Juan Luis Arsuaga

• Professor
• Complutense University of Madrid

Understanding taphonomic processes is essential for reconstructing past environmental dynamics and interpreting mixed sites, where successive occupations by different biological agents have occurred and, in many cases, have been modified by post-depositional processes. Such is the case in the western part of Buena Pinta Cave (Pinilla del Valle, Mad...

This study presents a statistical approach that combines quantitative and qualitative variables, using features of modern and fossil samples beyond their size (volume content, texture and more), to improve the taxonomic identification of coprolite producers.

Located in the upper valley of the Lozoya River, Cueva del Camino (Madrid, Spain) is one of the richest Early Pleistocene paleontological sites in the Iberian Peninsula. The results of the work carried out over the last three decades have led to the interpretation of the site as a hyena den with intermittent human presence. The faunal assemblage of...

Cattle ( Bos taurus ) play an important role in the life of humans in the Iberian Peninsula not just as a food source but also in cultural events. When domestic cattle were first introduced to Iberia, wild aurochs ( Bos primigenius ) were still present, leaving ample opportunity for mating (whether intended by farmers or not). Using a temporal bioa...

Who the first inhabitants of Western Europe were, what their physical characteristics were, and when and where they lived are some of the pending questions in the study of the settlement of Eurasia during the Early Pleistocene epoch. The available palaeoanthropological information from Western Europe is limited and confined to the Iberian Peninsula...

The Cueva Des-Cubierta site (Pinilla del Valle, Madrid, Spain) contains a peculiar association of crania from large horned herbivores, which were deposited over generations by Neanderthals. To understand the anthropogenic modification of the crania, an experimental program has been carried out from multiple perspectives, with functionality being on...

Revealing the evolutionary processes which resulted in the derived morphologies that characterize the Neanderthal clade has been an important task for paleoanthropologists. One critical method to quantify evolutionary changes in the morphology of hominin populations is through evaluating morphological phenotypic diversity (i.e., disparity) in phylo...

Cattle have been a valuable economic resource and cultural icon since prehistory. From the initial expansion of domestic cattle into Europe during the Neolithic period, taurine cattle ( Bos taurus ) and their wild ancestor, the aurochs ( B. primigenius ), had overlapping ranges, leading to ample opportunities for mating (whether intended by farmers...

Se realiza el estudio preliminar de los fósiles de proboscídeo del Mioceno medio recuperados durante los trabajos de control de tierras en las obras realizadas en el polígono industrial de La Atalayuela 33 (Vallecas, Madrid, España). Se presenta por primera vez la secuencia estratigráfica del yacimiento y el análisis preliminar de los elementos den...

The extremely rich palaeontological record of the horse family, also known as equids, has provided many examples of macroevolutionary change over the last ~55 Mya. This family is also one of the most documented at the palaeogenomic level, with hundreds of ancient genomes sequenced. While these data have advanced understanding of the domestication h...

Two main hypotheses have been proposed to explain the evolution of the derived Neanderthal cranial morphology, with some scholars arguing for a gradual emergence and others proposing distinct phases. However, several Middle Pleistocene specimens show an incipient Neanderthal morphology characterized by a mosaic of derived and primitive characters....

We describe the shape variability of nine dog hemimandibles recovered from two Holocene archaeological sites on the Iberian Peninsula. In this study we mainly focus on the Chalcolithic age dog remains recovered from Barrio del Castillo (Torrejón de Ardoz, Madrid), but also, make comparisons to dog specimens from El Portalón (Sierra de Atapuerca, Bu...

Caregiving for disabled individuals among Neanderthals has been known for a long time, and there is a debate about the implications of this behavior. Some authors believe that caregiving took place between individuals able to reciprocate the favor, while others argue that caregiving was produced by a feeling of compassion related to other highly ad...

Here, we provide a complete, updated, and illustrated inventory, as well as a comprehensive study, of the tarsals (rearfoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossil. The minimum number of individuals (MNI) estimated from the tarsa...

Information on the evolution of the thorax and lumbar spine in the genus Homo is hampered by a limited fossil record due to the inherent fragility of vertebrae and ribs. Neandertals show significant metric and morphological differences in these two anatomical regions, when compared to Homo sapiens. Thus, the important fossil record from the Middle...

