Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia

The Early Cretaceous of Colombia presents a rich, abundant, and well-preserved fossil vertebrate record of marine reptiles related to the Alto Ricaurte Marine Reptile Lagerstätte. However, despite the growing number of studies on soft tissue preservation worldwide in recent years, few have explored...

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Autores:
Martinez Motta, Manuel Francisco
Tipo de recurso:
Trabajo de grado de pregrado
Fecha de publicación:
2025
Institución:
Universidad de los Andes
Repositorio:
Séneca: repositorio Uniandes
Idioma:
eng
OAI Identifier:
oai:repositorio.uniandes.edu.co:1992/75501
Acceso en línea:
https://hdl.handle.net/1992/75501
Palabra clave:
Paja formation
Marine reptile lagerstätte
Ichthyosaur
Soft tissues
Geociencias
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https://repositorio.uniandes.edu.co/static/pdf/aceptacion_uso_es.pdf
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oai_identifier_str oai:repositorio.uniandes.edu.co:1992/75501
network_acronym_str UNIANDES2
network_name_str Séneca: repositorio Uniandes
repository_id_str
dc.title.eng.fl_str_mv Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
title Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
spellingShingle Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
Paja formation
Marine reptile lagerstätte
Ichthyosaur
Soft tissues
Geociencias
title_short Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
title_full Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
title_fullStr Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
title_full_unstemmed Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
title_sort Recognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia
dc.creator.fl_str_mv Martinez Motta, Manuel Francisco
dc.contributor.advisor.none.fl_str_mv Cadena Rueda, Edwin Alberto
Noe, Leslie Francis
dc.contributor.author.none.fl_str_mv Martinez Motta, Manuel Francisco
dc.contributor.jury.none.fl_str_mv Eickmann, Benjamin
dc.subject.keyword.eng.fl_str_mv Paja formation
Marine reptile lagerstätte
Ichthyosaur
Soft tissues
topic Paja formation
Marine reptile lagerstätte
Ichthyosaur
Soft tissues
Geociencias
dc.subject.themes.spa.fl_str_mv Geociencias
description The Early Cretaceous of Colombia presents a rich, abundant, and well-preserved fossil vertebrate record of marine reptiles related to the Alto Ricaurte Marine Reptile Lagerstätte. However, despite the growing number of studies on soft tissue preservation worldwide in recent years, few have explored this remarkable preservation in this region. This study aims to identify and characterize the potentially preserved soft-tissue remains in the ichthyosaur specimen CIP-0107, addressing the question of how the taphonomic conditions of the Paja Formation contributed to exceptional soft-tissue preservation. In this context, samples of an Aptian ichthyosaur from the Paja Formation of Colombia were analyzed using techniques such as EDTA demineralization, thin section preparation, transmitted and polarized light microscopy, SEM-EDS, and ATR-FTIR spectroscopy. These analyses revealed net-like structures interpreted as potential keratinized epithelium in the stratum spinosum of epidermis, and skin-like tissues. The findings indicate preservation facilitated by rapid burial in “soupy” sediments, microbial activity, and dysoxic-anoxic marine conditions, and highlights the potential of Colombia as a key region for paleontological studies.
publishDate 2025
dc.date.accessioned.none.fl_str_mv 2025-01-20T19:32:30Z
dc.date.issued.none.fl_str_mv 2025-01-20
dc.date.accepted.none.fl_str_mv 2025-01-20
dc.date.available.none.fl_str_mv 2026-01-30
dc.type.none.fl_str_mv Trabajo de grado - Pregrado
dc.type.driver.none.fl_str_mv info:eu-repo/semantics/bachelorThesis
dc.type.version.none.fl_str_mv info:eu-repo/semantics/acceptedVersion
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dc.type.content.none.fl_str_mv Text
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dc.identifier.uri.none.fl_str_mv https://hdl.handle.net/1992/75501
dc.identifier.instname.none.fl_str_mv instname:Universidad de los Andes
dc.identifier.reponame.none.fl_str_mv reponame:Repositorio Institucional Séneca
dc.identifier.repourl.none.fl_str_mv repourl:https://repositorio.uniandes.edu.co/
url https://hdl.handle.net/1992/75501
identifier_str_mv instname:Universidad de los Andes
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dc.language.iso.none.fl_str_mv eng
language eng
dc.relation.references.none.fl_str_mv Alfonso-Rojas, A., & Cadena, E. A. (2020). Exceptionally preserved ‘skin’ in an Early Cretaceous fish from Colombia. PeerJ, 8, e9479.
Bardet, N., & Fernández, M. (2000). A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria. Journal of Paleontology, 74(3), 503-511.
Benavides-Cabra, C. D., Páramo-Fonseca, M. E., Palma-Castro, H. D., Narváez-Rincón, J. A., & Ramos-Clavijo, M. P. (2023). Stratigraphic distribution of marine vertebrates from the Arcillolitas Abigarradas Member (Paja Formation) of the Villa de Leiva–Sáchica-Sutamarchán region, Boyacá, Colombia. Earth Sciences Research Journal, 27(3), 211-226.
Bonnevier Wallstedt, I., Sjövall, P., Thuy, B., De La Garza, R. G., Eriksson, M. E., & Lindgren, J. (2024). Skin anatomy, bone histology and taphonomy of a Toarcian (Lower Jurassic) ichthyosaur (Reptilia: Ichthyopterygia) from Luxembourg, with Implications for Paleobiology. Diversity, 16(8), 492.
