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

1 Vol.:(0123456789) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports Aurochs roamed along the SW coast of Andalusia (Spain) during Late Pleistocene Carlos Neto de Carvalho1,2, Fernando Muñiz3, Luis M. Cáceres4*, Zain Belaústegui5, Joaquín Rodríguez‑Vidal4, João Belo6, Noel Moreira7, Mário Cachão2,8, Pedro P. Cunha9, Silvério Figueiredo10,11, José María Galán12, Yilu Zhang13, Paula Gómez4, Antonio Toscano4, Francisco Ruiz4, Samuel Ramírez‑Cruzado4, Francisco Giles‑Guzmán14, Geraldine Finlayson14,15,16, Stewart Finlayson14,15 & Clive Finlayson14,15,16,17 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. The artiodactyls are even-toed ungulates belonging to a diversified group of large land mammals that can be found nowadays all over the world, except Antarctica. The oldest fossils of even-toed ungulates date back to the early Eocene and were found in Europe, Asia and North America1. The oldest bone remains known in Europe were found in the Iberian Peninsula and are dated from this period2,3. However, their record of living activities, known through the study of trace fossils, such as tracks or excrements, is much more scattered, and known only since the late Eocene to early Oligocene in North America and Europe, including Spain4–7 (see the recent attribution of late Paleocene tracks to possible basal artiodactyls or tapiroids8). Fossil tracks, when safely attributed to a specific producer, can be complementary to the osteological record, providing additional data on geographical distribution of species, their paleoecology and biological behaviour. OPEN 1Naturtejo UNESCO Global Geopark, Geology Office of the Municipality of Idanha-a-Nova, Idanha a Nova, Portugal. 2Instituto D. Luiz, University of Lisbon, Lisbon, Portugal. 3Departamento de Cristalografía, Mineralogía y Química Agrícola, Universidad de Sevilla, Sevilla, Spain. 4Departamento de Ciencias de la Tierra, Universidad de Huelva, Huelva, Spain. 5Departament de Dinàmica de la Terra i de l’Oceà, Facultat de Ciències de la Terra, Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona (UB), Barcelona, Spain. 6Geosciences Center, FlyGIS-UAV Surveys, University of Coimbra, Coimbra, Portugal. 7Instituto de Investigação e Formação Avançada, Institute of Earth Sciences (ICT)-Pole of Évora, University of Évora, Évora, Portugal. 8Department of Geology, Faculty of Sciences, University of Lisbon, 1749-016 Lisboa, Portugal. 9Department of Earth Sciences, MARE-Marine and Environmental Sciences Centre, University of Coimbra, Coimbra, Portugal. 10Department of Archeology, Conservation and Heritage, CGeo-UC, Polytechnical Institute of Tomar, Tomar, Portugal. 11Centro Português de Geo-História e Pré-História, São Caetano, Portugal. 12Centro Administrativo del Acebuche, Parque Nacional de Doñana, Matalascañas, Spain. 13Academy of Natural Resources of Henan, Zhengzhou, China. 14The Gibraltar National Museum, Gibraltar, UK. 15Institute of Life and Earth Sciences, University of Gibraltar, Gibraltar, UK. 16Department of Life Sciences, Liverpool John Moores University, Liverpool, UK. 17Department of Anthropology, University of Toronto, Scarborough Campus, Toronto, Canada. *email: [email protected] 2 Vol:.(1234567890) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ The body fossil record of the bovine tribe is relatively common in Quaternary deposits, but rarely have tracks been attributed to these animals. In the Iberian Peninsula, artiodactyl tracks from Pleistocene beds were mostly attributed to goats, cervids, and wild boar9–16. Here we formally describe, for the first time, large artiodactyl tracks from two different sites in SW Spain ascribed to the early Late Pleistocene, safely attributed to aurochs (Bos primigenius). During the Late Pleistocene, the aurochs were widespread and the only wild cattle living along the large river valleys in southern Iberia. Artiodactyls are cloven-hoofed animals, which in most cases bear the body weight equally on two of their five toes: the third and fourth. We compare the morphology of very large cloven-hoofed tracks found during the study of Late Pleistocene eolianites in SW Iberia, with the ones of cervids and recent tracks made by Bos taurus. Although commonly found in fossil sites and especially in cave bone assemblages17–19, the trace fossil record of aurochs was only known from the Holocene of Great Britain, especially in tidal flat deposits from estuarine areas20,21. In SW Spain, the Cape Trafalgar and Asperillo cliff new tracksites (Fig.1) provide an opportunity to endorse paleoecological interpretations regarding the recurrent use of the coastal habitats by these massive grazers during the Quaternary. The track fossil record of artiodactyls in the Iberian Peninsula and the presence of aurochs