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

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RESEARCH PAPER

Three‑dimensional analysis of the titanosaurian limb skeleton: implications for systematic analysis Adrián Páramo1   · Pedro Mocho1,2   · Francisco Ortega1  Received: 2 March 2020 / Accepted: 15 September 2020 © Universidad Complutense de Madrid 2020

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. Keywords  Titanosauria · Lo Hueco · Geometric morphometrics · Cretaceous · High density surface semilandmarks 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