Publications by authors named "William L Jungers"

39 Publications

Complete permanent mandibular dentition of early Homo from the upper Burgi Member of the Koobi Fora Formation, Ileret, Kenya.

J Hum Evol 2019 06 16;131:152-175. Epub 2019 Apr 16.

Turkana Basin Institute, Stony Brook University, Stony Brook, NY 11794, USA.

The KNM-ER 64060 dentition derives from a horizon that most likely dates to between 2.02 and 2.03 Ma. A proximate series of postcranial bones (designated KNM-ER 64061) derives from the same siltstone unit and may be associated with the dentition, but their separation on the surface of the site leaves some room for doubt. KNM-ER 64060 is one of fewer than ten hominin specimens from the Early Pleistocene of East Africa that comprises a full or nearly complete mandibular dentition. Its taxonomic attribution is potentially significant, especially if the postcranial elements are related. At least three, and probably four hominin species, including Paranthropus boisei and Homo erectus (= H. ergaster), are known at about this time in East Africa. Other penecontemporaneous fossils have been referred to a single, highly variable species, H. habilis, or two taxa, namely H. habilis and H. rudolfensis. Although the weight of evidence supports the attribution of these specimens to two species, there is notable lack of agreement over the assignation of individual fossils. We take a conservative approach and group all such specimens under the designation "early Homo sp." for comparative purposes. KNM-ER 64060 is clearly attributable to Homo rather than Paranthropus. The preponderance of the evidence suggests that the affinities of KNM-ER 64060 are with fossils assigned to the early Homo sp. category rather than with H. erectus. This is indicated by the overall sizes of the KNM-ER 64060 canine, premolar and molar crowns, the size relationships of the P to P, the relative narrowness of its premolar crowns, the cusp proportions of the M and especially those of the M and M, and seemingly the possession of a two-rooted P. Some of these comparisons suggest further that among the fossils comprising the early Homo sp. sample, the KNM-ER 64060 dentition exhibits greater overall similarity to specimens such as OH 7 and OH 16 that represent Homo habilis sensu stricto.
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http://dx.doi.org/10.1016/j.jhevol.2019.03.017DOI Listing
June 2019

Evolution and function of the hominin forefoot.

Proc Natl Acad Sci U S A 2018 08 13;115(35):8746-8751. Epub 2018 Aug 13.

Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794.

The primate foot functions as a grasping organ. As such, its bones, soft tissues, and joints evolved to maximize power and stability in a variety of grasping configurations. Humans are the obvious exception to this primate pattern, with feet that evolved to support the unique biomechanical demands of bipedal locomotion. Of key functional importance to bipedalism is the morphology of the joints at the forefoot, known as the metatarsophalangeal joints (MTPJs), but a comprehensive analysis of hominin MTPJ morphology is currently lacking. Here we present the results of a multivariate shape and Bayesian phylogenetic comparative analyses of metatarsals (MTs) from a broad selection of anthropoid primates (including fossil apes and stem catarrhines) and most of the early hominin pedal fossil record, including the oldest hominin for which good pedal remains exist, Results corroborate the importance of specific bony morphologies such as dorsal MT head expansion and "doming" to the evolution of terrestrial bipedalism in hominins. Further, our evolutionary models reveal that the MT1 of shifts away from the reconstructed optimum of our last common ancestor with apes, but not necessarily in the direction of modern humans. However, the lateral rays of are transformed in a more human-like direction, suggesting that they were the digits first recruited by hominins into the primary role of terrestrial propulsion. This pattern of evolutionary change is seen consistently throughout the evolution of the foot, highlighting the mosaic nature of pedal evolution and the emergence of a derived, modern hallux relatively late in human evolution.
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http://dx.doi.org/10.1073/pnas.1800818115DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6126759PMC
August 2018

Body mass estimates of the earliest possible hominins and implications for the last common ancestor.

J Hum Evol 2018 09 15;122:84-92. Epub 2018 Jun 15.

Association Vahatra, BP 3972, Antananarivo 101, Madagascar; Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USA.

