Human Evolution

A new interpretation of Madagascar's megafaunal decline: The "Subsistence Shift Hypothesis" Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Laurie R. Godfrey, Nick Scroxton, Brooke E. Crowley, Stephen J. Burns, Michael R. Sutherland, Ventura R. Pérez, Peterson Faina, David McGee, Lovasoa Ranivoharimanana Abstract Fundamental disagreements remain regarding the relative importance of climate change and human activities as triggers for Madagascar's Holocene megafaunal extinction. We use stable isotope data from stalagmites from northwest Madagascar coupled with radiocarbon and butchery records from subfossil bones across the island to investigate relationships between megafaunal decline, climate change, and habitat modification. Archaeological and genetic evidence support human presence by 2000 years Before Common Era (BCE). Megafaunal decline was at first slow; it hastened at ∼700 Common Era (CE) and peaked between 750 and 850 CE, just before a dramatic vegetation transformation in the northwest that resulted in the replacement of C3 woodland habitat with C4 grasslands, during a period of heightened monsoonal activity. Cut and chop marks on subfossil lemur bones reveal a shift in primary hunting targets from larger, now-extinct species prior to ∼900 CE, to smaller, still-extant species afterwards. By 1050 CE, megafaunal populations had essentially collapsed. Neither the rapid megafaunal decline beginning ∼700 CE, nor the dramatic vegetation transformation in the northwest beginning ∼890 CE, was influenced by aridification. However, both roughly coincide with a major transition in human subsistence on the island from hunting/foraging to herding/farming. We offer a new hypothesis, which we call the "Subsistence Shift Hypothesis," to explain megafaunal decline and extinction in Madagascar. This hypothesis acknowledges the importance of wild-animal hunting by early hunter/foragers, but more critically highlights negative impacts of the shift from hunting/foraging to herding/farming, settlement by new immigrant groups, and the concomitant expansion of the island's human population. The interval between 700 and 900 CE, when the pace of megafaunal decline quickened and peaked, coincided with this economic transition. While early megafaunal decline through hunting may have helped to trigger the transition, there is strong evidence that the economic shift itself hastened the crash of megafaunal populations.

Tools, trails and time: Debating Acheulian group size at Attirampakkam, India Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Shanti Pappu, Kumar Akhilesh Abstract Estimating Acheulian group sizes based on a fragmentary archaeological record is fraught with difficulties, more so in regions like India, where lithics form the primary source of information. Here, we review current approaches towards modeling group size in Indian archaeology. We then examine to what extent one may address issues related to seasonality, fission-fusion strategies and group size in the context of Acheulian sites, drawing on our research along the southeastern coast of India. We move between multiple scales of analysis: from the regional Acheulian archaeological record to specific studies at the site of Attirampakkam (ATM). We consider aspects of site distribution, sizes, artefact densities and Acheulian lithic reduction strategies, factoring in issues related to geomorphology, taphonomy and chronology. Acheulian hominins occupied the study region over the early to middle Pleistocene, and the fragmented lithic reduction sequence noted on landscape scales suggests diverse site functions structured by ease of access to quartzite raw material for large flake production in addition to other resources. In contrast to most sites, the absence of raw material at ATM necessitated groups to anticipate this, and organize their behavior on landscape scales, and on-site, to resolve this issue. We show how successive groups were attracted to the site over the early Pleistocene, potentially aiming at exploiting seasonally predictable biological resources in a riparian environment, knowledge of which was transmitted across generations. Considerations of the spatial and temporal variability in artefact densities across a vast site area, along with aspects of the lithic reduction sequences suggests a short-duration occupation by a potentially large group, possibly resulting from aggregation of several small groups as noted in some ethnographic examples of hunter-gatherer fission-fusion strategies. We show drastic changes in behavioral organization in the succeeding Middle Palaeolithic phases at the site and in the region.

