A new giraffid (Mammalia, Ruminantia, Pecora) from the late Miocene of Spain, and the evolution of the sivathere-samothere lineage

Giraffids include the only living giraffomorph ruminants and are diagnosed by the presence of bi-lobed canines and a special type of epiphyseal cranial appendages called ossicones. The family Giraffidae ranges from the latest early Miocene until today. However they are currently extant relics with only two living representatives, the African genera Okapia and Giraffa. Giraffids were much more diverse and widespread in the past, with more than 30 fossil species described. For the past decades a number of studies intended to resolve the phylogenetic relationships of the family, but due to the lack of really good cranial material no clear consensus was reached regarding the phylogenetic relationships amongst the different members of the group. The exceptionally complete remains of a new large giraffid from the late Miocene of Spain, Decennatherium rex sp. nov., allows us to improve and reassess giraffid systematics, offering a lot of new data, both anatomic and phylogenetic, on the large late Miocene giraffids of Eurasia. The results of our cladistic analysis show Decennatherium as a basal offshoot of a clade containing the gigantic samotheres and sivatheres, characterized by the presence of a Sivatherium-like ossicone-plan among other features. Decennatherium thus offers the most ancient evidence of this Sivatherium-plan and firmly establishes the early morphological patterns of evolution of a sivathere / samothere-clade that is defined as the less inclusive clade that contains Decennatherium and Sivatherium. Finally, this large group of four-ossiconed giraffids evolutionarily links Miocene Europe and Africa indicating vicariance / migration processes among the giraffid genetic pools separated by the Mediterranean Sea.MR acknowledges a FPI predoctoral grant (2012-2016) (Spanish Government MINECO http://www.mineco.gob.es/ codes: BES-2012-052589, EEBB-I-14-07947, EEBB-I-15-09349, EEBB-I-16-10617) as well as the EEBB-FPI grant program 2013, 2014 and 2015. IMS acknowledges the CERCA Program (Generalitat de Catalunya) and the CGL2016-76431-P research project by the Spanish Ministerio de Economía y Competitividad (http://www.mineco.gob.es/). This study was supported by the Spanish Ministerio de Economía y Competitividad (research projects CGL2011-25754 (2012-2015) and CGL2015-68333-P (2016-2019), http://www.mineco.gob.es/).Peer Reviewe