Many hypotheses regarding the paleobiology of the earliest possible hominins, Orrorin tugenensis and Ardipithecus ramidus, are dependent upon accurate body mass estimates for these taxa. While we have previously published body mass predictions for Orrorin and Ardipithecus, the accuracies of those estimates depend on the assumption that the postcranial skeletal dimensions and body masses of these taxa followed scaling patterns that were similar to those observed in modern humans. This assumption may not be correct because certain aspects of postcranial morphology in Orrorin and Ardipithecus differ from modern humans, and suggest that their overall body plans might be unique but more similar to modern non-human great apes than to modern humans. Here we present individual body mass predictions for O. tugenensis and Ar. ramidus assuming that they followed postcranial scaling patterns similar to those of chimpanzees. All estimates include individual prediction intervals as measures of uncertainty. In addition, we provide equations for predicting body mass from univariate postcranial measurements based on the largest sample (n = 25) yet compiled of common chimpanzee skeletons with known body masses, which is vital for calculating prediction intervals for individual fossils. Our results show that estimated body masses in Orrorin and Ardipithecus are generally larger when derived from a chimpanzee-like scaling pattern compared to estimates that assume a human-like pattern, though the prediction intervals of the two sets of estimates overlap. In addition, the more complete of the two known Orrorin femora has an overall scaling pattern that is more similar to common chimpanzees than to modern humans, supporting the application of a non-human great ape comparative model. Our new estimates fall near the male (Ardipithecus) average and in between the male and female averages (Orrorin) for wild-caught common chimpanzees. If a chimpanzee-like pattern of scaling between postcranial dimensions and body mass did exist in these earliest hominins, our results suggest the large body masses found in some early australopiths were already present in taxa near the origins of our lineage, and perhaps also in the Pan-Homo last common ancestor.
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http://dx.doi.org/10.1016/j.jhevol.2018.05.001DOI Listing
September 2018

Evidence of a chimpanzee-sized ancestor of humans but a gibbon-sized ancestor of apes.

Nat Commun 2017 10 12;8(1):880. Epub 2017 Oct 12.

Association Vahatra, Antananarivo 101, BP, 3972, Madagascar.

Body mass directly affects how an animal relates to its environment and has a wide range of biological implications. However, little is known about the mass of the last common ancestor (LCA) of humans and chimpanzees, hominids (great apes and humans), or hominoids (all apes and humans), which is needed to evaluate numerous paleobiological hypotheses at and prior to the root of our lineage. Here we use phylogenetic comparative methods and data from primates including humans, fossil hominins, and a wide sample of fossil primates including Miocene apes from Africa, Europe, and Asia to test alternative hypotheses of body mass evolution. Our results suggest, contrary to previous suggestions, that the LCA of all hominoids lived in an environment that favored a gibbon-like size, but a series of selective regime shifts, possibly due to resource availability, led to a decrease and then increase in body mass in early hominins from a chimpanzee-sized LCA.The pattern of body size evolution in hominids can provide insight into historical human ecology. Here, Grabowski and Jungers use comparative phylogenetic analysis to reconstruct the likely size of the ancestor of humans and chimpanzees and the evolutionary history of selection on body size in primates.
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http://dx.doi.org/10.1038/s41467-017-00997-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5638852PMC
October 2017

Comment on relative brain size in early primates and the use of encephalization quotients in primate evolution.

J Hum Evol 2017 08 3;109:79-87. Epub 2017 Jun 3.

Department of Anatomical Sciences, School of Medicine, Stony Brook University, Stony Brook, NY 11794, USA; Association Vahatra, Antananarivo 101, Madagascar.

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http://dx.doi.org/10.1016/j.jhevol.2017.04.007DOI Listing
August 2017

The affinities of Homo floresiensis based on phylogenetic analyses of cranial, dental, and postcranial characters.

J Hum Evol 2017 06 21;107:107-133. Epub 2017 Apr 21.

Department of Anatomical Sciences, Stony Brook University Medical Center, Stony Brook, NY, USA; Association Vahatra, BP 3972, Antananarivo 101, Madagascar.

