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Marugán-Lobón, J., Martín-Abad. H., Buscalioni, A.D. 2023. The Las Hoyas Lagerstätte: a palaeontological look to an Early Cretaceous wetland. JSG (in the press).

Marugán‐Lobón, J., Gómez-Recio, M., Nebreda, S.M. 2022. The geometry of Synapsid skull disparity. Historical Biology 34(8):1692-1700. 

Herrera-Castillo C.M., Moratalla J.J., Belaústegui Z., Marugán-Lobón J., Martín-Abad H., et al. (2022) A theropod trackway providing evidence of a pathological foot from the exceptional locality of Las Hoyas (upper Barremian, Serranía de Cuenca, Spain). PLOS ONE 17(4): e0264406.

Marugán‐Lobón, J., Nebreda, S.M., Navalón, G., Benson, R.J.B. 2021. Beyond the beak: Brain size and allometry in avian craniofacial evolution. Journal of Anatomy 240(2):197:209. 

Nebreda, S.M., Hernández Fernández, M., Marugán-Lobón, J. 2021.‘Dinosaur-bird’macroevolution, locomotor modules and the origins of flight. Journal of Iberian Geology 47 (3), 565-574.

Navalón, G., Nebreda, S.M., Bright, J.A., Fabbri, M., Benson, R.J.B., Bhullar, B-A., Marugán-Lobón, J., Rayfield, E.J. 2021. Craniofacial development illuminates the evolution of nightbirds (Strisores). Proceedings of the Royal Society B 288 (1948), 20210181.                

Bell, A., Marugán-Lobón, J., Navalón, G., Nebreda, S.M., DiGuildo, J., Chiappe, L.M. 2021. Quantitative analysis of morphometric data of pre-modern birds: phylogenetic versus ecologic signal. Frontiers in Earth Science 9, 534.

Ksepka, D.T., A.M.Balanoff, N.A. Smith, G.S.Bever. B.A.S. Bhullar, E. Bourdon, E.L. Braun, J.G. Burleigh, J.A.Clarke, M.W. Colbert, J.R. Corfield, F.J.Degrange, V.L.De Pietri, C.M. Early, D.J.Field, P.M. Gignac, M.E. L. Gold, R.T. Kimball, S. Kawabe, L. Lefebvre, J. Marugán-Lobón, C.S. Mongle, A. Morhardt, M.A. Norell, R.C.Ridgely, R.S. Rothman, R.P. Scofield, C.P. Tambussi, C.R.Torres, M. van Tuinen, S.A. Walsh, A. Watanabe, L.M. Witmer, A.K. Wright, L.E. Zanno, E.D.Jarvis, and J.B.Smaers. 2020. Tempo and pattern of avian brain size evolution. Current Biology 30: 2026-2036.

Nebreda, S.M., Navalón, G., Menéndez, I., Sigurdsen, T., Chiappe, L.M., Marugán-Lobón, J. 2020. Disparity and macroevolutionary transformation of the maniraptoran manus. In Pittman, M. and X. Xu (editors), Pennaraptoran theropod dinosaurs: past progress and new frontiers: 183-203. New York: Bulletin of the American Museum of Natural History.

Navalón, G., Marugán-Lobón, J., Bright, J. Clooney, C.R., Reyfield, EJ. 2020. The consequences of craniofacial integration for the adaptive radiations of Darwin’s finches and Hawaiian honeycreepers. Nature Ecology and Evolution.


Chinsamy, A, Marugán-Lobón, J., Serrano, F., Chiappe, L.M. 2019. Osteohistology and Life History of the Basal Pygostylian, Confuciusornis sanctus. Anatomical Record,

Costa-Pérez, M., Moratalla, J.J., Marugán-Lobón, J. 2019. Studying bipedal dinosaur trackways using Geometric Morphometrics. Palaeontologia Electronica, 22.3.pvc-3.

Blanco-Moreno, C., Gomez, B., Marugán-Lobón, J., Daviero-Gomez, V., Buscalioni, A.D. 2019. A novel approach for the metric analysis of fern fronds: Growth and architecture of the Mesozoic fern Weischelia reticulata in the light of modern ferns. PloSONE

Bright, J., Marugán-Lobón, J., Cob, S., Rayfield, W. 2019. The multifactorial nature of beak and skull shape evolution in parrots and cockatoos (Psittaciformes). BMC Evolutionary Biology 19:104

Kaye, TG., Pittman, M, Marugán-Lobón, J, Martín-Abad. H, Sanz, JL, Buscalioni AD. 2019. Fully fledged enantiornithine hatchling revealed by Laser-Stimulated Fluorescence supports precocial nesting behavior Scientific Reports 9: 5006.