Brown, C. M., Henderson, D. M., Vinther, J., Fletcher, I., Sistiaga, A., Herrera, J., & Summons, R. E. (2017). An exceptionally preserved three-dimensional armored dinosaur reveals insights into coloration and Cretaceous predator-prey dynamics. Current Biology, 27(16), 2514-2521.
Cadena, E. (2015). The first South American sandownid turtle from the Lower Cretaceous of Colombia. PeerJ, 3, e1431.
Cadena, E. (2016). Microscopical and elemental FESEM and Phenom ProX-SEM-EDS analysis of osteocyte-and blood vessel-like microstructures obtained from fossil vertebrates of the Eocene Messel Pit, Germany. PeerJ, 4, e1618.
Cadena, E. A. & Parham, J. F. (2015). Oldest known marine turtle? A new protostegid from the Lower Cretaceous of Colombia. PaleoBios, 32,1-42.
Cadena, E. A. (2020). In situ SEM/EDS compositional characterization of osteocytes and blood vessels in fossil and extant turtles on untreated bone surfaces; different preservational pathways microns away. PeerJ, 8, e9833.
Cadena, E. A., Parra-Ruge, M. L., Parra-Ruge, J. D. & Padilla-Bernal, S. (2019). A gravid fossil turtle from the Early Cretaceous reveals a different egg development strategy to that of extant marine turtles. Palaeontology, 62, 533-545.
Carballido, J. L., Pol, D., Parra Ruge, M. L., Padilla Bernal, S., Páramo-Fonseca, M. E., & Etayo-Serna, F. (2015). A new Early Cretaceous brachiosaurid (Dinosauria, Neosauropoda) from northwestern Gondwana (Villa de Leiva, Colombia). Journal of Vertebrate Paleontology, 35(5), e980505.
Cleland T. P., Schroeter E. R., Zamdborg. L., Zheng. W., Lee J. E., Tran J. C., Bern M., Duncan M. B., Lebleu V. S., Ahlf D. R., Thomas P. M., Kalluri R., Kelleher N. L., Schweitzer M. H. (2015). Mass spectrometry and antibody-based characterization of blood vessels from Brachylophosaurus canadensis. Journal of Proteome Research 14:5252–5262.
Cohen, K. M., Finney, S. C., Gibbard, P. L., & Fan, J. X. (2013; updated 2023/09). The ICS International Chronostratigraphic Chart. Episodes Journal of International Geoscience, 36(3), 199-204.
Cortés, D., Larsson, H. C., Maxwell, E. E., Ruge, M. L. P., Patarroyo, P., & Wilson, J. A. (2019). An early Cretaceous teleosauroid (Crocodylomorpha: Thalattosuchia) from Colombia. Ameghiniana, 56(5), 365-379.
Cortés, D., Maxwell, E. E. & Larsson, H. C. E. (2021). Re-appearance of hypercarnivore ichthyosaurs in the Cretaceous with differentiated dentition: revision of ‘Platypterygius’ sachicarum (Reptilia: Ichthyosauria, Ophthalmosauridae) from Colombia. Journal of Systematic Palaeontology, 19(14), 969-1002.
Cortés, D., Parra-Ruge, M. L., Parra-Ruge, J. D. D., Deers-Potvin, A., Smith, A., Hoai-Nam, B., & Larsson, H. (2023). Theropod dinosaur footprints from the Early Cretaceous of Colombia: walking dynamics and paleoenvironmental implications. Revista Facies, 9, 1-17.
Cozzi, B., Huggenberger, S., & Oelschläger, H. A. (2017). Anatomy of dolphins: insights into body structure and function. Academic Press.
Cremona, J. (2014). Extreme close-up photography and focus stacking. Crowood.
De La Garza, R. G., Sjövall, P., Hauff, R., & Lindgren, J. (2023). Preservational modes of some ichthyosaur soft tissues (Reptilia, Ichthyopterygia) from the Jurassic Posidonia Shale of Germany. Palaeontology, 66(4), e12668.
Delair, J. B. (1966). Unusual preservation of fibrous elements in an ichthyosaur skull. Nature, 212(5062), 575-576.
Eriksson, M. E., De La Garza, R., Horn, E., & Lindgren, J. (2022). A review of ichthyosaur (Reptilia, Ichthyopterygia) soft tissues with implications for life reconstructions. Earth-Science Reviews, 226, 103965.
Etayo-Serna, F. (1965). Sinopsis estratigráfica de la región de Villa de Leiva y zonas próximas. Boletín de Geología, 21, 19-32.
Etayo-Serna, F. (1968). El Sistema Cretáceo en la región de Villa de Leiva y zonas próximas. Geología Colombiana, 5, 5-74.
Frandsen, A. F. (2016). Polarized light microscopy. NASA, Kennedy Space Center. (No. KSC-E-DAA-TN37401).
Gaona-Narvaez, T., Maurrasse, F. J. M., & Etayo-Serna, F. (2013). Geochemistry, palaeoenvironments and timing of Aptian organic-rich beds of the Paja Formation (Curití, Eastern Cordillera, Colombia). Geological Society, London, Special Publications, 382(1), 31-48.
