Homo antecessor

Pleistocene level TD6-2 of the Gran Dolina site (Sierra de Atapuerca, Spain) is the result of anthropogenic accumulation. Hominin groups occupied the cave as a home base, where they brought in, butchered and consumed the carcasses of ungulates and other hominins. In this paper, we reassess the role of carnivores in the formation and/or modification of the assemblage. We employed different methods to explore the scenario in which the TD6-2 assemblage was formed: (1) identifying the actor responsible for tooth marks; (2) determining the frequency of carnivore tooth marks and their distribution; (3) identifying the co-occurrence of modifications (butchering marks and carnivore tooth marks); (4) calculating the percentage of change and the epiphysis to shaft ratio. Carnivore tooth marks are scarce, as is the co-occurrence of hominin and carnivore modifications. However, not all tooth marks have been attributed to a particular agent due to the high equifinality between human and carnivore tooth marks. For these reasons, the frequency of tooth marks and the co-occurrence of modifications have been of little help in interpreting the role of carnivores.

Axial skeletal remains and the epiphyses of the long bones are in large part missing. The percentage of change and the epiphysis to shaft ratio suggest moderate carnivore ravaging activity. Our data indicate that the role of carnivores in TD6-2 seems to have had an impact on the original assemblage after hominins had extracted a large amount of nutrients from the carcasses. Cannibalized hominin remains showed no carnivore tooth marks and have a greater presence of low survival bones compared to ungulate remains. These findings point to a different taphonomic history suggesting that TD6-2 represents a succession of settlements having different characteristics.

Summary: The establishment of the new species ‘Homo antecessor’ was mainly based on the remains of an immature individual ATD6-69, found at the TD6 litostratigraphic unit (LU) of the Gran Dolina site in the Sierra de Atapuerca—Burgos, northern Spain , along with cranial, mandibular and dental traits from other fossil hominid specimens recovered in the same level. TD6 LU was initially dated about 0.78 Ma using ESR-U-series; but recent redatings, employing TL and IRSL methods, suggest that its age is between 0,9-0,95 Ma. It seems that there is general consensus that TD6 LU corresponds to Marine Isotope Stage (MIS) 25. TD6-69 specimen, a partial face with R P3, M1, and germs of M2 and M3; and L I2-M1, was found in the so-called “Aurora stratum”, ~1.5 m below the Brunhes–Matuyama (B–M) geomagnetic boundary. This “stratum” was in fact, an excavation area of only 6 m2 for archaeological test pit purposes. There is no real basis, from an earnest stratigraphic point of view, to consider such sedimentary level as a new litostratigraphic unit and, therefore, “Aurora stratum” should be take into account as an informal archaeo-sedimentary small area whose name was established to gain public attention from the media.

摘要:人属先驱种(Homo antecessor)，是根据西班牙阿塔普尔卡 Gran Dolina 洞穴堆积中发现的人类 化石厘定的。出产人类化石的地层年代测定距今约 90 万年到 78 万年之间。之所以定为先驱种，是因为 最初研究者认为这个西班牙的古人类群是较晚的海德堡人(H. heidelbergensis)和尼安德特人的共同祖 先(H. neanderthalensis )，而在非洲和欧洲发现的海德堡人则已被认为是现代人的直接祖先。目前对 先驱种的争论还没有明确的结论。之所以会形成这种局面，与“先驱种”模式化石的生物学年龄和保存 状况有极大的关系。本文对当初厘定“先驱种”的头骨和面骨的 11 个特定性状做了一个系统的回顾， 尤其是和各类南方古猿 (Australopithecus 和 Praeanthropus)，最早的人属成员能人种 (H. habilis) 和鲁道 夫种 (H. rudolfensis)，人属匠人种 (H. ergaster)，以及人属直立人种 (H. erectus) 面部梨状孔周边和上颌骨功 能区的特征的比较研究。这 11 个头面骨的特征包括:1) 眶下骨骼的冠状面朝向、2) 犬齿窝的存在、3) 上 颌骨下缘外侧端向下的转折切迹、4) 弧状颧骨下缘、5) 梨状孔外侧上颌骨的转折和朝向、6) 梨状孔外缘和 颧骨起点的相对位置、7) 梨状孔下缘鼻棘和侧棘的位置和组合形态、8) 颧骨和上颌骨结节、9) 类现代人的 骨表面重塑模式、10) 突出的鼻骨、和 11) 颞骨的上凸的颞鳞上缘。结果表明“先驱种”的头面骨的定性特 征没有特异性，其组合与周口店的直立人面部基本形态也非常相似，因而作者倾向于认为“先驱种”就是 直立人种的欧洲变异群体。所谓“人属先驱种”可能只是广义的人属直立人种(H. erectus sensu lato)欧洲 代表。目前的化石材料显示欧洲和亚洲的直立人种是非洲匠人种的演变而来。
关键词 : 阿塔普尔卡;直立人;头面部;人类进化;更新世

