A Late Pleistocene hominin footprint site on the North African coast of Morocco.

Footprints represent a relevant vestige providing direct information on the biology, locomotion, and behaviour of the individuals who left them. However, the spatiotemporal distribution of hominin footprints is heterogeneous, particularly in North Africa, where no footprint sites were known before the Holocene. This region is important in the evolution of hominins. It notably includes the earliest currently known Homo sapiens (Jebel Irhoud) and the oldest and richest African Middle Stone Age hominin sites. In this fragmented ichnological record, we report the discovery of 85 human footprints on a Late Pleistocene now indurated beach surface of about 2800 m2 at Larache (Northwest coast of Morocco). The wide range of sizes of the footprints suggests that several individuals from different age groups made the tracks while moving landward and seaward across a semi-dissipative bar-trough sandy beach foreshore. A geological investigation and an optically stimulated luminescence dating of a rock sample extracted from the tracksite places this hominin footprint surface at 90.3 ± 7.6 ka (MIS 5, Late Pleistocene). The Larache footprints are, therefore, the oldest attributed to Homo sapiens in Northern Africa and the Southern Mediterranean.

New dating of the Matalascañas footprints provides new evidence of the Middle Pleistocene (MIS 9-8) hominin paleoecology in southern Europe.

Hominin footprints were recently discovered at Matalascañas (Huelva; South of Iberian Peninsula). They were dated thanks to a previous study in deposits of the Asperillo cliff to 106 ± 19 ka, Upper Pleistocene, making Neandertals the most likely track-makers. In this paper, we report new Optically Stimulated Luminescence dating that places the hominin footprints surface in the range of 295.8 ± 17 ka (MIS 9-MIS 8 transition, Middle Pleistocene). This new age implies that the possible track-makers are individuals more likely from the Neandertal evolutionary lineage. Regardless of the taxon attributed to the Matalascañas footprints, they supplement the existing partial fossil record for the European Middle Pleistocene Hominins being notably the first palaeoanthropological evidence (hominin skeleton or footprints) from the MIS 9 and MIS 8 transition discovered in the Iberian Peninsula, a moment of climatic evolution from warm to cool. Thus, the Matalascañas footprints represent a crucial record for understanding human occupations in Europe in the Pleistocene.

Fast-running theropods tracks from the Early Cretaceous of La Rioja, Spain.

Theropod behaviour and biodynamics are intriguing questions that paleontology has been trying to resolve for a long time. The lack of extant groups with similar bipedalism has made it hard to answer some of the questions on the matter, yet theoretical biomechanical models have shed some light on the question of how fast theropods could run and what kind of movement they showed. The study of dinosaur tracks can help answer some of these questions due to the very nature of tracks as a product of the interaction of these animals with the environment. Two trackways belonging to fast-running theropods from the Lower Cretaceous Enciso Group of Igea (La Rioja) are presented here and compared with other fast-running theropod trackways published to date. The Lower Cretaceous Iberian fossil record and some features present in these footprints and trackways suggest a basal tetanuran, probably a carcharodontosaurid or spinosaurid, as a plausible trackmaker. Speed analysis shows that these trackways, with speed ranges of 6.5-10.3 and 8.8-12.4 ms-1, testify to some of the top speeds ever calculated for theropod tracks, shedding light on the question of dinosaur biodynamics and how these animals moved.

Tracking late Pleistocene Neandertals on the Iberian coast.

Here, we report the recent discovery of 87 Neandertal footprints on the Southwest of the Iberian Peninsula (Doñana shoreline, Spain) located on an upper Pleistocene aeolian littoral setting (about 106 ± 19 kyr). Morphometric comparisons, high resolution digital photogrammetric 3D models and detailed sedimentary analysis have been provided to characterized the footprints and the palaeoenvironment. The footprints were impressed in the shoreline of a hypersaline swamped area related to benthic microbial mats, close to the coastline. They have a rounded heel, a longitudinal arch, relatively short toes, and adducted hallux, and represent the oldest upper Pleistocene record of Neandertal footprints in the world. Among these 87 footprints, 31 are longitudinally complete and measure from 14 to 29 cm. The calculated statures range from 104 to 188 cm, with half of the data between 130 and 150 cm. The wide range of sizes of the footprints suggests the existence of a social group integrated by individuals of different age classes but dominated, however, by non-adult individuals. The footprints, which are outside the flooded area are oriented perpendicular to the shoreline. These 87 footprints reinforce the ecological scenario of Neandertal groups established in coastal areas.