The identification of anthropogenically-modified carnivoran bones in archaeological sites is rare in Pleistocene contexts, especially in the most ancient periods. Neanderthal groups have clearly shown a great variety of subsistence activities and the use of carnivoran resources, until rare, is also present in some archaeological sites. However, th...

This study provides a complete, updated and illustrated inventory, as well as a comprehensive study, of the metatarsals and foot phalanges (forefoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossils. This current updated review has establ...

Cattle have been a valuable economic resource and cultural icon since prehistory. From the initial expansion of domestic cattle into Europe during the Neolithic period, taurine cattle (Bos taurus) and their wild ancestor, the aurochs (B. primigenius), had overlapping ranges leading to ample opportunities for intentional and unintentional hybridizat...

Cattle have been a valuable economic resource and cultural icon since prehistory. From the initial expansion of domestic cattle into Europe during the Neolithic period, taurine cattle (Bos taurus) and their wild ancestor, the aurochs (B. primigenius), had overlapping ranges leading to ample opportunities for intentional and unintentional hybridizat...

Buena Pinta Cave (Pinilla del Valle, Madrid) has been interpreted as a hyena den with sporadic occupations of Homo neanderthalensis in the western part of the site (level 23). In order to identify the different formation processes in this area of the site, spatial analyses have been carried out with GIS and spatial statistics based on the taphonomi...

Los retocadores óseos son considerados como parte de la industria ósea de los grupos humanos pleistocenos. Normalmente se caracterizan por ser fragmentos de diáfisis de ungulados empleados para retocar el filo de una herramienta lítica. Estos objetos son relativamente abundantes en contextos del Paleolítico Medio, aunque hay una amplia variabilidad...

The basicranium contains multiple synchondroses potentially informative for estimating the developmental stage of individuals. The basilar synchondrosis has been routinely used for this purpose in bioarchaeological, forensic and paleoanthropological research, and studies carried out in modern human populations have shown a close relationship betwee...

The analysis of the locomotor anatomy of Late Pleistocene Homo has largely focused on changes in proximal femur and pelvic morphologies, with much attention centered on the emergence of modern humans. Although much of the focus has been on changes in the proximal femur, some research has also been conducted on tibiae and, to a lesser extent, fibula...

The analysis of the locomotor anatomy of Late Pleistocene Homo has largely focused on changes in proximal femur and pelvic morphologies, with much attention centered on the emergence of modern humans. Although much of the focus has been on changes in the proximal femur, some research has also been conducted on tibiae and, to a lesser extent, fibula...

Ancient DNA analyses have provided key insights into the phylogenetic relationships of our closest hominin relatives and paleoproteomic research is promising to do the same for our ape ancestors. Nevertheless, several aspects of our evolutionary history remain contentious due to the fragmentary nature of the fossil record, the recurrent independent...

We present new datings and a new anthropological study of Early Neolithic human remains found in Galería del Sílex in 1979. This gallery is part of the Cueva Mayor system in the Sierra de Atapuerca. The human fossils attributed to the Neolithic period correspond to a minimum number of three individuals that have been radiocarbon dated to the last t...

Cattle have been a valuable economic resource and cultural icon since prehistory. From the initial expansion of domestic cattle into Europe during the Neolithic period, taurine cattle (Bos taurus) and their wild ancestor, the aurochs (B. primigenius), had overlapping ranges leading to ample opportunities for intentional and unintentional hybridizat...

The forearm skeleton is composed of two bones: the radius and the ulna. This is closely related to manipulative movements. The ulna is part of the elbow joint, whereas the radius and ulna together with the scaphoid and lunate bones, form the wrist joints. Thus, morphofunctional analysis of the adult Sima de los Huesos (SH) forearm bones, provides c...

The shape, size, and content of droppings has long been used to identify their producers. Mammalian scats are very varied, but it is sometimes possible to differentiate faecal morphotypes and to link them to at least family level. Furthermore, since the diameter of the droppings is related to the size of the producer, it is sometimes even possible...

Information regarding the evolution of the neck in genus Homo is hampered owing to a limited fossil record. Neandertals display significant metric and/or morphological differences in all the cervical vertebrae, when compared to Homo sapiens. Thus, the important fossil record from the Middle Pleistocene site of Sima de los Huesos (SH) not only offer...