Present paper briefly reviews the mean archaeological record proposed as evidence of human presence during Lower Pleistocene in the Iberian Peninsula. A critical review questions the ancient chronologies assigned to surface lithic collections recorded in high fluvial terraces. Nowadays, archaeological data that most likely traces the Iberian earliest human occupation are those recorded in Barranco León 5, Fuentenueva 3 (Granada), Gran Dolina, and Sima del Elefante (Sierra de Atapuerca, Burgos). Paying attention to this archaeological and palaeontological record, some hominid groups inhabited some Iberian areas between the normal polarity episode Gilsá (OIS 34) and the paleomagnetic boundary Brunhes/Matuyama (OIS 20), Lower/Middle Pleistocene conventional limit. This chronological proposal for the earliest human occupation on the Iberian Peninsula introduces us in the broader view of the earliest human occupation of Eurasia, giving some reflections about this phenomena and the state of the art.

El presente trabajo pasa somera revista al registro arqueológico propuesto como evidencia de actividad antrópica en la Península Ibérica durante el Pleistoceno inferior. Una revisión crítica permite cuestionar las cronologías antiguas asignadas a concentraciones superficiales localizadas en terrazas fluviales altas, atendiendo a precisiones geomorfológicas y arqueológicas.
Hasta la fecha, los datos con un grado de fiabilidad mínima que podrían registrar el poblamiento peninsular más antiguo son los procedentes de Barranco León 5 y Fuentenueva 3 (Granada) y Gran Dolina y Sima del Elefante (Sierra de Atapuerca, Burgos). Atendiendo a los mismos, algunos grupos de homínidos ocuparon algunas zonas meridionales y centroseptentrionales de la Península Ibérica entre el episodio de polaridad positiva Gilsá (OIS 34) y poco antes del tránsito paleomagnético Brunhes/Matuyama (OIS 20), límite convencional Pleistoceno inferior/medio. Esta propuesta cronoestratigráfica es aprovechada para enmarcar el primer poblamiento peninsular en el contexto de Eurasia realizando una serie de reflexiones en torno a este proceso y el estado actual de su investigación.

The phylogenetic affinities of Homo antecessor, a hominin dating from the early Middle Pleistocene of Europe, are still unclear. In this study we conducted a comprehensive review of the TD6 hypodigm within the context of the historical development of paleoanthropological issues concerning this species. H. antecessor, based on all available craniofacial features to date, displays a midfacial morphology very similar to specimens attributed to Classic Homo erectus, suggesting that H. antecessor is the geographical European variant of Classic H. erectus.

La revisión del registro inferopaleolítico europeo permite identificar tres episodios de poblamiento. El segundo de los mismos, datado entre OIS25/OIS24-OIS19/OIS18, ofrece industrias líticas y fósiles humanos que indican un posible origen esteasiático. Se ofrece una síntesis sobre este episodio de dispersión, relacionado con el reemplazo de faunas galerienses. Se analizan las posibles vías migratorias que siguió y los motivos que pudieron ocasionar su interrupción. También se plantean las relaciones filogenéticas de estas poblaciones, insertándolas en el marco evolutivo del Pleistoceno inferior y medio.

HOMINID DISPERSAL INTO EUROPE AROUND THE MATUYAMA/BRUNHES BOUNDARY: ORIGIN AND SIGNIFICANCE IN HUMAN EVOLUTION

A review of the European Lower Palaeolithic record shows three human dispersal episodes. The second one is dated between OIS25/OIS24-OIS19/OIS18. It is recorded by lithic assemblages and human remains that point to an eastern Asian origin. This paper reviews this hominid dispersal,
related to the Galerian faunal event. Also we analyse the most feasible migratory routes, and the reasons of its abrupt ending. The phylogenetic relationships of these populations are also proposed, putting their context into the human evolutionary sequences of the Lower and Middle Pleistocene.

La revisión del registro inferopaleolítico europeo permite identificar tres episodios de poblamiento. El segundo de los mismos, datado entre OIS25/OIS24-OIS19/OIS18, ofrece industrias líticas y fósiles humanos que indican un posible origen esteasiático. Se ofrece una síntesis sobre este
episodio de dispersión, relacionado con el reemplazo de faunas galerienses. Se analizan las posibles vías migratorias que siguió y los motivos que pudieron ocasionar su interrupción. También se plantean las relaciones filogenéticas de estas poblaciones, insertándolas en el marco evolutivo del
Pleistoceno inferior y medio.