Differential locomotor and predatory strategies of Gondwanan and derived Laurasian dromaeosaurids (Dinosauria, Theropoda, Paraves): Inferences from morphometric and comparative anatomical studies.

Tetrapod limbs morphology is a reliable proxy of locomotor capacities. Beyond this, other aspects of life habits, such as predation abilities, can also be relevant to determine main morphofunctional appendicular properties, which ultimately reflect a compromise between different factors of the biological role. Dromaeosauridae is a dinosaur clade belonging to Theropoda, a group of bipedal predators. Dromaeosaurids represent an interesting study case, in which the hindlimbs have been proposed to be involved in both locomotion and predation activity. A peculiar feature characterizing all dromaeosaurids is a modified second pedal digit, which is typically related to predation. This theropod group is closely related to birds and diversified during the Cretaceous Period, mainly in the Northern Hemisphere (Laurasia). However, a subclade of dromaeosaurids, the Unenlagiinae, was recently recognized for Gondwana. Nevertheless, there are morphological differences between derived Laurasian dromaeosaurids (eudromaeosaurs) and unenlagiines. Such differences are observed in the proportions between hindlimb bones and in the presence of a subarctometatarsalian condition in unenlagiines, which is mainly characterized by a proximally constricted metatarsal III. To evaluate the function of these divergent morphologies, we conducted morphometric analyses and comparisons of qualitative morphological aspects, encompassing unenlagiines, other dromaeosaurids, as well as taxa from other theropod groups, including extant birds. The former approach consisted of two phylogenetic principal component analyses, one based on the main measurements of the hindlimb, and the other focused on the lengths of the pedal phalanges. The first analysis drew the unenlagiines close to taxa with long tibiae, as well as long and slender metatarsi. Instead, eudromaeosaurs are closer to taxa with shorter tibiae, and shorter and wider metatarsi. The second analysis showed that eudromaeosaurs and unenlagiines have similar phalangeal proportions, including the elongation of distal phalanges. However, the shorter second phalanx of the pedal digit II of eudromaeosaurs could have increased the force generated by this digit, which was the main predatory tool of the autopodium. This, together with a shorter and wider metatarsus, and a marked hinge-like morphology of the articular surfaces of metatarsals and phalanges, possibly allowed eudromaeosaurs to exert a great gripping strength and hunt large prey. Conversely, the longer and slender subarctometatarsus, and less well-marked hinge joints of unenlagiines possibly gave them greater cursorial capacities. Additionally, the longer second phalanx of digit II allowed unenlagiines fast movements of this digit to hunt smaller and elusive prey. Thus, the distinctive morphological evolutionary pathways of these two dromaeosaurid clades seem to have been influenced by the particular locomotor and predatory specializations that characterized each of these lineages.

Multi-aged social behaviour based on artiodactyl tracks in an early Miocene palustrine wetland (Ebro Basin, Spain).

We present a new locality with at least 880 vertebrate tracks found at the top of a limestone bed from the lower Miocene Tudela Formation (Spain). The trampled surface was formed by artiodactyls that crossed a muddy carbonate accumulated under the influence of water level variations in a palustrine environment. The tracks reflect different types of morphological preservation. The well-preserved tracks have tetradactyl digit impressions caused by both manus and pes, and are the type series of a new artiodactyl ichnotaxon, Fustinianapodus arriazui ichnogen. nov. and ichnosp. nov. The rest of the tracks, which are not as well preserved, are didactyl and were classified as undetermined artiodactyl tracks. According to their preservation, morphology, size, arrangement and orientation, we propose that this tracksite is the product of a social behaviour, particularly gregariousness, of a multi-age group of artiodactyls ~19 Ma ago. The morphologic and palaeoecologic data presented here suggest that the trackmakers were a group of anthracotheres with a livelihood similar to current hippos. They crossed, periodically, a fresh water palustrine area along some preferential pathways (trails).

Triassic pentadactyl tracks from the Los Menucos Group (Río Negro province, Patagonia Argentina): possible constraints on the autopodial posture of Gondwanan trackmakers.

