Adrián Pablos | Universidad Complutense de Madrid (original) (raw)

Papers by Adrián Pablos

Research paper thumbnail of Metatarsals and foot phalanges from the Sima de los Huesos Middle Pleistocene site (Atapuerca, Burgos, Spain

The Anatomical Record, 2024

This study provides a complete, updated and illustrated inventory, as well as a comprehensive stu... more 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 established a minimum number of individuals (MNI) of 17, which represent 58.6% of the 29 dental individuals identified within the SH sample. An exclusive or autoapomorphic combination of traits can be recognized within the SH hominin foot sample. A few traits appear primitive or plesiomorphic when compared with earlier Homo individuals and other recent modern humans. There are other metrical and morphological traits that SH hominins and Neandertals have in common that
sometimes represent shared derived traits in this evolutionary line, most of which are probably related to robusticity. Furthermore, some exclusive autoapomorphic traits are observed in the SH sample: a very broad first metatarsal, long and broad hallucal proximal foot phalanges and possibly extremely robust lateral distal foot phalanges compared to those of Neandertals and modern humans. In these last traits, the SH metatarsals and pedal phalanges are even more robust than in Neandertals. They are herein named as “hyper-Neandertal” traits, which could suggest a slight gracilization process in this evolutionary line, at least in the hallux toe. Finally, some paleobiological inferences are made in relation to body size (stature and body mass) and some associations are proposed within the SH sample.

Research paper thumbnail of Stress in a Middle Pleistocene hominid (Atapuerca, Spain): periosteal reaction compatible with fatigue fracture in a metatarsal bone

Sima de los Huesos (SH) site (Atapuerca, Spain) has yielded a large collection of Middle Pleistoc... more Sima de los Huesos (SH) site (Atapuerca, Spain) has yielded a large collection of Middle Pleistocene human fossils that represent the ancestors of the Neandertals. Among these fossils there are some metatarsal bones, AT-534 being one of them. This IV right metatarsal bone preserves the proximal epiphysis, 2/3 of the diaphysis, and lacks the distal epiphysis; its dimensions are compatible with those of an adult individual. New bone formation was clearly distinguished on the diaphysis of the bone, affecting mainly the dorso-lateral surface. It is the nature and distribution of the diaphyseal periostitis that is of primary interest here. Radiographs of the area show that the reaction is not invasive of the underlying cortical bone and there are no visible fracture-lines. Periostitis is commonly observed as a secondary response to a pathogen and can be part of, or a reaction to, a pathological condition in the underlying bone. It may also be a primary response to local or systemic infec...

Research paper thumbnail of Cueva de los Torrejones revisited. New insights on the paleoecology inland Iberia during the Late Pleistocene

Quaternary Science Reviews, 2021

The interior of the Iberian Peninsula has orographic conditions that make this territory especial... more The interior of the Iberian Peninsula has orographic conditions that make this territory especially vulnerable to Quaternary climate oscillations and which actually could have made it decisive for Paleolithic human populations at critical points. For this reason, the information provided by paleon-tological sites is important for reconstructing climatic and environmental conditions during the Late Pleistocene and understanding how they influenced the species that inhabited them, including humans. Nevertheless, the archaeo-paleontological record is scarce in central Iberia for the Late Pleistocene. A central Iberian site that is key to addressing this issue is Cueva de los Torrejones, which was discovered and excavated during the nineties. Clues indicating the presence of Neandertal populations near the cave site were announced during prior field excavations, including Neandertal remains, Middle Paleolithic artifacts, and evidence of anthropic exploitation of faunal resources at the site. Here we report the new results from the recent excavations and research, including detailed studies on stratigraphy, micro-morphology, macro and microvertebrate paleontology, physical and molecular anthropology, taphonomy and zooarchaeology, and analysis of lithic and pottery remains. Our research has led to the detection of three Prehistoric chronologies recorded at the site. The oldest episode corresponds to between MIS 5 and MIS 4 in which the cave was used by carnivores. The second episode is represented by a faunal association dated to 30.0 ka cal BP and is indicative of cooler and more arid environmental conditions and, therefore, compatible with the worsening climate detected previously for MIS 3 in this area. The last episode corresponds to the Chalcolithic, directly dated to~5000 cal BP in which humans used the cavity for funerary purposes. The DNA analysis of the human remain was assigned to mtDNA haplogroup K, which was originated in the Near East and reached western Europe through the Neolithic expansion. Human occupation during the Paleolithic has been ruled out, including Paleolithic human remains and any kind of anthropic intervention on the Hermann’s tortoise and leopard as was previously proposed at the site.

Research paper thumbnail of Direct dating of the human navicular from the Cueva de los Torrejones (Guadalajara, Spain)

Radiocarbon, 2020

Pleistocene human remains are rare inland on the Iberian Peninsula. Most are considered Neanderta... more Pleistocene human remains are rare inland on the Iberian Peninsula. Most are considered Neandertals, but anthropological analyses and direct dating are rare. Recently, we published a study of a navicular from this region found in the Torrejones Cave. The results showed it differed from that of Neandertals and it was re-identified as Homo sapiens. Following the previous stratigraphic and biochronologic descriptions, we suggested that it could correspond to an Upper Paleolithic human, since the navicular was apparently recovered in the Late Pleistocene from an in situ unit. Direct radiocarbon dating from this fossil (4855-5036 cal BP), believed to be the only Paleolithic Homo sapiens from inland Iberia, as well as other hominin and faunal remains from the site, show that the human bones actually date to the Chalcolithic. The unexpectedly recent chronology for the navicular implies that there is no evidence of human fossils from the Upper Paleolithic in Torrejones Cave. Thus, any date from the Middle/Upper Paleolithic human record should be taken with caution until in-depth paleoanthropological, stratigraphical and/or direct dating studies are conducted. Extraordinary caution is recommended when human remains are recovered from apparently Paleolithic units in contexts bearing Holocene sepulchral units on the uppermost levels and/or some evidence of bioturbation.

