The endocast morphology of LES1, Homo naledi.

OBJECTIVES: Homo naledi is near the extreme of small brain size within Homo but is easily recognized as Homo in other aspects of endocast morphology. This study adds new evidence of the endocast morphology of H. naledi by describing the Lesedi Hominin 1 (LES1) endocranium from the Lesedi Chamber and compares it to the previously known H. naledi individual Dinaledi Hominin 3 (DH3) as well as other hominin taxa. MATERIALS AND METHODS: We examined interlandmark distances with both univariate and multivariate methods in multiple hominin taxa and both species of Pan. For each distance, we compared groups using adjusted Z-scores (Azs). Our multivariate analyses included both principal component analyses (PCA) and linear discriminant analyses (LDA). RESULTS: DH3 and LES1 each have absolute third frontal convolution measures that enter the upper half of the variation for Homo sapiens, Homo erectus, and Homo neanderthalensis. Examined relative to the cube root of endocranial volume, H. naledi ranks among the highest values in these samples of Homo. Both absolute and relative values for H. naledi specimens are far above Pan, Australopithecus, and Paranthropus, suggesting an expanded Broca's area. CONCLUSIONS: Both qualitative and quantitative analyses show consistency between LES1 and other H. naledi endocasts and confirm the shared morphology of H. naledi with H. sapiens, H. neanderthalensis, and some specimens of H. erectus.

Personal identification using frontal sinus coding methods: The effect of mixed image modality comparisons.

Several code-based methods have been created for comparing the frontal sinus in skeletal identification scenarios. However, little is known regarding matched-pair accuracy rates of these methods or how varying image modalities may affect these rates. The goals of this study were to validate the exclusion rates and to establish matched-pair accuracy rates of two well-cited coding methods, Cameriere et al. [23] and Tatlisumak et al. [24]. Additionally, individual variables were assessed for consistency in scoring between image modalities. Using a sample of U.S. African American, Native American, and European American females and males (n = 225), we examined individual variable scoring and string codes between two different image modalities (radiographs and CT-based 3D models). Arcades showed poor scoring consistency between modalities (p < 0.001). Although exclusion rates were similar to those reported in the original studies (93%-96%), matched-pair accuracy rates were low (13%-18%). None of the demographics (collection, sex, age, ancestry, and orientation) had an effect on the odds of a match. Interobserver and intraobserver analyses showed moderate to near-perfect agreement for all variables except supraorbital cells, which had minimal to no agreement. Currently, we do not recommend the application of these frontal sinus coding methods independent of other supporting identification methods given low variable consistency and accuracy rates. Visual identification should still be used to include or exclude an identification when using the frontal sinus.

Investigating error in saw mark minimum kerf width measurements.

Minimum kerf width (MKW) is often assessed in medicolegal cases of skeletal sharp force trauma and can provide information about the tool used. Previously published saw MKW research focuses on the relationship between MKW and saw blade set width (width of the saw blade including tooth set). Despite these publications using various measurement tools and methods to collect MKW, error in MKW measurements is not well-documented. The goal of this research was to investigate MKW measurement precision in terms of intraobserver error, interobserver error, and the effects of measurement modality (e.g., calipers versus stereo microscope). The study sample consisted of 351 incomplete cuts made with 27 diverse saws. MKW was collected using three methods: (1) inserting the internal caliper jaws into the incomplete kerf, (2) placing the external caliper jaws on the cortical surface, and (3) using a digital stereo microscope and associated measurement software. For each method, intraobserver error and interobserver error were assessed. Differences in measurements taken with the three measurement modalities were assessed for each observer as well. Relative technical error of measurement (rTEM) and coefficients of reliability indicate that internal caliper and stereo microscope measurements had the lowest intra- and interobserver error (rTEM = 3.72% to 6.15%; r = 0.98-0.996). External caliper measurements performed the worst (rTEM: intra = 8.53% and inter = 21.32%). There was higher precision between internal caliper and stereo microscope measurements than with the external caliper measurements. This research highlights the need for measurement standardization.

Ontogenetic patterns in human frontal sinus shape: A longitudinal study using elliptical Fourier analysis.