We have carefully read the article by Bermúdez de Castro & Martinón-Torres (2019) titled "What does Homo antecessor tell us about the origin of the 'emergent humanity' that gave rise to Homo sapiens?". The authors update their thoughts on the human remains of the TD6.2 level of Gran Dolina de Atapuerca (Burgos, Spain), and then support their research with previous work by members of the Atapuerca team. This work includes: a) the validity of the Homo antecessor taxon; b) the phylogenetic relationships between this taxon and the line that leads to H. sapiens and H. neanderthalensis; and c) the geographic area of the origin of H. antecessor. The first two points pivot, fundamentally, on the midfacial morphology of the subadult individual ATD6-69 (Hominid 3), a specimen that is said to exhibit modern type facial morphology (e.g.,

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124). The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70þ107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck.

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124).
The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70þ107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the
specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest
of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck.

Olduvai Hominid 8 (OH 8), an articulating set of fossil hominin tarsal and metatarsal bones, is critical to interpretations of the evolution of hominin pedal morphology and bipedal locomotion. It has been suggested that OH 8 may represent the foot of a subadult and may be associated with the OH 7 mandible, the type specimen of Homo habilis. This assertion is based on the presence of what may be unfused distal metatarsal epiphyses. Accurately assessing the skeletal maturity of the OH 8 foot is important for interpretations of the functional morphology and locomotor behavior of Plio-Pleistocene hominins. In this study, we compare metatarsal fusion patterns and internal bone morphology of the lateral metatarsals among subadult hominines (85 modern humans, 48 Pan, and 25 Gorilla) to assess the likelihood that OH 8 belonged to either an adult or subadult hominin. Our results suggest that if OH 8 is indeed from a subadult, then it displays a metatarsal developmental pattern that is unobserved in our comparative sample. In OH 8, the fully fused base of the first metatarsal and the presence of trabecular bone at the distal ends of the second and third metatarsal shafts make it highly improbable that it belonged to a subadult, let alone a subadult that matches the developmental age of the OH 7 mandible. In total, the results of this study suggest that the OH 8 foot most likely belonged to an adult hominin.

The Gran Dolina cave site is famous for having delivered some of the oldest hominin remains of Western Europe (Homo antecessor, ca. 960 ka). Moreover, the evidence of lithic industries throughout the long vertical section suggests occupation on the part of hominins from the latest early Pleistocene (levels TD3/4, TD5, and TD6) to the late middle Pleistocene (level TD10). The Gran Dolina Sondeo Sur (TDS) has furnished a great number of small-vertebrate remains; among them some 40,000 bones are attributed to amphibians and squamates. Although they do not differ specifically from the extant herpetofauna of the Iberian Peninsula, the overlap of their current distribution areas (¼ mutual climatic range method) in Spain can provide mean annual temperatures (MAT), the mean temperatures of the coldest (MTC) and warmest (MTW) months, and mean annual precipitation (MAP) estimations for each sub-level, and their change can be studied throughout the sequence. Results from the squamate and amphibian study indicate that during hominin occupation the MAT (10-13 C) was always slightly warmer than at present in the vicinity of the Gran Dolina Cave, and the MAP (800-1000 mm) was greater than today in the Burgos area. Climatic differences between ''glacial'' and ''interglacial'' phases are poorly marked. Summer temperatures (MTW) show stronger oscillations than winter temperatures (MTC), but seasonality remains almost unchanged throughout the sequence. These results are compared with those for large mammals, small mammals, and pollen analysis, giving a scenario for the palaeoclimatic conditions that occurred during the early to middle Pleistocene in Atapuerca, and hence a scenario for the hominins that once lived in the Sierra de Atapuerca.

Reconstruction of extinct hominin diets is currently a topic of much interest and debate, facilitated by new methods such as the analysis of dental calculus. It has been proposed, based on chemical analyses of calculus, that Neanderthals self-medicated, yet this conclusion has been questioned. Gastrophagy has been suggested as an alternative explanation for the Neanderthal data, based on ethnographic analogies, which show this practice to have been widespread in traditional extant Homo sapiens diets, and nutritional evidence for its benefits at high latitudes. Here we expand the discussion of the potential importance of gastrophagy in human evolution by considering its role for an extant group of tropical foragers, the Hadza of Tanzania, and questioning its role in the diets of extinct tropical hominin species. Gastrophagy is frequently practiced among the Hadza and adult men in particular consume substantial, seasonally variable, amounts of prey guts. In addition to the important fact that gastrophagy is not a rare event, this demographic information may be useful in interpreting evidence from archaeological samples. The consumption of semi-digested chyme would have allowed extinct hominins to gain calories from plant sources without the cost of digesting them, possibly contributing to the encephalisation and shrinking of the gut in genus Homo. As an easy to process food-source, chyme could have likewise been an important food source for the old and the young, potentially playing a part in reducing inter-birth intervals and increasing reproductive success in our lineage. Thus gastrophagy may have played a key part in human evolution and its potentially confounding signal should be considered in future dietary reconstructions.