Goldstein, J. I., Newbury, D. E., Michael, J. R., Ritchie, N. W., Scott, J. H. J., & Joy, D. C. (2018). Scanning electron microscopy and X-ray microanalysis. springer.
Gómez-Pérez, M., & Noè, L. F. (2017). Cranial anatomy of a new pliosaurid Acostasaurus pavachoquensis from the Lower Cretaceous of Colombia, South America. Palaeontographica Abteilung A, 310, 5-42.
Hampe, O. (1992). Ein großwüchsiger Pliosauride (Reptilia: Plesiosauria) aus-der Unterkreide (oberes Aptium) von Kolumbien. Courier Forschungsinstitut Senckenberg, 145, 1–25.
IUGS [International Union of Geological Sciences]. (2022). The first 100 IUGS geological heritage sites. Edited and published by IUGS.
Lee, Y. C., Chiang, C. C., Huang, P. Y., Chung, C. Y., Huang, T. D., Wang, C. C., Chen, C., Chang, R. S., Liao, C. H., & Reisz, R. R. (2017). Evidence of preserved collagen in an Early Jurassic sauropodomorph dinosaur revealed by synchrotron FTIR microspectroscopy. Nature communications, 8(1), 14220.
Lindgren, J., Sjövall, P., Thiel, V., Zheng, W., Ito, S., Wakamatsu, K., Hauff, R., Kear, B. P., Engdahl, A., Alwmark, C., Eriksson, M. E., Jarenmark, M., Sachs, S.,
Ahlberg, P. E., Marone, F., Kuriyama, T., Gustafsson, O., Malmberg, P., Thomen, A., Rodríguez-Meizoso, I., Uvdal, P., Ojika, M., Schweitzer, M. H. (2018). Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur. Nature, 564, 359–365.
Lingham-Soliar, T. (1999). Rare soft tissue preservation showing fibrous structures in an ichthyosaur from the Lower Lias (Jurassic) of England. Proceedings of the Royal Society of London. Series B: Biological Sciences, 266(1436), 2367-2373.
Lingham‐Soliar, T. (2001). The ichthyosaur integument: skin fibers, a means for a strong, flexible and smooth skin. Lethaia, 34(4), 287-302.
Lingham-Soliar, T. (2015). Integumental Taphonomy. In: The Vertebrate Integument Volume 2: Structure, Design and Function, 263-293.
Lingham-Soliar, T., & Plodowski, G. (2010). The integument of Psittacosaurus from Liaoning Province, China: taphonomy, epidermal patterns and color of a ceratopsian dinosaur. Naturwissenschaften, 97, 479-486.
Martill, D. M. (1988). Preservation of fish in the Cretaceous Santana Formation Brazil. Palaeontology, 31(1), 1–18.
Martill, D. M. (1993). Soupy substrates: a medium for the exceptional preservation of ichthyosaurs of the Posidonia Shale (Lower Jurassic) of Germany. Kaupia, 2, 77-97.
Martill, D. M. (1995). An ichthyosaur with preserved soft tissue from the Sinemurian of southern England. Palaeontology, 38(4), 897-904.
Martin, T., Marugán-Lobón, J., Vullo, R., Martín-Abad, H., Luo, Z. X., & Buscalioni, A. D. (2015). A Cretaceous eutriconodont and integument evolution in early mammals. Nature, 526(7573), 380-384.
Maxwell, E. E., Cortés, D., Patarroyo, P., & Parra Ruge, M. L. (2019). A new specimen of Platypterygius sachicarum (Reptilia, Ichthyosauria) from the Early Cretaceous of Colombia and its phylogenetic implications. Journal of Vertebrate Paleontology, 39(1), e1577875.
Maxwell, E. E., Dick, D., Padilla, S., & Parra, M. L. (2016). A new ophthalmosaurid ichthyosaur from the Early Cretaceous of Colombia. Papers in Palaeontology, 2(1), 59-70.
Montoya-Arenas, D. M., & Etayo-Serna, F. (2019). Formación La Paja: Descripción de la sección tipo. Influencia de los tapices microbiales en su génesis. In Estudios geológicos y paleontológicos sobre el Cretácico en la región del embalse del río Sogamoso, Valle Medio del Magdalena, 51-67.
Morales‐Guerrero, B., Barragán‐Vargas, C., Silva‐Rosales, G. R., Ortega‐Ortiz, C. D., Gendron, D., Martinez‐Levasseur, L. M., & Acevedo‐Whitehouse, K. (2017). Melanin granules melanophages and a fully‐melanized epidermis are common traits of odontocete and mysticete cetaceans. Veterinary dermatology, 28(2), 213-e50.
Motani, R., You, H., & McGowan, C. (1996). Eel-like swimming in the earliest ichthyosaurs. Nature, 382(6589), 347-348.
NIST [National Institute of Standards and Technology, U.S. Department of Commerce]. (last accessed 26/11/2024). Calcium carbonate (precipitated). NIST Chemistry
WebBook, SRD 69. webbook.nist.gov/cgi/cbook.cgi?ID=C471341&Units=SI&Mask=80#IR-Spec
Noè, L. F., & Gómez–Pérez, M. (2020). Plesiosaurs, palaeoenvironments, and the Paja Formation Lagerstätte of central Colombia: An overview. In: Gómez J. & Pinilla–Pachon A. O. (Editors). The Geology of Colombia, Volume 2 Mesozoic. Publicaciones Geológicas Especiales 36, Servicio Geológico Colombiano, 441-483.