Research paper thumbnail of A Partial Neandertal Foot From the Late Middle Paleolithic of Amud Cave, Israel

PaleoAnthropology, 2020

Excavations of Amud Cave in 1991-1994 yielded 14 hominin skeletal specimens (Amud 5-19) in additi... more Excavations of Amud Cave in 1991-1994 yielded 14 hominin skeletal specimens (Amud 5-19) in addition to those recovered in the 1960s. Amud 9 is a partial right distal leg and foot that preserves portions of the distal tibia, talus, first metatarsal, first proximal phalanx, and a middle and distal phalanx of digit II-IV. The bones are fairly small and likely belonged to a female. The talus features a strongly projecting fibular articular facet in common with Neandertals and many tali from Sima de los Huesos. Discriminant analysis of the talus shows that its nearest match lies among tali from Sima de los Huesos, a result primarily attributable to its moderately enlarged posterior trochlear articular breadth. The first metatarsal falls among Neandertals in discriminant space. The pedal phalan-ges are short and broad, in common with other Neandertals. The length of the first metatarsal and talus predict a female's stature of 160-166cm and the width of the talar trochlea predicts a body mass of 59.9kg. The bones were found within anthropogenic deposits dated date to 55 ka, very close in time to the proposed main pulse of Neandertal interbreeding, as inferred from living people's DNA, and slightly before the first appearance of Upper Paleolithic industries.

Research paper thumbnail of Neandertal foot remains from Regourdou 1 (Montignac-sur-Vézère, Dordogne, France)

Journal of Human Evolution, 2019

Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minim... more Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minimum of two Neandertal individuals. The first individual (Regourdou 1) is represented by a partial skeleton while the second one is represented by a calcaneus. The foot remains of Regourdou 1 have been used in a number of comparative studies, but to date a full description and comparison of all the foot remains from the Regourdou 1 Neandertal, coming from the old excavations and from the recent reanalysis of the faunal remains, does not exist. Here, we describe and comparatively assess the Regourdou 1 tarsals, metatarsals and phalanges. They display traits observed in other Neandertal feet, which are different from some traits of the Sima de los Huesos (Atapuerca) hominins and of Middle Paleolithic, Upper Paleolithic and recent modern humans. These Neandertal features are: a rectangular talar trochlea with a large lateral malleolar facet, a broad talar head, a broad calcaneus with a projecting sustentaculum tali, a wide and wedged navicular with a projecting medial tubercle, large and wide bases of the lateral metatarsals, and mediolaterally expanded and robust phalanges that also show hallux valgus in a strongly built hallux.

Research paper thumbnail of A Neandertal foot phalanx from the Galería de las Estatuas site (Sierra de Atapuerca, Spain)

American Journal of Physical Anthropology, 2018

Objectives: The Galería de las Estatuas site (GE), a new Mousterian site at the Sierra de Atapuer... more Objectives: The Galería de las Estatuas site (GE), a new Mousterian site at the Sierra de Atapuerca site complex (Spain), has revealed a Late Pleistocene detrital sequence with at least five lithostratigraphic units. These units have yielded evidence of Mousterian occupations with sporadic carnivore activity, and have provided datings of 80–112 ka BP using single-grain optically stimulated luminescence. This places the sequence at the end of MIS5 and beginning of the MIS4. We described here a complete adult human distal foot phalanx (GE-1573) recovered during the 2017 field season in the interface between lithostratigraphic units 3 and 4 (107–112 ka BP) in the GE-I test pit.
Materials and method: This phalanx (GE-1573) probably corresponds to the fifth toe from the right side due to the medial deviation of the distal tuberosity. We compared the metric variables of this phalanx to several fossil and recent Homo samples.
Results: Neandertals display foot phalanges that are broader and more robust than those of recent humans. Despite the scarcity of well-identified distal phalanges in the Homo fossil record, the GE-1573 phalanx is broad, long and robust when compared with recent and Upper Paleolithic modern humans.
Discussion: These traits, which align the GE-1573 foot phalanx with the Neandertal morphology, are consistent with the stratigraphic context, likely corresponding to one of the oldest Late Neandertals found inland on the Iberian Peninsula. Additionally, it provides the first evidence of a Neandertal human fossil in a stratigraphic context in the Sierra de Atapuerca.

Research paper thumbnail of Long-term patterns of body mass and stature evolution within the hominin lineage

Royal Society Open Science, 2017

Body size is a central determinant of a species' biology and adaptive strategy, but the number of... more Body size is a central determinant of a species' biology and adaptive strategy, but the number of reliable estimates of hominin body mass and stature have been insufficient to determine long-term patterns and subtle interactions in these size components within our lineage. Here, we analyse 254 body mass and 204 stature estimates from a total of 311 hominin specimens dating from 4.4 Ma to the Holocene using multi-level chronological and taxonomic analytical categories. The results demonstrate complex temporal patterns of body size variation with phases of relative stasis intermitted by periods of rapid increases. The observed trajectories could result from punctuated increases at speciation events, but also differential proliferation of large-bodied taxa or the extinction of small-bodied populations. Combined taxonomic and temporal analyses show that in relation to australopithecines, early Homo is characterized by significantly larger average body mass and stature but retains considerable diversity, including small body sizes. Within later Homo, stature and body mass evolution follow different trajectories: average modern stature is maintained from ca 1.6 Ma, while consistently higher body masses are not established until the Middle Pleistocene at ca 0.5–0.4 Ma, likely caused by directional selection related to colonizing higher latitudes. Selection against small-bodied individuals (less than 40 kg; less than 140 cm) after 1.4 Ma is associated with a decrease in relative size variability in later Homo species compared with earlier Homo and australopithecines. The isolated small-bodied individuals of Homo naledi (ca 0.3 Ma) and Homo floresiensis (ca 100–60 ka) constitute important exceptions to these general patterns, adding further layers of complexity to the evolution of body size within the genus Homo. At the end of the Late Pleistocene and Holocene, body size in Homo sapiens declines on average, but also extends to lower limits not seen in comparable frequency since early Homo.