Frontal sinus morphology is highly variable across individuals, but little is known regarding how or at what age that variation is reached. Existing ontogenetic studies are conflicting and often cross-sectional in nature, limiting understanding of individualistic growth. Studies investigating sinus growth with longitudinal series often focus on lateral cephalograms and consequently do not capture the sinus morphological features that are most relevant to clinical and medicolegal settings (e.g., arcade/scalloping, width-to-height dimensions, asymmetry). Longitudinal analysis of sinus morphology from frontal radiographs is important to understand when sinus morphology stabilizes. The purpose of this study was to investigate at what age the frontal sinus attains its final shape, and whether sex-based differences in ontogeny are evident, using a longitudinal sample of posterior-anterior (PA) frontal radiographs from the AAOF Legacy Collection. Frontal sinus outlines were manually traced in 935 radiographs from 111 individuals (55F/56M) spanning 8-29 years of age. Outlines were subjected to elliptical Fourier analysis (EFA) and underwent principal components analysis (PCA). PC1 (51.02% of variation) appears to represent the relative height and breadth of the sinus, PC2 (11.73%) and PC3 (10.03%) captures the degree of relative complexity in the outlines. Individual PC scores were plotted against age-in-months with individual Loess growth curves. Overall, younger individuals typically display relatively shorter, flatter sinuses, increasing in vertical complexity with age. Mixed-effect models on PC1 indicate significant effects for the repeated measure of years (p < 0.001). Within individuals, Euclidean distances of PCs between each sinus outline and their oldest-age outline (i.e., final morphology) were calculated and plotted against age-in-months with Loess growth curves. The results indicate that final frontal sinus morphology is mostly attained by 20 yoa regardless of sex. There is sexual dimorphism in ontogenetic trajectories: females attain frontal sinus shape earlier than males. Specifically, Loess growth curves of the Euclidean distances to final sinus shape indicate that female shape shows decreased development at 14-16 yoa, with males approaching stabilization at 18-20 yoa. These trends were supported by paired t-tests on PC1 between each year and the oldest age, whereby significant differences end for females starting at 15 and 18 yoa for males. The timing of shape-stabilization in the current study closely aligns with previous studies on linear and size dimensions, indicating a close relationship between the ontogeny of frontal sinus shape and size. This research has several implications in diverse fields. Documenting ontogenetic patterns in modern humans could lead to more accurate interpretations of frontal sinus variation in hominin lineages. Understanding the age at which frontal sinus shape and size stabilizes in pediatric populations has important clinical implications, with future studies needed to investigate if/how sinus development directly relates to sinonasal disease susceptibility (e.g., sinusitis), surgical complications, and/or expected trauma patterns. For forensic practitioners utilizing frontal sinus comparisons for decedent identifications, it is important to know at what age these features stabilize to understand how much change may be expected between antemortem and postmortem radiographs.

The Effects of Cranial Orientation on Forensic Frontal Sinus Identification as Assessed by Outline Analyses.

The utility of frontal sinuses for personal identification is widely recognized, but potential factors affecting its reliability remain uncertain. Deviations in cranial position between antemortem and postmortem radiographs may affect sinus appearance. This study investigates how slight deviations in orientations affect sinus size and outline shape and potentially impact identification. Frontal sinus models were created from CT scans of 21 individuals and digitally oriented to represent three clinically relevant radiographic views. From each standard view, model orientations were deviated at 5° intervals in horizontal, vertical, and diagonal (e.g., left-up) directions (27 orientations per individual). For each orientation, sinus dimensions were obtained, and outline shape was assessed by elliptical Fourier analyses and principal component (PC) analyses. Wilcoxon sign rank tests indicated that sinus breadth remained relatively stable (p > 0.05), while sinus height was significantly affected with vertical deviations (p < 0.006). Mann-Whitney U tests on Euclidean distances from the PC scores indicated consistently lower intra- versus inter-individual distances (p < 0.05). Two of the three orientations maintained perfect (100%) outline identification matches, while the third had a 98% match rate. Smaller and/or discontinuous sinuses were most problematic, and although match rates are high, practitioners should be aware of possible alterations in sinus variables when conducting frontal sinus identifications.