Noè, L. F. & Gómez–Pérez, M. (2022). Giant pliosaurids (Sauropterygia; Plesiosauria) from the Lower Cretaceous peri-Gondwanan seas of Colombia and Australia. Cretaceous Research, 132, e105122.
Paik, I. S., Kim, H. J., & Huh, M. (2010). Impressions of dinosaur skin from the Cretaceous Haman Formation in Korea. Journal of Asian Earth Sciences, 39(4), 270-274.
Palma-Castro, H. D., Cómbita-Romero, D. A., Cadena, E. A., Carvalho, M. R., & Herrera, F. (2023). An Early Cretaceous Sphenophyllum or a hatchling turtle?. Palaeontologia Electronica, 26(3), 1-10.
Páramo-Fonseca, M. E. (1997). Platypterygius sachicarum (Reptilia, Ichthyosauria) nueva especie del Cretácico de Colombia. Revista Ingeominas, 6, 1-12.
Páramo-Fonseca, M. E., Benavides-Cabra, C. D., & Garavito-Camacho, R. A. (2024). A new species of Platypterygius (Ophthalmosauridae) from the lower Barremian of Colombia and assessment of the species composition of the genus. Earth Sciences Research Journal, 28(2), 103-126.
Páramo-Fonseca, M. E., Benavides-Cabra, C. D., & Gutiérrez, I. E. (2018). A new large Pliosaurid from the Barremian (Lower Cretaceous) of Sáchica, Boyacá, Colombia. Earth Sciences Research Journal, 22(4), 223-238.
Páramo-Fonseca, M. E., García-Guerrero, J., Benavides-Cabra, C. D., Padilla-Bernal, S., & Castañeda-Gómez, A. J. (2021). A benchmark specimen of Muiscasaurus catheti from the upper Aptian of Villa de Leiva, Colombia: New anatomical features and phylogenetic implications. Cretaceous Research, 119, 104685.
Páramo-Fonseca, M. E., Gómez-Pérez, M., Noé, L. F., & Etayo-Serna, F. (2016). Stenorhynchosaurus munozi, gen. et sp. Nov. a new pliosaurid from the Upper Barremian (Lower Cretaceous) of Villa de Leiva, Colombia, South America. Revista de la Academia Colombiana de Ciencias Exactas Físicas y Naturales, 40(154), 84-103.
Páramo-Fonseca, M. E., O’Gorman, J. P., Gasparini, Z., Padilla, S., & Parra-Ruge, M. L. (2019). A new late Aptian elasmosaurid from the Paja Formation, Villa de Leiva, Colombia. Cretaceous Research, 99, 30-40.
Reeb, D., Best, P. B., & Kidson, S. H. (2007). Structure of the integument of southern right whales, Eubalaena australis. The Anatomical Record, 290(6), 596-613.
Renesto, S., Dal Sasso, C., Fogliazza, F., & Ragni, C. (2020). New findings reveal that the Middle Triassic ichthyosaur Mixosaurus cornalianus is the oldest amniote with a dorsal fin. Acta Palaeontologica Polonica, 65(3), 511-522.
Rossi, V., Bernardi, M., Fornasiero, M., Nestola, F., Unitt, R., Castelli, S., & Kustatscher, E. (2024). Forged soft tissues revealed in the oldest fossil reptile from the early Permian of the Alps. Palaeontology, 67(1), e12690.
Schäfer, W. (1972). Ecology and palaeoecology of marine environments. University of Chicago Press.
Schroeder, A. B., Dobson, E. T., Rueden, C. T., Tomancak, P., Jug, F., & Eliceiri, K. W. (2021). The ImageJ ecosystem: Open‐source software for image visualization, processing, and analysis. Protein Science, 30(1), 234-249.
Schultze, H. P., & Stöhr, D. (1996). Vinctifer (Pisces, Aspidorhynchidae) aus der unterkreide (oberes Aptium) von Kolumbien. Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen, 199(3), 395-415.
Schuttlefield, J. D., & Grassian, V. H. (2008). ATR–FTIR spectroscopy in the undergraduate chemistry laboratory. Part I: Fundamentals and examples. Journal of Chemical Education, 85(2), 279.
Schweitzer, M. H., Schroeter, E. R., Cleland, T. P., & Zheng, W. (2019). Paleoproteomics of Mesozoic dinosaurs and other Mesozoic fossils. Proteomics, 19(16), 1800251.
Schweitzer, M. H., Zheng, W., Cleland, T. P., & Bern, M. (2013). Molecular analyses of dinosaur osteocytes support the presence of endogenous molecules. Bone, 52(1), 414-423.
Scotese, C. R. (2021). An atlas of Phanerozoic paleogeographic maps: the seas come in and the seas go out. Annual Review of Earth and Planetary Sciences, 49(1), 679-728.
Seilacher, A., Reif, W. E., & Westphal, F. (1985). Sedimentological, ecological and temporal patterns of fossil Lagerstätten. Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 311(1148), 5-24.
Signore, M., Bucci, E., Pede, C., & Barbera, C. (2005). A new ichthyodectid fish from the Lower Cretaceous of Pietraroja (Southern Italy). PalArch, 5, 25-29.