The kipunji, a recently discovered primate endemic to Tanzania's Southern Highlands and Udzungwa Mountains, was initially referred to the mangabey genus Lophocebus (Cercopithecinae: Papionini), but subsequent molecular analyses showed it to be more closely related to Papio. Its consequent referral to a new genus, Rungwecebus, has met with skepticism among papionin researchers, who have questioned both the robustness of the phylogenetic results and the kipunji's morphological distinctiveness. This circumstance has been exacerbated by the immaturity of the single available specimen (FMNH 187122), an M1-stage juvenile. Therefore, a geometric morphometric analysis of juvenile papionin cranial shape was used to explore the kipunji's phenetic affinities and evaluate morphological support for its separation from Lophocebus. Three-dimensional craniometric landmarks and semi-landmarks were collected on a sample of 124 subadult (dp4-M2 stage) cercopithecid crania. Traditional interlandmark distances were compared and a variety of multivariate statistical shape analyses were performed for the zygomaxillary region (diagnostic in mangabeys) and the cranium as a whole. Raw and size-adjusted interlandmark distances show the kipunji to have a relatively taller, shorter neurocranium and broader face and cranial base than is seen in M1-stage Lophocebus. Principal components and cluster analyses consistently unite the two Lophocebus species but group the kipunji with Cercocebus and/or Macaca. Morphological distances (Mahalanobis D 2 ) between the kipunji and Lophocebus species are comparable to distances between recognized papionin genera. Discriminant function analyses suggest phenetic affinities between the kipunji and Cercocebus/Macaca and do not support the kipunji's classification to Lophocebus or to any other papionin taxon. In canonical plots, the kipunji occupies a region intermediate between macaques and African papionins or groups with Cercocebus, suggesting that it retains basal papionin shape characteristics. In shape comparisons among M1-stage papionins, the kipunji cranium is distinguished from Lophocebus by its relatively unrestricted suborbital fossa, more parasagittally oriented zygomatic arches, and longer auditory tube and from all papionins by its relatively tall, short neurocranium, broad face and cranial base, short nasals, dished nasal profile, and dorsally oriented rostrum. The kipunji is thus a cranially diagnosable phenon with a unique combination of cranial traits that cannot be accommodated within Lophocebus as currently defined. Based upon these results, Rungwecebus appears to be a valid and useful nomen that accurately reflects the morphological diversity of African papionins.

Fecha de aceptación de la versión definitiva: 09-06-05 BIBLID [0514-7336 (2005) 58; 23-36] RESUMEN: El registro inferopaleolítico de Europa permite establecer un mínimo de tres fases de poblamiento del subcontinente. Las evidencias seguras de actividad antrópica más antigua, localizadas todas ellas en la Península Ibérica, son escasas y se datan entre 1,5 y 1,25 Ma BP. No existen datos que indiquen su continuidad. Las similitudes de estos conjuntos líticos con las colecciones documentadas circa 1,8 Ma BP en Dmanisi (Georgia) invitan a considerar la posibilidad de que todos estos yacimientos manifiesten una dispersión de homínidos con origen en África oriental. La misma parece estar relacionada con las transformaciones de faunas apreciadas en el Cáucaso, Próximo Oriente y Europa en torno al límite Plio-Pleistoceno convencional, episodio que implicó la difusión de especies africanas en estos ecosistemas. Una segunda fase de poblamiento del continente, restringido al área mediterránea, está representada por una nómina de yacimientos algo más nutrida. La misma parece no prolongarse más allá de la transición Pleistoceno inferior/Pleistoceno medio, datándose entre OIS 25-OIS 24 y la transición OIS 19-OIS 18. Los datos paleoantropológicos y arqueológicos de estos yacimientos parecen señalar un origen asiático para estas poblaciones, cuya penetración en Europa coincide con el episodio de reemplazo faunístico Galeriense, que implicó la difusión de taxa asiáticos en los ecosistemas europeos. Tras un aparente hiato del registro europeo, a partir de la transición OIS 16-OIS 15, se cuenta con evidencia de actividad antrópica más abundante. El rango geográfico ocupado por grupos de homínidos se amplió hasta alcanzar áreas tan septentrionales como el sur de Gran Bretaña y el norte de Alemania durante las fases isotópicas pares. En términos generales, los yacimientos correspondientes a esta tercera fase manifiestan la aparición en Europa de nuevas tecnologías y pautas de comportamiento, así como un mejor conocimiento del medio por parte de sus protagonistas. Mientras los ensayos de colonización anteriores no parecen tener éxito, los datos paleoantropológicos y arqueológicos del registro documentado desde circa 550 Ka BP en adelante indican claramente signos de continuidad de este poblamiento, enlazando con las primeras manifestaciones de las tecnologías y los tipos humanos característicos del Paleolítico medio.