Applying posterior probability informed thresholds to traditional cranial trait sex estimation methods.

Sex estimation methods using traditional cranial nonmetric traits utilize predictive models to produce a final sex estimation, using the resulting model's score to classify the individual. When sex estimations are assigned from discriminant scoring alone, statistical confidence in the resultant estimate is not always assessed or reported. Although some forensic anthropologists may qualitatively report their confidence in the assessment (e.g., "probable male"), these statements are subjective, not standardized, and not necessarily based on statistical results in a uniform way. The goals of this study were to evaluate how posterior probability-informed thresholds (PPITs) impacted accuracy rates, assess the balance between sample inclusion and accuracy for the proposed PPIT approach, and make recommendations for the use and interpretation of specific thresholds in casework. Using a sample of U.S. Black and White females and males (n = 292), we examined how PPITs can standardize the decision-making process of inferring sex for two methods using nonmetric cranial traits. We found that using PPITs of at least 0.85 increased accuracy (over 92% for some PPITs) yet remained highly inclusive of the sample. PPITs < 0.75 did not produce classification accuracy rates significantly higher than chance, and when using these cranial trait sex estimation methods, cases with posterior probabilities (PPs) <0.75 should be reported as "indeterminate." The 0.75-0.84 PPIT interval had an accuracy rate of 76%, which was both statistically significantly different from chance as well as from the higher (>0.85) groups, suggesting that although sex estimation at this level may be acceptable, the results hold lower confidence.

Sex estimation of the subadult ilium prior to acetabular fusion.

Previous studies evaluating sexual dimorphism in subadult pelvic features present variable, and at times conflicting, conclusions. As a result, there is yet to be a consensus on whether the subadult pelvis can be used in sex estimation methods. This study aims to assess the forensic utility of ilium shape and greater sciatic notch morphology in sexing subadult pelves prior to acetabular fusion. A sample of 397 modern U.S. individuals with unfused acetabula (i.e., tri-radiate cartilages) aged birth to 14 years was queried from a larger sample of postmortem computed tomography scans. Elliptical Fourier analyses were performed on ilium and greater sciatic notch outlines and resultant PCs were evaluated for significant effects of sex and age. Greater sciatic notch metrics were also collected. Stepwise linear discriminant function analyses with leave-one-out cross-validation were performed on the PCs and metric variables. Analyses were performed on pooled samples, on age-specific cohort samples, and on samples that iteratively removed the youngest one-year cohort. Cross-validated correct classification rates ranged from 57% to 65% and no patterns were observed to support an appearance and/or consistent expression of sexually diagnostic traits. Based on the results, sex estimation using these features is not recommended in pelvic remains prior to acetabular fusion, although the sample sizes of individuals over 5 years of age were limited in this study. Future studies should focus on the sexually diagnostic ability of pelvic traits in subadult samples post-fusion of the acetabulum.

Forensic Tools for Species Identification of Skeletal Remains: Metrics, Statistics, and OsteoID.

Although nonhuman remains constitute a significant portion of forensic anthropological casework, the potential use of bone metrics to assess the human origin and to classify species of skeletal remains has not been thoroughly investigated. This study aimed to assess the utility of quantitative methods in distinguishing human from nonhuman remains and present additional resources for species identification. Over 50,000 measurements were compiled from humans and 27 nonhuman (mostly North American) species. Decision trees developed from the long bone data can differentiate human from nonhuman remains with over 90% accuracy (>98% accuracy for the human sample), even if all long bones are pooled. Stepwise discriminant function results were slightly lower (>87.4% overall accuracy). The quantitative models can be used to support visual identifications or preliminarily assess forensic significance at scenes. For species classification, bone-specific discriminant functions returned accuracies between 77.7% and 89.1%, but classification results varied highly across species. From the study data, we developed a web tool, OsteoID, for users who can input measurements and be shown photographs of potential bones/species to aid in visual identification. OsteoID also includes supplementary images (e.g., 3D scans), creating an additional resource for forensic anthropologists and others involved in skeletal species identification and comparative osteology.