Mandibular molar root and pulp cavity morphology in Homo naledi and other Plio-Pleistocene hominins Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Kornelius Kupczik, Lucas K. Delezene, Matthew M. Skinner Abstract The craniomandibular morphology of Homo naledi shows variable resemblances with species across Homo, which confounds an easy assessment of its phylogenetic position. In terms of skull shape, H. naledi has its closest affinities with Homo erectus, while mandibular shape places it closer to early Homo. From a tooth crown perspective, the smaller molars of H. naledi make it distinct from early Homo and H. erectus. Here, we compare the mandibular molar root morphology of six H. naledi individuals from the Dinaledi Chamber to those of African and Eurasian Plio-Pleistocene fossil hominins (totalling 183 mandibular first, second and third molars). The analysis of five root metric variables (cervical plane area, root length, root cervix volume, root branch volume, and root surface area) derived from microCT reconstructions reveals that the molar roots of H. naledi are smaller than those of Homo habilis, Homo rudolfensis, and H. erectus, but that they resemble those of three Homo sp. specimens from Swartkrans and Koobi Fora in size and overall appearance. Moreover, though H. naledi molar roots are similar in absolute size to Pleistocene Homo sapiens, they differ from H. sapiens in having a larger root volume for a given cervical plane area and less taurodont roots; the root cervix-to-branch proportions of H. naledi are comparable to those of Australopithecus africanus and species of Paranthropus. H. naledi also shares a metameric root volume pattern (M2 > M3 > M1) with Australopithecus and Paranthropus but not with any of the other Homo species (M2 > M1 > M3). Our findings therefore concur with previous studies that found that H. naledi shares plesiomorphic features with early Homo, Australopithecus, and Paranthropus. While absolute molar root size aligns H. naledi with Homo from North and South Africa, it is distinguishable from these in terms of root volumetric proportions.

Brain size growth in Australopithecus Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Zachary Cofran Abstract Postnatal growth is one of the proximate means by which humans attain massive adult brain size. Humans are characterized by the maintenance of prenatal brain growth rates into the first postnatal year, as well as an overall extended period of growth. The evolution of this pattern is difficult to assess due to its relatively brief duration and the underrepresentation of well-preserved fossil individuals who died during this short period. In this study, I use Monte Carlo methods to reconstruct postnatal brain growth rates in Australopithecus afarensis and Australopithecus africanus, based on estimates of neonatal brain size and of likely brain size and age at death of infant specimens (A.L. 333-105, DIK-1-1, and Taung). Neonatal brain size is reconstructed from the empirical scaling relationship among catarrhines which humans follow, and conservative estimates of fossils' chronological ages and brain sizes are drawn from the literature. Simulated distributions of these values are used to calculate average annual rates (ARs) of brain growth and proportional size change from birth (PSC), which are compared to resampled statistics from humans, chimpanzees and gorillas of known age and sex. Simulated ARs and PSCs for A. afarensis are significantly lower than those of chimpanzees and gorillas. Both ARs and PSCs for A. africanus are similar to chimpanzee and gorilla values. These results indicate that although these early hominins were derived in some aspects of brain anatomy, high rates of brain growth did not appear until later in human evolution. Moreover, findings also imply that brain growth rates are not a simple function of adult brain size. This study provides important new information about the evolution of brain growth, despite limitations inherent in fossil samples.

The mechanical origins of arm-swinging Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Michael C. Granatosky, Daniel Schmitt Abstract Arm-swinging is a locomotor mode observed only in primates, in which the hindlimbs no longer have a weight bearing function and the forelimbs must propel the body forward and support the entirety of the animal's mass. It has been suggested that the evolution of arm-swinging was preceded by a shift to inverted quadrupedal walking for purposes of feeding and balance, yet little is known about the mechanics of limb use during inverted quadrupedal walking. In this study, we test whether the mechanics of inverted quadrupedal walking make sense as precursors to arm-swinging and whether there are fundamental differences in inverted quadrupedal walking in primates compared to non-primate mammals that would explain the evolution of arm-swinging in primates only. Based on kinetic limb-loading data collected during inverted quadrupedal walking in primates (seven species) and non-primate mammals (three species), we observe that in primates the forelimb serves as the primary propulsive and weight bearing limb. Additionally, heavier individuals tend to support a greater distribution of body weight on their forelimbs than lighter ones. These kinetic patterns are not observed in non-primate mammals. Based on these findings, we propose that the ability to adopt arm-swinging is fairly simple for relatively large-bodied primates and merely requires the animal to release its grasping foot from the substrate. This study fills an important gap concerning the origins of arm-swinging and illuminates previously unknown patterns of primate locomotor evolution.

