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

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 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 boar [9][10][11][12][13][14][15][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 assemblages [17][18][19] , the trace fossil record of aurochs was only known from the Holocene of Great Britain, especially in tidal flat deposits from estuarine areas 20,21 . 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 Miocene 22 . The first tracksites were described in the Late Pleistocene (MIS 3) of the Balearic Islands in Mallorca, Menorca and Formentera 9,11,23 , 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][13][14] , and in the Late Pleistocene of the Catalan Bay eolianite, in Gibraltar 24 . 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 Pleniglacial [25][26][27] . Recently, the Matalascañas Trampled Surface (MTS) 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 aurochs 15,16 . Aurochs tracks have been studied especially in coastal deposits in England and Wales associated with early Mesolithic to early Bronze Age industries 20,[28][29][30][31][32][33][34][35][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 BP 37 . 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 periods 38 . This is exemplified in Gibraltar, where the deposits of Genista I Cave provided remains of Cervus elaphus, Bos primigenius and Sus scrofa (reviewed in 39 ). Coeval to this site is Solana del Zamborino, in Granada (Spain), where Bos primigenius was also found 40 . 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 aurochs 34,41 , 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 deposits 42 (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 m-thick 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 ka 42 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 ka 42 .
When comparing this with the Asperillo cliff reference section in the Bay of Cadiz, according to the few OSL ages presently available 43 , 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.
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.
Type horizon: Trafalgar Trampled Surface (TTS): Fig. 5a-c. Etymology: In Latin, the prefix uri-is 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 MTS 16 . They are now synonymized under B. uripeda despite showing smaller dimensions (Fig. 3c,d).
The morphotype A1 described in the MTS as Cervipeda 15 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. www.nature.com/scientificreports/   45 (Fig. 6e).
Remarks: Sarjeant and Langston 6 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 variations 6,22 . In this regard, there were attempts to revise the ichnotaxonomy of the artiodactyl tracks 6 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 valid 22  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 Spain 22 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 hindprint 16 . 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 Bianco 6 . Besides that, the didactyl shape of these tracks attributed to camelids clearly distinguishes it from Bovinichnus. Therefore, and following the suggestion of Sarjeant and Langston 6 , Bovinichnus uripeda is considered a new artiodactyl ichnotaxon of bovoid morphology.
Slow walking usually keeps manus and pes separated in the artiodactyls 46 , 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 mm 37 , which are also comparable in shape with Bovinichnus uripeda and must be included in the new ichnospecies (see also 34 ). 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 tracks 34,41 . 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 MTS 15 , 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 Britain 34,47 (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). 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 MTS 10,13-15 . 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 Flor 10 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 length 37 , while modern average of a full-grown red deer stag's fore print is 80-90 mm long and 60-70 mm wide 46 . 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 cm 46 . Barr and Bell 20 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 aurochs 48 . www.nature.com/scientificreports/ Despite being broadly bovoid rounded tracks, there is a marked difference in size, especially in length, between Bos taurus and Bos primigenius. Tracks of extant, fully grown cattle are ~ 100 × 95 mm in size 20 , which is less than half the size of the very large bovine tracks found in Cape Trafalgar. For the purpose of comparing the morphology (but not taphonomic variation) of extant domestic cattle hoofprints with Cape Trafalgar and Matalascañas tracks, a bull 1128 kg in weight was invited to walk in a dry fine sand substrate. He left rounded tracks measuring 180 mm long which are nevertheless more rounded than aurochs tracks. Moreover, the hooves outside are convex and the inner edge is concave at the front and convex at the back 46 .

Aurochs vs. Bisons as producers of Bovinichnus in SW Iberia. Aurochs and Steppe Bisons coexisted
in Iberia during the Late Pleistocene. Since these two bovines were in size and morphologically very similar, these would rise the possibility that steppe bisons could be likely producers of Bovinichnus. Therefore, it is vital to understand the biogeographical distribution of the two species in order to be sure about the most likely producer of Bovinichnus uripeda in SW Spain.
The origin of the aurochs is not clear, although it is thought to have Indian ancestry, dating back to about 2 Ma 49 . The aurochs were first reported in Europe during the Middle Pleistocene at Venosa-Notarchirico, Italy, and finally became extinct in Poland in 1627 AD 38 . Aurochs finds are less numerous during the Pleistocene than during the Holocene, but a large distribution area is represented nonetheless, from SW Iberia to southern Scandinavia, North Africa to Korean peninsula and Japan 50 . In the Eemian (MIS 5e), they expanded towards northern territories, where their skeletons were found, e.g., in excavations made in the Trafalgar Square at central London 51 . During the early Holocene (Mesolithic and Neolithic cultural phases in Europe) the species seems to have increased in number, probably due to increasingly favorable mild and wet conditions after the end of the Last Ice Age 50 . Its early appearance in Spain dates back to 0.7 Ma 52 . Remains of aurochs dated from the Middle or Late Pleistocene have been uncovered in Great Britain, France, Spain, Portugal, Italy, and Germany. The distribution of the aurochs has fluctuated with the changing climate during the Pleistocene, being more widely dispersed in interglacials and interstadials 50 . During the Pleniglacial, with the European northern ice cap boundary having a southerly course, and with the Iberian and Italian peninsulas being used as refugial areas 53 , aurochs became rare or completely disappeared during colder periods in many areas of Europe. At Gibraltar, taking advantage of the milder climate in the southernmost Europe during the MIS 3, Cervus elaphus, Sus scrofa and Bos primigenius were found related to habitats of stone pine/juniper woodland, savannah, shrubland with patches of grassland 54 .