in the SW In the Iberian Peninsula, the oldest cloven-hoofed tracks were identified in the Lower Miocene22. The first tracksites were described in the Late Pleistocene (MIS 3) of the Balearic Islands in Mallorca, Menorca and Formentera9,11,23, and attributed to the insular goat Myotragus balearicus Bate. Large cervid tracks were described in the Late Pleistocene eolianites of Gorliz (Basque coast, north Spain)10, in the Late Pleistocene (MIS 5–4) eolianites of SW Alentejo (Portugal)12–14, and in the Late Pleistocene of the Catalan Bay eolianite, in Gibraltar24. However, if the eolianites are confirmed to be MIS 3 in age, the more rounded cervid didactyl tracks from Gorliz could also be compared to those of the reindeer Rangifer tarandus, a species commonly found from the bone record in the northern Spain during the Pleniglacial25–27. Recently, the Matalascañas Trampled Surface (MTS) 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. corel draw. com/ la/). 3 Vol.:(0123456789) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ in the province of Huelva (SW Spain), dated from the early Late Pleistocene (MIS 5), was described showing abundant artiodactyl tracks which were attributed to wild boar, red deer and possibly aurochs15,16. Aurochs tracks have been studied especially in coastal deposits in England and Wales associated with early Mesolithic to early Bronze Age industries20,28–36. These tracks have been important to understanding the ecology of Homo sapiens and its predator–prey interactions, and in determining the onset of cattle domestication. However, they were never formally described from an ichnotaxonomical point of view, and only rarely figured in discussions. At Formby, Sefton Coast (Liverpool) in England, tracks of aurochs, red deer, roe deer, and wild boar have been found in marshland deposits dated from 7.4 to 4.3ka BP37. The co-existence of red deer, wild boar and aurochs seems to have been a common feature especially in the southern latitudes of Europe during the interglacial periods38. This is exemplified in Gibraltar, where the deposits of Genista I Cave provided remains of Cervus elaphus, Bos primigenius and Sus scrofa (reviewed in39). Coeval to this site is Solana del Zamborino, in Granada (Spain), where Bos primigenius was also found40. Dated as late Pleistocene (MIS 3), the painting of the Aurochs’ Hoofprint of Lascaux, with the clear representation of the dewclaws (digits II and V) in the posterior part of the track as being an important identifying feature when tracking aurochs34,41, is relevant to this study. Geological setting of the Trafalgar Cape The Pleistocene coastal deposits of SW Spain are distributed in short sections along the shores of the Gulf of Cadiz (Fig.2). The Asperillo cliff in the Province of Huelva is one of the longest continuous outcrops of Late Pleistocene-to-Holocene deposits of the region, mostly composed of poorly-cemented eolianites and revealing at the base a paleosol developed over interdune pond deposits with a large trampled surface recently described as the Matalascañas Trampled Surface (MTS)15,16. About 100km SE of the MTS is Cape Trafalgar, a tombolo connecting the 17m-high paleodune, oriented WNW-ESE, with the mainland. The last interglacial-Holocene succession between Barbate and Cape Trafalgar rests unconformably on Upper Miocene deposits42 (Fig.2a). The outcrop extends by a maximum area of 600m by 300m during low-tide, beneath the lighthouse. The succession (Fig.2b) is composed of coarse-grained bioclastic sandstones, mostly massive but with a fine, almost horizontal lamination with a NE dip. This unit is truncated by an erosional discontinuity dipping south where abundant large artiodactyl tracks and rhizolith clusters can be found: the Trafalgar Trampled Surface or the TTS. Overlying the TTS and filling the tracks is a 40cm-thick bed of very coarse, bioclastic sandstones with large broken bivalve shells and balanids, but also with occasional branched rhizoliths, mostly horizontal, up to 4cm thick (Fig.2c). The remaining succession is composed of eolianites made of medium-to-fine grained sandstones with planar cross-lamination. Reactivation surfaces are marked by the development of rhizolith horizons, and four (I-IV) highly bioturbated levels with large artiodactyl cloven-hoofed tracks were recognized in a 1.5mthick sequence composed of foresets dipping over 20º towards N (Fig.2a). The succession is then partially covered by the Holocene dune system, but still shows around 2m of eolianites with similar features to the ones mentioned below, including sparse bioturbation by isolated tracks, horizontal thin rhizoliths and insect burrows (Fig.2d). This unit ends with a paleosol with a dense vertical rhizoturbation. The basal beach unit from Cape Trafalgar was dated with U-Th