Temporal shifts in the distribution of murine rodent body size classes at Liang Bua (Flores, Indonesia) reveal new insights into the paleoecology of Homo floresiensis and associated fauna Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): E. Grace Veatch, Matthew W. Tocheri, Thomas Sutikna, Kate McGrath, E. Wahyu Saptomo, Jatmiko, Kristofer M. Helgen Abstract Liang Bua, the type locality of Homo floresiensis, is a limestone cave located in the western part of the Indonesian island of Flores. The relatively continuous stratigraphic sequence of the site spans the past ∼190 kyr and contains ∼275,000 taxonomically identifiable vertebrate skeletal elements, ∼80% of which belong to murine rodent taxa (i.e., rats). Six described genera are present at Liang Bua (Papagomys, Spelaeomys, Hooijeromys, Komodomys, Paulamys, and Rattus), one of which, Hooijeromys, is newly recorded in the site deposits, being previously known only from Early to Middle Pleistocene sites in central Flores. Measurements of the proximal femur (n = 10,212) and distal humerus (n = 1186) indicate five murine body size classes ranging from small (mouse-sized) to giant (common rabbit-sized) are present. The proportions of these five classes across successive stratigraphic units reveal two major changes in murine body size distribution due to significant shifts in the abundances of more open habitat-adapted medium-sized murines versus more closed habitat-adapted smaller-sized ones. One of these changes suggests a modest increase in available open habitats occurred ∼3 ka, likely the result of anthropogenic changes to the landscape related to farming by modern human populations. The other and more significant change occurred ∼60 ka suggesting a rapid shift from more open habitats to more closed conditions at this time. The abrupt reduction of medium-sized murines, along with the disappearance of H. floresiensis, Stegodon florensis insularis (an extinct proboscidean), Varanus komodoensis (Komodo dragon), Leptoptilos robustus (giant marabou stork), and Trigonoceps sp. (vulture) at Liang Bua ∼60–50 ka, is likely the consequence of these animals preferring and tracking more open habitats to elsewhere on the island. If correct, then the precise timing and nature of the extinction of H. floresiensis and its contemporaries must await new discoveries at Liang Bua or other as yet unexcavated sites on Flores.

Speeding in the slow lane: Phylogenetic comparative analyses reveal that not all human life history traits are exceptional Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Ian F. Miller, Steven E. Churchill, Charles L. Nunn Abstract Humans are thought to exhibit an unusual suite of life history traits relative to other primates, with a longer lifespan, later age at first reproduction, and shorter interbirth interval. These assumptions are key components of popular hypotheses about human life history evolution, but they have yet to be investigated phylogenetically. We applied two phylogenetic comparative methods to investigate whether these human life history traits differ from expectations based on other primates: one fits and selects between Brownian and Ornstein-Uhlenbeck models of trait evolution; the other tests for phylogenetic outliers by predicting phenotypic characteristics based on trait covariation and phylogeny for a species of interest. We found that humans have exceptionally short interbirth intervals, long lifespans, and high birth masses. We failed to find evidence that humans have a delayed age at first reproduction relative to body mass or other covariates. Overall, our results support several previous assertions about the uniqueness of human life history characteristics and the importance of cooperative breeding and socioecology in human life history evolution. However, we suggest that several hypotheses about human life history need to be revised in light of our finding that humans do not have a delayed age at first reproduction.

Efficacy of diffeomorphic surface matching and 3D geometric morphometrics for taxonomic discrimination of Early Pleistocene hominin mandibular molars Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): José Braga, Veronika Zimmer, Jean Dumoncel, Chafik Samir, Frikkie de Beer, Clément Zanolli, Deborah Pinto, F. James Rohlf, Frederick E. Grine Abstract Morphometric assessments of the dentition have played significant roles in hypotheses relating to taxonomic diversity among extinct hominins. In this regard, emphasis has been placed on the statistical appraisal of intraspecific variation to identify morphological criteria that convey maximum discriminatory power. Three-dimensional geometric morphometric (3D GM) approaches that utilize landmarks and semi-landmarks to quantify shape variation have enjoyed increasingly popular use over the past twenty-five years in assessments of the outer enamel surface (OES) and enamel–dentine junction (EDJ) of fossil molars. Recently developed diffeomorphic surface matching (DSM) methods that model the deformation between shapes have drastically reduced if not altogether eliminated potential methodological inconsistencies associated with the a priori identification of landmarks and delineation of semi-landmarks. As such, DSM has the potential to better capture the geometric details that describe tooth shape by accounting for both homologous and non-homologous (i.e., discrete) features, and permitting the statistical determination of geometric correspondence. We compare the discriminatory power of 3D GM and DSM in the evaluation of the OES and EDJ of mandibular permanent molars attributed to Australopithecus africanus, Paranthropus robustus and early Homo sp. from the sites of Sterkfontein and Swartkrans. For all three molars, classification and clustering scores demonstrate that DSM performs better at separating the A. africanus and P. robustus samples than does 3D GM. The EDJ provided the best results. P. robustus evinces greater morphological variability than A. africanus. The DSM assessment of the early Homo molar from Swartkrans reveals its distinctiveness from either australopith sample, and the "unknown" specimen from Sterkfontein (Stw 151) is notably more similar to Homo than to A. africanus.

