadrodus hewletti and described by Applegate (1970), who considered it as an acipenseriform. This material also includes some lepidotrichia, dorsal scutes, and vertebrae with arcocentra and apparently also autocentra. This fact, and the great resemblance of the skull roof and the dorsal scutes of Hadrodus hewletti with those of the Acipenseridae (e.g., compare Applegate 1970: figs 181, 184E with Hilton & Bemis 1999: figs 5, 6) indicate that these remains do not belong to a pycnodontiform (already suggested by Lambers 1991: 509). This genus is in need of revision, and is not included in the analysis.Published as part of Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1) on page 147, DOI: 10.5281/zenodo.537164

A new insight into pycnodontiform fishes

Poyato-Ariza F. J., Wenz S. (2002): A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139-248, DOI: http://doi.org/10.5281/zenodo.537164

Coccodontidae Berg 1940

Family COCCODONTIDAE Berg, 1940 <p> TYPE GENUS. — <i>Coccodus</i> Pictet, 1850.</p> <p> INCLUDED GENERA. — The type genus, <i>Ichthyoceros</i> Gayet, 1984, and <i>Trewavasia</i> White & Moy-Thomas, 1941.</p> <p>REVISED DIAGNOSIS. — Pycnodontoid fishes with the following unique combination of primitive and derived characters: extrascapular not fused to parietal; neural and haemal corresponding arcocentra partially surrounding notochord; six to eight epichordal elements in caudal endoskeleton; hypochordal elements in caudal endoskeleton enlarged; less than 29 dorsal axonosts; if present, first dorsal ridge scale larger than subsequent dorsal ridge scales; two or less dorsal ridge scales; three or less ventral keel scales; at most one post-cloacal ventral keel scale.</p>Published as part of <i>Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1)</i> on page 220, DOI: <a href="http://zenodo.org/record/5371649">10.5281/zenodo.5371649</a&gt

Coelodus Heckel 1854, sensu stricto

Genus Coelodus Heckel, 1854 sensu stricto TYPE SPECIES. — Coelodus saturnus Heckel, 1854. INCLUDED SPECIES. — Monotypic genus. REVISED DIAGNOSIS. — Derived pycnodontid fish with the following autapomorphies: prearticular teeth extremely elongated, up to five times longer perpendicularly to the longitudinal axis of the bone; diastema in caudal endoskeleton present, separating hypochordal elements 7 and 8. Unique combination of reverted primitive and derived characters: mouth gape subhorizontal; body ovoid in contour, maximum body height about 60% of standard length; caudal pedicle long, well differentiated; 11 teeth on main prearticular tooth row; grooves on vomerine and prearticular teeth present; about 31 vertebrae; 10 or 11 hypochordal elements in caudal endoskeleton, some of them hypertrophied; caudal fin double emarginated in distal contour; dorsal ridge scales separated from each other; no spines on dorsal ridge scales.Published as part of Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1) on page 222, DOI: 10.5281/zenodo.537164

Mesturus Wagner 1862

Genus Mesturus Wagner, 1862 TYPE SPECIES. — By monotypy: Mesturus verrucosus Wagner, 1862. Lower Tithonian from the “Solnhofener Plattenkalke” of Bavaria, Germany. Holotype of Mesturus verrucosus: specimen figured by Wagner (1862: pl. 3, fig. 1; caudal region only). Currently housed at the Bayerischen Staatssamlung für Paleontologie und Historische Geologie, München, Germany, specimen AS V 508. OTHER SPECIES. — M. leedsi, Woodward 1895, Oxfordian from the Peterborough Member of the Oxford Clay Formation, Peterborough, Cambridgeshire, United Kingdom; “ Mesturus cordillera ”, see under Gyrodus sp.; Mesturus sp., Tithonian from Canjuers, Var, France. MATERIAL EXAMINED. — Mesturus verrucosus: JM SOS 2366; NHML 49147 (cast of holotype), P. 37023a-b, 8656; M. leedsi: holotype, NHML 6834 (isolated bones from a single skull). Other specimens: NHML 8382, 8383 (remains of skull), 8384 (head with dentition plus some scales), 8385 (skull with dentition); Mesturus sp., MNHN CNJ 130 a-b (nearly complete, partially disarticulated specimen).Published as part of Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1) on page 148, DOI: 10.5281/zenodo.537164

Mesturidae Nursall 1996

Family MESTURIDAE Nursall, 1996 TYPE GENUS. — Mesturus Wagner, 1862. INCLUDED GENERA. — The type genus, and Micropycnodon Hibbard & Graffham, 1945. REVISED DIAGNOSIS. — Primitive pycnodontiform fishes with the following unique combination of derived characters: maxilla ornamented and elongated; grooves and strong crenulations present on most vomerine and prearticular teeth; anal fin at 60-69% of standard length; first dorsal ridge scale larger than subsequent dorsal ridge scales.Published as part of Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1) on page 218, DOI: 10.5281/zenodo.537164

Anomoeodus Forir 1887

Genus Anomoeodus Forir, 1887 TYPE SPECIES. — Pycnodus subclavatus Agassiz, 1833. INCLUDED SPECIES. — The type species, A. angustus (Agassiz, 1837), A. willetti Woodward, 1893, and A. nursalli Kriwet, 1999; plus numerous nominal species based on isolated dentitions.Published as part of Poyato-Ariza F. J. & Wenz S., 2002, A new insight into pycnodontiform fishes, pp. 139-248 in Geodiversitas 24 (1) on page 221, DOI: 10.5281/zenodo.537164