First record of a fossil monkey (Primates, Cercopithecidae) from the Late Pliocene of Serbia Publication date: December 2019 Source: Journal of Human Evolution, Volume 137 Author(s): Predrag Radović, Joshua Lindal, Zoran Marković, Sanja Alaburić, Mirjana Roksandic Abstract The cercopithecid fossil record of the Balkan Peninsula extends from the Late Miocene to the Early Pleistocene, but to date no fossils of non-human primates have been identified in Serbia. Here we report the identification of two primate teeth from Ridjake, a rich paleontological site in western Serbia. NHMBEO 042501 is an upper third molar with heavy occlusal wear and taphonomic weathering. NHMBEO 042502 is a well-preserved lower third molar with only minor damage to the cusps and root apices. We performed an analysis of non-metric traits and made bivariate comparisons of crown linear measurements in order to assess the taxonomic affinity of the molars. Both show typical papionin occlusal patterns and relatively large overall sizes. In combination with the early Villafranchian (MN16) age of the site, we attribute both Ridjake primate fossils to cf. Paradolichopithecus sp. This represents the first identification of a non-human primate in Serbia, and the first identification of any primate in the Neogene period of Serbia. Along with recent hominin discoveries, the Ridjake fossils contribute to the growing primate fossil record in Serbia, and indicate the need for increased research into fossil primates in the country.

New approaches to modeling primate socioecology: Does small female group size BEGET loyal males? Publication date: December 2019 Source: Journal of Human Evolution, Volume 137 Author(s): Kristin N. Crouse, Carrie M. Miller, Michael L. Wilson Abstract Humans are unusual in having stable male-female breeding bonds within multi-level societies. Such societies are not found in other apes, but have evolved independently in other primates, including several African papionins: hamadryas and Guinea baboons and gelada monkeys. Stable breeding bonds have been proposed to evolve either (1) because males can monopolize females when food distribution forces females to forage in small groups or (2) because females exchange exclusive mating for male services, such as protection from infanticide. Comparative studies are needed to test these hypotheses. In the meantime, we used an agent-based computer model to test the plausibility of these hypotheses. We simulated primates living in social groups within a larger population using a model we call BEGET (Behavior, Ecology, Genetics, Evolution, and Tradeoffs), which employed decision vectors, experimental evolution, realistic trade-offs, and phenotypic plasticity. We employed experimental evolution to generate male genotypes that varied in their competitive ability and in their long-term mating strategy. “Rover” males searched for and mated with any sexually receptive females whereas “Loyalist” males formed stable associations with particular groups of females. Much like living primates, the virtual primates exhibited tradeoffs between contest and scramble competition for access to females: Loyalists evolved larger body size than Rovers. We tested the effect of female foraging group size and the presence of infanticide and infant protection on the relative success of these strategies. We found that Loyalists achieved greater reproductive success than Rovers only when females were in groups smaller than four. Both Rovers and Loyalists sometimes evolved infanticidal behavior, but the presence of infanticide benefited Rovers rather than Loyalists, suggesting that the evolution of stable breeding bonds depends on the spatial distribution of females, rather than the risk of infanticide.

Social evolution in Plio-Pleistocene hominins: Insights from hamadryas baboons and paleoecology Publication date: December 2019 Source: Journal of Human Evolution, Volume 137 Author(s): Larissa Swedell, Thomas Plummer Abstract Reconstructions of hominin evolution have long benefited from comparisons with nonhuman primates, especially baboons and chimpanzees. The hamadryas baboon (Papio hamadryas) is arguably one of the best such models, as it exhibits both the male kin bonding and the cross-sex pair bonding thought to have been important in hominin evolution. Here we link processes of behavioral evolution in hamadryas baboons with those in a Plio-Pleistocene hominin, provisionally identified as Homo erectus (sensu lato) – a pivotal species in that its larger body and brain size and wider ranging patterns increased female costs of reproduction, increasing the importance of sociality. The combination of these higher costs of reproduction and shifts in diet and food acquisition have previously been argued to have been alleviated either via strengthening of male-female bonds (involving male provisioning and the evolution of monogamy) or via the assistance of older, post-reproductive females (leading to post-reproductive longevity in females, i.e., the grandmother hypothesis). We suggest that both arrangements could have been present in Plio-Pleistocene hominins if they lived in multilevel societies. Here we expand on our earlier scenario with two sets of recent data in support of it, (1) archaeological data from the 2 million year old Oldowan site of Kanjera South, Kenya and other sites that are suggestive of tool dependent foraging on nutrient dense resources (animal carcasses and plant underground storage organs), cooperation, and food sharing; and (2) a pattern of genetic variation in hamadryas baboons that suggests the operation of kin selection among both males and females at multiple levels of society. Taken together, these two sets of data strengthen our model and support the idea of a complex society linked by male-male, male-female, and female-female bonds at multiple levels of social organization in Plio-Pleistocene hominins.

