Estimating sexual size dimorphism in fossil species from posterior probability densities.

Accurate characterization of sexual dimorphism is crucial in evolutionary biology because of its significance in understanding present and past adaptations involving reproductive and resource use strategies of species. However, inferring dimorphism in fossil assemblages is difficult, particularly with relatively low dimorphism. Commonly used methods of estimating dimorphism levels in fossils include the mean method, the binomial dimorphism index, and the coefficient of variation method. These methods have been reported to overestimate low levels of dimorphism, which is problematic when investigating issues such as canine size dimorphism in primates and its relation to reproductive strategies. Here, we introduce the posterior density peak (pdPeak) method that utilizes the Bayesian inference to provide posterior probability densities of dimorphism levels and within-sex variance. The highest posterior density point is termed the pdPeak. We investigated performance of the pdPeak method and made comparisons with the above-mentioned conventional methods via 1) computer-generated samples simulating a range of conditions and 2) application to canine crown-diameter datasets of extant known-sex anthropoids. Results showed that the pdPeak method is capable of unbiased estimates in a broader range of dimorphism levels than the other methods and uniquely provides reliable interval estimates. Although attention is required to its underestimation tendency when some of the distributional assumptions are violated, we demonstrate that the pdPeak method enables a more accurate dimorphism estimate at lower dimorphism levels than previously possible, which is important to illuminating human evolution.

Canine sexual dimorphism in Ardipithecus ramidus was nearly human-like.

Body and canine size dimorphism in fossils inform sociobehavioral hypotheses on human evolution and have been of interest since Darwin's famous reflections on the subject. Here, we assemble a large dataset of fossil canines of the human clade, including all available Ardipithecus ramidus fossils recovered from the Middle Awash and Gona research areas in Ethiopia, and systematically examine canine dimorphism through evolutionary time. In particular, we apply a Bayesian probabilistic method that reduces bias when estimating weak and moderate levels of dimorphism. Our results show that Ar. ramidus canine dimorphism was significantly weaker than in the bonobo, the least dimorphic and behaviorally least aggressive among extant great apes. Average male-to-female size ratios of the canine in Ar. ramidus are estimated as 1.06 and 1.13 in the upper and lower canines, respectively, within modern human population ranges of variation. The slightly greater magnitude of canine size dimorphism in the lower than in the upper canines of Ar. ramidus appears to be shared with early Australopithecus, suggesting that male canine reduction was initially more advanced in the behaviorally important upper canine. The available fossil evidence suggests a drastic size reduction of the male canine prior to Ar. ramidus and the earliest known members of the human clade, with little change in canine dimorphism levels thereafter. This evolutionary pattern indicates a profound behavioral shift associated with comparatively weak levels of male aggression early in human evolution, a pattern that was subsequently shared by Australopithecus and Homo.

Reconstruction and analysis of the DAN5/P1 and BSN12/P1 Gona Early Pleistocene Homo fossils.

Two Early Pleistocene fossils from Gona, Ethiopia, were originally assigned to Homo erectus, and their differences in size and robusticity were attributed to either sexual dimorphism or anagenetic evolution. In the current study, we both revisit the taxonomic affinities of these fossils and assess whether morphological differences between them reflect temporal evolution or sexual variation. We generated virtual reconstructions of the mostly complete ∼1.55 Ma DAN5/P1 calvaria and the less complete 1.26 Ma BSN12/P1 fossil, allowing us to directly compare their anterior vault shapes using landmark-based shape analysis. The two fossils are similar in calvaria shape to H. erectus and also to other Early Pleistocene Homo species based on a geometric morphometric analysis of calvaria landmarks and semilandmarks. The DAN5/P1 fossil bears a particularly close affinity to the Georgian H. erectus fossils and to KNM-ER 1813 (H. habilis), probably reflecting allometric influences on vault shape. Combined with species-specific traits of the neurocranium (e.g., midline keeling, angular torus), we confirm that these fossils are likely early African H. erectus. We calculated regression-based estimates of endocranial volume for BSN12/P1 of 882-910 cm3 based on three virtual reconstructions. Although BSN12/P1 is markedly larger than DAN5/P1 (598 cm3), both fossils represent the smallest adult H. erectus known from their respective time periods in Africa. Some of the difference in endocranial volume between the two Gona fossils reflects broader species-level brain expansion from 1.77 to 0.01 Ma, confirmed here using a large sample (n = 38) of H. erectus. However, shape differences between these fossils did not reflect species-level changes to calvaria shape. Moreover, the analysis failed to recover a clear pattern of sexually patterned size or shape differences within H. erectus based on our current assessments of sex for individual fossils.