Revisiting global patterns of frontal sinus aplasia utilizing computed tomography.

While frontal sinus aplasia (agenesis, absence) has been proposed as a potential marker in forensic positive identifications, frequency rates are likely dependent upon how presence is defined. This study investigates how two methods of defining frontal sinus presence affects aplasia frequency rates. Using CT scans of 772 adult individuals from diverse geographic regions, frontal sinus presence was assessed two ways: 1) the XR-method- coded present if the sinus extended above the supra-orbital line, and 2) the CT-method- coded present with any indication of the frontal sinus. The XR-method consistently provided higher aplasia frequencies, averaging an 18.31% discrepancy with the CT-method. Method discrepancies were higher in females (averaged-sides: 24.6%) than males (averaged-sides: 13.82%). Oceanian individuals displayed the highest aplasia rates using either method, and the highest discrepancy between methods (averaged-sides: 31.30%); Europeans, displaying the lowest aplasia rates in either method, also displayed the lowest method discrepancy (average-sides: 7.37%). Fisher's Exact tests on the biologically-defined CT aplasia rates indicate females are significantly different from males for unilateral aplasia (p=0.0035); Arctic populations are significantly different from most groups (all p<0.005), exception being Oceanian. Results suggest the lower, biologically-defined CT aplasia rates are more useful in corroborating identifications than the inflated XR frequencies and illustrate the importance of citing aplasia rates from similarly-composed samples. However, due to practicality and resource availability, the XR-method may be more appropriate. Most importantly, reported aplasia rates from one method should be cited in casework utilizing the other method.

The importance of processing procedures and threshold values in CT scan segmentation of skeletal elements: An example using the immature os coxa.

As the accessibility and utility of virtual databases of skeletal collections continues to grow, the impact that scan processing procedures has on the accuracy of data obtained from virtual databases remains relatively unknown. This study quantifies the intra- and inter-observer error generated from varying computed tomography (CT) scan processing protocols, including re-segmentation, incrementally varying thresholding value, and data collectors' selection of the threshold value on a set of virtual subadult pelves. Four observers segmented the subadult ossa coxarum from postmortem CT scans of the fully-fleshed bodies of eleven individuals of varying ages. Segmentation protocol was set, with the exception of each observer selecting their own thresholding value for each scan. The resulting smoothed pelvic surfaces were then compared using deviation analyses. Root mean square error (RMSE), average distance deviation, and maximum deviation distances demonstrated that thresholding values of ∼50 HU (Hounsfield units) are easily tolerated, the surfaces generated are robust to error, and threshold value selection does not systematically vary with user experience. The importance of consistent methodology during segmentation protocol is highlighted here, especially with regards to consistency in both selected thresholding value as well as smoothing protocol, as these variables can affect subsequent measurements of the resultant surfaces.

A Quantitative Assessment of Zygomatic Projection for Ancestry Estimation.

Anterior zygomatic projection (ZP) is historically referenced as a useful trait in ancestry estimation, particularly when differentiating between Native Americans and U.S. Whites and Blacks. However, methods of assessing ZP vary, are susceptible to multiple interpretations, and have not been quantitatively validated. This study uses 228 3D surface scans of U.S. Whites, U.S. Blacks, and Native Americans to quantitatively test the ZP methods published by Rhine in 1990 (Skeletal attribution of race: methods for forensic anthropology, Albuquerque, NM, Maxwell Museum of Anthropology, 1990) and Bass in 1995 (Human osteology: a laboratory and field manual, Columbia, MO: Missouri Archaeological Society, 1995). Two ZP angles and two distances, representing method interpretations, were collected and analyzed via ANOVA and discriminant function analyses. Although significant ancestry differences were found across all variables, only the Bass inferior zygomatic distance successfully differentiated the pooled Native American group from pooled U.S. Whites/Blacks (73.7% correct). Arctic Native Americans, displaying the most projecting zygomas, are driving group differences. Significant overlap in measurement distributions were observed between groups in all variables, indicating limited forensic utility.

An Alternative Method to Using a Mandibulometer.