The Lower Pleistocene TD6 level at the Gran Dolina site in the Sierra de Atapuerca (Burgos, Spain) has yielded nine ribs that represent a minimum of three individuals of the species, Homo antecessor. We present a detailed morphological and metric study of these costal elements, including the siding and anatomical position of all of the rib remains. The adult or nearly adult ribs are also metrically compared with other fossil hominins and with modern comparative samples. The costal elements recovered to date from the TD6 level at Gran Dolina can neither confirm nor reject the hypothesis that H. antecessor had a large thorax, similar to that of Neandertals. However, the fragmentary evidence of the H. antecessor thoracic skeleton is not inconsistent with this suggestion based on other skeletal elements, such as clavicles.

Efectivamente, son las coordenadas de longitud y latitud de la localización de los yacimientos de Atapuerca, y así figuran en la inscripción como bien Patrimonio de la Humanidad de la UNESCO (World Heritage) desde el 2 de diciembre del año 2000.

The kipunji, a recently discovered primate endemic to Tanzania's Southern Highlands and Udzungwa Mountains, was initially referred to the mangabey genus Lophocebus (Cercopithecinae: Papionini), but subsequent molecular analyses showed it to be more closely related to Papio. Its consequent referral to a new genus, Rungwecebus, has met with skepticism among papionin researchers, who have questioned both the robustness of the phylogenetic results and the kipunji's morphological distinctiveness. This circumstance has been exacerbated by the immaturity of the single available specimen (FMNH 187122), an M1-stage juvenile. Therefore, a geometric morphometric analysis of juvenile papionin cranial shape was used to explore the kipunji's phenetic affinities and evaluate morphological support for its separation from Lophocebus. Three-dimensional craniometric landmarks and semi-landmarks were collected on a sample of 124 subadult (dp4-M2 stage) cercopithecid crania. Traditional interlandmark distances were compared and a variety of multivariate statistical shape analyses were performed for the zygomaxillary region (diagnostic in mangabeys) and the cranium as a whole. Raw and size-adjusted interlandmark distances show the kipunji to have a relatively taller, shorter neurocranium and broader face and cranial base than is seen in M1-stage Lophocebus. Principal components and cluster analyses consistently unite the two Lophocebus species but group the kipunji with Cercocebus and/or Macaca. Morphological distances (Mahalanobis D 2 ) between the kipunji and Lophocebus species are comparable to distances between recognized papionin genera. Discriminant function analyses suggest phenetic affinities between the kipunji and Cercocebus/Macaca and do not support the kipunji's classification to Lophocebus or to any other papionin taxon. In canonical plots, the kipunji occupies a region intermediate between macaques and African papionins or groups with Cercocebus, suggesting that it retains basal papionin shape characteristics. In shape comparisons among M1-stage papionins, the kipunji cranium is distinguished from Lophocebus by its relatively unrestricted suborbital fossa, more parasagittally oriented zygomatic arches, and longer auditory tube and from all papionins by its relatively tall, short neurocranium, broad face and cranial base, short nasals, dished nasal profile, and dorsally oriented rostrum. The kipunji is thus a cranially diagnosable phenon with a unique combination of cranial traits that cannot be accommodated within Lophocebus as currently defined. Based upon these results, Rungwecebus appears to be a valid and useful nomen that accurately reflects the morphological diversity of African papionins.