Fossil Cercopithecidae from the Early Pliocene Sagantole Formation at Gona, Ethiopia.

The Early Pliocene Sagantole Fm. in the Gona Project area, Afar State, Ethiopia, is noted for discoveries of the early hominin Ardipithecus ramidus. A large series of fossil cercopithecid primates dated to between 4.8 and 4.3 Ma has also been collected from these sediments. In this paper, we use qualitative analysis and standard dental and postcranial measures to systematically describe the craniodental remains and tentatively allocate postcrania to taxa where we are able to. We then use these data to compare these specimens to fossil assemblages from contemporary sites, interpret their paleobiology, and discuss implications for the paleoecology of the Gona Sagantole Fm. We recognize three cercopithecid species in the Gona Sagantole Fm. Pliopapio alemui makes up approximately two-thirds of the identifiable specimens; nearly all of the rest are allocated to Kuseracolobus aramisi, and a single molar indicates the presence of a second, somewhat larger but morphologically distinct papionin. Among the Early Pliocene cercopithecids from Gona are also a number of postcranial elements. None of the postcranial remains are directly associated with any of the cranial material. Nonetheless, some of the distal humeri and proximal femora can be tentatively allocated to either Pl. alemui or K. aramisi based on a combination of size, as the latter is approximately 50% larger than the former, and morphology. If these assignments are correct, they suggest K. aramisi was primarily arboreal and similar to most extant colobines, whereas Pl. alemui was more mixed in its substrate use, being more terrestrially adapted than K. aramisi, but less so than extant Papio or Theropithecus. Thus, we interpret the predominance of Pl. alemui over K. aramisi is consistent with a somewhat more open environment at Gona than at Aramis.

Ardipithecus ramidus postcrania from the Gona Project area, Afar Regional State, Ethiopia.

Functional analyses of the 4.4 Ma hominin Ardipithecus ramidus postcrania revealed a previously unknown and unpredicted locomotor pattern combining arboreal clambering and a form of terrestrial bipedality. To date, all of the fossil evidence of Ar. ramidus locomotion has been collected from the Aramis area of the Middle Awash Research Project in Ethiopia. Here, we present the results of an analysis of additional early Pliocene Ar. ramidus fossils from the Gona Project study area, Ethiopia, that includes a fragmentary but informative partial skeleton (GWM67/P2) and additional isolated manual remains. While we reinforce the original functional interpretations of Ar. ramidus of having a mixed locomotor adaptation of terrestrial bipedality and arboreal clambering, we broaden our understanding of the nature of its locomotor pattern by documenting better the function of the hip, ankle, and foot. The newly recovered fossils document a greater adaptation to bipedality in the Ar. ramidus ankle and hallux than previously recognized. In addition, a newly discovered scaphoid bone with a fusing os centrale provides further evidence about the nature of hominin hand evolution.

Late Miocene hominin teeth from the Gona Paleoanthropological Research Project area, Afar, Ethiopia.

Since 2000, significant collections of Latest Miocene hominin fossils have been recovered from Chad, Kenya, and Ethiopia. These fossils have provided a better understanding of earliest hominin biology and context. Here, we describe five hominin teeth from two periods (ca. 5.4 Million-years-ago and ca. 6.3 Ma) that were recovered from the Adu-Asa Formation in the Gona Paleoanthropological Research Project area in the Afar, Ethiopia that we assign to either Hominina, gen. et sp. indet. or Ardipithecus kadabba. These specimens are compared with extant African ape and other Latest Miocene and Early Pliocene hominin teeth. The derived morphology of the large, non-sectorial maxillary canine and mandibular third premolar links them with later hominins and they are phenetically distinguishable and thus phyletically distinct from extant apes.