The long-horned Bison priscus (Bojanus), or steppe bison, is the earlier form of Bison, thought to have become extinct in Europe at the end of the Pleistocene 55 . As the aurochs, the steppe bison had also a wide distribution across Europe, from the Iberian Peninsula, through central and eastern Europe and into Siberia, crossing the Bering strait into North America during the early Holocene 56,57 . The steppe bison was able to adapt to a wide range of environments, both glacial and temperate 53 , by performing long migrations. Of the two species, Bison priscus is thought to be particularly problematic in terms of its skeletal distinction from Bos primigenius 58,59 . The problem is enhanced by the fact that Bison priscus was relatively abundant during the Pleistocene, and seems to appear alongside Bos primigenius on several sites. Nevertheless, the steppe bison is known only from MIS 3-2 from northern and central Spain 18,26,60 during which it was found together with B. primigenius 25 . During this period, the expanding territories of B. priscus related to colder and steppe environments may have reached the southern latitudes of Granada 61 . B. priscus appears in the cave art from North (Altamira, Altxerri, Santimamiñe and Ekain) and Central Spain (La Hoz and Siega Verde) during the Last Glacial Maximum, where representations of aurochs are rare 18 . Bison are absent from the long and well-studied Late Pleistocene sequences in Gibraltar, and elsewhere in SW Iberia. Thus, the aurochs were the only bovines existing during the MIS 5 in SW Iberia and we can confidently attribute the Bovinichnus uripeda tracks from Cape Trafalgar and Matalascañas to aurochs.
The only fossil tracks related to Bison sensu lato that are known were described 62 in dune and interdune deposits from Nebraska (USA). However, they are found mainly in cross sections, as concave-up deformation structures or undertracks, circular to oval in plane, ranging from 70 to 160 mm in diameter. Due to this kind of preservation, it is not possible to compare the Nebraska tracks with Bovinichnus uripeda. However, if we compare the tracks produced by the extant Bison bonasus with Bovinichnus uripeda we realize they are wider than long and, therefore, do not present the more rectangular morphology (slightly longer than wide) evidenced in Bovinichnus. Therefore, the track length/width ratio can be very useful to distinguish between the presence of aurochs and steppe bisons in the track fossil record. "The Beach Bulls": large aurochs tracks in the TTS. Aurochs have reached their maximum body size in the early Late Pleistocene interglacial 38,61 . There is an increase in size of the metapodial bones preserved in levels dated as Middle Pleistocene to early Late Pleistocene and a decrease of the same during the Late Pleistocene to early Holocene 38 . This trend is also found for other large mammals in Europe such as the red deer 63 and the wild boar 64 (see evidence from tracks in 16 ). After that, there is a general size decrease between the Late Pleistocene and the early Holocene 52,58,64,65 , with a size increase recorded after beginning of the Holocene 66 . The Last Glacial was responsible for a decrease in size of the aurochs 47,67 . There is also clear evidence of a size reduction in red deer at the end of the Pleistocene in Iberia 67,68 . In the case of the aurochs, the body size increase during the early Late Pleistocene may be related to predation pressure since during this period these animals were prey for a large array of large carnivores, including Neanderthals. The large track size recorded in the early Late Pleistocene from Cape Trafalgar (Fig. 8) is comparable with the records of up to 280 mm in length from the lower Holocene of western England 37,47 , which correspond to the largest track record attributed to aurochs. This increase in size of aurochs recorded in tracks during the Holocene has been attributed to predation pressure and husbandry 20 www.nature.com/scientificreports/ Aurochs visiting the beach. Aurochs was typically a "wetland specialist" which preferred to live in marshy forests, near estuarine areas and floodplains for grazing during the day 50,69 . According to these authors, aurochs lived in small herds and were grazers, not browsers like many deer species, thus requiring open landscapes composed of grasslands. For this main reason, herds of aurochs were roaming the coastal areas of southern Europe during the Late Pleistocene. Pandolfi et al. 38 20,34,47 where they were preserved in intertidal deposits 30,70 . Glades may have attracted the animals to graze not far from the dense coastal woodland 37,70 . The TTS and the MTS are located not far from the marshlands of Cadiz, and the Guadalquivir and Odiel rivers, respectively. But their association to beach and dune-interdune deposits, not in direct relation with fluvial environments, requires further explanation. This explanation may be found in a modern analogue example from the Doñana National Park, by comparing the behaviour of wild cattle (Bos taurus) regarding resources partitioning between seasons.