Smithwick, F. (2019). A taphonomic and palaeoecological approach to the study of palaeocolour (Doctoral dissertation, University of Bristol).
Welles, S. P. (1962). A new species of elasmosaur from the Aptian of Colombia and a review of the Cretaceous plesiosaurs. University of California Publications in the Geological Sciences, 44, 1-96.
Xu, X., Zhou, Z., Wang, Y., & Wang, M. (2020). Study on the Jehol Biota: recent advances and future prospects. Science China Earth Sciences, 63, 757-773.
Yang, Z., Jiang, B., Xu, J., & McNamara, M. E. (2024). Cellular structure of dinosaur scales reveals retention of reptile-type skin during the evolutionary transition to feathers. Nature Communications, 15(1), 4063.
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spelling Cadena Rueda, Edwin AlbertoNoe, Leslie Francisvirtual::22237-1Martinez Motta, Manuel FranciscoEickmann, Benjamin2025-01-20T19:32:30Z2026-01-302025-01-202025-01-20https://hdl.handle.net/1992/75501instname:Universidad de los Andesreponame:Repositorio Institucional Sénecarepourl:https://repositorio.uniandes.edu.co/The Early Cretaceous of Colombia presents a rich, abundant, and well-preserved fossil vertebrate record of marine reptiles related to the Alto Ricaurte Marine Reptile Lagerstätte. However, despite the growing number of studies on soft tissue preservation worldwide in recent years, few have explored this remarkable preservation in this region. This study aims to identify and characterize the potentially preserved soft-tissue remains in the ichthyosaur specimen CIP-0107, addressing the question of how the taphonomic conditions of the Paja Formation contributed to exceptional soft-tissue preservation. In this context, samples of an Aptian ichthyosaur from the Paja Formation of Colombia were analyzed using techniques such as EDTA demineralization, thin section preparation, transmitted and polarized light microscopy, SEM-EDS, and ATR-FTIR spectroscopy. These analyses revealed net-like structures interpreted as potential keratinized epithelium in the stratum spinosum of epidermis, and skin-like tissues. The findings indicate preservation facilitated by rapid burial in “soupy” sediments, microbial activity, and dysoxic-anoxic marine conditions, and highlights the potential of Colombia as a key region for paleontological studies.Pregrado40 páginasapplication/pdfengUniversidad de los AndesGeocienciasFacultad de CienciasDepartamento de Geocienciashttps://repositorio.uniandes.edu.co/static/pdf/aceptacion_uso_es.pdfhttp://purl.org/coar/access_right/c_f1cf http://purl.org/coar/access_right/c_f1cfRecognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of ColombiaTrabajo de grado - Pregradoinfo:eu-repo/semantics/bachelorThesisinfo:eu-repo/semantics/acceptedVersionhttp://purl.org/coar/resource_type/c_7a1fTexthttp://purl.org/redcol/resource_type/TPPaja formationMarine reptile lagerstätteIchthyosaurSoft tissuesGeocienciasAlfonso-Rojas, A., & Cadena, E. A. (2020). Exceptionally preserved ‘skin’ in an Early Cretaceous fish from Colombia. PeerJ, 8, e9479.Bardet, N., & Fernández, M. (2000). A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria. Journal of Paleontology, 74(3), 503-511.Benavides-Cabra, C. D., Páramo-Fonseca, M. E., Palma-Castro, H. D., Narváez-Rincón, J. A., & Ramos-Clavijo, M. P. (2023). Stratigraphic distribution of marine vertebrates from the Arcillolitas Abigarradas Member (Paja Formation) of the Villa de Leiva–Sáchica-Sutamarchán region, Boyacá, Colombia. Earth Sciences Research Journal, 27(3), 211-226.Bonnevier Wallstedt, I., Sjövall, P., Thuy, B., De La Garza, R. G., Eriksson, M. E., & Lindgren, J. (2024). Skin anatomy, bone histology and taphonomy of a Toarcian (Lower Jurassic) ichthyosaur (Reptilia: Ichthyopterygia) from Luxembourg, with Implications for Paleobiology. Diversity, 16(8), 492.Brown, C. M., Henderson, D. M., Vinther, J., Fletcher, I., Sistiaga, A., Herrera, J., & Summons, R. E. (2017). An exceptionally preserved three-dimensional armored dinosaur reveals insights into coloration and Cretaceous predator-prey dynamics. Current Biology, 27(16), 2514-2521.Cadena, E. (2015). The first South American sandownid turtle from the Lower Cretaceous of Colombia. PeerJ, 3, e1431.Cadena, E. (2016). Microscopical and elemental FESEM and Phenom ProX-SEM-EDS analysis of osteocyte-and blood vessel-like microstructures obtained from fossil vertebrates of the Eocene Messel Pit, Germany. PeerJ, 4, e1618.Cadena, E. A. & Parham, J. F. (2015). Oldest