A missing piece of the Papio puzzle: Gorongosa baboon phenostructure and intrageneric relationships Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Felipe I. Martinez, Cristian Capelli, Maria J. Ferreira da Silva, Vera Aldeias, Zeresenay Alemseged, William Archer, Marion Bamford, Dora Biro, René Bobe, David R. Braun, Jörg M. Habermann, Tina Lüdecke, Hilário Madiquida, Jacinto Mathe, Enquye Negash, Luis M. Paulo, Maria Pinto, Marc Stalmans, Frederico Tátá, Susana Carvalho Abstract Most authors recognize six baboon species: hamadryas (Papio hamadryas), Guinea (Papio papio), olive (Papio anubis), yellow (Papio cynocephalus), chacma (Papio ursinus), and Kinda (Papio kindae). However, there is still debate regarding the taxonomic status, phylogenetic relationships, and the amount of gene flow occurring between species. Here, we present ongoing research on baboon morphological diversity in Gorongosa National Park (GNP), located in central Mozambique, south of the Zambezi River, at the southern end of the East African Rift System. The park exhibits outstanding ecological diversity and hosts more than 200 baboon troops. Gorongosa National Park baboons have previously been classified as chacma baboons (P. ursinus). In accordance with this, two mtDNA samples from the park have been placed in the same mtDNA clade as the northern chacma baboons. However, GNP baboons exhibit morphological features common in yellow baboons (e.g., yellow fur color), suggesting that parapatric gene flow between chacma and yellow baboons might have occurred in the past or could be ongoing. We investigated the phenostructure of the Gorongosa baboons using two approaches: 1) description of external phenotypic features, such as coloration and body size, and 2) 3D geometric morphometric analysis of 43 craniofacial landmarks on 11 specimens from Gorongosa compared to a pan-African sample of 352 baboons. The results show that Gorongosa baboons exhibit a mosaic of features shared with southern P. cynocephalus and P. ursinus griseipes. The GNP baboon phenotype fits within a geographic clinal pattern of replacing allotaxa. We put forward the hypothesis of either past and/or ongoing hybridization between the gray-footed chacma and southern yellow baboons in Gorongosa or an isolation-by-distance scenario in which the GNP baboons are geographically and morphologically intermediate. These two scenarios are not mutually exclusive. We highlight the potential of baboons as a useful model to understand speciation and hybridization in early human evolution.

Late Middle Pleistocene hominin teeth from Tongzi, southern China Publication date: May 2019 Source: Journal of Human Evolution, Volume 130 Author(s): Song Xing, María Martinón-Torres, José María Bermúdez de Castro Abstract In 1972 and 1983, four hominin teeth were recovered from Yanhui Cave, Tongzi, southern China and assigned to later Homo erectus or archaic Homo sapiens. The teeth can be dated to approximately 172,000–240,000 years before present. Here, in addition to the standard morphological comparisons, we reevaluate the morphology of the hominin teeth using geometric morphometric analyses and micro-computed tomography (micro-CT) scans. The Tongzi teeth were primarily compared to hominins from the same chronological period (late Middle Pleistocene) and/or the same geographic area (East Asia), although a wide range of hominins were included in the comparative sample. This study reveals that the Tongzi teeth do not fit the morphological pattern of classic H. erectus, and expands our understanding of the morphological diversity of the Asian Middle Pleistocene hominins. Overall, our results point to the existence of more than one paleodeme in East Asia during this period: one that can be taxonomically classified as H. erectus sensu stricto (represented by fossils such as Zhoukoudian, Hexian, and Yiyuan) and a second that is characterized by the expression of derived traits more commonly found in later Homo (such as the crown symmetry, lingual reduction, and simplified EDJ surface of the third premolar). More fossil and genetic findings will help assess the taxonomy of the "non-erectus" Asian Middle Pleistocene populations, like the Tongzi hominins.