The kipunji, a recently discovered primate endemic to Tanzania's Southern Highlands and Udzungwa Mountains, was initially referred to the mangabey genus Lophocebus (Cercopithecinae: Papionini), but subsequent molecular analyses showed it to be more closely related to Papio. Its consequent referral to a new genus, Rungwecebus, has met with skepticism among papionin researchers, who have questioned both the robustness of the phylogenetic results and the kipunji's morphological distinctiveness. This circumstance has been exacerbated by the immaturity of the single available specimen (FMNH 187122), an M1-stage juvenile. Therefore, a geometric morphometric analysis of juvenile papionin cranial shape was used to explore the kipunji's phenetic affinities and evaluate morphological support for its separation from Lophocebus. Three-dimensional craniometric landmarks and semi-landmarks were collected on a sample of 124 subadult (dp4-M2 stage) cercopithecid crania. Traditional interlandmark distances were compared and a variety of multivariate statistical shape analyses were performed for the zygomaxillary region (diagnostic in mangabeys) and the cranium as a whole. Raw and size-adjusted interlandmark distances show the kipunji to have a relatively taller, shorter neurocranium and broader face and cranial base than is seen in M1-stage Lophocebus. Principal components and cluster analyses consistently unite the two Lophocebus species but group the kipunji with Cercocebus and/or Macaca. Morphological distances (Mahalanobis D 2 ) between the kipunji and Lophocebus species are comparable to distances between recognized papionin genera. Discriminant function analyses suggest phenetic affinities between the kipunji and Cercocebus/Macaca and do not support the kipunji's classification to Lophocebus or to any other papionin taxon. In canonical plots, the kipunji occupies a region intermediate between macaques and African papionins or groups with Cercocebus, suggesting that it retains basal papionin shape characteristics. In shape comparisons among M1-stage papionins, the kipunji cranium is distinguished from Lophocebus by its relatively unrestricted suborbital fossa, more parasagittally oriented zygomatic arches, and longer auditory tube and from all papionins by its relatively tall, short neurocranium, broad face and cranial base, short nasals, dished nasal profile, and dorsally oriented rostrum. The kipunji is thus a cranially diagnosable phenon with a unique combination of cranial traits that cannot be accommodated within Lophocebus as currently defined. Based upon these results, Rungwecebus appears to be a valid and useful nomen that accurately reflects the morphological diversity of African papionins.

Kidnaping Europe... one time and another

European Early Palaeolithic's record of Europe shows, at least, three human occupation episodes. The oldest one is only recorded in the Iberian Peninsula. The sites are scarce and dated between 1,5 and 1,25 Ma BP. The lithic assemblages are very similar to those recorded in Dmanisi (Georgia) circa 1,8 Ma BP and may represent the same hominid dispersal event. This is related to the faunal changes detected in Caucasus, Middle East and European ecosystems by the Plio-Pleistocene conventional limit.

This event involved the diffusion of African taxa in those regions. A second human occupation of Europe episode, restricted into the Mediterranean area, it is recorded by a slightly more abundant number of archaeological sites. They are dated between late Early Pleistocene and Matuyama/Brunhes limit, berween OIS 25-OIS 24 and OIS 19-OIS 18 transition. Paleoantropological and archaeological record points to an Asian origin of this hominid dispersal, related with the Galerian faunal event. After a new archaeological record hiatus, there are new evidences of anthropic activity in Europe since OIS 16-OIS 15. At this time the European geographic range populated by hominids increases, reaching as northern latitudes as South Great Britain and North Germany during the warm isotopic stages. In general terms, locations representing this third phase involved the introduction in Europe of new technologies and behavioural trends, as a better knowledge of the environment. While the previous human occupation events don't seem to be successful, paleoanthropological and archaeological data recorded from circa 550 Ka BP points to demographic continuity, linking with the earliest manifestations of the technologies and human populations characteristic of the European Middle Paleolithic.

El registro inferopaleolítico de Europa permite establecer un mínimo de tres fases de poblamiento del subcontinente. Las evidencias seguras de actividad antrópica más antigua, localizadas todas ellas en la Península Ibérica, son escasas y se datan entre 1,5 y 1,25 Ma BP. No existen datos que indiquen su continuidad. Las similitudes de estos conjuntos líticos con las colecciones documentadas circa 1,8 Ma BP en Dmanisi (Georgia) invitan a considerar la posibilidad de que todos estos yacimientos manifiesten una dispersión de homínidos con origen en África oriental. La misma parece estar relacionada con las transformaciones de faunas apreciadas en el Cáucaso, Próximo Oriente y Europa en torno al límite Plio-Pleistoceno convencional, episodio que implicó la difusión de especies africanas en estos ecosistemas. Una segunda fase de poblamiento del continente, restringido al área mediterránea, está representada por una nómina de yacimientos algo más nutrida. La misma parece no prolongarse más allá de la transición Pleistoceno inferior/ Pleistoceno medio, datándose entre OIS 25-OIS 24 y la transición OIS 19-OIS 18. Los datos paleoantropológicos y arqueológicos de estos yacimientos parecen señalar un origen asiático para estas poblaciones, cuya penetración en Europa coincide con el episodio de reemplazo faunístico Galeriense, que implicó la difusión de taxa asiáticos en los ecosistemas europeos. Tras un aparente hiato del registro europeo, a partir de la transición OIS 16-OIS 15, se cuenta con evidencia de actividad antrópica más abundante. El rango geográfico ocupado por grupos de homínidos se amplió hasta alcanzar áreas tan septentrionales como el sur de Gran Bretaña y el norte de Alemania durante las fases isotópicas pares. En términos generales, los yacimientos correspondientes a esta tercera fase manifiestan la aparición en Europa de nuevas tecnologías y pautas de comportamiento, así como un mejor conocimiento del medio por parte de sus protagonistas.