The recovery of additional mandibular fossils from the Atapuerca Sima de los Huesos (SH) site provides new insights into the evolutionary significance of this sample. In particular, morphological descriptions of the new adult specimens are provided, along with standardized metric data and phylogenetically relevant morphological features for the exp...

Complete Neanderthal skeletons are almost unique findings. A very well-preserved specimen of this kind was discovered in 1993 in the deepest recesses of a karstic system near the town of Altamura in Southern Italy. We present here a detailed description of the cranium, after we virtually extracted it from the surrounding stalagmites and stalactites...

Spatial analysis has been much used to examine the distribution of archaeological remains at Pleistocene sites. However, little is known about the distribution patterns at sites identified as hunting camps, i.e., places occupied over multiple short periods for the capture of animals later transported to a base camp. The present work examines a Nean...

The Sima de los Huesos site in the Sierra de Atapuerca (Burgos, northern Spain) is a Middle Pleistocene locality with the most important accumulation of humans in the European record for this age. In addition to the hominin collection, the Sima has provided numerous faunal remains. Here, we update the carnivoran mammal fauna from the Sima de los Hu...

The postcranial skeleton of fossil hominins is crucial for reconstructing the processes that occurred between the time of death and the recovery of the bones. Thousands of postcranial skeletal fragments from at least 29 hominin individuals have been recovered from the Sima de los Huesos Middle Pleistocene site in Spain. This study's primary objecti...

Since their discovery in 1978, the hominin fossil footprints from Laetoli have been the focus, of extensive research on the locomotion, speed, body size, and behavior of the responsible track-makers (nominally Australopithecus afarensis). In this work, we show that careful examination of walking speed and displacement yields valuable information re...

Previous studies on the morphology of the inner ear (semicircular canals and cochlea) in the Sima de los Huesos hominin sample have provided important results on the evolution of these structures in the Neandertal lineage. Similarly, studies of the anatomy of the external and middle ear cavities of the Sima de los Huesos hominins have also provided...

This work examines the possible behaviour of Neanderthal groups at the Cueva Des-Cubierta (central Spain) via the analysis of the latter’s archaeological assemblage. Alongside evidence of Mousterian lithic industry, Level 3 of the cave infill was found to contain an assemblage of mammalian bone remains dominated by the crania of large ungulates, so...

Here we present an updated inventory and study of pectoral girdle remains recovered from the Sima de los Huesos (SH) site. Here, we describe the key morphological traits of adults and, for the first time, subadult specimens. Because morphological traits can change with age, we also discuss some shortcomings related to age estimation in postcranial...

The bony labyrinth contains phylogenetic information that can be used to determine interspecific differences between fossil hominins. The present study conducted a comparative 3D geometric morphometric analysis on the bony labyrinth of the Middle Pleistocene Sima de los Huesos (SH) hominins. The findings of this study corroborate previous multivari...

Since the discovery of a human mandible in 1887 near the present-day city of Banyoles, northeastern Spain, researchers have generally emphasized its archaic features, including the lack of chin structures, and suggested affinities with the Neandertals or European Middle Pleistocene (Chibanian) specimens. Uranium-series and electron spin resonance d...

The study and interpretation of the evidence that demonstrates the controlled use of fire by Neanderthal groups in an archaeological context, i.e. pyroarchaeology, relies on various disciplines which, when put together, could reveal the existence of intentional combustion processes in situ. In the case of the Mousterian site of Cueva Des-Cubierta (...

The identification of anthropogenic signatures in archaeological contexts through preserved molecular markers requires an exhaustive previous experimental work to characterise the specific biomarkers involved in different activities from the anthropic occupation of natural spaces. The site of the Cueva Des-Cubierta Cave (Pinilla del Valle, Madrid),...

Paleolithic assemblages preserved in cave deposits are taphonomically complex due to the effect of alternating hominin and carnivore occupations and the impact of post-depositional processes. Quantifying the effect of post-depositional disturbance, identifying the factors operating in these cave deposits, and defining individual depositional events...