known marine turtle? A new protostegid from the Lower Cretaceous of Colombia. PaleoBios, 32,1-42.Cadena, E. A. (2020). In situ SEM/EDS compositional characterization of osteocytes and blood vessels in fossil and extant turtles on untreated bone surfaces; different preservational pathways microns away. PeerJ, 8, e9833.Cadena, E. A., Parra-Ruge, M. L., Parra-Ruge, J. D. & Padilla-Bernal, S. (2019). A gravid fossil turtle from the Early Cretaceous reveals a different egg development strategy to that of extant marine turtles. Palaeontology, 62, 533-545.Carballido, J. L., Pol, D., Parra Ruge, M. L., Padilla Bernal, S., Páramo-Fonseca, M. E., & Etayo-Serna, F. (2015). A new Early Cretaceous brachiosaurid (Dinosauria, Neosauropoda) from northwestern Gondwana (Villa de Leiva, Colombia). Journal of Vertebrate Paleontology, 35(5), e980505.Cleland T. P., Schroeter E. R., Zamdborg. L., Zheng. W., Lee J. E., Tran J. C., Bern M., Duncan M. B., Lebleu V. S., Ahlf D. R., Thomas P. M., Kalluri R., Kelleher N. L., Schweitzer M. H. (2015). Mass spectrometry and antibody-based characterization of blood vessels from Brachylophosaurus canadensis. Journal of Proteome Research 14:5252–5262.Cohen, K. M., Finney, S. C., Gibbard, P. L., & Fan, J. X. (2013; updated 2023/09). The ICS International Chronostratigraphic Chart. Episodes Journal of International Geoscience, 36(3), 199-204.Cortés, D., Larsson, H. C., Maxwell, E. E., Ruge, M. L. P., Patarroyo, P., & Wilson, J. A. (2019). An early Cretaceous teleosauroid (Crocodylomorpha: Thalattosuchia) from Colombia. Ameghiniana, 56(5), 365-379.Cortés, D., Maxwell, E. E. & Larsson, H. C. E. (2021). Re-appearance of hypercarnivore ichthyosaurs in the Cretaceous with differentiated dentition: revision of ‘Platypterygius’ sachicarum (Reptilia: Ichthyosauria, Ophthalmosauridae) from Colombia. Journal of Systematic Palaeontology, 19(14), 969-1002.Cortés, D., Parra-Ruge, M. L., Parra-Ruge, J. D. D., Deers-Potvin, A., Smith, A., Hoai-Nam, B., & Larsson, H. (2023). Theropod dinosaur footprints from the Early Cretaceous of Colombia: walking dynamics and paleoenvironmental implications. Revista Facies, 9, 1-17.Cozzi, B., Huggenberger, S., & Oelschläger, H. A. (2017). Anatomy of dolphins: insights into body structure and function. Academic Press.Cremona, J. (2014). Extreme close-up photography and focus stacking. Crowood.De La Garza, R. G., Sjövall, P., Hauff, R., & Lindgren, J. (2023). Preservational modes of some ichthyosaur soft tissues (Reptilia, Ichthyopterygia) from the Jurassic Posidonia Shale of Germany. Palaeontology, 66(4), e12668.Delair, J. B. (1966). Unusual preservation of fibrous elements in an ichthyosaur skull. Nature, 212(5062), 575-576.Eriksson, M. E., De La Garza, R., Horn, E., & Lindgren, J. (2022). A review of ichthyosaur (Reptilia, Ichthyopterygia) soft tissues with implications for life reconstructions. Earth-Science Reviews, 226, 103965.Etayo-Serna, F. (1965). Sinopsis estratigráfica de la región de Villa de Leiva y zonas próximas. Boletín de Geología, 21, 19-32.Etayo-Serna, F. (1968). El Sistema Cretáceo en la región de Villa de Leiva y zonas próximas. Geología Colombiana, 5, 5-74.Frandsen, A. F. (2016). Polarized light microscopy. NASA, Kennedy Space Center. (No. KSC-E-DAA-TN37401).Gaona-Narvaez, T., Maurrasse, F. J. M., & Etayo-Serna, F. (2013). Geochemistry, palaeoenvironments and timing of Aptian organic-rich beds of the Paja Formation (Curití, Eastern Cordillera, Colombia). Geological Society, London, Special Publications, 382(1), 31-48.Goldstein, J. I., Newbury, D. E., Michael, J. R., Ritchie, N. W., Scott, J. H. J., & Joy, D. C. (2018). Scanning electron microscopy and X-ray microanalysis. springer.Gómez-Pérez, M., & Noè, L. F. (2017). Cranial anatomy of a new pliosaurid Acostasaurus pavachoquensis from the Lower Cretaceous of Colombia, South America. Palaeontographica Abteilung A, 310, 5-42.Hampe, O. (1992). Ein großwüchsiger Pliosauride (Reptilia: Plesiosauria) aus-der Unterkreide (oberes Aptium) von Kolumbien. Courier Forschungsinstitut Senckenberg, 145, 1–25.IUGS [International Union of Geological Sciences]. (2022). The first 100 IUGS geological heritage sites. Edited and published by IUGS.Lee, Y. C., Chiang, C. C., Huang, P. Y., Chung, C. Y., Huang, T. D., Wang, C. C., Chen, C., Chang, R. S., Liao, C. H., & Reisz, R. R. (2017). Evidence of preserved collagen in an Early Jurassic sauropodomorph dinosaur revealed by synchrotron FTIR microspectroscopy. Nature communications, 8(1), 14220.Lindgren, J., Sjövall, P., Thiel, V., Zheng, W., Ito, S., Wakamatsu, K., Hauff, R., Kear, B. P., Engdahl, A., Alwmark, C., Eriksson, M. E., Jarenmark, M., Sachs, S.,Ahlberg, P. E., Marone, F., Kuriyama, T., Gustafsson, O., Malmberg, P., Thomen, A., Rodríguez-Meizoso, I., Uvdal, P., Ojika, M., Schweitzer, M. H. (2018). Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur. Nature, 564, 359–365.Lingham-Soliar, T. (1999). Rare soft tissue preservation showing fibrous structures in an ichthyosaur from the Lower Lias (Jurassic) of England. Proceedings of the Royal Society of London. Series B: Biological Sciences, 266(1436), 2367-2373.Lingham‐Soliar, T. (2001). The ichthyosaur integument: skin fibers, a means for a strong, flexible and smooth skin. Lethaia, 34(4), 287-302.Lingham-Soliar, T. (2015). Integumental Taphonomy. In: The Vertebrate Integument Volume 2: Structure, Design and Function, 263-293.Lingham-Soliar, T., & Plodowski, G. (2010). The integument of Psittacosaurus from Liaoning Province, China: taphonomy, epidermal patterns and color of a ceratopsian dinosaur. Naturwissenschaften, 97, 479-486.Martill, D. M. (1988). Preservation of fish in the Cretaceous Santana Formation Brazil. Palaeontology, 31(1), 1–18.Martill, D. M. (1993). Soupy substrates: a medium for the exceptional preservation of ichthyosaurs of the Posidonia Shale (Lower Jurassic) of Germany. Kaupia, 2, 77-97.Martill, D. M. (1995). An ichthyosaur with preserved soft tissue from the Sinemurian of southern England. Palaeontology, 38(4), 897-904.Martin, T., Marugán-Lobón, J., Vullo, R., Martín-Abad, H., Luo, Z. X., & Buscalioni, A. D. (2015). A Cretaceous eutriconodont and integument evolution in early mammals. Nature, 526(7573), 380-384.Maxwell, E. E., Cortés, D., Patarroyo, P., & Parra Ruge, M. L. (2019). A new specimen of Platypterygius sachicarum (Reptilia, Ichthyosauria) from the Early Cretaceous of Colombia and its phylogenetic implications. Journal of Vertebrate Paleontology, 39(1), e1577875.Maxwell, E. E., Dick, D., Padilla, S., & Parra, M. L. (2016). A new ophthalmosaurid ichthyosaur from the Early Cretaceous of Colombia. Papers in Palaeontology, 2(1), 59-70.Montoya-Arenas, D. M., & Etayo-Serna, F. (2019). Formación La Paja: Descripción de la sección tipo. Influencia de los tapices microbiales en su génesis. In Estudios geológicos y paleontológicos sobre el Cretácico en la región del embalse del río Sogamoso, Valle Medio del Magdalena, 51-67.Morales‐Guerrero, B., Barragán‐Vargas, C., Silva‐Rosales, G. R., Ortega‐Ortiz, C. D., Gendron, D., Martinez‐Levasseur, L. M., & Acevedo‐Whitehouse, K. (2017). Melanin granules melanophages and a fully‐melanized epidermis are common traits of odontocete and mysticete cetaceans. Veterinary dermatology, 28(2), 213-e50.Motani, R., You, H., & McGowan, C. (1996). Eel-like swimming in the earliest ichthyosaurs. Nature, 382(6589), 347-348.NIST [National Institute of Standards and Technology, U.S. Department of Commerce]. (last accessed 26/11/2024). Calcium carbonate (precipitated). NIST ChemistryWebBook, SRD 69. webbook.nist.gov/cgi/cbook.cgi?ID=C471341&Units=SI&Mask=80#IR-SpecNoè, L. F., & Gómez–Pérez, M. (2020). Plesiosaurs, palaeoenvironments, and the Paja Formation Lagerstätte of central Colombia: An overview. In: Gómez J. & Pinilla–Pachon A. O. (Editors). The Geology of Colombia, Volume 2 Mesozoic. Publicaciones Geológicas Especiales 36, Servicio Geológico Colombiano, 441-483.Noè, L. F. & Gómez–Pérez, M. (2022). Giant pliosaurids (Sauropterygia; Plesiosauria) from the Lower Cretaceous peri-Gondwanan seas of Colombia and Australia. Cretaceous Research, 132, e105122.Paik, I. S., Kim, H. J., & Huh, M. (2010). Impressions of dinosaur skin from the Cretaceous Haman Formation in Korea. Journal of Asian Earth Sciences, 39(4), 270-274.Palma-Castro, H. D., Cómbita-Romero, D. A., Cadena, E. A., Carvalho, M. R., & Herrera, F. (2023). An Early Cretaceous Sphenophyllum or a hatchling turtle?. Palaeontologia Electronica, 26(3), 1-10.Páramo-Fonseca, M. E. (1997). Platypterygius sachicarum (Reptilia, Ichthyosauria) nueva especie del Cretácico de Colombia. Revista Ingeominas, 6, 1-12.Páramo-Fonseca, M. E., Benavides-Cabra, C. D., & Garavito-Camacho, R. A. (2024). A new species of Platypterygius (Ophthalmosauridae) from the lower Barremian of Colombia and assessment of the species composition of the genus. Earth Sciences Research Journal, 28(2), 