Research paper thumbnail of Taxonomic reassignment of the Paleolithic human navicular from Cueva de los Torrejones (Guadalajara, Spain)

Taxonomic reassignment of the Paleolithic human navicular from Cueva de los Torrejones (Guadalajara, Spain)

Evidence of human activity and hominin remains are very scarce inland on the Iberian Peninsula. T... more Evidence of human activity and hominin remains are very scarce inland on the Iberian Peninsula. This fact raises the issue of the scarcity of evidence that Paleolithic Homo sapiens occupied this area outside of the littoral margins (Atlantic, Cantabrian, and Mediterranean coasts). Here, we comparatively describe a human right adult navicular bone recovered in the Cueva de los Torrejones site, located in the village of Tamajón (Guadalajara, Spain). This fossil was preliminarily established as belonging to Homo cf. neanderthalensis, due to the late Pleistocene faunal association, mainly because of the presence of Crocuta crocuta and Panthera pardus. The metrical and morphological study of the navicular T93-S3.27 from Cueva de los Torrejones clearly differentiates it from Neandertals and their ancestors, the hominins from Sima de los Huesos, allowing for this fossil to be taxonomically assigned with confidence as H. sapiens. The navicular from the Cueva de los Torrejones is absolutely and relatively medio-laterally narrow with a low wedging index as those of fossil and modern H. sapiens, and clearly different of Neandertals. The increased discoveries and publications of new naviculars belonging to genus Homo, together with the findings of P. pardus and C. crocuta in more recent chronologies in the Iberian Peninsula, are compatible with this reevaluation.We propose a probable chronology for this fossil between 12 and 15 ka and ca. 25 ka, based on the biostratigraphy and the oldest presence of H. sapiens in the Iberian
Peninsula. This work confirms the human presence within the Iberian Peninsula during the Upper Paleolithic and reopens the question of the peopling of the inner Peninsula during this period.

Research paper thumbnail of The Sima de los Huesos Crania: Analysis of the cranial breakage patterns

Research paper thumbnail of The foot in the Homo fossil record

In this article, the foot in the Homo fossil record throughout the world is reviewed. The main pr... more In this article, the foot in the Homo fossil record throughout the world is reviewed. The main problem with the study of foot remains is the paucity of fossils from this anatomical area, in particular from the earlier members of the genus Homo. In spite of this, a comprehensive review of the morphology of the entire fossil record for the foot has been achieved. All the fossils belonging to the genus Homo are proposed to be biped due to the presence of longitudinal and transversal arches, the robusticity pattern of the metatarsals and an adducted hallux. Even in the early members of the genus Homo, the morphology
of the foot is modern-like, with size being practically the only variation observed. Of the foot remains attributed to the genus Homo, two morphotypes become apparent: small-sized and large-sized individuals. It is important though to take into account that the earliest Homo feet belonging to smaller individuals could not belong to the genus Homo. Later, a new robust bauplan appears in the Homo fossil record for the
foot represented by Homo erectus/ergaster, Homo antecessor, the hominins from Sima de los Huesos and Neandertals. Finally, modern humans display long feet that are gracile compared with their ancestors. An examination of the morphology of the Neandertal foot and of the foot from Sima de los Huesos confirms the evolutionary relationship between these two populations. However, enough differences exist between the
two samples to indicate that they are in fact morphologically distinct. A parallel gracilization process is proposed in both modern humans and Neandertals.

Research paper thumbnail of Metric and morphological analysis of the foot in the Middle Pleistocene population of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

The preservation of Homo fossil foot remains prior to modern humans and Neandertals is very scarc... more The preservation of Homo fossil foot remains prior to modern humans and Neandertals is very scarce throughout the fossil record. Understanding foot morphology in human evolution is essential to know taxonomic processes and to acquiring information about corporal size. Here, we present a comprehensive study of the large foot remains sample recovered from the Middle Pleistocene site of Sima de los Huesos (SH). The minimum number of individuals (MNI) has been established at 16 by the foot remains. As is the case with other elements of the foot, metric and morphological differences in the feet between Middle and Late Pleistocene hominins tend to be subtle. However, an exclusive combination of traits can be recognized in the feet of the SH hominins. The SH hominins and Neandertals display tali with short necks and broad lateral malleolar facets, broad calcanei with long bodies and projected sustentaculum tali, stout naviculars, as well as robust metatarsals and phalanges. Nevertheless, the feet from SH are characterized by a very
broad lateral malleolar facet in the talus and a very broad sustentaculum tali in the calcaneus, even more so than in Neandertals with respect to both traits. More importantly, the head of the talus from SH is narrower than the Neandertal's broad talus head, and the short intermediate cuneiforms found at SH distinguish them from the other comparative samples. The bodymass estimation based on the trochlear breadth of the talus provides a mean of 69.7 ± 10.0 kg for the SH hominins, similar to that determined by the femoral head
and significantly lower than that obtained from the bi-iliac breadth.

Research paper thumbnail of Lethal interpersonal violence in the Middle Pleistocene

Evidence of interpersonal violence has been documented previously in Pleistocene members of the ... more Evidence of interpersonal violence has been documented previously in Pleistocene members
of the genus Homo, but only very rarely has this been posited as the possible manner
of death. Here we report the earliest evidence of lethal interpersonal violence in the hominin
fossil record. Cranium 17 recovered from the Sima de los Huesos Middle Pleistocene site
shows two clear perimortem depression fractures on the frontal bone, interpreted as being
produced by two episodes of localized blunt force trauma. The type of injuries, their location,
the strong similarity of the fractures in shape and size, and the different orientations and implied
trajectories of the two fractures suggest they were produced with the same object in
face-to-face interpersonal conflict. Given that either of the two traumatic events was likely
lethal, the presence of multiple blows implies an intention to kill. This finding shows that the
lethal interpersonal violence is an ancient human behavior and has important implications
for the accumulation of bodies at the site, supporting an anthropic origin.

Research paper thumbnail of Regreso a la Cueva de los Casares (Guadalajara). Un nuevo proyecto de investigación para el yacimiento del Seno A.

ARPI. Arqueología y Prehistoria del Interior peninsular, 2: 68-89. , 2015

Research paper thumbnail of Early Pleistocene human hand phalanx from the Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain)

In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante ... more In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave site (TE - Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting greater overall body robusticity in these populations or a higher gracility in modern humans. The age of level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the morphology of the proximal hand phalanges and, thus, the morphology of the hand could have remained stable over the last 1.2e1.3 Ma. Taking into account the evidence recently provided by a metacarpal from Kaitio (Kenya) from around 1.42 Ma, we argue that modern hand morphology is present in the genus Homo subsequent to Homo habilis.

