The taxonomy of Sahelanthropus tchadensis from a craniometric perspective.

Sahelanthropus tchadensis has raised much debate since its initial discovery in Chad in 2001, given its controversial classification as the earliest representative of the hominin lineage. This debate extends beyond the phylogenetic position of the species, and includes several aspects of its habitual behavior, especially in what regards its locomotion. The combination of ancestral and derived traits observed in the fossils associated with the species has been used to defend different hypotheses related to its relationship to hominins. Here, the cranial morphology of Sahelanthropus tchadensis was assessed through 16 linear craniometric measurements, and compared to great apes and hominins through Principal Component Analysis based on size and shape and shape information alone. The results show that S. tchadensis share stronger morphological affinities with hominins than with apes for both the analysis that include size information and the one that evaluates shape alone. Since TM 266-01-060-1 shows a strong morphological affinity with the remaining hominins represented in the analysis, our results support the initial interpretations that S. tchadensis represents an early specimen of our lineage or a stem basal lineage more closely related to hominins than to Panini.

Ledi-Geraru strikes again: Morphological affinities of the LD 350-1 mandible with early Homo.

The origins of the genus Homo have been a focus of much debate in the paleoanthropological literature due to its importance in understanding the evolutionary trajectories that led to the appearance of archaic humans and our species. On the level of taxonomic classification, the controversies surrounding the origins of Homo are the result of lack of clear classification criteria that separate our genus from australopiths, given the general similarities observed between fossils ascribed to late australopiths and early Homo. The challenge in finding clear autapomorphies for Homo has even led to debates about the classification of Homo habilis and Homo rudolfensis as part of our genus. These debates are further complicated by the scarcity of fossils in the timeframe of appearance of our genus, making any fossils dated to between 3.0 and 2.5 Ma of particular relevance in the context of this discussion. The Ledi-Geraru mandible is one such fossils, which has called the attention of researchers due to its combination of primitive traits seen in Australopithecus and derived traits observed in later Homo. Despite being fragmented and poorly preserved, it is one of the key fossil specimens available from the period mentioned above.

Modularity of the wrist in extant hominines.

Wrist shape varies greatly across primates and previous studies indicate that the numerous morphological differences among them are related to a complex mixture of phylogeny and function. However, little is known about whether the variation in these various anatomical differences is linked and to what extent the wrist bones vary independently. Here, we used 3D geometric morphometrics on a sample of extant hominines (Homo sapiens, Pan troglodytes, Gorilla gorilla, and Gorilla beringei), to find the model that best describes the covariation patterns among four of the eight carpals (i.e., capitate, lunate, scaphoid, and trapezium). For this purpose, 15 modular hypotheses were tested using the Covariance Ratio. Results indicate that there is a covariation structure common to all hominines, which corresponds to stronger covariation within each carpal as compared to the covariation between carpals. However, the results also indicate that that there is a degree of codependence in the variation of some carpals, which is unique in humans, chimpanzees, and gorillas, respectively. In humans there is evidence of associated shape changes between the lunate and capitate, and between the scaphoid and trapezium. This covariation between lunate and capitate is also apparent in gorillas, while chimpanzees display the greatest disassociation among carpals, showing low covariation values in all pairwise comparisons. Our analyses indicate that carpals have an important level of variational independence which might suggest a high degree of independent evolvability in the wrists of hominines, and that although weak, the structure of associated changes of these four carpals varies across genera. To our knowledge this is the first report on the patterns of modularity between these four wrist bones in the Homininae and future studies might attempt to investigate whether the anatomical shape associations among carpals are functionally related to locomotion and manipulation.

Von Economo neurons: Cellular specialization of human limbic cortices?

Von Economo neurons (VENs) have been mentioned in the medical literature since the second half of the 19th century; however, it was not until the second decade of the 20th century that their cytomorphology was described in detail. To date, VENs have been found in limbic sectors of the frontal, temporal and insular lobes. In humans, their density seems to decrease in the caudo-rostral and ventro-dorsal direction; that is, from the anterior regions of the cingulate and insular cortices towards the frontal pole and the superior frontal gyrus. Several studies have provided similar descriptions of the shape of the VEN soma, but the size of the soma varies from one cortical region to another. There is consensus among different authors about the selective vulnerability of VENs in certain pathologies, in which a deterioration of the capacities involved in social behaviour is observed. In this review, we propose that the restriction of VENs towards the sectors linked to limbic information processing in Homo sapiens gives them a possible functional role in relation to the structures in which they are located. However, given the divergence in characteristics such as location, density, size and biochemical profile among VENs of different cortical sectors, the activities in which they participate could allow them to partake in a wide spectrum of neurological functions, including autonomic responses and executive functions.