Doñana feral cattle has been living under free-ranging conditions since the thirteenth century 76 . High quality habitats in Doñana National Park include lagoon meadows and the lower grassland of the ecotone zone between the scrubland and the marshland, which are most used by the active habitat selection of the feral cow-herds 76 . Land sand dunes found in large areas in the coast of Doñana (primary or covered by a xerophytic community of shrubs) are considered a low-quality habitat where the availability of the forage is low and therefore avoided by the cattle. Most of the tracks in the MTS likely attributed to aurochs are smaller in size than Bovinichnus uripeda described in the TTS. This track size difference in coeval deposits may be explained by habitat difference/preference and the strong sexual dimorphism known for aurochs. Lagoon meadows related with interdune drying ponds was the most likely environment described in the MTS 15,16 . Aurochs, especially cow-herds would have centred their movements around food patches of high availability and quality of food, such as the MTS. This could happen especially at the end of spring and during summer, providing enough food for the offspring. However, in the coastal beach and dunes of Trafalgar Cape the aurochs presence recorded by larger-sized tracks and trackways, in the TTS showing a preferential orientation of movement towards the shoreline. The well-drained sandy substrate certainly resulted in the virtual absence of an herbaceous layer in this beach area, where also the eolianites show a sparse rhizoturbation.
Among dimorphic mammals, males have higher metabolic needs than females and are usually relegated to low quality habitats, resulting in male greater seasonal home range 76 . The tracks in the TTS are mostly from large bulls and may indicate this wide dispersion to poor-quality coastal habitats. But there are few tracks 100 mm in length that can be related with the presence of cows. In Doñana National Park, this occasional presence of both cows and bulls together in coastal habitats usually happens in autumn, when rut is in its peak and the bulls were associated with the cow-herds 76 . Nevertheless, cow-size tracks in the TTS are rare comparing with the large tracks and trackways, showing mostly the same orientation, which can only be attributed to bulls walking together in herd. Therefore, while the MTS may record the presence of a cow-herd, the five trampled horizons in Cape Trafalgar may have resulted from the occasional but recurrent procurement of the shore by small herds of bulls.
It is relatively common nowadays to find feral cattle visiting the shores across the world. The Xhosa beach cattle, the "beach-loving cows" in South Africa, even became a tourist attraction. They come down to the shores usually to curl up on the sand and chew the cud. Other examples of recurrent visits of herds to the shores across the world may be found in Hong Kong or Corsica. The Cape Trafalgar's TTS might represent the arrival of a

Materials and methods
Field campaigns in the Cape Trafalgar section, made between 2020 and 2021, produced 2D cartography, 3D photogrammetric models of the most representative tracks and a detailed stratigraphic log. This data was then compared qualitatively with the large artiodactyl tracks that have been described in the MTS 15,16 .
The ichnological material. The Cape Trafalgar section provides at least five stratigraphic horizons with large cloven-hoofed tracks. The upper four horizons can be seen mostly in section as undertracks and preserved as true tracks in bedding soles. There, it was not possible to identify trackways. The TTS is exposed for an area of at least 70 m 2 and 36 tracks were positively identified. Tracks are concentrated in the eastern sector of the TTS, but at least 9 tracks can be found towards west, with at least two of them belonging to the same trackway (T1). Six short trackways, composed of two to seven tracks sets, were determined in total. Each track was provided with a code (TTSx) and measured (Table 1). Well-preserved hoofprints were measured regarding length, the breadth of the widest part of the print, length and breadth of the dewclaws whenever present, orientation, pace and stride when organized in trackways. Track length/width ratio was calculated and may be used to distinguish artiodactyl tracks in the fossil record with comparable morphologies, e.g., Bison. Track depth was indicated only for the deep tracks since most of the tracks show no more than 20-30 mm in depth. However, several tracks are just rounded prints without evidence of toes. In these cases, only rarely measurements were obtained using the pressure pads to orientate the track. The track length/width ratio and orientation patterns were examined by plotting, in bivariate and boxplot graphs, and in a rose diagram, respectively, the information of individual tracks (Fig. 3).
The storm surge of the spring 2020 exposed for a short period large area of the MTS, cleaning over 1.5 m of beach. Since the outcrop is located in the intertidal zone of Matalascañas beach, i.e., an area affected by intense tidal gradients, data recovery was limited in time and mainly consisted of taking photographs for photogrammetry and making casts for some of the tracks. The recognized tracks and trackways have been grouped in four morphotypes for mammals and three for birds. Likewise, insect traces have been recognized, as well as rhizoliths.
Photogrammetry. Digital photogrammetry and GIS techniques have been used to produce safeguard records and highlight the morphological features of some of the tracks in both the TTS and the MTS. Bovinichnus uripeda holotype and paratypes were interpreted and described based on the field observations, measurements and analysis of the digital 3D models. Photos were made with a Canon PowerShot SX5OHS camera with a zoom lens 50xls 4.3-215 mm 1: 3.4-6.5 VSM. The photogrammetric models were processed using the software Meshroom 2021.1.0 77 and ODM 2.1.0 78 , post-processed, analysed and highlighted in Meshlab v2020.12 79 and CloudCompare v2.11.0 80 . The results obtained enabled to highlight with false colors the 3D morphology of the tracks 81,82 , making visual interpretation more effective.