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Three-dimensional analysis of the titanosaurian limb skeleton: implications for systematic analysis

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Abstract

The titanosaurian appendicular skeleton exhibits morphological similarities among different clades and its osteological information is usually less taxonomically meaningful than those of other regions of the skeleton. There is a probable morphological convergence due to morphofunctional similarities between members of different titanosaurian groups. In addition, higher intraspecific variability has hindered the assessment of similar forms such as the Late Cretaceous titanosaurians of the Ibero-Armorican domain. The use of 3D-geometric morphometrics and discriminant analyses on an abundant sample of titanosaurian limb elements from the Lo Hueco site and other titanosaurian taxa has been able to characterize the differences between several sauropod clades with the control of the intraspecific variability. Similar methods with the use of surface analyses of other titanosaurs enabled the recognition of morphological similarities congruent with morphofunctional convergences between one of the lithostrotian morphotypes identified in Lo Hueco (the Morphotype II) and some gracile colossosaurs such as Mendozasaurus neguyelap. In contrast, Morphotype I at Lo Hueco present a more typical titanosaurian morphology, resembling Jainosaurus cf. septentrionalis. The use of discriminant analyses allowed us to distinguish Colossosauria on the basis of limb morphology for the first time. We also observed that Colossosauria and Saltasauridae diverge from a more typical titanosaurian non-autopodial limb skeleton. Our results also suggest a highly different and specialized limb skeleton for the Saltasauridae in comparison with other sampled titanosaurians. The use of surface semilandmarks and discriminant analyses also allowed us to propose several new potential osteological characters for use in phylogenetic analyses through the maximization of differences between the sampled clades.

Resumen

El esqueleto apendicular de los titanosaurios presenta similitudes morfológicas entre clados muy diversos y su información osteológica es de menor utilidad para taxonomía respecto a otras regiones del esqueleto. Es possible que se produzca una convergencia morfológica debido a similitudes morfofuncionales entre miembros de los diferentes grupos de titanosaurios. Además, la elevada variabilidad intraespecífica ha dificultado la diferenciación entre formas de titanosaurio morfológicamente similares en el Cretácico tardío del dominio Ibero-Armoricano. El uso de técnicas de morfometría geométrica 3D así como análisis discriminante en una muestra abundante de ejemplares apendiculares del yacimiento de Lo Hueco y otros taxones de titanosaurio ha permitido caracterizar algunas diferencias entre los distintos clados de titanosaurios con el control de la variabilidad intraespecífica. Estos métodos se aplicaron también en análisis de superficies permitiendo reconocer similitudes morfológicas congruentes con convergencias morfofuncionales entre uno de los morfotipos de lithostrotios identificados en Lo Hueco (Morfortipo II) y formas gráciles de colossosaurios como Mendozasaurus neguyelap. Sin embargo, el Morfotipo I de Lo Hueco presentaría una morfología titanosauriana más típica, asemejándose a Jainosaurus cf. septentrionalis. El uso de análisis discriminantes permitió distinguir Colossosauria en base a la morfología de sus extremidades por primera vez. También se puede observar que la morfología de Colossosauria y Saltasauridae divergen de morfologías apendicular no autopodiales más típicamente titanosaurianas. Nuestros resultados sugieren que las grandes diferencias morfológicas de las extremidades de Saltasauridae con otros clados se debería a su especialización en comparación a otros titanosaurios muestrados en este estudio. El uso de semilandmarks de superficie y análisis discriminante permitió proponer una serie de potenciales caracteres osteológicos nuevos, útiles para el análisis filogenético, gracias a maximizar las diferencias existentes entre los diferentes clados muestreados.