Mandibular length, mandibular angle, and maximum ramus height measurements used during forensic evaluation of skeletal remains require use of a mandibulometer. This study presents a new method of taking these measurements from images, so that practitioners without access to an expensive mandibulometer or those working with 3D models (e.g., CT scans) can collect and utilize these measurements. Ten trials performed on a sample of 45 mandibles were used to compare measurements collected from photographs and images extracted from 3D surface scans to those collected with a mandibulometer, including intra- and inter-observer analyses. All technical error of measurement (TEM) values were less than 2 mm regardless of observer, trial, or method. Relative TEM values were less than 2% for all except mandibular length (2.10%) and ramus height (2.32%) for the right versus left photographs. Results are comparable with mandibulometer error rates, indicating that the proposed method is accurate and reliable.

Endocast morphology of Homo naledi from the Dinaledi Chamber, South Africa.

Hominin cranial remains from the Dinaledi Chamber, South Africa, represent multiple individuals of the species Homo naledi This species exhibits a small endocranial volume comparable to Australopithecus, combined with several aspects of external cranial anatomy similar to larger-brained species of Homo such as Homo habilis and Homo erectus Here, we describe the endocast anatomy of this recently discovered species. Despite the small size of the H. naledi endocasts, they share several aspects of structure in common with other species of Homo, not found in other hominins or great apes, notably in the organization of the inferior frontal and lateral orbital gyri. The presence of such structural innovations in a small-brained hominin may have relevance to behavioral evolution within the genus Homo.

A Morphometric Outline Analysis of Ancestry and Sex Differences in Cranial Shape.

Forensic anthropological techniques that utilize nonmetric cranial traits to estimate sex and ancestry have historically been criticized for their subjectivity and qualitative nature. Nonmetric traits, however, continue to be valuable tools in identifying remains in forensic investigations. In this study, geometric morphometric analyses of cranial outlines were performed to quantitatively assess population and sex variation in modern human cranial shapes and to verify group differences in previously reported qualitative traits. Elliptical Fourier analysis was conducted on two-dimensional images of the left lateral, posterior, and superior cranial views from 198 U.S. Black and White crania. Results reveal significant sex and population differences in cranial shape that generally coincide with traditional qualitative descriptions. Although sex classification was relatively low (70.2%), ancestry classification was higher (92.4%), indicating that outline analyses that incorporate multiple nonmetric traits into a single statistical analysis may provide a more objective and accurate means of ancestry classification.

A Validation Study of the Langley et al. (2017) Decision Tree Model for Sex Estimation.

Langley et al. (2017) developed a sex estimation decision tree utilizing two traditional cranial traits (glabella and mastoid) and a new trait: zygomatic extension. This study aimed to test the reliability of their zygomatic extension scoring method and validate their sex estimation method. Ordinal score data were collected from 281 male and female U.S. White and Black individuals. The five traditional cranial traits were collected from physical specimens, while zygomatic extension was scored from 3D cranial models. Intra- and interobserver analyses carried out on a subsample of 30 individuals indicate good agreement between zygomatic scores. The decision tree correctly sexed 71.5% of the sample, but a strong sex bias (94.2% correct for females, 49.3% correct for males) severely limits the utility of this method. The Walker (2008) and Stevenson et al. (2009) methods produced higher accuracy rates (80.8% and 82.6%, respectively), although these methods also produced sex and ancestry biases.

An Investigation into the Relationship between Human Cranial and Pelvic Sexual Dimorphism.

When faced with commingled remains, it might be assumed that a more "masculine" pelvis is associated with a more "masculine" cranium, but this relationship has not been specifically tested. This study uses geometric morphometric analyses of pelvic and cranial landmarks to assess whether there is an intra-individual relationship between the degrees of sexual expression in these two skeletal regions. Principal component and discriminant function scores were used to assess sexual dimorphism in 113 U.S. Black individuals. Correlation values and partial least squares regression (PLS) were used to evaluate intra-individual relationships. Results indicate that the os coxae is more sexually dimorphic than the cranium, with element shape being more sexually dimorphic than size. PLS and correlation results suggest no significant intra-individual relationship between pelvic and cranial sexual size or shape expression. Thus, in commingled situations, associations between these skeletal elements cannot be inferred based on degree of "masculinity."