from the last interglacial, MIS 5c (107 ± 2ka)42. This sequence shows a disconformity where the TTS was developed, mostly representing the foreshore. The sand barrier retreat has exposed the TTS to the activity of large artiodactyls in the shoreline. The first eolianite unit was dated by OSL of 99 ± 12 ka42 and reveals the progradation of the dune system in a regressive context, including the development of rhizoliths downward to the disconformity and intersecting the tracks. The aggradation deposition of eolian dune sands is marked by high bioturbated levels and the formation of incipient soils with intense rhizoturbation. The second eolianite unit was deposited circa 78 ± 8 ka42. When comparing this with the Asperillo cliff reference section in the Bay of Cadiz, according to the few OSL ages presently available43, the MTS was formed during the Last Interglacial warm and seasonal humid period (106 ± 19ka) as a reddish paleosol (MIS 5), when the sea-level was at highstand position but with frequent fluctuations. Therefore, the TTS may be contemporary of the MTS, and both highlight the presence in the shore of large artiodactyls (see below). Systematic ichnology Nomenclatural acts: This study follows the requirements of the International Code of Zoological Nomenclature regarding the registration of the nomenclatural act in ZooBank. Ichnogenus: Bovinichnus igen. nov. Type ichnospecies: Bovinichnus uripeda isp. nov. Etymology: referring to traces (= ichnus) with the morphology of tracks shared by the species of the Tribe Bovini, subfamily Bovinae, which includes besides Bos, buffalos, bisons, among others. Diagnosis: Tetradactyl, paraxonic and subsymmetrical tracks that are as wide as long; central (III–IV) digit impressions are subparallel, with broad distal end pointing forward, and a narrow, linear interdigital space; internal surfaces of hoofprints flat to slightly concave, external surfaces slightly to markedly convex; hoofprints widest near the heel, but tapering only to minor degree between the heel and the apex; rounded-to-rectangular posterior dewclaw (digits II and V) prints that are much smaller than central digits, printed next to or close to the heel; manus and pes of closely similar form and size. Bovinichnus uripeda ichnosp. nov. ZooBank—urn:lsid:zoobank.org:pub:E834D014-3A20-4721-9B2D-4577B2E10A52. Koenigswald etal., 1996: Fig.3E. Roberts, 2009: Figs.13, 14. Neto de Carvalho etal., 2020: Fig.2A,C,D,E. 4 Vol:.(1234567890) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ 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. corel draw. com/ la/). 5 Vol.:(0123456789) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ Holotype and Paratypes: TTS10 (Fig.5a), and TTS8, TTS13, respectively. They were left insitu but photogrammetric 3D models were produced (Fig.4). Type horizon: Trafalgar Trampled Surface (TTS): Fig.5a–c. Etymology: In Latin, the prefix uriis the plural word for aurochs and the suffix -peda refers to feet. Type locality: Cape Trafalgar, GPS: 36°10′54.92′′N–6°02′05.87′′W. Diagnosis: The same as for the ichnogenus. Description: Tracks occur in the TTS in concave epirelief, mostly isolated, or in small manus-pes sets up to seven in the same trackway (T3) (Fig.3a,b). The breadth size range varies between 100 and 270mm, but with most of the tracks measuring 200–240mm (Fig.3c,d). They are mostly oriented towards WSW (Fig.3e) determined by the tipping of the hoofprints or the location of the dewclaws (Fig.4a,b). Usually, well-preserved true tracks are tetradactyl and show evidence of pressure pads. The digits III and IV are by far the largest and correspond to crescent-shaped impressions. The posterior part of the prints is deeper than the cleave or digit tips and may reveal rounded-to-rectangular small dew claw imprints proximal to the heel. The outer width measured between dew claw impressions corresponds in size to the posterior width of the hoofprints (Fig.4). The termination of each hoof impression is blunt and directed forward. In convex hyporelief preservation and cross sections, hoof imprints are separated by a central ridge (Fig.5b). At least four trampled levels can be seen in cross section in the eolianite beds. The indentation produced by the large rigid hoof over the well-laminated sandstone show soft deformation, composed of marginal upfolds and microfaults, of the underlying laminae and a concave track infill (Fig.5c,d). The stride length, or distance between footprints from the same foot, calculated for the trackway T3 varies from 210 to 100cm and is 150cm in the trackway T1. The pace length, or the distance between two successive footprints, varies from 110 to 45cm in T3 and is 96cm in the trackway T2. There is no evidence in trackways for manus-pes overprinting and ’direct register’, in which pes is