A quantitative approach for analysing bone modelling patterns from craniofacial surfaces in hominins.

Bone size and shape arise throughout ontogeny as a result of the coordinated activity of osteoblasts and osteoclasts, responsible for bone deposition and resorption, and growth displacements. The modelling processes leave specific microstructural features on the bone surface, which can be used to infer the mechanisms shaping craniofacial traits in extinct and extant species. However, the analysis of bone surfaces from fossils and archaeological samples faces some difficulties related to the bone loss caused by taphonomic factors, and the lack of formal methods for estimating missing information and comparing the patterns of bone modelling among several specimens and samples. The present study provides a new approach for the quantitative analysis of bone formation and resorption patterns obtained from craniofacial surfaces. First, interpolation techniques were used to estimate missing data on high-resolution replicas of the left maxilla in a sample of sub-adult and adult modern humans and sub-adult fossil hominins. The performance of this approach was assessed by simulating variable amounts of missing data. Then, we applied measures of dispersion and central tendency to represent the variation and average pattern of bone modelling within samples. The spatial interpolation resulted in reliable estimations of the type of cell activity (deposition or resorption) in the missing areas, even when large extensions of the bone surface were lost. The quantification of the histological data allowed us to integrate the information of different specimens and depict the areas with higher and lower variation in the bone modelling pattern of the maxilla among specimens. Overall, the main advantages of the quantitative approach used here for generating bone modelling patterns are the high replicability and the possibility of incorporating variation among specimens into the comparisons among samples.

Morphological affinities of Homo naledi with other Plio-Pleistocene hominins: a phenetic approach.

Recent fossil material found in Dinaledi Chamber, South Africa, was initially described as a new species of genus Homo, namely Homo naledi. The original study of this new material has pointed to a close proximity with Homo erectus. More recent investigations have, to some extent, confirmed this assignment. Here we present a phenetic analysis based on dentocranial metric variables through Principal Components Analysis and Cluster Analysis based on these fossils and other Plio-Pleistocene hominins. Our results concur that the Dinaledi fossil hominins pertain to genus Homo. However, in our case, their nearest neighbors are Homo habilis and Australopithecus sediba. We suggest that Homo naledi is in fact a South African version of Homo habilis, and not a new species. This can also be applied to Australopithecus sediba.

Morphological affinities of Homo naledi with other Plio-Pleistocene hominins: a phenetic approach

ABSTRACT Recent fossil material found in Dinaledi Chamber, South Africa, was initially described as a new species of genus Homo, namely Homo naledi. The original study of this new material has pointed to a close proximity with Homo erectus. More recent investigations have, to some extent, confirmed this assignment. Here we present a phenetic analysis based on dentocranial metric variables through Principal Components Analysis and Cluster Analysis based on these fossils and other Plio-Pleistocene hominins. Our results concur that the Dinaledi fossil hominins pertain to genus Homo. However, in our case, their nearest neighbors are Homo habilis and Australopithecus sediba. We suggest that Homo naledi is in fact a South African version of Homo habilis, and not a new species. This can also be applied to Australopithecus sediba.

Developmental identity versus typology: Lucy has only four sacral segments.

OBJECTIVES: Both interspecific and intraspecific variation in vertebral counts reflect the action of patterning control mechanisms such as Hox. The preserved A.L. 288-1 ("Lucy") sacrum contains five fused elements. However, the transverse processes of the most caudal element do not contact those of the segment immediately craniad to it, leaving incomplete sacral foramina on both sides. This conforms to the traditional definition of four-segmented sacra, which are very rare in humans and African apes. It was recently suggested that fossilization damage precludes interpretation of this specimen and that additional sacral-like features of its last segment (e.g., the extent of the sacral hiatus) suggest a general Australopithecus pattern of five sacral vertebrae. METHODS: We provide updated descriptions of the original Lucy sacrum. We evaluate sacral/coccygeal variation in a large sample of extant hominoids and place it within the context of developmental variation in the mammalian vertebral column. RESULTS: We report that fossilization damage did not shorten the transverse processes of the fifth segment of Lucy's sacrum. In addition, we find that the extent of the sacral hiatus is too variable in apes and hominids to provide meaningful information on segment identity. Most importantly, a combination of sacral and coccygeal features is to be expected in vertebrae at regional boundaries. DISCUSSION: The sacral/caudal boundary appears to be displaced cranially in early hominids relative to extant African apes and humans, a condition consistent with the likely ancestral condition for Miocene hominoids. While not definitive in itself, a four-segmented sacrum accords well with the "long-back" model for the Pan/Homo last common ancestor. Am J Phys Anthropol 160:729-739, 2016. © 2016 Wiley Periodicals, Inc.