103-126.Páramo-Fonseca, M. E., Benavides-Cabra, C. D., & Gutiérrez, I. E. (2018). A new large Pliosaurid from the Barremian (Lower Cretaceous) of Sáchica, Boyacá, Colombia. Earth Sciences Research Journal, 22(4), 223-238.Páramo-Fonseca, M. E., García-Guerrero, J., Benavides-Cabra, C. D., Padilla-Bernal, S., & Castañeda-Gómez, A. J. (2021). A benchmark specimen of Muiscasaurus catheti from the upper Aptian of Villa de Leiva, Colombia: New anatomical features and phylogenetic implications. Cretaceous Research, 119, 104685.Páramo-Fonseca, M. E., Gómez-Pérez, M., Noé, L. F., & Etayo-Serna, F. (2016). Stenorhynchosaurus munozi, gen. et sp. Nov. a new pliosaurid from the Upper Barremian (Lower Cretaceous) of Villa de Leiva, Colombia, South America. Revista de la Academia Colombiana de Ciencias Exactas Físicas y Naturales, 40(154), 84-103.Páramo-Fonseca, M. E., O’Gorman, J. P., Gasparini, Z., Padilla, S., & Parra-Ruge, M. L. (2019). A new late Aptian elasmosaurid from the Paja Formation, Villa de Leiva, Colombia. Cretaceous Research, 99, 30-40.Reeb, D., Best, P. B., & Kidson, S. H. (2007). Structure of the integument of southern right whales, Eubalaena australis. The Anatomical Record, 290(6), 596-613.Renesto, S., Dal Sasso, C., Fogliazza, F., & Ragni, C. (2020). New findings reveal that the Middle Triassic ichthyosaur Mixosaurus cornalianus is the oldest amniote with a dorsal fin. Acta Palaeontologica Polonica, 65(3), 511-522.Rossi, V., Bernardi, M., Fornasiero, M., Nestola, F., Unitt, R., Castelli, S., & Kustatscher, E. (2024). Forged soft tissues revealed in the oldest fossil reptile from the early Permian of the Alps. Palaeontology, 67(1), e12690.Schäfer, W. (1972). Ecology and palaeoecology of marine environments. University of Chicago Press.Schroeder, A. B., Dobson, E. T., Rueden, C. T., Tomancak, P., Jug, F., & Eliceiri, K. W. (2021). The ImageJ ecosystem: Open‐source software for image visualization, processing, and analysis. Protein Science, 30(1), 234-249.Schultze, H. P., & Stöhr, D. (1996). Vinctifer (Pisces, Aspidorhynchidae) aus der unterkreide (oberes Aptium) von Kolumbien. Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen, 199(3), 395-415.Schuttlefield, J. D., & Grassian, V. H. (2008). ATR–FTIR spectroscopy in the undergraduate chemistry laboratory. Part I: Fundamentals and examples. Journal of Chemical Education, 85(2), 279.Schweitzer, M. H., Schroeter, E. R., Cleland, T. P., & Zheng, W. (2019). Paleoproteomics of Mesozoic dinosaurs and other Mesozoic fossils. Proteomics, 19(16), 1800251.Schweitzer, M. H., Zheng, W., Cleland, T. P., & Bern, M. (2013). Molecular analyses of dinosaur osteocytes support the presence of endogenous molecules. Bone, 52(1), 414-423.Scotese, C. R. (2021). An atlas of Phanerozoic paleogeographic maps: the seas come in and the seas go out. Annual Review of Earth and Planetary Sciences, 49(1), 679-728.Seilacher, A., Reif, W. E., & Westphal, F. (1985). Sedimentological, ecological and temporal patterns of fossil Lagerstätten. Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 311(1148), 5-24.Signore, M., Bucci, E., Pede, C., & Barbera, C. (2005). A new ichthyodectid fish from the Lower Cretaceous of Pietraroja (Southern Italy). PalArch, 5, 25-29.Smithwick, F. (2019). A taphonomic and palaeoecological approach to the study of palaeocolour (Doctoral dissertation, University of Bristol).Welles, S. P. (1962). A new species of elasmosaur from the Aptian of Colombia and a review of the Cretaceous plesiosaurs. University of California Publications in the Geological Sciences, 44, 1-96.Xu, X., Zhou, Z., Wang, Y., & Wang, M. (2020). Study on the Jehol Biota: recent advances and future prospects. Science China Earth Sciences, 63, 757-773.Yang, Z., Jiang, B., Xu, J., & McNamara, M. E. (2024). Cellular structure of dinosaur scales reveals retention of reptile-type skin during the evolutionary transition to feathers. Nature Communications, 15(1), 4063.202012622Publicationhttps://scholar.google.es/citations?user=gYgnH7QAAAAJvirtual::22237-1https://scienti.minciencias.gov.co/cvlac/visualizador/generarCurriculoCv.do?cod_rh=0001439204virtual::22237-168838014-e95b-41af-819e-1a5e692f3a9bvirtual::22237-168838014-e95b-41af-819e-1a5e692f3a9bvirtual::22237-1ORIGINALRecognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of Colombia.pdfRecognition and characterization of potential soft tissue from a fossil ichthyosaur found in the Early Cretaceous (Barremian - Aptian) of 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