The Los Menucos locality in Patagonia, Argentina, bears a well-known ichnofauna mostly documented by small therapsid footprints. Within this ichnofauna, large pentadactyl footprints are also represented but to date were relatively underinvestigated. These footprints are here analyzed and discussed based on palaeobiological indications (i.e., trackmaker identification). High resolution digital photogrammetry method was performed to achieve a more objective representation of footprint three-dimensional morphologies. The footprints under study are compared with Pentasauropus from the Upper Triassic lower Elliot Formation (Stormberg Group) of the Karoo Basin (Lesotho, southern Africa). Some track features suggest a therapsid-grade synapsid as the potential trackmaker, to be sought among anomodont dicynodonts (probably Kannemeyeriiformes). While the interpretation of limb posture in the producer of Pentasauropus tracks from the Los Menucos locality agrees with those described from the dicynodont body fossil record, the autopodial posture does not completely agree. The relative distance between the impression of the digital (ungual) bases and the distal edge of the pad trace characterizing the studied tracks likely indicates a subunguligrade foot posture (i.e., standing on the last and penultimate phalanges) in static stance, but plantiportal (i.e., the whole foot skeleton and related soft tissues are weight-bearing) during the dynamics of locomotion. The reconstructed posture might have implied an arched configuration of the articulated metapodials and at least of the proximal phalanges, as well as little movement capabilities of the metapodials. Usually, a subunguligrade-plantiportal autopod has been described for gigantic animals (over six hundreds kilograms of body weight) to obtain an efficient management of body weight. Nevertheless, this kind of autopod is described here for large but not gigantic animals, as the putative trackmakers of Pentasauropus were. This attribution implies that such an autopodial structure was promoted independently from the body size in the putative trackmakers. From an evolutionary point of view, subunguligrade-plantiportal autopods not necessarily must be related with an increase in body size, but rather the increase in body size requires a subunguligrade or unguligrade, plantiportal foot. Chronostratigraphically, Pentasauropus was reported from Upper Triassic deposits of South Africa and United States, and from late Middle Triassic and Upper Triassic deposits of Argentina. Based on the stratigraphic distribution of the ichnogenus currently accepted, a Late Triassic age is here proposed for the Pentasauropus-bearing levels of the Los Menucos Group.

An Unexpected Early Rhabdodontid from Europe (Lower Cretaceous of Salas de los Infantes, Burgos Province, Spain) and a Re-Examination of Basal Iguanodontian Relationships.

Disarticulated and incomplete remains from a new diminutive ornithopod are described. They come from the Cameros Basin in the north of Spain and were collected from the red clays of the Castrillo de la Reina Formation, ranging from Upper Barremian to Lower Aptian. The new ornithopod described here is slender and one of the smallest ever reported. An up-to-date phylogenetic analysis recovers this taxon as a basal iguanodontian. Its unique combination of characters makes it more derived than slender ornithopods like Hyphilophodon and Gasparinisaura, and bring very interesting insights into the basal iguanodontian phylogeny. Though possessing a minimum of three premaxillary teeth, this taxon also bears an extensor ilio-tibialis groove on the distal part of its femur. Moreover, its dentary and maxillary teeth are unique, remarkably similar to those regarded as having a "rhabdomorphan" affinity. This unknown taxon is suggested to be a stem taxon within Rhabdodontidae, a successful clade of basal iguanodonts from the Late Cretaceous of Europe. The Gondwanan ornithopods share the strongest affinities with this family, and we confirm Muttaburrasaurus as a sister taxon of the Rhabdodontidae within a newly defined clade, the Rhabdodontomorpha.

A reappraisal of the Middle Triassic chirotheriid Chirotherium ibericus Navás, 1906 (Iberian Range NE Spain), with comments on the Triassic tetrapod track biochronology of the Iberian Peninsula.

Triassic vertebrate tracks are known from the beginning of the 19th century and have a worldwide distribution. Several Triassic track ichnoassemblages and ichnotaxa have a restricted stratigraphic range and are useful in biochronology and biostratigraphy. The record of Triassic tracks in the Iberian Peninsula has gone almost unnoticed although more than 25 localities have been described since 1897. In one of these localities, the naturalist Longinos Navás described the ichnotaxon Chirotherium ibericus in 1906.The vertebrate tracks are in two sandy slabs from the Anisian (Middle Triassic) of the Moncayo massif (Zaragoza, Spain). In a recent revision, new, previously undescribed vertebrate tracks have been identified. The tracks considered to be C. ibericus as well as other tracks with the same morphology from both slabs have been classified as Chirotherium barthii. The rest of the tracks have been assigned to Chirotheriidae indet., Rhynchosauroides isp. and undetermined material. This new identification of C. barthii at the Navás site adds new data to the Iberian record of this ichnotaxon, which is characterized by the small size of the tracks when compared with the main occurrences of this ichnotaxon elsewhere. As at the Navás tracksite, the Anisian C. barthii-Rhynchosauroides ichnoassemblage has been found in other coeval localities in Iberia and worldwide. This ichnoassemblage belongs to the upper Olenekian-lower Anisian interval according to previous biochronological proposals. Analysis of the Triassic Iberian record of tetrapod tracks is uneven in terms of abundance over time. From the earliest Triassic to the latest Lower Triassic the record is very scarce, with Rhynchosauroides being the only known ichnotaxon. Rhynchosauroides covers a wide temporal range and gives poor information for biochronology. The record from the uppermost Lower Triassic to the Middle Triassic is abundant. The highest ichnodiversity has been reported for the Anisian with an assemblage composed of Dicynodontipus, Procolophonichnium, Rhynchosauroides, Rotodactylus, Chirotherium, Isochirotherium, Coelurosaurichnus and Paratrisauropus. The Iberian track record from the Anisian is coherent with the global biochronology proposed for Triassic tetrapod tracks. Nevertheless, the scarcity of track occurrences during the late Olenekian and Ladinian prevents analysis of the corresponding biochrons. Finally, although the Iberian record for the Upper Triassic is not abundant, the presence of Eubrontes, Anchisauripus and probably Brachychirotherium is coherent with the global track biochronology as well. Thus, the Triassic track record in the Iberian Peninsula matches the expected record for this age on the basis of a global biochronological approach, supporting the idea that vertebrate Triassic tracks are a useful tool in biochronology.