The early Middle Pleistocene human material from Boxgrove (West Sussex, UK) consists of a partial left tibia and two lower incisors from a separate adult individual. These remains derive from deposits assigned to the MIS 13 interglacial at about 480 ka and have been referred to as Homo cf. heidelbergensis. The much larger skeletal sample from the S...

El Portalón de Cueva Mayor cave (Atapuerca, Spain) is a settlement site at the entrance of a natural cave. This Holocene archaeological site shows a record of a long archaeological sequence that includes a Chalcolithic occupation starting from 3090 to 2240 cal. BC 2σ. During this phase, different human activities have been identified: habitational...

The study of dental morphology can be a very useful tool to understand the origin and evolution of Neanderthals in Europe during the Middle Pleistocene (MP). At present, the earliest evidence, ca. 430 ka, of a pre-Neanderthal population in Europe is the hominin sample from Atapuerca-Sima de los Huesos (SH) that present clear dental affinities with...

Santiago de Compostela is, together with Rome and Jerusalem, one of the three main pilgrimage and religious centres for Catholicism. The belief that the remains of St James the Great, one of the twelve apostles of Jesus Christ, is buried there has stimulated, since their reported discovery in the 9th century AD, a significant flow of people from ac...

To assess the phenotypic affinities of the Sima de los Huesos (SH) mandibular incisors dental tissue proportions, and radicular dimensions, relative to Neandertals, recent modern humans (RMH), and a large comparative sample of Pleistocene hominins. Two dimensional (2D) and three dimensional (3D) data were extracted from SH (n = 22) incisors, and co...

El Portalón de Cueva Mayor cave (Atapuerca, Spain) is a settlement site at the entrance of a natural cave. This Holocene archaeological site shows a record of a long archaeological sequence that includes a Chalcolithic occupation starting from 3090 to 2240 cal. BC 20. During this phase, different human activities have been identified: habitational...

Neanderthals have been claimed to have had a selective adaptation to rugged, wooded landscapes that would have partially compensate their high basal metabolic rate and locomotor energetic costs through reducing search time and increasing diet breadth. The archaeological site of the Navalmaíllo rockshelter (Pinilla del Valle, Madrid), located in a m...

The Galería de las Estatuas is a Mousterian site located within the Cueva Mayor-Cueva del Silo karstic system at the Sierra de Atapuerca (Burgos, Spain). This site is characterised by an important Upper Pleistocene stratigraphic sequence in which a large number of lithic artefacts exhibiting clear Mousterian affinities and a rich assemblage of faun...

Evidence of dog consumption at the El Portalón de Cueva Mayor site (Sierra de Atapuerca, Spain) from the Holocene is revealed for the first time. The taxonomical and taphonomical studies of the animal bones from the El Portalón site have been carried out. The morphological and metrical analyses indicate that 130 dog bone remains have been identifie...

The Sima del Elefante site is located within the Sierra de Atapuerca karst system (Burgos, northern Spain), and forms part of a series of important Early, Middle and Late Pleistocene archaeological complexes that have been dated previously with luminescence techniques (Gran Dolina, Galería Complex, Sima de los Huesos, Galería de las Estatuas). This...

Tres son los principales debates en los estudios de evolución humana en Europa: primer poblamiento de nuestro continente, patrón evolutivo durante el Pleistoceno medio, origen del Hombre moderno y su relación con los neandertales. Hasta la fecha, no existe consenso en admitir la presencia humana en Europa antes del episodio isotópico 13, pero el re...

The Sima de los Huesos (SH) hominin assemblage is composed of thousands of fossil fragments, including pieces of crania and mandibles. The main objective of this work is to address the main taphonomic features of the cranial and mandibular remains from the SH sample, including antemortem, perimortem, and postmortem skeletal disturbances. We present...

The Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth‐by‐tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here, we show its application on five hominins from the TD6.2 l...

The Neanderthal site of Cueva Des-Cubierta Cave (CDC, Pinilla del Valle, Madrid, Spain) has yielded an anomalous archaeological assemblage in level 3, characterized by the presence of Mousterian industry, an anthropically modified faunal record dominated by cranial remains of large ungulates with defensive cranial appendages [1-2], and the existenc...

The Middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, northern Spain) has yielded a considerable number of human fossils during the period 1984–2020. Among them, up to 253 maxillary teeth have been recovered. In this article, we present the description of the eight dental classes of the maxilla following the Arizona State Univ...

The Middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, northern Spain) has yielded a considerable number of human fossils during the period 1984–2020. Among them, up to 314 mandibular teeth have been identified. In this second paper dedicated to the dentition we present the description of the eight dental classes of the mandibl...

Los yacimientos del Calvero de la Higuera (Pinilla del Valle, Madrid) son un referente para comprender la forma de vida, relación con el medio y capacidad simbólica de los grupos de neandertales que habitaron este territorio durante el Pleistoceno Superior. Los trabajos arqueológicos que, sobre todo desde 2002, se desarrollan en dicho enclave, situ...

Domestication of horses fundamentally transformed long-range mobility and warfare¹. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling2–4 at Botai, Central Asia around 3500 bc³. Other longstanding candidate regions for horse domestic...

The two- and three-dimensional assessment of dental tissues has become routine in human taxonomic studies throughout the years. Nonetheless, most of our knowledge of the variability of the enamel and dentine dimensions of the human evolutionary lineage comes from the study of permanent dentition, and particularly from molars. This leads to a biased...

The interior of the Iberian Peninsula has few Middle Palaeolithic sites, especially when compared to other areas of the Mediterranean Basin and the northern Spanish region. Few in number too are the zooarchaeological and taphonomic studies that throw light on the relationships between Neanderthal groups, their environment, and the use they made of...

Brown bears (Ursus arctos) diverged from the cave bear lineage c. 1.2 million years ago and likely originated in Asia, where the oldest fossils belong to a Middle Pleistocene chronology. Brown bear fossils from the Middle Pleistocene are scarce in the Iberian Peninsula, especially when compared to the cave bear record and they are mainly located in...

Middle Pleistocene Homo in the Levant Our understanding of the origin, distribution, and evolution of early humans and their close relatives has been greatly refined by recent new information. Adding to this trend, Hershkovitz et al. have uncovered evidence of a previously unknown archaic Homo population, the “Nesher Ramla Homo ” (see the Perspecti...

Circum‐nasal and nasal cavity morphology add to the picture of the Sima de los Huesos specimens as, at one level, representing a distinct morph and, at another, displaying individual variation. They developed a robust, midline‐grooved, three‐dimensional spinal ridge lying anteriorly in the nasal cavity floor that was distended posteriorly over the...

The Chalcolithic levels of El Portalón de Cueva Mayor (Atapuerca, Burgos, Spain) offer a good opportunity to test whether the small-mammal contents of different archaeo-stratigraphical units may be useful to characterize them as independent entities. With that purpose, we studied representative samples of small-mammal remains from the two main cont...

The origin and evolution of hominin mortuary practices are topics of intense interest and debate1–3. Human burials dated to the Middle Stone Age (MSA) are exceedingly rare in Africa and unknown in East Africa1–6. Here we describe the partial skeleton of a roughly 2.5- to 3.0-year-old child dating to 78.3 ± 4.1 thousand years ago, which was recovere...

The study of audition in fossil hominins is of great interest given its relationship with intraspecific vocal communication. While the auditory capacities have been studied in early hominins and in the Middle Pleistocene Sima de los Huesos hominins, less is known about the hearing abilities of the Neanderthals. Here, we provide a detailed approach...

Speleothems are a recognized source of palaeoclimatic information, but their value as a source of signals from human activities in caves with an archaeological record has rarely been explored. Previous studies of speleothems in the Sierra de Atapuerca karst system (Burgos, northern Spain) revealed an important human fossil record, provided informat...

The TD6 unit of the Gran Dolina contains an assemblage of the Early Pleistocene, interpreted firstly as a home base. More recently has been proposed a transported origin of the remains according to the sedimentology. Following this model, the remains should be dragged or lagged in a predictable pattern related to their weight, density, shape, and s...

After 34 years of research and findings in the Middle Pleistocene site of the Sima de los Huesos (SH) of the Sierra de Atapuerca (Burgos, Spain), we present an update of the estimation of the number of individuals (ENI) identified in the SH hominin assemblage. The last ENI, published in 2004, was 28. Although the number of specimens recovered has a...