The endocast from Dana Aoule North (DAN5/P1): A 1.5 million year-old human braincase from Gona, Afar, Ethiopia.

The nearly complete cranium DAN5/P1 was found at Gona (Afar, Ethiopia), dated to 1.5-1.6 Ma, and assigned to the species Homo erectus. Its size is, nonetheless, particularly small for the known range of variation of this taxon, and the cranial capacity has been estimated as 598 cc. In this study, we analyzed a reconstruction of its endocranial cast, to investigate its paleoneurological features. The main anatomical traits of the endocast were described, and its morphology was compared with other fossil and modern human samples. The endocast shows most of the traits associated with less encephalized human taxa, like narrow frontal lobes and a simple meningeal vascular network with posterior parietal branches. The parietal region is relatively tall and rounded, although not especially large. Based on our set of measures, the general endocranial proportions are within the range of fossils included in the species Homo habilis or in the genus Australopithecus. Similarities with the genus Homo include a more posterior position of the frontal lobe relative to the cranial bones, and the general endocranial length and width when size is taken into account. This new specimen extends the known brain size variability of Homo ergaster/erectus, while suggesting that differences in gross brain proportions among early human species, or even between early humans and australopiths, were absent or subtle.

Early Pliocene hominids from Gona, Ethiopia.

Comparative biomolecular studies suggest that the last common ancestor of humans and chimpanzees, our closest living relatives, lived during the Late Miocene-Early Pliocene. Fossil evidence of Late Miocene-Early Pliocene hominid evolution is rare and limited to a few sites in Ethiopia, Kenya and Chad. Here we report new Early Pliocene hominid discoveries and their palaeoenvironmental context from the fossiliferous deposits of As Duma, Gona Western Margin (GWM), Afar, Ethiopia. The hominid dental anatomy (occlusal enamel thickness, absolute and relative size of the first and second lower molar crowns, and premolar crown and radicular anatomy) indicates attribution to Ardipithecus ramidus. The combined radioisotopic and palaeomagnetic data suggest an age of between 4.51 and 4.32 million years for the hominid finds at As Duma. Diverse sources of data (sedimentology, faunal composition, ecomorphological variables and stable carbon isotopic evidence from the palaeosols and fossil tooth enamel) indicate that the Early Pliocene As Duma sediments sample a moderate rainfall woodland and woodland/grassland.

Technological variation in the earliest Oldowan from Gona, Afar, Ethiopia.

Inter-site technological variation in the archaeological record is one of the richest potential sources of information about Plio-Pleistocene hominid behavior and evolution. However, appropriate methods for describing and comparing Oldowan assemblages have yet to be agreed upon, and interpretation of the early record remains highly controversial. Particularly salient is disagreement over whether the Oldowan is a single technological phenomenon or is more accurately divided into multiple regional and/or chronological traditions, perhaps including a less developed Pre-Oldowan phase in the late Pliocene. Some of this disagreement reflects theoretical and methodological differences between research traditions and some is more directly evidential. Here we present a framework for describing and interpreting Oldowan variation and apply it to three Pliocene assemblages (EG-10, EG-12, and OGS-7) from Gona, all dated to c. 2.6 million years (Ma). Results indicate proficient knapping and a full range of Oldowan reduction strategies in these earliest known occurrences, consistent with the idea of an Oldowan "technological stasis" from 2.6-1.6 Ma. Patterns of variation in raw material selection and predominant reduction strategy at each site clearly indicate the importance of cultural transmission in the Oldowan, but confounding ecological and economic variation continue to render interpretation in terms of multiple tool making traditions or species inappropriate. We propose that cultural transmission and ecological adaptation should be recognized as complementary, rather than mutually exclusive, mechanisms in future attempts to explain Oldowan technological variation.

Co-occurrence of Acheulian and Oldowan artifacts with Homo erectus cranial fossils from Gona, Afar, Ethiopia.