placed directly in manus impression, or in the cross sections of the trampled surfaces in the eolianite. Tetradactyl tracks with the morphology of Bovinichnus uripeda were previously described in the MTS16. They are now synonymized under B. uripeda despite showing smaller dimensions (Fig.3c,d). The morphotype A1 described in the MTS as Cervipeda15 corresponds to large hoofprints with an average size range of 12–13, 5cm in length and 7.5cm broad and some few trackways with measured stride lengths between 100 and 132cm. However, some tracks are up to 17.8cm in length and show pace lengths of 100cm, which are now integrated in the new Bovinichnus uripeda (Figs.3c,d and 6a). The tracks can be rectangular or squared to oval in outline. Impressions of third and fourth toes are central, well-developed mirror images with internal surfaces sometimes in median or posterior contact, strait or concave at the front (Fig.6b,c); in some examples, they are separated by a continuous interdigital space, usually broadest at the front (Fig.6d,e). They can be widest near the heel or in the central part, tapering to an apex of angular or sharply parabolic shape, directed forward, or rotated inwards or outwards, to the midline of the trackway. The second and fifth toe impressions (dew claw imprints) can be seen in some tracks, are smaller and subtriangular or rectangular, located immediately behind the hoofprints (Fig.6e) or separated from them, extending the tracks for extra 30mm (Fig.6c). The trackways show no major differences in size between fore feet and hind feet (Fig.6a). Pressure pads surrounding the track indicate they were imprinted directly on the surface. 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. corel draw. com/ la/). 6 Vol:.(1234567890) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ 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/ OpenD roneM ap/ WebODM; MeshLab 2020.12, (https:// www. meshl ab. net/); CloudCompare 2.11.0, (https:// www. danie lgm. net/ cc/) and the 3D model of the images (e,f) were produced with the software Meshroom 2021.1.0, (https:// alice vision. org/# meshr oom); MeshLab 2020.12, (https:// www. meshl ab. net/); CloudCompare 2.11.0, (https:// www. danie lgm. net/ cc/). 7 Vol.:(0123456789) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ When not showing dew claw prints, the Matalascañas hoofprints are similar to ichnogenus Bifidipes Demathieu etal. They are didactyl footprints organized in homopod trackways with cleaves pointed and hooves diverging and clearly separated, although not so much closer, and wider, to the heel45 (Fig.6e). Remarks: Sarjeant and Langston6 provide an early revision of artiodactyl fossil tracks emphasizing the lack of ichnological description of kidney-shape to crescentic hoofprints of many bovoids. The ichnotaxonomy of artiodactyl tracks is problematic since the hooves are characterised by a wide range of small morphological variations6,22. In this regard, there were attempts to revise the ichnotaxonomy of the artiodactyl tracks6 with the last made by Neto de Carvalho etal.16, and completed in the present work (Fig.7). In this study the number of digit impressions in the well-preserved true tracks has been considered as the most important ichnotaxobase, thus Bovinichnus tracks were compared only with artiodactyl tetradactyl ichnotaxa that are presently valid22. Tetradactyl tracks attributed to artiodactyls are resumed to Cervipeda (Vialov), Bothriodontipus Casanovas- Cladellas and Santafé-Llopis, Cervipus Matsukawa etal., Fustinianapodus Díaz-Martínez etal., and Suidichnus Neto de Carvalho etal.. Bothriodontipus has four forward-direct ungual prints while Bovinichnus shows two large hoofprints and two posteriorly-located dewclaws. Both Cervipeda and Cervipus shows slender hoofprints and their largest size is in average much smaller than Bovinichnus uripeda. Fustinianapodus described recently in the Lower Miocene of Spain22 is recognized by subtriangular dew claw impressions in the pes tracks, and elongate ones in the manus tracks. Finally, Suidichnus described in the MTS are tracks longer than wide, with a general trapezoidal shape, and dew claw impressions conical to comma-shaped, smaller than the main toes, longer than wide, and projected laterally from behind the main toes, with a wider angle in the foreprint comparing with the hindprint16. Artiodactyl tracks with hooves parallel but with hoof apices almost as rounded as the hind portions, the track having an oval to rounded-rectangular outline, are placed into Lamaichnium Aramayo and Bianco6. Besides that, the didactyl shape of these tracks attributed to camelids clearly distinguishes it from Bovinichnus. Therefore, and following the suggestion of Sarjeant and Langston6, Bovinichnus uripeda is considered a new artiodactyl