Sexual dimorphism and regional variation in human frontal bone inclination measured via digital 3D models.

The frontal bone is one of the most sexually dimorphic elements of the human skull, due to features such as the glabella, frontal eminences, and frontal inclination. While glabella is frequently evaluated in procedures to estimate sex in unknown human skeletal remains, frontal inclination has received less attention. In this study we present a straightforward, quick, and reproducible method for measuring frontal inclination angles from glabella and supraglabella. Using a sample of 413 human crania from four different populations (U.S. Whites, U.S. Blacks, Portuguese, and Chinese), we test the usefulness of the inclination angles for sex estimation and compare their performance to traditional methods of frontal inclination assessment. Accuracy rates in the range 75-81% were achieved for the U.S. White, U.S. Black, and Portuguese groups. For Chinese the overall accuracy was lower, i.e. 66%. Although some regional variation was observed, a cut-off value of 78.2° for glabellar inclination angles separates female and male crania from all studied populations with good accuracy. As inclination angles measured from glabella captures two sexually dimorphic features (i.e. glabellar prominence and frontal inclination) in a single measure, the observed clear male/female difference is not unexpected. Being continuous variables, inclination angles are suitable for use in statistical methods for sex estimations.

Body size, brain size, and sexual dimorphism in Homo naledi from the Dinaledi Chamber.

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus, together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo, including Homo habilis, Homo rudolfensis, and some early H. erectus, suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus, with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates. H. naledi endocranial volumes (465-560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus, but similar to derived members of genus Homo, the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

New fossil remains of Homo naledi from the Lesedi Chamber, South Africa.

The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated from the Dinaledi Chamber within the Rising Star cave system, and represents a second depositional context for hominin remains. In each of three collection areas within the Lesedi Chamber, diagnostic skeletal material allows a clear attribution to H. naledi. Both adult and immature material is present. The hominin remains represent at least three individuals based upon duplication of elements, but more individuals are likely present based upon the spatial context. The most significant specimen is the near-complete cranium of a large individual, designated LES1, with an endocranial volume of approximately 610 ml and associated postcranial remains. The Lesedi Chamber skeletal sample extends our knowledge of the morphology and variation of H. naledi, and evidence of H. naledi from both recovery localities shows a consistent pattern of differentiation from other hominin species.

Skull diversity in the Homo lineage and the relative position of Homo naledi.

The discovery of Homo naledi has expanded the range of phenotypic variation in Homo, leading to new questions surrounding the mosaic nature of morphological evolution. Though currently undated, its unique morphological pattern and possible phylogenetic relationships to other hominin taxa suggest a complex evolutionary scenario. Here, we perform geometric morphometric analyses on H. naledi cranial and mandibular remains to investigate its morphological relationship with species of Homo and Australopithecus. We use Generalized Procrustes analysis to place H. naledi within the pattern of known hominin skull diversity, distributions of Procrustes distances among individuals to compare H. naledi and Homo erectus, and neighbor joining trees to investigate the potential phenetic relationships between groups. Our goal is to address a set of hypotheses relating to the uniqueness of H. naledi, its affinity with H. erectus, and the age of the fossils based on skull morphology. Our results indicate that, cranially, H. naledi aligns with members of the genus Homo, with closest affiliations to H. erectus. The mandibular results are less clear; H. naledi closely associates with a number of taxa, including some australopiths. However, results also show that although H. naledi shares similarities with H. erectus, some distances from this taxon - especially small-brained members of this taxon - are extreme. The neighbor joining trees place H. naledi firmly within Homo. The trees based on cranial morphology again indicate a close relationship between H. naledi and H. erectus, whereas the mandibular tree places H. naledi closer to basal Homo, suggesting a deeper antiquity. Altogether, these results emphasize the unique combination of features (H. erectus-like cranium, less derived mandible) defining H. naledi. Our results also highlight the variability within Homo, calling for a greater focus on the cause of this variability, and emphasizing the importance of using the total morphological package for species diagnoses.