Aurochs roamed along the SW coast of Andalusia (Spain) during Late Pleistocene

Abstract

In the Iberian Peninsula the fossil record of artiodactyls spans over 53 million years. During the Pleistocene, wild cattle species such as Bison and especially Bos became common. In Late Pleistocene, the aurochs (Bos primigenius) was widespread and the only bovine living along the large river valleys of southern Iberia. Although commonly found in fossil sites and especially in cave bone assemblages, the trace fossil record of aurochs was known worldwide only from the Holocene. Large bovine and roe deer/caprine tracks were found in at least five horizons of the early Late Pleistocene (MIS 5) beach and eolian deposits of Cape Trafalgar (Cadiz Province, South of Spain). The large bovine tracks are formally described as Bovinichnus uripeda igen. et isp. nov. and compared with the record of aurochs tracks, large red deer tracks and steppe bison biogeographical distribution in Iberia. Aurochs were the most likely producers of the newly described Trafalgar Trampled Surface (TTS) and some of the large artiodactyl tracks in the Matalascañas Trampled Surface, representing the oldest aurochs track record known. This new evidence, together with comparisons with the record of possible aurochs tracks in the Mid-Late Pleistocene coastal deposits from the Asperillo cliff section in Matalascañas (Huelva Province, SW Spain) and bone assemblages known in Gibraltar, point to a recurrent use of the coastal habitat by these large artiodactyls in SW Iberia.

Figure 1

(a) Location of Cape Trafalgar and Matalascañas tracksites in SW Iberian Peninsula, Image adapted from GoogleEarth©; (b) The MTS at Matalascañas, Province of Huelva (SW Spain); (c) the new TTS in Cape Trafalgar (Province of Cadiz). Map in (a) (left) and modifications to the photograph in (c) have been drawn with CorelDraw 12 (https://www.coreldraw.com/la/).

Figure 2

Stratigraphy of the TTS in Cape Trafalgar (Province of Cadiz). (a) Geological map of the Cap Trafalgar area (adapted from the Geological Map of Spain 1:50,000, Map no.1073 Vejer de la Frontera44, source: “© Geological Survey of Spain (IGME)”; (b) detailed stratigraphic column of Cape Trafalgar transition between beach and dune facies and location of the TTS and other trampled horizons (I–IV); rhizolith horizons corresponding to incipient paleosols are indicated (legend: 1—possible hominin footprint; 2—Pecoripeda isp.; 3—bioturbated lamina; 4—large footprint related to Palaeoloxodon antiquus; 5—rhizoliths; 6—Bovinichnus uripeda); (c) thin horizontal and densely branched rhizoliths from the second eolianite unit; (d) insect burrows in linear clusters preserved as convex hyporelief; ruler is 15 cm. Legend in (a), figure (b) and modifications to the photograph (d) have been made with CorelDraw 12 (https://www.coreldraw.com/la/).

Figure 3

Graphical representation and data on measurements taken in the TTS and the MTS: (a,b) Distribution of tracks in the main area of the TTS; (c) bivariate plot of track length/track width ratio (n = 18 in the TTS and n = 17 in the MTS). The track length included dew claw impressions if present; (d) boxplot of track width variation (n = 18 for the TTS and n = 17 for the MTS); (e) rose diagram with the orientation of the hoofprints in the TTS (n = 14). Figure (b) has been drawn with CorelDraw 12. (https://www.coreldraw.com/la/).

Figure 4

3D models of the Bovinichnus uripeda holotype (TTS10) and paratype TTS8: (a) Orthogonal view of the natural colours textured 3D model of the TTS10 with the general morphology depicted from the contour lines map, equidistance of 1 mm (b); (c) false colour DSM in orthogonal view showing deeper areas in the heel side of the hooves and the tips resulting from the impulse created in the foot-off event. The circular-to-elliptical dew claw imprints are also evident behind the hooves; (d) oblique view of the previous DSM to complement previous observations; (e) oblique view of the textured natural colour 3D model of the paratype TTS 8; (f) orthogonal view of the paratype TTS 8 where the deepest part of the hoofprints is displaced to their anterior part reflecting the pressure angle exerted by the limb on the substrate during the foot-off event. Scale bar is 150 mm. The 3D model of the images (a–d) were produced by the software WebODM 2.1.0, https://github.com/OpenDroneMap/WebODM; MeshLab 2020.12, (https://www.meshlab.net/); CloudCompare 2.11.0, (https://www.danielgm.net/cc/) and the 3D model of the images (e,f) were produced with the software Meshroom 2021.1.0, (https://alicevision.org/#meshroom); MeshLab 2020.12, (https://www.meshlab.net/); CloudCompare 2.11.0, (https://www.danielgm.net/cc/).

Figure 5

Bovinichnus uripeda igen. nov. et isp nov. from the Late Pleistocene (MIS 5c) of Cape Trafalgar: (a) well-preserved track as concave epirelief (holotype TTS10) showing the dewclaws; ruler is 15 cm; (b) convex hyporelief preservation in the trampled horizon I, evidencing the narrow ridge between toes; (c) tramped level IV with large bifid tracks attributed to aurochs in section (above) and interpretation of the sedimentary deformation (below). Scale bar is 15 cm. The interpretation of (c) (below) was drawn with CorelDraw X7 (https://www.coreldraw.com/la/).

Figure 6

Large artiodactyl tracks from the MTS. (a) Trackway T1 attributed to aurochs based on the large size of the squared tracks and the preservation of the dewclaws as rectangular imprints; (b) 3D model of a tetradactyl track from the MTS interpreted as Bovinichnus uripeda, vertical view of the texturized 3D model in natural colours; (c) vertical view from the previous track made of the false colours DSM (cold and hot colours represent depth gradient, from deeper to shallower, respectively); (d) Bovinichnus uripeda, large track; (e) manus-pes couple, note the dewclaws; (f) large cervid track found in the MTS for comparison, with slender toes with sharp tips, well splayed, with almost linear outlines. Scale bar is 10 cm. The 3D model of the images (b,c) were produced by the software Meshroom 2021.1.0 (https://alicevision.org/#meshroom); MeshLab 2020.12 (https://www.meshlab.net/); CloudCompare 2.11.0 (https://www.danielgm.net/cc/).

Figure 7

Revision of the valid artiodactyl ichnogenera with visual comparison of morphologies, age and locality (country). All tracks have been redrawn from the original references where they were described for the first time. (m) manus, (p) pes. Anoplotheriipus left pes; Diplartiopus left pes; Bothriodontipus pes (not indicated which side); Lamaichnum manus (not indicated which side); Paracamelichnum right manus; Fustinianapodus left manus and pes; Suidichnus overprinting of left manus by corresponding pes; Bovinichnus right track; for other ichnogenera autopods were not specified/shown no morphological distinction in the references. This figure has been drawn with CorelDraw X6 (https://www.coreldraw.com/la/).

Figure 8

Reconstitution of the size of the aurochs based on the maximum size of the tracks found in the TTS, in a context of beach environment where they were produced (artwork of José María Galán).