ichnotaxon of bovoid morphology. Slow walking usually keeps manus and pes separated in the artiodactyls46, with manus impression lying ahead of the pes impression. Stride over 2 m together with the shape of cleaves and the large width of the prints indicate that the TTS were produced by large bovines. In the Sefton Coast and at Peterstone (England), aurochs tracks associated with Chalcolithic industries, show lengths of 220–280 mm37, which are also comparable in shape with Bovinichnus uripeda and must be included in the new ichnospecies (see also34). Finally, the Late Paleolithic rock art painting of the Aurochs’ Hoofprint of Lascaux (France), with the clear representation of the dewclaws in the posterior part of the track, recognizes the tetradactyl shape of aurochs tracks34,41. 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. corel draw. com/ la/). 8 Vol:.(1234567890) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ The tetradactyl A1 morphotype described in the MTS shows some differences that allow to separate into two different morphotypes (Fig.6). Despite the preservational variation in morphology that can be found in the MTS15, and the fact that in average A1 tracks have the same size, there are subtle differences that suggest different producers. The larger tracks, blunt shape of cleaves without separation in the proximal area and their convex outline, making many of them rounded or squared, with circular to oval dew claw imprints, can be compared with Bovinichnus uripeda from Cape Trafalgar and the tracks of the aurochs known in the Holocene deposits of Great Britain34,47 (Fig.6a–e). On the other hand, cervid prints are slenderer with the previous half of each foot tapering to a point, for example, in red deer (Cervus elaphus) the forefoot generally creates splayed tracks, especially in soft and/or sandy grounds (Fig.6f). 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:// alice vision. org/# meshr oom); MeshLab 2020.12 (https:// www. meshl ab. net/); CloudCompare 2.11.0 (https:// www. danie lgm. net/ cc/). 9 Vol.:(0123456789) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ Discussion The difference between red deer, domestic cattle and aurochs tracks. Despite red deer, domestic cattle and aurochs tracks may show a tetradactyl preservation, red deer tracks are slenderer, with sharp cleave anteriorly, and cattle leave rounded tracks with convex outlines, when compared to aurochs. Large artiodactyl hoofprints attributed to cervids have been described in the Late Pleistocene of the Iberian Peninsula, including Gorliz in northern Spain, SW Portugal and the MTS10,13–15. These longer-than-wide tracks composed of hoofprints that are more slender than those of Bovinichnus uripeda, can reach up to 150mm in length; the tracks from Gorliz attributed by Flor10 to Cervus elaphus (see its Fig.5) reach 130mm in length but are rounded and could have been produced by the reindeer (with the widest and roundest tracks of all deers) or the European Bison. In the Sefton Coast, the red deer tracks dated from the Neolithic are 100–150mm in length37, while modern average of a full-grown red deer stag’s fore print is 80–90mm long and 60–70mm wide46. Pointed cleaves of the red deer contrast in this area to the more rounded and larger hooves of the aurochs. Also, the outer edge of the cleaves curve evenly towards the tip. In red deer, the dewclaws are rounded impressions. Straddle is relatively narrow, with a stride length for adult varying between 80 and 150 cm46. Barr and Bell20 interpreted rectangular hoof prints, ranging from 120 to 180mm in length, as made by red deer and the larger square hoof prints, ranging from 200 to 260mm in length, as made by aurochs. When showing similar sizes, aurochs differ from cervid tracks by the blunt tips and wider hooves, besides the smaller space enclosed between the two hooves in aurochs48. Figure7. Revision of the valid artiodactyl ichnogenera16 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. corel draw. com/ la/). 16 Vol:.(1234567890) Scientific Reports | (2022) 12:9911 | https://doi.org/10.1038/s41598-022-14137-6 www.nature.com/scientificreports/ S.F. and Y.Z. analysed the tracks. J.R.V. designed the project. F.G.G., G.F., S.F. and C.F. provided contextual data. All authors discussed and commented on the manuscript. Competing interests The authors declare no competing interests. Additional information Supplementary Information The online version contains supplementary material available at https:// doi. org/ 10. 1038/ s41598- 022- 14137-6. Correspondence and requests for materials should be addressed to L.M.C. Reprints and permissions information is available at www.nature.com/reprints. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. 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