Research paper thumbnail of Human calcanei from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

The existence of calcanei in the fossil record prior to modern humans and Neandertals is very sca... more The existence of calcanei in the fossil record prior to modern humans and Neandertals is very scarce. This skeletal element is fundamental to understanding the evolution of the morphology of the foot in human evolution. Here we present and metrically and comparatively describe 29 calcaneus remains from the Middle Pleistocene site of Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These calcanei belong to 15 individuals (nine adults, two adolescents and four immature individuals). The metric and morphological differences in the calcanei among Middle and Late Pleistocene hominins tend to be subtle. However, the calcanei from SH are broad and robust with large articular surfaces and most significantly, exhibit a very projected sustentaculum tali. A biomechanical and phylogenetic interpretation is proffered to explain the observed morphology of these calcanei. It has been possible to propose tentative sex assignments for the SH calcanei based on size, using methods similar to those used to establish sex from the talus bones from SH. The estimation of stature based on the calcaneus provides a mean of 175.3 cm for males and 160.6 for females, which is similar to that obtained using other skeletal parts from the site. In sum, the SH calcanei are robust with a proportionally long tubercle and a projected sustentaculum tali, which are traits shared by Neandertals.

Research paper thumbnail of Neandertal roots: Cranial and chronological evidence from Sima de los Huesos

Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analy... more Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.

Research paper thumbnail of Nuevos restos humanos procedentes de la Cueva de la Zarzamora (Segovia, España)

Munibe, 2013

""The present paper reports the recently recovered human bone remains from the Cueva de la Zarzam... more ""The present paper reports the recently recovered human bone remains from the Cueva de la Zarzamora in the southernmost limits of the Castilian Plateau of the Iberian Peninsula (Segovia, Spain). A total of two teeth and nine human bones from the trunk, pelvis and foot regions have been recovered. A complete inventory, metrical and morphological study has been carried out. All this remains could correspond to a single individual and its age-of-death was very likely around 10 years old. The direct radiocarbon dating of one of the rib fragments using ultrafiltration pre-treatment protocols has provided a date of 4749 ± 34 years BP which would place these remains in the Chalcolithic archaeological period. This piece of evidence is of great relevance, taking into account the scarcity of human bone remains of this age in this region of the Iberian Peninsula."

Research paper thumbnail of Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the... more Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima
de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and
three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this
study has identified unique morphological characteristics of the SH tali. They are vertically shorter than
those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet
than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the
hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a
broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene
H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for
the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for
males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.

Research paper thumbnail of From toe to head: Use of robust regression methods in stature estimation based

Stature estimation is a standard procedure in the fields of forensic and biological anthropology,... more Stature estimation is a standard procedure in the fields of forensic and biological anthropology, bioarchaeology and paleoanthropology, in order to gain biological insights into the individuals/populations studied. The most accurate stature estimation method is based on anatomical reconstruction (i.e., the
Fully method), followed by type I regression equations (e.g., ordinary least squares – OLS) based on long bones, preferably from the lower limb. In some cases, due to the fragmentary nature of the osseous material recovered, stature estimates have to rely on other elements, such as foot remains. In this study, we explore stature estimation based on different foot bones: the talus, calcaneus, and metatarsals 1–4 in Afro- and Euroamericans of both sexes. The approach undertaken in this study is novel for two reasons. First, individual estimates for each bone are provided, and tarsals and metatarsals are combined in order to obtain more accurate estimates. Second, robust statistical methods based on type I regression equations are used, namely least trimmed squares (LTS). Our results show that the best individual bones
for estimating stature are the first and second metatarsal and both the talus and the calcaneus. The combination of a tarsal and a metatarsal bone slightly improves the accuracy of the stature estimate.

Research paper thumbnail of Metatarsals and foot phalanges from the Sima de los Huesos Middle Pleistocene site (Atapuerca, Burgos, Spain

The Anatomical Record, 2024

This study provides a complete, updated and illustrated inventory, as well as a comprehensive stu... more 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 established a minimum number of individuals (MNI) of 17, which represent 58.6% of the 29 dental individuals identified within the SH sample. An exclusive or autoapomorphic combination of traits can be recognized within the SH hominin foot sample. A few traits appear primitive or plesiomorphic when compared with earlier Homo individuals and other recent modern humans. There are other metrical and morphological traits that SH hominins and Neandertals have in common that
sometimes represent shared derived traits in this evolutionary line, most of which are probably related to robusticity. Furthermore, some exclusive autoapomorphic traits are observed in the SH sample: a very broad first metatarsal, long and broad hallucal proximal foot phalanges and possibly extremely robust lateral distal foot phalanges compared to those of Neandertals and modern humans. In these last traits, the SH metatarsals and pedal phalanges are even more robust than in Neandertals. They are herein named as “hyper-Neandertal” traits, which could suggest a slight gracilization process in this evolutionary line, at least in the hallux toe. Finally, some paleobiological inferences are made in relation to body size (stature and body mass) and some associations are proposed within the SH sample.

Research paper thumbnail of Stress in a Middle Pleistocene hominid (Atapuerca, Spain): periosteal reaction compatible with fatigue fracture in a metatarsal bone

Sima de los Huesos (SH) site (Atapuerca, Spain) has yielded a large collection of Middle Pleistoc... more Sima de los Huesos (SH) site (Atapuerca, Spain) has yielded a large collection of Middle Pleistocene human fossils that represent the ancestors of the Neandertals. Among these fossils there are some metatarsal bones, AT-534 being one of them. This IV right metatarsal bone preserves the proximal epiphysis, 2/3 of the diaphysis, and lacks the distal epiphysis; its dimensions are compatible with those of an adult individual. New bone formation was clearly distinguished on the diaphysis of the bone, affecting mainly the dorso-lateral surface. It is the nature and distribution of the diaphyseal periostitis that is of primary interest here. Radiographs of the area show that the reaction is not invasive of the underlying cortical bone and there are no visible fracture-lines. Periostitis is commonly observed as a secondary response to a pathogen and can be part of, or a reaction to, a pathological condition in the underlying bone. It may also be a primary response to local or systemic infec...