Mientras los ensayos de colonización anteriores no parecen tener éxito, los datos paleoantropológicos y arqueológicos del registro documentado desde circa 550 Ka BP en adelante indican claramente signos de continuidad de este poblamiento, enlazando con las primeras manifestaciones de las tecnologías y los tipos humanos característicos del Paleolítico medio.

El trabajo aborda una revisión del registro europeo correspondiente al Paleolítico inferior europeo hasta el estadio isotópico 8, a fin de determinar cuándo y como se produjo el proceso de primer poblamiento del continente, así como las rutas de dispersión hacia el continente; su relación con el primer poblamiento de Asia, en el que debe contextualizarse el de Europa, y la especie humanas implicadas en el proceso. Una vez diagnosticada la validez de los diferentes yacimientos considerados, el registro inferopaleolítico de Europa permite establecer un mínimo de tres fases de poblamiento del subcontinente. La evidencias seguras de actividad antrópica más antigua, localizadas todas ellas en la Península Ibérica, son escasas y se datan entre 1,5 y 1,25 ma. BP. No existen datos que indiquen su continuidad. Las similitudes de estos conjuntos líticos con las colecciones documentadas circa 1,8 ma. BP en Dmanisi (Georgia) invitan a considerar la posibilidad de que todos estos yacimientos manifiestan una dispersión de homínidos con origen en África oriental. La mixta parece estar relacionada con las transformaciones de faunas apreciadas en el Cáucaso, Próximo Oriente y Europa en tramo al límite Plio-Pleistoceno convencional, episodio que implicó la difusión de especies africanas en estos ecosistemas. Una segunda fase de poblacimiento del continente, restringido al área mediterráneas, está representada por una nómina de yacimientos algo más nutrida. La misma parece no prolongarse más allá de la transición Pleistoceno inferior/Pleistoceno, datándose entre las trasiciones OIS 25/OIS 24 y OIS 19/OIS 18. Los datos paleoantropológicos y arqueológicos y paleontológicos de estos yacimientos parecen señalar un origen asiático para estas poblaciones, cuya penetración en Europa coincide con el episodio de reemplazo faunístico Galeriense, que implicó la difusión de taxa asiáticos en los ecosistemas europeos.

The earliest hominin occupation of Europe is one of the most debated topics in palaeoanthropology. However, the purportedly oldest of the Early Pleistocene sites in Eurasia lack precise age control and contain stone tools rather than human fossil remains. Here we report the discovery of a human mandible associated with an assemblage of Mode 1 lithic tools and faunal remains bearing traces of hominin processing, in stratigraphic level TE9 at the site of the Sima del Elefante, Atapuerca, Spain. Level TE9 has been dated to the Early Pleistocene (approximately 1.2-1.1 Myr), based on a combination of palaeomagnetism, cosmogenic nuclides and biostratigraphy. The Sima del Elefante site thus emerges as the oldest, most accurately dated record of human occupation in Europe, to our knowledge. The study of the human mandible suggests that the first settlement of Western Europe could be related to an early demographic expansion out of Africa. The new evidence, with previous findings in other Atapuerca sites (level TD6 from Gran Dolina), also suggests that a speciation event occurred in this extreme area of the Eurasian continent during the Early Pleistocene, initiating the hominin lineage represented by the TE9 and TD6 hominins

We have carefully read the article by Bermúdez de Castro & Martinón-Torres (2019) titled "What does Homo antecessor tell us about the origin of the 'emergent humanity' that gave rise to Homo sapiens?". The authors update their thoughts on the human remains of the TD6.2 level of Gran Dolina de Atapuerca (Burgos, Spain), and then support their research with previous work by members of the Atapuerca team. This work includes: a) the validity of the Homo antecessor taxon; b) the phylogenetic relationships between this taxon and the line that leads to H. sapiens and H. neanderthalensis; and c) the geographic area of the origin of H. antecessor. The first two points pivot, fundamentally, on the midfacial morphology of the subadult individual ATD6-69 (Hominid 3), a specimen that is said to exhibit modern type facial morphology (e.g.,