Navalón, Bright. J., Marugán-Lobón, J., Rayfield E. J. 2018 The evolutionary relationship between beak shape, biting performance, and feeding ecology in modern birds. Evolution.

Knoll, F., Chiappe, L.M., Sanchez, S., Garwood, R.J., Edwards, N.P., Wogelius, R.A., Sellers, W.I., Manning, P.L., Ortega, F., Serrano, F.J., Marugán-Lobón, J., Cuesta, E., Escaso, F., Sanz, J.L. 2018. A diminutive perinate European Enantiornithes reveals an asynchronous ossification pattern in early birds. Nature Communications. 9, 937 doi:10.1038/s41467-018-03295-9.


Serrano, F. J., Chiappe, L.M., Palmqvist, P., Figueirido, B., Marugán-Lobón, J., Sanz, L. 2018. Flight reconstruction of two European Enantionithines (Aves, Pygostylia) and the achievement of bounding flight in Early Cretaceous birds. Paleontology. doi: 10.1111/pala.12351


Navalón, G., Meng, Q., Marugán-Lobón, J., Zhang, Y. Wang, B., Xing, H., Liu, D., Chiappe, L.M. 2018. Diversity and evolution of the Confuciuornithidae: Evidence from a new 131-million-year-old specimen from the Huajiying Formation in NE China. Journal of Asian Earth Sciences. 152:12-22.


Moratalla J.J, Marugán-Lobón, J., Martín-Abad, H., Cuesta, E., Buscalioni, A.D. 2017. A new trackway made by a trotting theropod at the Las Hoyas fossil site (Early Cretaceous, Cuenca Province, Spain): Identification, bio-dynamics and palaeoenvironmental implications. Paleontologica Electronica. 20.3.59A 

Martín Abad, H., Blanco Moreno, C., Barrios de Pedro, S., Marugán-Lobón, J., Poyato Ariza, F.J., Delvene, G., Moratalla, J.J., Fregenal Martínez, M., Vullo, R., Cuesta, E., Iniesto, M., Barral, A., Gomez, B., Buscalioni, A.D. 2017. The exceptional fossil site of Las Hoyas (SPAIN) from an educational perspective. Geoheritage.

Fernández Montraveta, C., Marugán-Lobón, J. 2017. Geometric morphometrics reveals sex-differential shape allometry in a spider. PeerJ 5:e3617

Davies, ImranT.G., Rahman A., Lautenschlager, S., Cunningham, J.A., Asher, R.J., Barrett, P.R., Bates, K.T., Bengtson, S., Benson, R.B.J., Boyer, D.M., Braga, J., Bright, J.A., Claessens, L.P.A.M., Cox, P.G., Dong, X-P., Evans, A.E., Falkingham, P.L., Friedman, M., Garwood, R.J., Goswami, A., Hutchinson, J.R., Jeffery, N.S., Johanson, Z., Lebrun, R., Martínez-Pérez, C., Marugán-Lobón, J., O'Higgins, P.M., Metscher, B., Orliac, M., Rowe, T.B., Rücklin, M., Sánchez-Villagra, M.R., Shubin, N.H., Smith, S.Y., Starck, J. Stringer, C., Summers, A.P., Sutton, M.D., Walsh, S.G., Weisbecker, V., Witmer, L.M., Wroe, S., Yin, A., Rayfield, E.J., Donoghue P.C.J. 2017. Open data an digital morphology. Proceedings of the Royal Society B, Biological Sciences 284,  DOI: 10.1098/rspb.2017.0194 

Marugán-Lobón, J. 2017. The woodcock’s head: Resolving a morphological oddity using geometric morphometrics. Zoological Systematics. 42:102-107.


Bright, J., Marugán-Lobón, J., Cob, S., Rayfield, E. 2016. The shapes of bird beaks are highly controlled by nondietary factors. PNAS 113(19):5352-5357 doi: 10.1073/pnas.1602683113.

