New Early Miocene Material of Iberictis, the Oldest Member of the Wolverine Lineage (Carnivora, Mustelidae, Guloninae)

  • Alberto ValencianoEmail author
  • Juan Abella
  • David M. Alba
  • Josep M. Robles
  • María A. Álvarez-Sierra
  • Jorge Morales
Original Paper


We describe new dental remains of the genus Iberictis (Carnivora: Mustelidae) from the late early Miocene of the Iberian Peninsula. The new fossils of Iberictis azanzae from Artesilla (16.5–16.3 Ma, MN4; Calatayud-Teruel Basin, Zaragoza, Spain) add important morphological information about this species. Material from another species, Iberictis buloti, is described from els Casots (16.5–16.3 Ma, MN4; Vallès-Penedès Basin, Barcelona, Spain). This material constitutes the most complete sample of Iberictis and the first record of I. buloti in the Iberian Peninsula. Our analyses confirm the taxonomic validity of both species, and confirm the more plesiomorphic status of I. buloti compared to I. azanzae. Re-examination of large mustelid Miocene genera (Dehmictis, Ekorus, Eomellivora, Hoplictis, Iberictis, Ischyrictis, and Plesiogulo) and their inclusion for the first time in a cladistic analysis indicate that Iberictis is the sister taxon of Plesiogulo, and that these genera constitute the sister group of the extant wolverine (Gulo gulo). Our analysis thus confirms a close relationship between the early Miocene Iberictis, the late Miocene Plesiogulo, and the Plio-Pleistocene Gulo. Iberictis is the oldest member of Gulonini, the total clade of wolverines, thereby tracking the fossil record of this clade back to the early Miocene. We further propose a new systematic arrangement for the aforementioned large Miocene mustelids into the subfamilies Guloninae, Mellivorinae, and Mustelinae.


Mustelidae Neogene Gulo Wolverine Plesiogulo Iberictis Guloninae 



We would like to thank the following curators and collection managers for access to comparative material under their care: E. Westwig and J. Galkin (AMNH), D. Kalthoff (NRM), A. Prieur (FSL), C. Argot (MNHN), J.O.R. Ebbestad and V. Berg-Madsen (PMU), D. Lunde (USNM), U.B. Göhlich (NHMW), J.I. Canudo (MPZ), and E. López Errasquin and S. Fraile (MNCN). We are grateful to L. Werdelin (NRM) for kindly allowing us to study the cast of the skull and mandibles of Ekorus ekakeran. We further thank M. March and J. Galindo for assistance with the ICP collections; G. Rössner (BSPG) for the pictures of Plesiogulo sp. from Paşalar (Turkey); S. Mayda (Ege University, Turkey) for the pictures of Paralutra jaegeri from Ravensburg and Steinheim (Germany), Ischyrictis mustelinus from La Grive-Saint-Alban (France) and Erkertshofen 2 (Germany), and Dehmictis vorax from Wintershof-West (Germany); B. Azanza (Universidad de Zaragoza) for the pictures of old material of I. azanzae; J.I. Canudo (MPZ) for cataloguing the new material of I. azanzae; S. Peigné (MNHN) and the photographer Philippe Loubry (UMR7207, MNHN) for the pictures of I. buloti from Pellecahus (France); S. Govender and R. Govender (SAM) for the pictures of Plesiogulo monspessulanus from Langebaanweg; and L. Costeur (NMB) for the pictures of Hoplictis noueli from Artenay (France). We also are grateful to M. Sotnikova (Geological Institute, Russian Academy of Sciences, Moscow, Russia) for her help concerning Gulo minor, and to M. Pina (ICP) for assistance with the pictures of I. buloti from els Casots. A.V. was researcher in formation in the CSIC program JAE-PRE_CP2011 (CSIC program “Junta para la ampliación de estudios”), co-funded by the European Social Fund. This research received support by A.V. from the SYNTHESYS3 Project (SYNTHESYS; AT-TAF-5457), which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Programme, and the European Union’s Seventh Framework Programme (FP7/2007-2013) under grant agreement n° 226506 (SYNTHESYS; SE-TAF-3637). A.V. also received support by an American Museum of Natural History Collection Study Grant Program 2014. The support of the DST-NFR Centre of Excellence in Palaeosciences (CoE-Pal) toward this research for A. V. (COE2018-09POST) is hereby acknowledged. Opinions expressed and conclusions arrived at, are those of the author and are not necessarily to be attributed to the CoE. This study was also supported by CGL2015-68333-P (MINECO/FEDER-UE), Agencia Estatal de Investigación–European Regional Development Fund of the European Union (CGL2016-76431-P, AEI/FEDER, EU), the Generalitat de Catalunya (CERCA Programme), and the research group UCM 910607. We are also indebted to the Editor-in-Chief J.R. Wible, and both J. Baskin and one anonymous reviewer for their useful comments and suggestions, which significantly improved the original manuscript.We want to express our deepest gratitude to the late Stéphane Peigné (MNHN), a colleague, friend and a recognized world expert in Neogene carnivorans, who passed away during the development of this project.

Supplementary material

10914_2018_9445_MOESM1_ESM.docx (58 kb)
ESM 1 Descriptions of dental characters used in the phylogenetic analysis performed in this work. (DOCX 58 kb)
10914_2018_9445_Fig11_ESM.png (388 kb)

Character-taxon matrix used in the phylogenetic analysis performed in this work. (PNG 388 kb)

10914_2018_9445_MOESM2_ESM.tif (29 mb)
High Resolution (TIF 29 mb)
10914_2018_9445_MOESM3_ESM.nex (7 kb)
ESM 3 Character-taxon matrix in nexus format. (NEX 6.78 kb)


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Copyright information

© Springer Science+Business Media, LLC, part of Springer Nature 2018
corrected publication August/2018

Authors and Affiliations

  1. 1.Departamento de Geología Sedimentaria y Cambio MedioambientalInstituto de Geociencias (CSIC, UCM)MadridSpain
  2. 2.Departamento de Geodinámica, Facultad de Ciencias Geológicas UCMEstratigrafía y Paleontología Universidad Complutense de Madrid (UCM)MadridSpain
  3. 3.Research and Exhibitions Department, Iziko Museums of South AfricaCape TownSouth Africa
  4. 4.Department of Biological Sciences, Palaeobiological Research GroupUniversity of Cape TownCape TownSouth Africa
  5. 5.Facultad Ciencias del Mar, Edificio Instituto de Investigacion Científica y Desarrollo Tecnológicon (INCYT), Segundo PisoUniversidad Estatal Península de Santa ElenaLa LibertadEcuador
  6. 6.Institut Català de Paleontologia Miquel Crusafont, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UABUniversitat Autònoma de BarcelonaBarcelonaSpain
  7. 7.Departamento de PaleobiologíaMuseo Nacional de Ciencias Naturales-CSICMadridSpain

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