Camp dels Ninots Skeletons Reveal a Much More Complex Evolutionary History of Bovids Than Previously Thought

An international study provides key evidence for understanding the evolution of European bovids around 4 million years ago. The research focuses on the species Parabos tigneresi, documented at the Camp dels Ninots fossil site (Caldes de Malavella, La Selva). This locality is crucial for investigating continental ecosystems in southern Europe during a period that remains poorly represented in the European fossil record.

The fossil assemblage analysed includes a minimum of 14 individuals, several of which are represented by nearly complete skeletons preserved in anatomical connection, making it an exceptional record within the European context. This remarkable preservation has enabled detailed anatomical and comparative analyses, including computed tomography (CT) scanning and morphometric studies, placing this animal in a strategic position within the evolutionary history of the Bovinae, the group that includes modern bison, buffalo and cattle.

It was a large-bodied bovid, with an estimated body mass of approximately 500 kg, exceeding that of any other comparable bovid currently known from this time period.

The research, published in the international journal PLOS One, was led by Dr Leonardo Sorbelli, a postdoctoral researcher at the Museum für Naturkunde Berlin, with the participation of Dr Bruno Gómez de Soler and Dr Gerard Campeny, researchers at IPHES-CERCA and co-directors of the Camp dels Ninots research project, as well as researchers from the Universitat Autònoma de Barcelona (UAB), the Università degli Studi di Firenze, the Universitat Rovira i Virgili (URV), and the Museum für Naturkunde Berlin.

The results indicate that Parabos tigneresi exhibits a combination of traits that challenges the classical evolutionary models established to date. The researchers suggest that it may represent either one of the earliest European members of the Bovini or one of the last representatives of the older Tragoportacini lineage. This duality demonstrates that the evolution of these mammals in Europe was not a linear process, but rather a much more gradual and complex one than previously assumed. In this regard, the study identifies the Early Pliocene as a key period in the emergence and expansion of the first large bovids across Eurasia.

A Revision That Reshapes Our Understanding of Pliocene Bovids

Beyond the description of the species itself, the study presents a comprehensive systematic revision of the genera Parabos and Alephis, two groups that have generated scientific debate for decades. Thanks to this research, fossils previously assigned to the genus Alephis have now been reassigned to Parabos, a decision that contributes to redefining the classification of European bovids from this period.

“The morphological characteristics of Parabos tigneresi are a mixture of primitive and derived traits and do not fit within the traditional framework. This species appears to occupy an evolutionary position midway between the Tragoportacini and the Bovini, pointing to a much more complex evolutionary history than previously recognised,” explains Dr Leonardo Sorbelli.

In this context, the material recovered from Camp dels Ninots plays a fundamental role: not only does it strengthen the taxonomic assignment of the species, but it also provides a unique window into how bovids diversified during this period.

An Exceptional Fossil Assemblage on a European Scale

This exceptional state of preservation has enabled a detailed study of the cranial anatomy, dentition and postcranial skeleton, as well as morphometric analyses and precise body-mass estimates. Such a level of detail provides a solid basis for addressing evolutionary questions that often remain unresolved at sites where fossil remains are highly fragmented.

“This study finally brings together one of the most emblematic palaeontological assemblages from Camp dels Ninots. We recovered the first bovid skeleton in 2004,” recalls Dr Bruno Gómez de Soler. According to the researcher, “this is a long-awaited publication that reaffirms the scientific significance of the site.”

An Adaptable Bovid in a Changing Environment

The research also provides unprecedented information about the palaeoecology of this species. The characteristics of its dentition indicate a mixed feeding strategy, while the anatomy of its limbs suggests considerable versatility in moving through different habitat types.

These findings confirm a remarkable degree of ecological plasticity: the animal was adapted both to humid, densely vegetated environments and to more open landscapes. This interpretation is consistent with the palaeoenvironmental reconstruction of Camp dels Ninots, which depicts an ancient lake system surrounded by heterogeneous and dynamic landscapes.

The researchers propose that this early increase in body size may have been linked to the climatic and environmental changes that transformed Europe during the Pliocene, a period characterised by significant ecological fluctuations and the reorganisation of continental ecosystems.

Camp dels Ninots: A Unique Window into the Pliocene

The Camp dels Ninots fossil site is located within the crater of a phreatomagmatic explosion volcano, known as a maar. Over time, the crater became filled with lacustrine sediments deposited under anoxic conditions (absence of oxygen), a key factor that enabled the exceptional preservation of fossil remains.

After more than two decades of systematic excavations, this ancient maar has yielded complete and articulated skeletons of both large and small vertebrates, establishing itself as one of the key sites for reconstructing continental ecosystems of the European Pliocene. Within this framework, the Parabos tigneresi collection stands out for both its abundance and its extraordinary state of preservation.

Declared a Cultural Asset of National Interest (BCIN) because of its scientific importance, the site also serves as a model for public outreach and knowledge transfer.

“After 23 years of continuous work, Camp dels Ninots has developed around a research team of international excellence, capable of addressing virtually every discipline within the Earth Sciences,” says Dr Gerard Campeny. According to the researcher, the success of the project lies in “a clear commitment to heritage valorisation and social impact, thanks to initiatives such as Espai Aquae and the collaboration of the Caldes de Malavella Town Council.”

Article Reference

Sorbelli, L., Bibi, F., Madurell-Malapeira, J., Grandi, F., Moreno, E., Oms, O., Campeny, G., & Gómez de Soler, B. (2026). First of a Line or Last of a Dynasty? Parabos tigneresi and the Evolution of Eurasian Bovinae in the Early Pliocene. PLOS One. https://doi.org/10.1371/journal.pone.0340256

Funding

This research was supported by the Departament de Cultura de la Generalitat de Catalunya through the four-year research project (CLT009/22/000043) and the research group 2021 SGR 01238 (AGAUR), as well as by the Caldes de Malavella Town Council. Additional funding was provided by the Spanish Ministry of Science, Innovation and Universities through projects PID2024-157622NB-I00 and PID2024-156295NB-I00, and by the Universitat Rovira i Virgili (2023-URV-01238). The research also received support through the María de Maeztu Excellence Accreditation (CEX2024-01485-M), funded by the Spanish Ministry of Science, Innovation and Universities and managed by the State Research Agency (AEI).