Marugán-Lobón J., Watanabe, A., Kawabe, S. 2016. Studying avian encephalization with geometric morphometrics. Journal of Anatomy, 229(2):191-203. doi: 10.1111/joa.12476.

Navalón, G., Marugán-Lobón, J., Chiappe, L.M., Sanz, J.L., Buscalioni, A.D. 2015. Soft-tissue and dermal arrangement in the wing of an Early Cretaceous bird: Implications for the evolution of avian flight. Scientific Reports 5:14864 | doi: 10.1038/srep14864.

Martin, T., Marugán-Lobón, J., Vullo, R., Martín-Abad, H., Zhe-Xi, L., Buscalioni, A.D. 2015. A Cretaceous Eutriconodont and integument evolution in Early mammals. Nature (526):380-384. doi:10.1038/nature14905

Chamero, B., Buscalioni, A.D., Marugán-Lobón, J., Sarris, I. 2014. 3D Geometry and Quantitative Variation of the Cervico-Thoracic Region in Crocodylia. Anatomical Record, 297(7):1278-1291. doi: 10.1002/ar.22926


Marugán-Lobón, J., Blanco-Miranda, D., Chamero, B., Martín Abad, H. 2014. On the importance of examining the relationship between shape data and biologically meaningful variables. Spanish Journal of Palaeontology, 28(2):139-148. ISSN 2255-0550 (Antigua revista Española de Paleontología).


Chiappe, L., Zhao B., O’Connor, JK., Chunling, G., Wang, C., Habib, M., Marugán-Lobón, J., Meng, Q. 2014. A new specimen of the Early Cretaceous bird : Insights into the aerodynamics and diet of a basal ornithuromorph, PeerJ 2:e234 doi: 10.7717/peerj.234

Esteve-Altava, B., Marugán-Lobón, J., Botella, H., Rasskin-Gutman, D. 2014. Random loss and selective fusion of bones originate morphological complexity trends in tetrapod skull networks. Evolutionary Biology, 41(1): 52-61. doi: 10.1007/s11692-013-9245-4

Esteve-Altava, B., Marugán-Lobón, J, Botella, H., Rasskin-Gutman, D. 2013. Grist for Riedl’s Mill: A Network Model Perspective on the Integration and Modularity of the Human Skull. Journal of Experimental Zoology: Part B, 320: 489-500. doi:10.1002/jez.b.22524


Marugán-Lobón, J., Chiappe, L.M., Farke, A. 2013. The variability of inner ear orientation in saurischian dinosaurs: Testing the use of semicircular canals as a reference system for comparative anatomy. PeerJ 1:e124 doi: 10.7717/peerj.124


Llandrés Serrano, M., Vullo, R. , Marugán-Lobón, J., Ortega, F., Buscalioni, A.D. 2013. An articulated hindlimb of a basal iguanodont (Dinosauria, Ornithopoda) from the Early Cretaceous Las Hoyas Lagerstätte (Spain). Geological Magazine, 150(3):572-576. doi:10.1017/S0016756813000095


Klingenberg, C., Marugán-Lobón, J. 2013. Evolutionary Covariation in Geometric Morphometric Data: Analyzing Integration, Modularity and Allometry in a Phylogenetic Context. Systematic Biology, 62:591–610. doi: 10.1093/sysbio/syt025


Chamero, B., Marugán-Lobón, J, Buscalioni, A.D., 2013. Pectoral girdle and forelimb variation in extant Crocodylia: the coracoid-humerus pair as an evolutionary module. Biological Journal of the Linnean Society. 108(3):600-618. doi: 10.1111/j.1095-8312.2012.02037.x


Chinsamy, A., Chiappe, L.M., Marugán-Lobón, J, Chunling G., Fengjiao, Z. 2013 Gender identification of the Mesozoic bird . Nature Communications, 4: 1381, doi: 10.1038/ncomms2377


Esteve-Altava, B., Marugán-Lobón, J, Botella, H., Rasskin-Gutman, D. 2013. Structural Constraints in the Evolution of the Tetrapod Skull Complexity: Williston’s Law Revisited using Network Models. Evolutionary Biology, 40(2):209-219. doi:10.1007/s11692-012-9200-9

Bravo A. M., Marugán-Lobón, J, 2012.Morphometric analysis of dinosaur eggshells: constraints of size on shape. Historical Biology, doi:10.1080/08912963.2012.744989

Sarris, I., Marugán-Lobón, J, Chamero, B., Buscalioni, A.D. 2012. Shape variation and allometry in the precloacal vertebral series of the snake Daboia russelli (Viperidae). International Journal of Morphology 4(30):1363-1368. 

