Elena Santos

New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal)

Significance We describe a recently discovered cranium from the Aroeira cave in Portugal dated to around 400 ka. This specimen is the westernmost Middle Pleistocene cranium of Europe and is one of the earliest fossils from this region associated with Acheulean tools. Unlike most other Middle Pleistocene finds, which are of uncertain chronology, the Aroeira 3 cranium is firmly dated to around 400 ka and was in direct association with abundant faunal remains and stone tools. In addition, the presence of burnt bones suggests a controlled use of fire. The Aroeira cranium represents a substantial contribution to the debate on the origin of the Neandertals and the pattern of human evolution in the Middle Pleistocene of Europe. The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. Nevertheless, European sites containing well-dated human remains associated with an Acheulean toolkit remain scarce. The earliest European hominin crania associated with Acheulean handaxes are at the sites of Arago, Atapuerca Sima de los Huesos (SH), and Swanscombe, dating to 400–500 ka (Marine Isotope Stage 11–12). The Atapuerca (SH) fossils and the Swanscombe cranium belong to the Neandertal clade, whereas the Arago hominins have been attributed to an incipient stage of Neandertal evolution, to Homo heidelbergensis, or to a subspecies of Homo erectus. A recently discovered cranium (Aroeira 3) from the Gruta da Aroeira (Almonda karst system, Portugal) dating to 390–436 ka provides important evidence on the earliest European Acheulean-bearing hominins. This cranium is represented by most of the right half of a calvarium (with the exception of the missing occipital bone) and a fragmentary right maxilla preserving part of the nasal floor and two fragmentary molars. The combination of traits in the Aroeira 3 cranium augments the previously documented diversity in the European Middle Pleistocene fossil record.

Endocranial morphology of the Ursus deningeri Von Reichenau 1904 from the Sima de Los Huesos (Sierra de Atapuerca) Middle Pleistocene site

Abstract A complete skull of Ursus deningeri recovered from the Middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca (Spain) was studied with computed tomography. This technique makes it possible to study new and relevant information in the endocranial structures and avoids any damage to the specimen. The previously undescribed endocranial cavity of U. deningeri is compared with related species such as Ursus spelaeus Rosenmüller 1794 and Ursus arctos Linné 1758. The characters analyzed include the braincase morphology and also quantitative and volumetric measurements, which are only attainable with this technique. In particular, the volume, surface area, and form of the frontal sinuses and brain cavity, the shape of the ethmoid bone, and the basioccipital pneumatization were studied. The resulting analysis supports an ancestor-descendant relationship for the two ursids included in the cave bear phylogenetic lineage (U. deningeri and U. spelaeus). In contrast, U. arctos exhibits a different endocranial morphology, one that seems to be mainly plesiomorphous. Finally, this study has identified several new endocranial characters which are useful in reconstructing ursid phylogeny.

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri. Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size. Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull. We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.

Taphonomic inferences about Middle Pleistocene hominins: The human cranium of Gruta da Aroeira (Portugal)

OBJECTIVES The aim of this work is to describe the taphonomic signatures of the Aroeira 3 cranium, with a specific focus on cranial breakage, comparing the cranium with other Middle and Upper Pleistocene hominin fossils in order to approximate the cause of death and the biological agencies and geologic processes involved in the taphonomic record of this specimen. Aroeira-3 was recovered from Acheulean layer X of Gruta da Aroeira (Portugal), dated to 390-436 ka. MATERIALS AND METHODS Taphonomic analyses noted surface modifications employing standard methods. The cranial breakage pattern of Aroeira 3 was analyzed to assess the presence/absence of perimortem (fresh bone) and postmortem (dry bone) fractures and the possible causes of perimortem skull bone fractures. RESULTS Aroeira 3 presents substantial bone loss of the left supraorbital arch and the outer cranial table of the frontal squama. Most of the fractures present features consistent with postmortem injuries. The fracture to the posterior region of the parietal bone, however, displays features more usually present in perimortem bone fractures. No evidence of anthropogenic activity or of carnivore modification has been identified. None of the expected features of interpersonal conflict are observed. Finally, the bone loss in the frontal squama and the supraorbital arch could be attributed to different agencies, and a traumatic event cannot be totally ruled out as origin of the bone alteration. DISCUSSION Cannibalism, secondary treatment of the corpse and accumulation induced by carnivores can all be discarded, making an accident the most plausible explanation for the cranial fracture.

The lameness of King Philip II and Royal Tomb I at Vergina, Macedonia

Significance The knee ankylosis and the hole through it ties perfectly with the penetrating wound and lameness suffered by Philip II and conclusively identifies him as the occupant of Tomb I in Vergina, Greece. The age estimates of the three occupants are consistent with those derived from the historical sources. Cleopatras (Philip’s wife) child was born a few days before Philip II’s assassination and both were murdered soon after Philip’s assassination. It follows that Tomb II belongs to King Arrhidaeus and his wife Eurydice and may well contain some of the armor of Alexander the Great. Thus, a nearly 40-y-old mystery concerning the Royal Tombs of Vergina has finally been solved that puzzled historians, archaeologists, and physical anthropologists. King Philip II was the father of Alexander the Great. He suffered a notorious penetrating wound by a lance through his leg that was nearly fatal and left him lame in 339 B.C.E. (i.e., 3 y before his assassination in 336 B.C.E.). In 1977 and 1978 two male skeletons were excavated in the Royal Tombs II and I of Vergina, Greece, respectively. Tomb I also contained another adult (likely a female) and a newborn skeleton. The current view is that Philip II was buried in Tomb II. However, the male skeleton of Tomb II bears no lesions to his legs that would indicate lameness. We investigated the skeletal material of Tomb I with modern forensic techniques. The male individual in Tomb I displays a conspicuous case of knee ankylosis that is conclusive evidence of lameness. Right through the overgrowth of the knee, there is a hole. There are no obvious signs that are characteristic of infection and osteomyelitis. This evidence indicates that the injury was likely caused by a severe penetrating wound to the knee, which resulted in an active inflammatory process that stopped years before death. Standard anthropological age-estimation techniques based on dry bone, epiphyseal lines, and tooth analysis gave very wide age ranges for the male, centered around 45 y. The female would be around 18-y-old and the infant would be a newborn. It is concluded that King Philip II, his wife Cleopatra, and their newborn child are the occupants of Tomb I.