Ichnotaxonomic review of large ornithopod dinosaur tracks: temporal and geographic implications.

BACKGROUND: Large ornithopod tracks are known from the Upper Jurassic to the uppermost Cretaceous rocks of all continents but Antarctica. They include the tracks historically called Iguanodon footprints, iguanodontid footprints, hadrosaur/hadrosaurid footprints, and other large ornithopod tracks that have been used to define ichnotaxa. More than 40 ichnospecies based on large ornithopod tracks have been defined, but the validity of many of them is questionable. METHODOLOGY/PRINCIPAL FINDINGS: 34 ichnogenera and 44 ichnospecies have been analysed in this work. Many of them are considered to be invalid because they have been defined on the basis of poorly preserved tracks without diagnostic features, have an inadequate diagnosis, or are based on temporal and/or geographical criteria. Only eight ichnospecies belonging to the ichnogenera Caririchnium, Iguanodontipus and Hadrosauropodus are here regarded as valid. CONCLUSIONS/SIGNIFICANCE: The monospecific ichnogenus Iguanodontipus (I. burreyi) is characterized by a small, rounded heel and elongate, narrow digit impressions. Its distribution is limited to the Berriasian-Valanginian of Europe. Caririchnium consists of four ichnospecies (C. magnificum [type ichnospecies], C. kortmeyeri, C. billsarjeanti and C. lotus) with a large, rounded heel and short, wide digit impressions. This ichnogenus ranges from the Berriasian-Hauterivian to the Aptian-Albian of South America, North America, Asia and Europe. Finally, Hadrosauropodus (three ichnospecies: H. langstoni [type ichnospecies], H. leonardii and H. kyoungsookimi) shows a large, bilobed heel and short, wide digit impressions. It is known from the Aptian-Albian to the Maastrichtian of North America, Asia and Europe. The ichnofamily Iguanodontipodidae includes large iguanodontian tracks characterized mainly by mesaxonic, tridactyl and subsymmetrical pes tracks that are as wide as (or wider than) long and have one pad impression in each digit and one in the heel. Its distribution is confidently limited to the Cretaceous of Europe, Asia, North America and South America.

Tetradactyl footprints of an unknown affinity theropod dinosaur from the Upper Jurassic of Morocco.

BACKGROUND: New tetradactyl theropod footprints from Upper Jurassic (Oxfordian-Kimmeridgian) have been found in the Iouaridène syncline (Morocco). The tracksites are at several layers in the intermediate lacustrine unit of Iouaridène Formation. The footprints were named informally in previous works "Eutynichnium atlasipodus". We consider as nomen nudum. METHODOLOGY/PRINCIPAL FINDINGS: Boutakioutichnium atlasicus ichnogen. et ichnosp. nov. is mainly characterized by the hallux impression. It is long, strong, directed medially or forward, with two digital pads and with the proximal part of the first pad in lateral position. More than 100 footprints in 15 trackways have been studied with these features. The footprints are large, 38-48 cm in length, and 26-31 cm in width. CONCLUSIONS/SIGNIFICANCE: Boutakioutichnium mainly differs from other ichnotaxa with hallux impression in lacking metatarsal marks and in not being a very deep footprint. The distinct morphology of the hallux of the Boutakioutichnium trackmaker -i.e. size and hallux position- are unique in the dinosaur autopodial record to date.