Vullo, R., Marugán-Lobón, J, Kellner, A.W.A., Buscalioni, A.D., Gomez, B., de la Fuente, M., Moratalla, J.J. 2012. A New Crested Pterosaur from the Early Cretaceous of Spain: The First European Tapejarid (Pterodactyloidea: Azhdarchoidea). PLoS ONE, 7(7): e38900. doi:10.1371/journal.pone.0038900.

Buhllar, A., Marugán-Lobón, J, Racimo, F., Bever, G.S., Rowe, T.B., Norell, M.A., Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. , 487:223-226. doi:10.1038/nature11146

Esteve-Altava, B., Marugán-Lobón, J, Botella, H., Rasskin-Gutman, D. 2011. Network models in anatomical systems. Journal of Anthropological Sciences, 89:175-184. doi: 10.4436/jass.89016

Marugán-Lobón. J., Vullo, R. 2011. Feather Diversity in the Barremian (Early Cretaceous) of Las Hoyas. Comptes Redus Palevol, 10:219-223.  doi: 10.1016/j.crpv.2011.02.002

Marugán-Lobón J., Chiappe L. M., Ji, S. Zhou, Z., Chunling, G., Hu, D., Meng, Q. 2011.  Quantitative patterns of morphological variation in the appendicular skeleton of the Early Cretaceous bird Confuciursonis. Journal of Systematic Palaeontology, 9(1):91-101. doi: 10.1080/14772019.2010.517786


Chiappe L.M., Marugán-Lobón, J, Chimsany, A. 2010. Paleobiology of the Cretaceous Bird : A Reply to Peters & Peters (2010). Biology Letters, 6:529-530. doi: 10.1098/rsbl.2009.1057


Vullo R., Buscalioni, A., Marugán-Lobón, J , Moratalla, J.J. 2009. First pterosaur remains from the Early Cretaceous Lagerstätte of Las Hoyas, Spain: palaeoecological significance. Geological Magazine, 146(6):931-936. doi: 10.1017/S0016756809990525

Amaral, A.R., Coelho, M.M., Marugán-Lobón, J, Rohlf, F.J. 2009. Cranial shape differentiation in three closely related delphinid cetacean species: insights into evolutionary history. Zoology, 112(1):38-47. doi: 10.1016/j.zool.2008.03.001


Marugán-Lobón J., Buscalioni A. D. 2009. New insight on the anatomy and cranial architecture of the avian cranium. Anatomical Record, 292:364-370. doi: 10.1002/ar.20865

Chiappe L.M., Marugán-Lobón, J, Ji, & S., Zhou, Z. 2008. Life-history of a basal bird: Morphometrics of the Early Cretaceous . , 4(6):719-723. doi: 0.1098/rsbl.2008.0409


Buscalioni D.A., Fregenal, M.A., Bravo, A., Poyato-Ariza, F.J. Sanchíz B., Báez, A.M., Cambra Moo, O. Martín Closas, C., Evans, S.E., Marugán Lobón, J. 2008. The vertebrate assemblage of Buenache de la Sierra (Upper Barremian of Serranía de Cuenca, Spain) with insights into its taphonomy and paleoecology. Cretaceous Research, 29(4): 687-710. doi: 10.1016/j.cretres.2008.02.004


Cambra Moo, O. Chamero B.,Marugán-Lobón, J , Martínez-Delclós, X. Poyato-Ariza, F, Buscalioni, A. D. 2006. Estimating the ontogenetic status of an enantiornithine bird from the Lower Barremian of El Montsec, Central Pirenees, Spain. Estudios Geológicos, 62(1):241-248. ISSN: 0367-0449


Marugán-Lobón, J., Buscalioni, A. D. 2006. Avian skull morphological evolution: exploring exo- and endocranial covariation with two-block partial least squares. Zoology, 109:217-230. doi: 0.1016/j.zool.2006.03.005