Although stone tools generally co-occur with early members of the genus Homo, they are rarely found in direct association with hominins. We report that both Acheulian and Oldowan artifacts and Homo erectus crania were found in close association at 1.26 million years (Ma) ago at Busidima North (BSN12), and ca. 1.6 to 1.5 Ma ago at Dana Aoule North (DAN5) archaeological sites at Gona, Afar, Ethiopia. The BSN12 partial cranium is robust and large, while the DAN5 cranium is smaller and more gracile, suggesting that H. erectus was probably a sexually dimorphic species. The evidence from Gona shows behavioral diversity and flexibility with a lengthy and concurrent use of both stone technologies by H. erectus, confounding a simple "single species/single technology" view of early Homo.

1.9-million- and 2.4-million-year-old artifacts and stone tool-cutmarked bones from Ain Boucherit, Algeria.

East Africa has provided the earliest known evidence for Oldowan stone artifacts and hominin-induced stone tool cutmarks dated to ~2.6 million years (Ma) ago. The ~1.8-million-year-old stone artifacts from Ain Hanech (Algeria) were considered to represent the oldest archaeological materials in North Africa. Here we report older stone artifacts and cutmarked bones excavated from two nearby deposits at Ain Boucherit estimated to ~1.9 Ma ago, and the older to ~2.4 Ma ago. Hence, the Ain Boucherit evidence shows that ancestral hominins inhabited the Mediterranean fringe in northern Africa much earlier than previously thought. The evidence strongly argues for early dispersal of stone tool manufacture and use from East Africa or a possible multiple-origin scenario of stone technology in both East and North Africa.

A female Homo erectus pelvis from Gona, Ethiopia.

Analyses of the KNM-WT 15000 Homo erectus juvenile male partial skeleton from Kenya concluded that this species had a tall thin body shape due to specialized locomotor and climatic adaptations. Moreover, it was concluded that H. erectus pelves were obstetrically restricted to birthing a small-brained altricial neonate. Here we describe a nearly complete early Pleistocene adult female H. erectus pelvis from the Busidima Formation of Gona, Afar, Ethiopia. This obstetrically capacious pelvis demonstrates that pelvic shape in H. erectus was evolving in response to increasing fetal brain size. This pelvis indicates that neither adaptations to tropical environments nor endurance running were primary selective factors in determining pelvis morphology in H. erectus during the early Pleistocene.

Raw material selectivity of the earliest stone toolmakers at Gona, Afar, Ethiopia.

Published evidence of Oldowan stone exploitation generally supports the conclusion that patterns of raw material use were determined by local availability. This is contradicted by the results of systematic studies of raw material availability and use among the earliest known archaeological sites from Gona, Afar, Ethiopia. Artifact assemblages from six Pliocene archaeological sites were compared with six random cobble samples taken from associated conglomerates that record pene-contemporaneous raw material availability. Artifacts and cobbles were evaluated according to four variables intended to capture major elements of material quality: rock type, phenocryst percentage, average phenocryst size, and groundmass texture. Analyses of these variables provide evidence of hominid selectivity for raw material quality. These results demonstrate that raw material selectivity was a potential component of Oldowan technological organization from its earliest appearance and document a level of technological sophistication that is not always attributed to Pliocene hominids.

Cutmarked bones from Pliocene archaeological sites at Gona, Afar, Ethiopia: implications for the function of the world's oldest stone tools.

Newly recorded archaeological sites at Gona (Afar, Ethiopia) preserve both stone tools and faunal remains. These sites have also yielded the largest sample of cutmarked bones known from the time interval 2.58-2.1 million years ago (Ma). Most of the cutmarks on the Gona fauna possess obvious macroscopic (e.g., deep V-shaped cross-sections) and microscopic (e.g., internal microstriations, Herzian cones, shoulder effects) features that allow us to identify them confidently as instances of stone tool-imparted damage caused by hominid butchery. In addition, preliminary observations of the anatomical placement of cutmarks on several of the recovered bone specimens suggest that Gona hominids may have eviscerated carcasses and defleshed the fully muscled upper and intermediate limb bones of ungulates--activities that further suggest that Late Pliocene hominids may have gained early access to large mammal carcasses. These observations support the hypothesis that the earliest stone artifacts functioned primarily as butchery tools and also imply that hunting and/or aggressive scavenging of large ungulate carcasses may have been part of the behavioral repertoire of hominids by c. 2.5 Ma, although a larger sample of cutmarked bone specimens is necessary to support the latter inference.