The cochlea of the Sima de los Huesos hominins (Sierra de Atapuerca, Spain): New insights into cochlear evolution in the genus Homo Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Mercedes Conde-Valverde, Ignacio Martínez, Rolf M. Quam, Alejandro Bonmatí, Carlos Lorenzo, Alex D. Velez, Carolina Martínez-Calvo, Juan Luis Arsuaga Abstract The cochlea contains taxonomic and phylogenetic information and its morphology is related with hearing abilities among fossil hominins. Data for the genus Homo is presently limited to early Homo and the early Neandertals from Krapina. The present study of the middle Pleistocene hominins from the Sima de los Huesos (SH) provides new evidence on cochlear evolution in the genus Homo. We compared the absolute length, proportional lengths of each turn, number of turns, size and shape of the cross-section of the basal turn, volume, curvature gradient, and thickness of the cochlea between extant Pan troglodytes, extant Homo sapiens, Homo neanderthalensis and the SH hominins. The SH hominins resemble P. troglodytes in the proportionally long basal turn, the small size and round shape of the cross-section of the basal turn, the small cochlear volume and the low cochlear thickness. The SH hominins resemble Neandertals and H. sapiens in their long cochlear length and in the proportionally short third turn. Homo neanderthalensis and H. sapiens share several features, not present in the SH hominins, and that likely represent homoplasies: a larger volume, larger size and oval shape of the cross-section of the basal turn and higher cochlear thickness. Later Neandertals show a derived proportionally shorter apical turn. Changes in cochlear volume in Homo cannot be fully explained by variation in body mass or cochlear length but are more directly related to changes in the cross-sectional area of the basal turn. Based on previous studies of the outer and middle ear in SH hominins, changes in the outer and middle ear preceded changes in the inner ear, and the cochlea and semicircular canals seem to have evolved independently in the Neandertal clade. Finally, the small cochlear volume in the SH hominins suggests a slightly higher upper limit of hearing compared with modern humans.

The deciduous dentition of Homo naledi: A comparative study Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Shara E. Bailey, Juliet K. Brophy, Jacopo Moggi-Cecchi, Lucas K. Delezene Abstract In 2013, 2014 new hominin remains were uncovered in the Dinaledi chamber of the Rising Star cave system in South Africa. In 2015 Berger and colleagues identified these remains as belonging to a new species Homo naledi (Berger et al., 2015). Subsequent comparative studies of the skull, postcrania and permanent dentition have supported this taxonomic affiliation (Harcourt-Smith et al., 2015; Kivell et al., 2015; Irish et al., 2018). The deciduous teeth can offer unique insights into hominin evolution. Due to their early onset and rapid development their morphology is thought to be under stronger genetic control and less influenced by environment than are the permanent teeth. In this study we compared the H. naledi deciduous teeth from the 2013–2014 excavations to samples representing much of the hominin clade including Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, early Homo, Homo antecessor, Homo erectus s.l., Homo floresiensis, Middle Pleistocene Homo, Homo neanderthalensis, early Homo sapiens and recent H. sapiens from Sub-Saharan Africa. By making such a broad morphological comparison, we aimed to contextualize the Dinaledi hominins and to further assess the validity of their taxonomic assignment. Our analysis of the deciduous teeth revealed a unique combination of features that mirror (but also expand) that found in the permanent teeth. This mosaic includes an asymmetrical lower canine with a distal tubercle, an upper first molar with a large hypocone and epicrista associated with a mesial cuspule, a molarized lower first molar resembling Paranthropus, and upper and lower second molars that resemble later Homo in their lack of accessory cusps. The unique combination of deciduous dental characters supports previous studies assigning H. naledi to a new species, although its phylogenetic position vis-à-vis other Homo species remains ambiguous.