Comparative anatomical study of the leg's nerves of Cebus (barbed capuchins) with baboons, chimpanzees and modern humans / Estudo anatômico comparativo dos nervos da perna de Cebus (macaco-prego) com babuínos, chimpanzés e humanos modernos

The anatomical comparative studies among the primates are important for the investigation of ethology, evolution, taxonomy, and comprehension of tools by hominoids. Especially the anatomical knowledge of Cebus contributes to conservation of the species, and to development of surgical procedures and clinical treatments of these animals, as they frequently are victims of automobile accidents. Recent anatomical studies came to a wrong conclusion regarding behavioral traits of Cebus, ascribed to few data available in previous literature. Therefore, to provide anatomical data and to support the other sciences related to anatomy, and to develop surgical and/or clinical procedures, we described the nerves of the legs of Cebus foccusing on their position and trajectory, as wll as innerved muscles, and compared these results with those of humans and other primates. Eight adult capuchin specimens were used for this study. The anatomical comparative study of the leg's nerves of Cebus demonstrated that, in general, structural organization of the nerves is similar among the four primates analyzed here (Cebus, chimpanzees, baboons and humans), which might be attributed to the fact that the all four primates have similar body structures. However, nerve trajectory and muscles innervation in Cebus was more similar to baboons.(AU)

Os estudos anatômicos comparativos entre os primatas são importantes para pesquisas associadas com a etologia, evolução, taxonomia e compreensão dos usos de ferramentas pelos hominídeos. Especificamente, o conhecimento anatômico sobre Cebus contribui para sua própria conservação e para o desenvolvimento de procedimentos cirúrgicos e tratamentos clínicos destes animais, pois são frequentemente vítimas de acidentes automobilísticos. Recentemente, estudos sobre características comportamentais de Cebus indicaram conclusões erradas sobre sua anatomia, o que pode ser atribuído aos poucos dados disponíveis sobre a anatomia desses animais na literatura especializada. Portanto, para fornecer dados anatômicos e fornecer suporte para as outras ciências relacionadas com a anatomia e também desenvolver procedimentos cirúrgicos e/ou clínicos, foram descritos os nervos das pernas de Cebus com enfoque sobre a posição, a trajetória e os músculos inervados, e comparar esses resultados com os dos humanos modernos e outros primatas. Oito espécimes adultos de macacos-prego foram usados para este estudo. O estudo anatômico comparativo dos nervos da perna de Cebus demonstrou que, em geral, a organização estrutural dos nervos é semelhante entre os quatro primatas aqui analisados (Cebus, chimpanzés, babuínos e humanos modernos), o que pode ser atribuído ao fato de que os quatro primatas terem estruturas corporais semelhantes. No entanto, a trajetória dos nervos e a inervação dos músculos em Cebus o aproximam dos babuínos.(AU)

Comparative anatomical study of the forearm extensor muscles of Cebus libidinosus (Rylands et al., 2000; Primates, Cebidae), modern humans, and other primates, with comments on primate evolution, phylogeny, and manipulatory behavior.

Despite its abundance in Latin America, and its remarkable ability to use tools, there are only a few myological studies on the capuchin monkey, Cebus libidinosus. In the present study, we dissected the forearm extensor muscles of six adult males and two adult females of this species. We describe these muscles and compare them with those of other primates dissected by us and by other authors. The forearm extensor muscles of Cebus monkeys are, in general, more similar to those of other platyrrhines than to distantly related taxa that use tools, such as chimpanzees and modern humans, with three main exceptions: contrary to most other platyrrhines, (1) in Cebus, chimpanzees and modern humans the extensor pollicis longus usually inserts onto Digit I, and not onto Digits I and II; (2) in Cebus the abductor pollicis longus has two separate tendons, as is the case in chimpanzees, and in modern humans (where one of these tendons is associated with a distinct belly, forming the muscle extensor pollicis brevis); (3) in Cebus, and in modern humans and chimpanzees, the extensor pollicis longus is not deeply blended with the extensor indicis. Therefore, the Cebus monkeys provide an illustrative example of how phylogenetic constrains and ecological adaptations have been combined to develop a specific myological configuration that, associated with their sophisticated neurological organization, allow them to easily navigate in their arboreal habitats and, at the same time, to finely manipulate objects in order to search for food and to prepare this food for ingestion.