Research paper thumbnail of Cueva de los Torrejones revisited. New insights on the paleoecology inland Iberia during the Late Pleistocene

Quaternary Science Reviews, 2021

The interior of the Iberian Peninsula has orographic conditions that make this territory especial... more The interior of the Iberian Peninsula has orographic conditions that make this territory especially vulnerable to Quaternary climate oscillations and which actually could have made it decisive for Paleolithic human populations at critical points. For this reason, the information provided by paleon-tological sites is important for reconstructing climatic and environmental conditions during the Late Pleistocene and understanding how they influenced the species that inhabited them, including humans. Nevertheless, the archaeo-paleontological record is scarce in central Iberia for the Late Pleistocene. A central Iberian site that is key to addressing this issue is Cueva de los Torrejones, which was discovered and excavated during the nineties. Clues indicating the presence of Neandertal populations near the cave site were announced during prior field excavations, including Neandertal remains, Middle Paleolithic artifacts, and evidence of anthropic exploitation of faunal resources at the site. Here we report the new results from the recent excavations and research, including detailed studies on stratigraphy, micro-morphology, macro and microvertebrate paleontology, physical and molecular anthropology, taphonomy and zooarchaeology, and analysis of lithic and pottery remains. Our research has led to the detection of three Prehistoric chronologies recorded at the site. The oldest episode corresponds to between MIS 5 and MIS 4 in which the cave was used by carnivores. The second episode is represented by a faunal association dated to 30.0 ka cal BP and is indicative of cooler and more arid environmental conditions and, therefore, compatible with the worsening climate detected previously for MIS 3 in this area. The last episode corresponds to the Chalcolithic, directly dated to~5000 cal BP in which humans used the cavity for funerary purposes. The DNA analysis of the human remain was assigned to mtDNA haplogroup K, which was originated in the Near East and reached western Europe through the Neolithic expansion. Human occupation during the Paleolithic has been ruled out, including Paleolithic human remains and any kind of anthropic intervention on the Hermann’s tortoise and leopard as was previously proposed at the site.

Research paper thumbnail of Direct dating of the human navicular from the Cueva de los Torrejones (Guadalajara, Spain)

Radiocarbon, 2020

Pleistocene human remains are rare inland on the Iberian Peninsula. Most are considered Neanderta... more Pleistocene human remains are rare inland on the Iberian Peninsula. Most are considered Neandertals, but anthropological analyses and direct dating are rare. Recently, we published a study of a navicular from this region found in the Torrejones Cave. The results showed it differed from that of Neandertals and it was re-identified as Homo sapiens. Following the previous stratigraphic and biochronologic descriptions, we suggested that it could correspond to an Upper Paleolithic human, since the navicular was apparently recovered in the Late Pleistocene from an in situ unit. Direct radiocarbon dating from this fossil (4855-5036 cal BP), believed to be the only Paleolithic Homo sapiens from inland Iberia, as well as other hominin and faunal remains from the site, show that the human bones actually date to the Chalcolithic. The unexpectedly recent chronology for the navicular implies that there is no evidence of human fossils from the Upper Paleolithic in Torrejones Cave. Thus, any date from the Middle/Upper Paleolithic human record should be taken with caution until in-depth paleoanthropological, stratigraphical and/or direct dating studies are conducted. Extraordinary caution is recommended when human remains are recovered from apparently Paleolithic units in contexts bearing Holocene sepulchral units on the uppermost levels and/or some evidence of bioturbation.

Research paper thumbnail of A Partial Neandertal Foot From the Late Middle Paleolithic of Amud Cave, Israel

PaleoAnthropology, 2020

Excavations of Amud Cave in 1991-1994 yielded 14 hominin skeletal specimens (Amud 5-19) in additi... more Excavations of Amud Cave in 1991-1994 yielded 14 hominin skeletal specimens (Amud 5-19) in addition to those recovered in the 1960s. Amud 9 is a partial right distal leg and foot that preserves portions of the distal tibia, talus, first metatarsal, first proximal phalanx, and a middle and distal phalanx of digit II-IV. The bones are fairly small and likely belonged to a female. The talus features a strongly projecting fibular articular facet in common with Neandertals and many tali from Sima de los Huesos. Discriminant analysis of the talus shows that its nearest match lies among tali from Sima de los Huesos, a result primarily attributable to its moderately enlarged posterior trochlear articular breadth. The first metatarsal falls among Neandertals in discriminant space. The pedal phalan-ges are short and broad, in common with other Neandertals. The length of the first metatarsal and talus predict a female's stature of 160-166cm and the width of the talar trochlea predicts a body mass of 59.9kg. The bones were found within anthropogenic deposits dated date to 55 ka, very close in time to the proposed main pulse of Neandertal interbreeding, as inferred from living people's DNA, and slightly before the first appearance of Upper Paleolithic industries.

Research paper thumbnail of Neandertal foot remains from Regourdou 1 (Montignac-sur-Vézère, Dordogne, France)

Journal of Human Evolution, 2019

Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minim... more Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minimum of two Neandertal individuals. The first individual (Regourdou 1) is represented by a partial skeleton while the second one is represented by a calcaneus. The foot remains of Regourdou 1 have been used in a number of comparative studies, but to date a full description and comparison of all the foot remains from the Regourdou 1 Neandertal, coming from the old excavations and from the recent reanalysis of the faunal remains, does not exist. Here, we describe and comparatively assess the Regourdou 1 tarsals, metatarsals and phalanges. They display traits observed in other Neandertal feet, which are different from some traits of the Sima de los Huesos (Atapuerca) hominins and of Middle Paleolithic, Upper Paleolithic and recent modern humans. These Neandertal features are: a rectangular talar trochlea with a large lateral malleolar facet, a broad talar head, a broad calcaneus with a projecting sustentaculum tali, a wide and wedged navicular with a projecting medial tubercle, large and wide bases of the lateral metatarsals, and mediolaterally expanded and robust phalanges that also show hallux valgus in a strongly built hallux.

Research paper thumbnail of A Neandertal foot phalanx from the Galería de las Estatuas site (Sierra de Atapuerca, Spain)

American Journal of Physical Anthropology, 2018

Objectives: The Galería de las Estatuas site (GE), a new Mousterian site at the Sierra de Atapuer... more Objectives: The Galería de las Estatuas site (GE), a new Mousterian site at the Sierra de Atapuerca site complex (Spain), has revealed a Late Pleistocene detrital sequence with at least five lithostratigraphic units. These units have yielded evidence of Mousterian occupations with sporadic carnivore activity, and have provided datings of 80–112 ka BP using single-grain optically stimulated luminescence. This places the sequence at the end of MIS5 and beginning of the MIS4. We described here a complete adult human distal foot phalanx (GE-1573) recovered during the 2017 field season in the interface between lithostratigraphic units 3 and 4 (107–112 ka BP) in the GE-I test pit.
Materials and method: This phalanx (GE-1573) probably corresponds to the fifth toe from the right side due to the medial deviation of the distal tuberosity. We compared the metric variables of this phalanx to several fossil and recent Homo samples.
Results: Neandertals display foot phalanges that are broader and more robust than those of recent humans. Despite the scarcity of well-identified distal phalanges in the Homo fossil record, the GE-1573 phalanx is broad, long and robust when compared with recent and Upper Paleolithic modern humans.
Discussion: These traits, which align the GE-1573 foot phalanx with the Neandertal morphology, are consistent with the stratigraphic context, likely corresponding to one of the oldest Late Neandertals found inland on the Iberian Peninsula. Additionally, it provides the first evidence of a Neandertal human fossil in a stratigraphic context in the Sierra de Atapuerca.