Enamel thickness and dental development in Rudapithecus hungaricus Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Tanya M. Smith, Paul Tafforeau, Joane Pouech, David R. Begun Abstract The fossil record of middle and late Miocene Eurasian hominoids has expanded considerably over the past few decades, particularly with the recovery of numerous isolated teeth and jaws. Scholars have turned to assessments of internal tooth structure and growth to make sense of the evolutionary radiations of these primates as well as their affinities to the living great apes (hominids). Here we characterize full-dentition enamel thickness and dental development in several juvenile Rudapithecus hungaricus individuals using multiple imaging modalities. Relative enamel thickness (RET) values for the anterior teeth and premolars of Rudapithecus are broadly akin to those of gorillas and chimpanzees and are thinner than those of orangutans. First molar RET values are most similar to chimpanzees, while posterior molar values are closer to thicker-enameled orangutans. When compared to Miocene hominoids, Rudapithecus shows an intermediate molar RET condition that is especially similar to other dryopithecines. Long-period line periodicity values are comparable to African apes and most Miocene hominoids, and lower than living and fossil orangutans. The mean cuspal daily secretion rate is similar to that of several other Miocene hominoids but is greater than extant great apes. Cusp-specific molar crown formation times generally exceed those of chimpanzees, are lower than those of orangutans, and are broadly like those of other Miocene apes. While Rudapithecus appears to have a somewhat unique pattern of enamel thickness and dental development relative to individual great ape genera, these structural and developmental features are consistent with its designation as a hominid.

Femoral neck cortical bone distribution of dryopithecin apes and the evolution of hominid locomotion Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Marta Pina, David M. Alba, Salvador Moyà-Solà, Sergio Almécija Abstract Only a few postcranial remains have been assigned to the Miocene great ape Dryopithecus fontani, leading to uncertainties in the reconstruction of its overall body plan and positional behavior. Here we shed light on the locomotor repertoire of this species through the study of the femoral neck cortical bone (FNCB) distribution of IPS41724, a partial proximal femur from the Abocador de Can Mata locality ACM/C3-Az (11.9 Ma, middle Miocene; Vallès-Penedès Basin, Spain) attributed to this taxon. This specimen was scanned through computed tomography to measure the superior (SUP) and inferior (INF) cortical thicknesses at the middle and the base of the femoral neck. Measurements were compared with a sample of extant primates and the femur IPS18800.29 from the younger great ape Hispanopithecus laietanus from Can Llobateres 2 (9.6 Ma, late Miocene; Vallès-Penedès Basin), previously shown to display a homogeneous FNCB distribution at the midneck section coupled with postcranial adaptations to below-branch suspensory behaviors. Our analyses indicate an asymmetric FNCB distribution for IPS41724 (SUP/INF index = ∼0.4 at the midneck and base of the neck sections), comparable with that of quadrupedal primates and bipedal hominins (including early australopiths), but contrasting with the homogeneous FNCB distribution of Hispanopithecus and extant great apes. An asymmetrical FNCB distribution has been associated with stereotyped loads at the hip joint (as in both quadrupedal and bipedal taxa). Our results therefore support a significant quadrupedal component of the positional behavior of Dryopithecus, thus strengthening the argument that plesiomorphic generalized quadrupedalism was still a major locomotor behavior for Miocene great apes. If that were the case, it could have deep implications for the origins of hominin bipedalism.

Feathers and food: Human-bird interactions at Middle Pleistocene Qesem Cave, Israel Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Ruth Blasco, Jordi Rosell, Antonio Sánchez-Marco, Avi Gopher, Ran Barkai Abstract The presence of fast-moving small game in the Paleolithic archaeological faunal record has long been considered a key variable to assess fundamental aspects of human behavior and subsistence. Birds occupy a prominent place in this debate not only due to their small size and to the difficulties in capturing them (essentially due to their ability to fly and their elusiveness), but also due to their possible role in the symbolic array in regard to non-nutritional elements (feathers, talons, etc.) and as reflectors of complex human–world relationships. In this study, we attempt to contribute to this topic by presenting taphonomical data of bird specimens from Qesem Cave (Israel), dated between 420 and 200 ka. Human-induced damage, including cut marks, peeling and human gnawing, has been identified on wing bones of Cygnus sp., Columba sp., Corvus ruficollis and Sturnus sp. Our evidence suggests that avian exploitation was not limited to food only—either to complement the human diet or as occasional food item—but also presumably for the use of feathers. While the consumption of birds as a dietary source seems to be evident as early as the Early Pleistocene, the non-alimentary use of inedible elements, such as feathers and talons, appears to be a practice from the Middle Paleolithic onwards. We argue that the combined nutritional and symbolic use of birds is one characteristic of the new mode of adaptation practiced already by the late Lower Paleolithic Acheulo-Yabrudian hominins in the Levant starting 400 ka. The Qesem findings point to the possible emergence of new cognitive and behavioral skills, which are followed in later periods in the Old World. Finally, we discuss the possible ontological and cosmological significance of human–bird interactions to illuminate our hypothesis regarding the emergence of a new perception of human relationships with the world as an integral part of the new Acheulo-Yabrudian mode of adaptation.