Cladistic analysis of continuous modularized traits provides phylogenetic signals in Homo evolution.

Evolutionary novelties in the skeleton are usually expressed as changes in the timing of growth of features intrinsically integrated at different hierarchical levels of development. As a consequence, most of the shape-traits observed across species do vary quantitatively rather than qualitatively, in a multivariate space and in a modularized way. Because most phylogenetic analyses normally use discrete, hypothetically independent characters, previous attempts have disregarded the phylogenetic signals potentially enclosed in the shape of morphological structures. When analysing low taxonomic levels, where most variation is quantitative in nature, solving basic requirements like the choice of characters and the capacity of using continuous, integrated traits is of crucial importance in recovering wider phylogenetic information. This is particularly relevant when analysing extinct lineages, where available data are limited to fossilized structures. Here we show that when continuous, multivariant and modularized characters are treated as such, cladistic analysis successfully solves relationships among main Homo taxa. Our attempt is based on a combination of cladistics, evolutionary-development-derived selection of characters, and geometric morphometrics methods. In contrast with previous cladistic analyses of hominid phylogeny, our method accounts for the quantitative nature of the traits, and respects their morphological integration patterns. Because complex phenotypes are observable across different taxonomic groups and are potentially informative about phylogenetic relationships, future analyses should point strongly to the incorporation of these types of trait.

Interlaminar astroglial processes in the cerebral cortex of great apes.

The present study was designed to document the architecture of neocortical astroglia in great apes, following glial fibrillary acidic protein immunohistochemistry. These anthropoid species were missing from previous phylogenetic descriptions of astroglia with interlaminar processes, a characteristic event of the cerebral cortex within the Primate Order. Pongo pygmaeus (orangutan), Gorilla gorilla (gorilla) and Pan troglodytes (chimpanzee) brain samples showed the typical "palisade" of interlaminar processes. Yet, those from Pan troglodytes were less uniform, showing extended cortical segments with astrocytic ("syncytial-type") appearance, intermingled with segments expressing the interlaminar "palisade". Present observations contribute to fill in missing data on the phylogenetic emergence of the cerebral cortex astroglial interlaminar processes. Considering the extreme consistency of the expression of astroglial interlaminar "palisades" among anthropoid species, this apparent variability among Pan individuals could be due to various possibilities, which are considered in this report.

Mapping and taphonomic analysis of the Homo erectus loci at Locality 1 Zhoukoudian, China.

From a detailed analysis of published and unpublished sources, we constructed a digitized three-dimensional, stratigraphically-controlled excavation grid of Zhoukoudian Locality 1 in order to assess the spatial relationships of the excavated materials. All 15 fossil Homo erectus loci were mapped on the grid. Meter cubes were used in excavation starting in 1934, and Loci H through O, established between 1934 and 1937, were mapped to within 1 m(3)vertical and horizontal provenience. Loci A through G, established between 1921 and 1933, were excavated in the northernmost part of Locality 1 by unmapped quarrying, but their stratigraphic levels were recorded. We could localize Loci A through G on the grid system by utilizing locations of remaining walls, stratigraphic sections, excavation reports, excavation maps, and photographs. Loci contained skeletal elements of Homo erectus individuals scattered over areas of the cave floor of up to 9 m in diameter. Scoring of taphonomic damage on the Homo erectus sample, as observed on casts and originals, demonstrates that 67% of the hominid sample shows bite marks or other modifications ascribed to large mammalian carnivores, particularly the large Pleistocene cave hyena, Pachycrocuta brevirostris. Virtually all of the remaining Homo erectus skeletal assemblage shows breakage consistent with this taphonomic pattern of fragmentation. Bioturbation by digging carnivores is the most likely explanation for a fragment of Homo erectus Skull XI discovered 1 m below its other conjoined portions in Locus L. Carbon on all the Homo erectus fossils from Locus G, a circumscribed area of 1-meter diameter, earlier taken to indicate burning, cooking, and cannibalism, is here interpreted as detrital carbon deposited under water, perhaps the result of hyaenid caching behavior. Locus G records the close stratigraphic and horizontal association of stone artifacts with Homo erectus and other vertebrate skeletal elements, an association that is seen at other loci as well. Layer 4 of the excavation contains equid cranial bone previously interpreted to have been burned while fresh. We here document that Locus B Homo erectus, including Skull I, is stratigraphically associated with this evidence, but at some 10-12 m distance. Even though the presence of wood-stoked fires and hearths is not supported by geochemical results, evidence of fire at Locality 1 in the form of burned bone is confirmed. Contextual relationships of fossil skeletal elements, relationships of carnivore damage and stone tool cutmarks on bone, and evidence of the burning of fresh bone associated with Homo erectus and stone tools support a model of transient hominid scavenging aided by the use of fire at the large hyenid den that became Zhoukoudian Locality 1. Although the original excavation catalogue from Locality 1, as well as a significant number of fossils and stone artifacts, were lost during World War II, catalogue numbers on the many surviving specimens can be used to locate fossils and artifacts within the three-dimensional grid provided in this paper.