Research paper thumbnail of Long-term patterns of body mass and stature evolution within the hominin lineage

Royal Society Open Science, 2017

Body size is a central determinant of a species' biology and adaptive strategy, but the number of... more Body size is a central determinant of a species' biology and adaptive strategy, but the number of reliable estimates of hominin body mass and stature have been insufficient to determine long-term patterns and subtle interactions in these size components within our lineage. Here, we analyse 254 body mass and 204 stature estimates from a total of 311 hominin specimens dating from 4.4 Ma to the Holocene using multi-level chronological and taxonomic analytical categories. The results demonstrate complex temporal patterns of body size variation with phases of relative stasis intermitted by periods of rapid increases. The observed trajectories could result from punctuated increases at speciation events, but also differential proliferation of large-bodied taxa or the extinction of small-bodied populations. Combined taxonomic and temporal analyses show that in relation to australopithecines, early Homo is characterized by significantly larger average body mass and stature but retains considerable diversity, including small body sizes. Within later Homo, stature and body mass evolution follow different trajectories: average modern stature is maintained from ca 1.6 Ma, while consistently higher body masses are not established until the Middle Pleistocene at ca 0.5–0.4 Ma, likely caused by directional selection related to colonizing higher latitudes. Selection against small-bodied individuals (less than 40 kg; less than 140 cm) after 1.4 Ma is associated with a decrease in relative size variability in later Homo species compared with earlier Homo and australopithecines. The isolated small-bodied individuals of Homo naledi (ca 0.3 Ma) and Homo floresiensis (ca 100–60 ka) constitute important exceptions to these general patterns, adding further layers of complexity to the evolution of body size within the genus Homo. At the end of the Late Pleistocene and Holocene, body size in Homo sapiens declines on average, but also extends to lower limits not seen in comparable frequency since early Homo.

Research paper thumbnail of Taxonomic reassignment of the Paleolithic human navicular from Cueva de los Torrejones (Guadalajara, Spain)

Taxonomic reassignment of the Paleolithic human navicular from Cueva de los Torrejones (Guadalajara, Spain)

Evidence of human activity and hominin remains are very scarce inland on the Iberian Peninsula. T... more Evidence of human activity and hominin remains are very scarce inland on the Iberian Peninsula. This fact raises the issue of the scarcity of evidence that Paleolithic Homo sapiens occupied this area outside of the littoral margins (Atlantic, Cantabrian, and Mediterranean coasts). Here, we comparatively describe a human right adult navicular bone recovered in the Cueva de los Torrejones site, located in the village of Tamajón (Guadalajara, Spain). This fossil was preliminarily established as belonging to Homo cf. neanderthalensis, due to the late Pleistocene faunal association, mainly because of the presence of Crocuta crocuta and Panthera pardus. The metrical and morphological study of the navicular T93-S3.27 from Cueva de los Torrejones clearly differentiates it from Neandertals and their ancestors, the hominins from Sima de los Huesos, allowing for this fossil to be taxonomically assigned with confidence as H. sapiens. The navicular from the Cueva de los Torrejones is absolutely and relatively medio-laterally narrow with a low wedging index as those of fossil and modern H. sapiens, and clearly different of Neandertals. The increased discoveries and publications of new naviculars belonging to genus Homo, together with the findings of P. pardus and C. crocuta in more recent chronologies in the Iberian Peninsula, are compatible with this reevaluation.We propose a probable chronology for this fossil between 12 and 15 ka and ca. 25 ka, based on the biostratigraphy and the oldest presence of H. sapiens in the Iberian
Peninsula. This work confirms the human presence within the Iberian Peninsula during the Upper Paleolithic and reopens the question of the peopling of the inner Peninsula during this period.

Research paper thumbnail of The Sima de los Huesos Crania: Analysis of the cranial breakage patterns

Research paper thumbnail of The foot in the Homo fossil record

In this article, the foot in the Homo fossil record throughout the world is reviewed. The main pr... more In this article, the foot in the Homo fossil record throughout the world is reviewed. The main problem with the study of foot remains is the paucity of fossils from this anatomical area, in particular from the earlier members of the genus Homo. In spite of this, a comprehensive review of the morphology of the entire fossil record for the foot has been achieved. All the fossils belonging to the genus Homo are proposed to be biped due to the presence of longitudinal and transversal arches, the robusticity pattern of the metatarsals and an adducted hallux. Even in the early members of the genus Homo, the morphology
of the foot is modern-like, with size being practically the only variation observed. Of the foot remains attributed to the genus Homo, two morphotypes become apparent: small-sized and large-sized individuals. It is important though to take into account that the earliest Homo feet belonging to smaller individuals could not belong to the genus Homo. Later, a new robust bauplan appears in the Homo fossil record for the
foot represented by Homo erectus/ergaster, Homo antecessor, the hominins from Sima de los Huesos and Neandertals. Finally, modern humans display long feet that are gracile compared with their ancestors. An examination of the morphology of the Neandertal foot and of the foot from Sima de los Huesos confirms the evolutionary relationship between these two populations. However, enough differences exist between the
two samples to indicate that they are in fact morphologically distinct. A parallel gracilization process is proposed in both modern humans and Neandertals.