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Code availability

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Acknowledgements

This research was funded by project CGL2012-35199 and CGL2015-68363-P from the Ministry of Economy and Competitiveness of Spain. Research of A. Páramo was funded by contract BES-2013-065509 granted by the Ministry of Economy and Competitiveness of Spain. Preparation and study of material from Lo Hueco was possible under projects SBPLY/15/180601/000045, SBPLY/16/180801/000017, SBPLY/17/ 180801/000063 of the Junta de Comunidades de Castilla-La Mancha, Spain. We want to thank also F. Marco-Fernández and the Conservacion GBE team as well as the alumni of Methodology of Archaeological Materials, Conservation and Restoration of Cultural Heritage degree (Fine Arts Faculty, UCM, Spain) and the alumni from the Taller de Restauración Paleontológica I, II & III, Cuenca (C-LM, Spain). Access to Argentinean titanosaurian material was possible thanks to the funding EEBB-I-16-11875 from the Ministry of Economy and Competitiveness of Spain. Access to NHM collections was possible thanks to funding of European Union SYNTHESYS program 2016. Ref. GB-TAF-6153. Research of P. Mocho was funded by FCT/MCTES for one CEEC Individual contract (CEECIND/00726/2017). We want to also thank the following people for access to the sampled specimens under their care to S. Langreo from the Museo de Paleontología de Castilla-La Mancha (Cuenca, Spain); J.A. Ramírez de la Peciña and C. Corral from the MCNA (Vitoria, Spain); R. Coria from the MCF (Plaza Huincul, Argentina); L. Filippi from the MRS (Rincón de los Sauces, Argentina); M. Reguero from the MLP (La Plata, Argentina) and A. Otero from the CONICET (La Plata, Argentina); S. Devincenzi from the IANIGLA (Mendoza, Argentina) and B. González Riga from the UNCUYO (Mendoza, Argentina); C. Muñoz from the MPCA (Cipolletti, Argentina) and I. Cerda from the CONICET (Cipolletti, Argentina); P. Ortiz from the PVL (Tucumán, Argentina); A.G. Kramarz and M. Ezcurra from the MACN (Buenos Aires, Argentina). We want to also thank P. Ullmann and an anonymous reviewer for their commentaries and suggestions that allowed to improve this article.

Funding

This research was funded by project CGL2012-35199 and CGL2015-68363-P from the Ministry of Economy and Competitiveness of Spain. Work from Páramo, A. was funded by contract BES-2013-065509 granted by the Ministry of Economy and Competitiveness of Spain. Preparation and study of Material from Lo Hueco was possible under projects SBPLY/15/180601/000045, SBPLY/16/180801/000017, SBPLY/17/ 180801/000063 of the Junta de Comunidades de Castilla-La Mancha, C-LM, Spain. Access to Argentinean titanosaurian material was possible thanks to the funding EEBB-I-16-11875 from the Ministry of Economy and Competitiveness of Spain. Access to NHM collections was possible thanks to the funding European Union SYNTHESYS program 2016. Ref. GB-TAF-6153. Work by Mocho, P. was funded by FCT/MCTES through one CEEC Individual contract (CEECIND/00726/2017).

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Authors and Affiliations

  1. Grupo de Biología Evolutiva (Ref. GI314), Departamento de Física Matemática y Fluidos, Facultad de Ciencias, Universidad Nacional de Educación a Distancia, Madrid, Spain

    Adrián Páramo, Pedro Mocho & Francisco Ortega

  2. Instituto Dom Luiz, Faculdade de Ciências da Universidade de Lisboa, Lisbon, Portugal

    Pedro Mocho

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  1. Adrián Páramo
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PA: conceptualization, formal analysis, investigation, methodology, resources, writing—original draft, writing—review and editing. MP: investigation, resources, writing—review and editing. OF: conceptualization, funding acquisition, investigation, project administration, supervision, writing—review and editing.

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Correspondence to Adrián Páramo.

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Páramo, A., Mocho, P. & Ortega, F. Three-dimensional analysis of the titanosaurian limb skeleton: implications for systematic analysis. J Iber Geol 46, 369–402 (2020). https://doi.org/10.1007/s41513-020-00139-8

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