Early Holocene human skeletal remains from Santana do Riacho, Brazil: implications for the settlement of the New World.

In this study we compare the cranial morphology of several late Paleoindian skeletons uncovered at Santana do Riacho, Central Brazil, with worldwide human cranial variation. Mahalanobis Distance and Principal Component Analysis are used to explore the extra-continental morphological affinities of the Brazilian Paleoindian sample. Santana do Riacho is a late Paleoindian burial site where approximately 40 individuals were recovered in varying states of preservation. The site is located at Lagoa Santa/Serra do Cipó, State of Minas Gerais. The first human activities in this rockshelter date back to the terminal Pleistocene, but the burials are bracketed between circa 8200 and 9500BP. The collection contains only six skulls well-enough preserved to be measured. The Santana do Riacho late Paleoindians present a cranial morphology characterized by long and narrow neurocrania, low and narrow faces, with low nasal apertures and orbits. The multivariate analyses show that they exhibit strong morphological affinities with present day Australians and Africans, showing no resemblance to recent Northern Asians and Native Americans. These findings confirm our long held opinion that the settlement of the Americas was more complicated in terms of biological input than has been widely assumed. The working hypothesis is that two very distinct populations entered the New World by the end of the Pleistocene, and that the transition between the cranial morphology of the Paleoindians and the morphology of later Native Americans, which occurred around 8-9ka, was abrupt. This, in our opinion, is a more parsimonious explanation for the diversity detected than a long, local microevolutionary process mediated by selection and drift. The similarities of the first South Americans with sub-Saharan Africans may result from the fact that the non-Mongoloid Southeast Asian ancestral population came, ultimately, from Africa, with no major modification in the original cranial bau plan of the first modern humans.

Evolución del pie / Foot evolution

Se realiza un análisis de la evolución del pie humano, desde los primates al hombre moderno, destacando los cambios que se produjeron en el pie para alcanzar la bipedestación. En función de entender cómo se desarrolló la estructura del pie humano se analizó su evolución en nuestro orden biológico; desde el pleisiádapis hace unos 60-65 millones de años, pasando por los distintos primates del Eocénico, Oligocénico, Miocénico, etc. hasta el hombre moderno, arribando a las siguientes conclusiones: A) La bipedestación, la marcha erguida, fue un hecho esencial en el proceso evolutivo desde los primeros primates al hombre moderno. B) El por qué de la bipedestación sigue siendo materia de discusión. C) De los cambios que se produjeron en el pie, me parecen los más importantes: el paralelismo del primer rayo con los otros metatarsianos; formación de los arcos del pie; aumento de la rigidez del ligamento calcáneoescafoideo; inserción del tendón tibial posterior en el escafoides, para el mantenimiento del arco longitudinal; rigidez en las estructuras del pie y alineamiento en los huesos del retropie (calcáneo y astrágalo). D) Creo que algunas patologías que se podrían explicar por la persistencia de caracteres ancestrales, como algunos casos de hallux-valgus y pie plano. (AU)

Odontometric microevolution in the Valley of Oaxaca, Mexico.

Tooth crown lengths, breadths, and areas were compared from five sequential human skeletal samples from the Valley of Oaxaca, Mexico, spanning the period from 1600 BC to AD 1521. Almost all of the measurements considered exhibited dramatic reduction over that period. Total crown area declined from 1320 mm2 to 1262 mm2, a change of 4.4% or 9.4 darwins, which is noticeably larger than any previously reported rate of dental reduction in a recent human population. This reduction was greatest in posterior tooth breadths, especially in the mandible. The patterning of the reduction, and of the variance of the measurements, indicates that natural selection was primarily responsible for the change. Of all the measurements, anterior breadths appear to have been the least subject to selection, and posterior lengths the next least. The degree of reduction suggests that selective pressures towards smaller teeth were greater in Mesoamerica than elsewhere. This case provides an example of the value of recent human skeletal series for the study of microevolutionary processes.