Structural analysis of premolar roots in Middle Pleistocene hominins from China Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Lei Pan, Jean Dumoncel, Arnaud Mazurier, Clément Zanolli Abstract This study investigates permanent maxillary and mandibular premolar root structural organization in East Asian Middle Pleistocene hominins. In addition to reporting and analyzing the linear and volumetric properties of the roots, we used a landmark-free approach to both qualify and quantify in 3D premolar root shape variation of Middle Pleistocene hominins in East Asia. Moreover, we focus on some mid-to late East Asian Middle Pleistocene hominin specimens whose taxonomic attribution is unclear. We find considerable cementum in this sample of hominins, similar to other fossil groups, but clearly different from modern humans which have a very small amount of cementum. Additionally, a smaller root pulp cavity is found in later Homo (Neanderthals and modern humans). Our analyses on the crown-root surface area ratio show that East Asian Middle Pleistocene Homo erectus as well as one late Middle Pleistocene Homo sp. specimen (PA 81 P4 from Changyang site) are distinguished from other fossil and extant groups by a relatively larger root surface, stout root branches and thick cementum deposits. This may represent a distinct East Asian H. erectus dental pattern. Geometric morphometric analyses on the external root surface reveal a general trend of shape simplification along the Homo lineage examined here, and distinguish Early Pleistocene Homo, Middle Pleistocene H. erectus, Neanderthals and modern human morphologies. The late Middle Pleistocene teeth from Changyang site (PA 76 P3 and PA 81 P4) are close to East Asian H. erectus and Neanderthals, while the mid-Middle Pleistocene P3 from Panxian Dadong falls within the modern human distribution. Combined with dental crown morphology and root number/form reported in previous studies, our results show that the external root shape can be considered a taxonomically relevant indicator. In general, an evolutionary tendency towards modern human morphology is observed in part of the East Asian Middle Pleistocene specimens, while a retention of primitive, H. erectus-like features is expressed in some late Middle Pleistocene specimens, supporting a multi-lineage and discontinuous scenario of human settlements in East Asia.

Endostructural morphology in hominoid mandibular third premolars: Discrete traits at the enamel-dentine junction Publication date: November 2019 Source: Journal of Human Evolution, Volume 136 Author(s): Thomas W. Davies, Lucas K. Delezene, Philipp Gunz, Jean-Jacques Hublin, Matthew M. Skinner Abstract The mandibular third premolar (P3) exhibits substantial differences in size and shape among hominoid taxa, and displays a number of discrete traits that have proven to be useful in studies of hominin taxonomy and phylogeny. Discrete traits at the enamel-dentine junction (EDJ) can be accurately assessed on moderately worn specimens, and often appear sharper than at the outer-enamel surface (OES). Here we use microtomography to image the P3 EDJ of a broad sample of extant apes, extinct hominins and modern humans (n = 100). We present typologies for three important premolar discrete traits at the EDJ (transverse crest, marginal ridge and buccal grooves), and score trait frequencies within our sample. We find that the transverse crest is variable in extant apes, while the majority of hominins display a transverse crest which runs directly between the two major premolar cusps. Some Neanderthals display a unique form in which the transverse crest fails to reach the protoconid. We find that mesial marginal ridge discontinuity is common in Australopithecus anamensis and Australopithecus afarensis while continuous marginal ridges largely characterize Australopithecus africanus and Paranthropus. Interrupted mesial and distal marginal ridges are again seen in Homo sapiens and Neanderthals. Premolar buccal grooves, previously identified at the OES as important for hominin systematics, are again found to show a number of taxon-specific patterns at the EDJ, including a clear difference between Australopithecus and Paranthropus specimens. However, their appearance may be dependent on the morphology of other parts of the crown such as the protoconid crest, and the presence of accessory dentine horns. Finally, we discuss rare variations in the form of dentine horns that underlie premolar cusps, and their potential homology to similar morphologies in other tooth positions.