Research paper thumbnail of Metric and morphological analysis of the foot in the Middle Pleistocene population of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

The preservation of Homo fossil foot remains prior to modern humans and Neandertals is very scarc... more The preservation of Homo fossil foot remains prior to modern humans and Neandertals is very scarce throughout the fossil record. Understanding foot morphology in human evolution is essential to know taxonomic processes and to acquiring information about corporal size. Here, we present a comprehensive study of the large foot remains sample recovered from the Middle Pleistocene site of Sima de los Huesos (SH). The minimum number of individuals (MNI) has been established at 16 by the foot remains. As is the case with other elements of the foot, metric and morphological differences in the feet between Middle and Late Pleistocene hominins tend to be subtle. However, an exclusive combination of traits can be recognized in the feet of the SH hominins. The SH hominins and Neandertals display tali with short necks and broad lateral malleolar facets, broad calcanei with long bodies and projected sustentaculum tali, stout naviculars, as well as robust metatarsals and phalanges. Nevertheless, the feet from SH are characterized by a very
broad lateral malleolar facet in the talus and a very broad sustentaculum tali in the calcaneus, even more so than in Neandertals with respect to both traits. More importantly, the head of the talus from SH is narrower than the Neandertal's broad talus head, and the short intermediate cuneiforms found at SH distinguish them from the other comparative samples. The bodymass estimation based on the trochlear breadth of the talus provides a mean of 69.7 ± 10.0 kg for the SH hominins, similar to that determined by the femoral head
and significantly lower than that obtained from the bi-iliac breadth.

Research paper thumbnail of Lethal interpersonal violence in the Middle Pleistocene

Evidence of interpersonal violence has been documented previously in Pleistocene members of the ... more Evidence of interpersonal violence has been documented previously in Pleistocene members
of the genus Homo, but only very rarely has this been posited as the possible manner
of death. Here we report the earliest evidence of lethal interpersonal violence in the hominin
fossil record. Cranium 17 recovered from the Sima de los Huesos Middle Pleistocene site
shows two clear perimortem depression fractures on the frontal bone, interpreted as being
produced by two episodes of localized blunt force trauma. The type of injuries, their location,
the strong similarity of the fractures in shape and size, and the different orientations and implied
trajectories of the two fractures suggest they were produced with the same object in
face-to-face interpersonal conflict. Given that either of the two traumatic events was likely
lethal, the presence of multiple blows implies an intention to kill. This finding shows that the
lethal interpersonal violence is an ancient human behavior and has important implications
for the accumulation of bodies at the site, supporting an anthropic origin.

Research paper thumbnail of Regreso a la Cueva de los Casares (Guadalajara). Un nuevo proyecto de investigación para el yacimiento del Seno A.

ARPI. Arqueología y Prehistoria del Interior peninsular, 2: 68-89. , 2015

Research paper thumbnail of Early Pleistocene human hand phalanx from the Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain)

In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante ... more In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave site (TE - Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting greater overall body robusticity in these populations or a higher gracility in modern humans. The age of level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the morphology of the proximal hand phalanges and, thus, the morphology of the hand could have remained stable over the last 1.2e1.3 Ma. Taking into account the evidence recently provided by a metacarpal from Kaitio (Kenya) from around 1.42 Ma, we argue that modern hand morphology is present in the genus Homo subsequent to Homo habilis.

Research paper thumbnail of Human calcanei from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

The existence of calcanei in the fossil record prior to modern humans and Neandertals is very sca... more The existence of calcanei in the fossil record prior to modern humans and Neandertals is very scarce. This skeletal element is fundamental to understanding the evolution of the morphology of the foot in human evolution. Here we present and metrically and comparatively describe 29 calcaneus remains from the Middle Pleistocene site of Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These calcanei belong to 15 individuals (nine adults, two adolescents and four immature individuals). The metric and morphological differences in the calcanei among Middle and Late Pleistocene hominins tend to be subtle. However, the calcanei from SH are broad and robust with large articular surfaces and most significantly, exhibit a very projected sustentaculum tali. A biomechanical and phylogenetic interpretation is proffered to explain the observed morphology of these calcanei. It has been possible to propose tentative sex assignments for the SH calcanei based on size, using methods similar to those used to establish sex from the talus bones from SH. The estimation of stature based on the calcaneus provides a mean of 175.3 cm for males and 160.6 for females, which is similar to that obtained using other skeletal parts from the site. In sum, the SH calcanei are robust with a proportionally long tubercle and a projected sustentaculum tali, which are traits shared by Neandertals.

Research paper thumbnail of Neandertal roots: Cranial and chronological evidence from Sima de los Huesos

Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analy... more Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.

Research paper thumbnail of Nuevos restos humanos procedentes de la Cueva de la Zarzamora (Segovia, España)

Munibe, 2013

""The present paper reports the recently recovered human bone remains from the Cueva de la Zarzam... more ""The present paper reports the recently recovered human bone remains from the Cueva de la Zarzamora in the southernmost limits of the Castilian Plateau of the Iberian Peninsula (Segovia, Spain). A total of two teeth and nine human bones from the trunk, pelvis and foot regions have been recovered. A complete inventory, metrical and morphological study has been carried out. All this remains could correspond to a single individual and its age-of-death was very likely around 10 years old. The direct radiocarbon dating of one of the rib fragments using ultrafiltration pre-treatment protocols has provided a date of 4749 ± 34 years BP which would place these remains in the Chalcolithic archaeological period. This piece of evidence is of great relevance, taking into account the scarcity of human bone remains of this age in this region of the Iberian Peninsula."

Research paper thumbnail of Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the... more Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima
de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and
three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this
study has identified unique morphological characteristics of the SH tali. They are vertically shorter than
those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet
than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the
hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a
broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene
H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for
the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for
males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.

Research paper thumbnail of From toe to head: Use of robust regression methods in stature estimation based

Stature estimation is a standard procedure in the fields of forensic and biological anthropology,... more Stature estimation is a standard procedure in the fields of forensic and biological anthropology, bioarchaeology and paleoanthropology, in order to gain biological insights into the individuals/populations studied. The most accurate stature estimation method is based on anatomical reconstruction (i.e., the
Fully method), followed by type I regression equations (e.g., ordinary least squares – OLS) based on long bones, preferably from the lower limb. In some cases, due to the fragmentary nature of the osseous material recovered, stature estimates have to rely on other elements, such as foot remains. In this study, we explore stature estimation based on different foot bones: the talus, calcaneus, and metatarsals 1–4 in Afro- and Euroamericans of both sexes. The approach undertaken in this study is novel for two reasons. First, individual estimates for each bone are provided, and tarsals and metatarsals are combined in order to obtain more accurate estimates. Second, robust statistical methods based on type I regression equations are used, namely least trimmed squares (LTS). Our results show that the best individual bones
for estimating stature are the first and second metatarsal and both the talus and the calcaneus. The combination of a tarsal and a metatarsal bone slightly improves the accuracy of the stature estimate.