In this paper we present new data pertaining to the paleo-landscape characteristics at the Acheulian site of Revadim, on the southern coastal plain of Israel. Sedimentological, isotopic, granulometric and micromorphological studies showed that the archaeological remains accumulated in an active fluvial environment where channel action, overbank flooding and episodic inundation occurred. Measurements of total organic matter and its carbon isotopic

The Lower Pleistocene TD6 level at the Gran Dolina site in the Sierra de Atapuerca (Burgos, Spain) has yielded nine ribs that represent a minimum of three individuals of the species, Homo antecessor. We present a detailed morphological and metric study of these costal elements, including the siding and anatomical position of all of the rib remains. The adult or nearly adult ribs are also metrically compared with other fossil hominins and with modern comparative samples. The costal elements recovered to date from the TD6 level at Gran Dolina can neither confirm nor reject the hypothesis that H. antecessor had a large thorax, similar to that of Neandertals. However, the fragmentary evidence of the H. antecessor thoracic skeleton is not inconsistent with this suggestion based on other skeletal elements, such as clavicles.

The kipunji, a recently discovered primate endemic to Tanzania's Southern Highlands and Udzungwa Mountains, was initially referred to the mangabey genus Lophocebus (Cercopithecinae: Papionini), but subsequent molecular analyses showed it to be more closely related to Papio. Its consequent referral to a new genus, Rungwecebus, has met with skepticism among papionin researchers, who have questioned both the robustness of the phylogenetic results and the kipunji's morphological distinctiveness. This circumstance has been exacerbated by the immaturity of the single available specimen (FMNH 187122), an M1-stage juvenile. Therefore, a geometric morphometric analysis of juvenile papionin cranial shape was used to explore the kipunji's phenetic affinities and evaluate morphological support for its separation from Lophocebus. Three-dimensional craniometric landmarks and semi-landmarks were collected on a sample of 124 subadult (dp4-M2 stage) cercopithecid crania. Traditional interlandmark distances were compared and a variety of multivariate statistical shape analyses were performed for the zygomaxillary region (diagnostic in mangabeys) and the cranium as a whole. Raw and size-adjusted interlandmark distances show the kipunji to have a relatively taller, shorter neurocranium and broader face and cranial base than is seen in M1-stage Lophocebus. Principal components and cluster analyses consistently unite the two Lophocebus species but group the kipunji with Cercocebus and/or Macaca. Morphological distances (Mahalanobis D 2 ) between the kipunji and Lophocebus species are comparable to distances between recognized papionin genera. Discriminant function analyses suggest phenetic affinities between the kipunji and Cercocebus/Macaca and do not support the kipunji's classification to Lophocebus or to any other papionin taxon. In canonical plots, the kipunji occupies a region intermediate between macaques and African papionins or groups with Cercocebus, suggesting that it retains basal papionin shape characteristics. In shape comparisons among M1-stage papionins, the kipunji cranium is distinguished from Lophocebus by its relatively unrestricted suborbital fossa, more parasagittally oriented zygomatic arches, and longer auditory tube and from all papionins by its relatively tall, short neurocranium, broad face and cranial base, short nasals, dished nasal profile, and dorsally oriented rostrum. The kipunji is thus a cranially diagnosable phenon with a unique combination of cranial traits that cannot be accommodated within Lophocebus as currently defined. Based upon these results, Rungwecebus appears to be a valid and useful nomen that accurately reflects the morphological diversity of African papionins.

This paper presents a geoarchaeological study of Middle and Upper Palaeolithic (Châtelperronian, Aurignacian and Solutrean) occupations preserved at the Bordes-Fitte rockshelter in Central France. The lithostratigraphic sequence is composed of near-surface sedimentary facies with vertical and lateral variations, in a context dominated by run-off and gravitational sedimentary processes. Field description and micromorphological analysis permit us to reconstruct several episodes of sediment slope-wash and endokarst dynamics, with hiatuses and erosional phases. The archaeostratigraphic succession includes Châtelperronian artefacts, inter-stratified between Middle Palaeolithic and Aurignacian occupations. Systematic refitting and spatial analysis reveal that the Châtelperronian point production and flake blanks retouched into denticulates, all recovered in the same stratigraphic unit, result from distinct and successive occupations and are not a 'transitional' Middle to Upper P...