Marugán-Lobón, J., Buscalioni, A. D. 2003. “Disparity and geometry of the skull in archosauria (reptilia: diapsida)”. Biological Journal of the Linnean Society, 80:67-88. doi: 10.1046/j.1095-8312.2003.00219.x

Chapters in Books


Buscalioni, A.D., Poyato-Ariza, F.J., Marugán-Lobón, J., Fregenal-Martínez, M., Sanisidro, O., Navalón, G. De Miguel, C. 2016. The Wetland of Las Hoyas. In: Las Hoyas: A Cretaceous Wetland. A multidisciplinary synthesis after 25 years of research on an exceptional fossil Lagerstatte from Spain. F. J. Poyato-Ariza & Buscalioni A.D. (eds.). Pfeil Verlag, Munich, Alemania. 262 pp.  ISBN 978-3-89937-153-6

Sanz J.L., Chiappe L.M., O’Connor J.K., Marugán-Lobón, J., Chamero B. 2016. Aves In: Las Hoyas: A Cretaceous Wetland. A multidisciplinary synthesis after 25 years of research on an exceptional fossil Lagerstatte from Spain. F. J. Poyato-Ariza & Buscalioni A.D. (eds.). Pfeil Verlag, Munich, Alemania. 262 pp. ISBN 978-3-89937-153-6

Marugán-Lobón, J., Vullo, R. 2016. Feathers. In: Las Hoyas: A Cretaceous Wetland. A multidisciplinary sythesis after 25 years of research on an exceptional fossil Lagerstatte from Spain. A multidisciplinary synthesis after 25 years of research on an exceptional fossil Lagerstatte from Spain. F. J. Poyato-Ariza & Buscalioni A.D. (eds.). Pfeil Verlag, Munich, Alemania. 262 pp. ISBN 978-3-89937-153-6

Vullo, R., Marugán-Lobón, J. 2016. The pterosaurs from Las Hoyas. In: Las Hoyas: A Cretaceous Wetland. A multidisciplinary synthesis after 25 years of research on an exceptional fossil Lagerstatte from Spain. F. J. Poyato-Ariza & Buscalioni A.D. (eds.). Pfeil Verlag, Munich. 262 pp. ISBN 978-3-89937-153-6

Marugán-Lobón, J. 2010. Combining Shape Data and Traditional Measurements with the 2B-PLS: Testing the Covariation Between Avian Brain Size and Cranial Shape Variation as an Example. In Eleewa, A. M. T. (ed), Morphometrics for Non-morphometricians. Lecture Notes in Earth Sciences 124. Springer-Verlag, Berlin Heidelberg, Alemania, pp.179-190. doi: 10.1007/978-3-540-95853-6_7.

Chamero, B., Marugán-Lobón, J. 2007. Aproximación al estudio de las proporciones de la extremidad posterior en aves y otros terópodos desde una perspectiva ontogenética. En: Cambra Moo, O. Martínez Pérez, C., Chamero Macho, B., Escaso Santos, F., Esteban de Trivigno, S., Marugán Lobón, J. Cantera Paleontológica. Diputación Provincial de Cuenca, Ediciones Provinciales, 53, pp. 111-121. I.S.B.N. 84-93557-37-4

Marugán-Lobón, J. 2005. Geometría craneal de aves modernas, morfometría, transformaciones y evolución. En: Meléndez G., Martínez-Pérez, C., Ros, S., Botella, H., y Plasencia, P. (eds.) Miscelánea Paleontológica, Publicaciones del Seminario de Paleontología de Zaragoza, 6, pp. 321-340. I.S.B.N: 84-96214-48-6

Marugán-Lobón J., Buscalioni A. D. 2004. Geometric Morphometrics in Macroevolution: morphological diversity of the skull in modern avian forms in contrast to some theropod dinosaurs. En: Elewa A.M.T. (ed.), Morphometrics in Paleontology and Biology. Springer-Verlag, Heildelber, Alemania, pp. 157-173. ISBN: 3-540-21429-1.

Marugán-Lobón, J. 2004. Diversidad y Disparidad en Macroevolución. En: Sanz García, J. L. (ed.): Aportaciones recientes en el conocimiento de la historia de la vida, Trabajos del VI Curso de Paleontología en Cuenca. Fundación de Cultura Ciudad de Cuenca. J. L. Sanz (Ed), Cuenca, pp.13-44. I.S.B.N. 97-88486-78-8353

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