Research paper thumbnail of The evolution of body size within the genus Homo: new empirical data and theoretical perspectives

Body size is one of the most important determinants of the biology of a species, as it correlates... more Body size is one of the most important determinants of the biology of a species, as it correlates with life history, energetic expenditure, diet, thermoregulation, and home range size, among other factors. Although the evolution of body size within the genus Homo is an important issue, the most influential large-scale studies have been performed over 20 years ago, with a recent interest in this issue only in the last few years. In the meantime, the widely accepted interpretation that there was a major shift in body size with the origin of Homo ergaster/erectus when compared Homo habilis, Homo rudolfensis, and australopithecines has come under criticism. Recent analyses have demonstrated that body size within early Homo is spatially and temporally variable, only showing significant increase in the Koobi Fora region after 1.7 Mya (Will & Stock, 2015). Broad temporal analyses of body size have highlighted the significant increase in body mass during the late Middle and Late Pleistocene (Ruff, 1997), and that brain size increases correspond closely with body mass increases throughout the Pleistocene (Grabowski, 2016). Even though new body size data is now accumulating rapidly for various parts of the hominin record (Arsuaga et al., 2015; Grabowski et al., 2015; Will & Stock, 2015), no study comparable to the scope of Ruff et al. (1997) has since been performed.
In this paper we investigate taxonomic, spatial and temporal variation in two components of body size within the genus Homo: body mass and stature. We combine size estimates of hominin fossils from our own studies with other published data, resulting in the largest sample for a single study so far (n=319). The body size estimates cover roughly four million years (4.1 Mya – 11 ka) and derive from African, European and Asian specimens, including several genera and species of hominins. This data set allows for a detailed assessment of body size evolution within the genus Homo and relative to earlier hominins.
Analyses of the body size estimates demonstrate that: a) the origins of the genus Homo are characterized by a significant increase in body size compared to australopithecines and paranthropines, but also feature abundant spatial and temporal variation within an enlarged size range; b) members of Homo erectus/ergaster are marked by a diversification in body mass and stature rather than directional increase; c) a consistent and universal increase in body size is only established in Middle Pleistocene hominins (e.g. Sima de los Huesos, Atapuerca; Homo heidelbergensis), Neanderthals and modern humans after ca. 0.5 Mya; d) selection against smaller body mass and stature occurred in the late Early and Middle Pleistocene, and; e) there are no simple latitudinal trends in the variation of body size estimates within Middle and Late Pleistocene Homo in Europe. These results have implications for studies concerned with human dispersal and encephalization, and more generally for how we interpret the evolution and biology of our genus. In light of the above, rather than focusing exclusively on species means and unidirectional models, perspectives that emphasize spatio-temporal variability and phenotypic plasticity might be more fruitful frameworks for interpreting the evolution of body size in our genus.

Research paper thumbnail of Linya. A new Late Glacial partial skeleton from Cova Gran de Santa Linya site (Lleida, Spain)

12th meeting of the European Society for the study of Human Evolution (ESHE), 2022

In the Iberian Peninsula there are many Upper Paleolithic archaeological sites with long sequence... more In the Iberian Peninsula there are many Upper Paleolithic archaeological sites with long sequences, especially in the northern, Mediterranean and Atlantic regions. However, it is remarkable the scarcity of human fossil record for this period. Most of the (scarce) fossils consist in cranio-dental remains, but just a few preserve postcranial Homo sapiens remains [1-3]. This is more relevant if we compare this human record with the wealth of human fossils and burials from France, Italy, Germany and Central Europe. In this work, we present, describe and comparatively study a new human partial skeleton from Cova Gran de Santa Linya site (Lleida, Catalunya, Spain) belonging to the Late Glacial period.
The Cova Gran de Santa Linya site is located in the first ranges of the SE Pre-Pyrenees, ca. 385 m a.s.l.. It represents a large archaeological rock shelter with a long chrono-cultural sequence spanning the Middle Paleolithic, Early Upper Paleolithic, Late Upper Paleolithic to Late Prehistory [4].
In a recent excavation of 9 m2 test-pit called “Sector V” several human remains were discovered corresponding to a human partial skeleton preserving fragmentary long bones, fragments of the pelvis, and some cranial, vertebral, hand and foot remains. Until now, it is remarkable the absence of teeth and the small representation of cranial, hand and foot fossils. Direct ultrafiltered radiocarbon dating of a human fibular fragment provided a date of 12,310 ± 40 BP (Beta-587946) deriving the chronometrical range 14,808-14,091 cal yr BP (2 ∂; 14,808-14,707 cal BP (14.3%) /14,468-14,091 cal BP (81.1%)) that frames the discovery in the Late Pleistocene.
Before the anthropological and taphonomic study, an in-depth restoration and conservation process was necessary due to the fragility and fragmentary nature of the bones. We provide a complete metric and morphological study of this individual for the first time within a chronologically and regionally comparative framework. The age at death determination for all the human remains suggests an adult age, based on the completely fused epiphysis on all the bones. There is no age at death or laterality incompatibilities, and therefore probably all the human sample belonged to the same individual, a likely adult female. Some probably pathological traumatic lesions and periosteal reactions have been observed in the radius, the ulna, the phalanges, the tibia and the fibula.
The taphonomic analysis of the skeletal remains reveals the total absence of anthropic intervention of the skeleton such as cut or percussion marks. This, rules out cannibalism as is the case with other individuals of Magdalenian chronology in Europe [5]. Tooth marks compatible with a small carnivore have been documented on both tibiae. This allows inferring punctual scavenging of the skeleton and could explain the absence of some anatomical elements. Finally, postmortem modifications typical of processes occurring after burial such as postmortem fractures or modification by plant roots are the most common alterations in this individual. These taphonomic characteristics together with the anatomical relationship of the elements of the same individual suggest a rapid burial of the corpse, compatible with funerary practices as a provisory scenario without discounting her natural or accidental death. In sum, the partial skeleton of Cova Gran, nicknamed Linya, constitutes an important addition to the Late Glacial human fossil record of South-Western Europe.