Although the diminutive Homo floresiensis has been known for a decade, its phylogenetic status remains highly contentious. A broad range of potential explanations for the evolution of this species has been explored. One view is that H. floresiensis is derived from Asian Homo erectus that arrived on Flores and subsequently evolved a smaller body size, perhaps to survive the constrained resources they faced in a new island environment. Fossil remains of H. erectus, well known from Java, have not yet been discovered on Flores. The second hypothesis is that H. floresiensis is directly descended from an early Homo lineage with roots in Africa, such as Homo habilis; the third is that it is Homo sapiens with pathology. We use parsimony and Bayesian phylogenetic methods to test these hypotheses. Our phylogenetic data build upon those characters previously presented in support of these hypotheses by broadening the range of traits to include the crania, mandibles, dentition, and postcrania of Homo and Australopithecus. The new data and analyses support the hypothesis that H. floresiensis is an early Homo lineage: H. floresiensis is sister either to H. habilis alone or to a clade consisting of at least H. habilis, H. erectus, Homo ergaster, and H. sapiens. A close phylogenetic relationship between H. floresiensis and H. erectus or H. sapiens can be rejected; furthermore, most of the traits separating H. floresiensis from H. sapiens are not readily attributable to pathology (e.g., Down syndrome). The results suggest H. floresiensis is a long-surviving relict of an early (>1.75 Ma) hominin lineage and a hitherto unknown migration out of Africa, and not a recent derivative of either H. erectus or H. sapiens.
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http://dx.doi.org/10.1016/j.jhevol.2017.02.006DOI Listing
June 2017

These feet were made for walking.

Elife 2016 12 14;5. Epub 2016 Dec 14.

Department of Anatomical Sciences, Stony Brook University School of Medicine, New York, United States.

New fossil footprints excavated at the famous Laetoli site in Tanzania suggest that our bipedal ancestors had a wide range of body sizes.
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http://dx.doi.org/10.7554/eLife.22886DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5156523PMC
December 2016

Form and function of the human and chimpanzee forefoot: implications for early hominin bipedalism.

Sci Rep 2016 07 28;6:30532. Epub 2016 Jul 28.

Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USA.

During bipedal walking, modern humans dorsiflex their forefoot at the metatarsophalangeal joints (MTPJs) prior to push off, which tightens the plantar soft tissues to convert the foot into a stiff propulsive lever. Particular features of metatarsal head morphology such as "dorsal doming" are thought to facilitate this stiffening mechanism. In contrast, chimpanzees are believed to possess MTPJ morphology that precludes high dorsiflexion excursions during terrestrial locomotion. The morphological affinity of the metatarsal heads has been used to reconstruct locomotor behavior in fossil hominins, but few studies have provided detailed empirical data to validate the assumed link between morphology and function at the MTPJs. Using three-dimensional kinematic and morphometric analyses, we show that humans push off with greater peak dorsiflexion angles at all MTPJs than do chimpanzees during bipedal and quadrupedal walking, with the greatest disparity occurring at MTPJ 1. Among MTPJs 2-5, both species exhibit decreasing peak angles from medial to lateral. This kinematic pattern is mirrored in the morphometric analyses of metatarsal head shape. Analyses of Australopithecus afarensis metatarsals reveal morphology intermediate between humans and chimpanzees, suggesting that this species used different bipedal push-off kinematics than modern humans, perhaps resulting in a less efficient form of bipedalism.
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http://dx.doi.org/10.1038/srep30532DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4964565PMC
July 2016

The evolution of body size and shape in the human career.

Philos Trans R Soc Lond B Biol Sci 2016 07;371(1698)

Division of Anthropology, American Museum of Natural History, New York, NY 10024, USA Department of Human Evolution, Max Plank Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.

Body size is a fundamental biological property of organisms, and documenting body size variation in hominin evolution is an important goal of palaeoanthropology. Estimating body mass appears deceptively simple but is laden with theoretical and pragmatic assumptions about best predictors and the most appropriate reference samples. Modern human training samples with known masses are arguably the 'best' for estimating size in early bipedal hominins such as the australopiths and all members of the genus Homo, but it is not clear if they are the most appropriate priors for reconstructing the size of the earliest putative hominins such as Orrorin and Ardipithecus The trajectory of body size evolution in the early part of the human career is reviewed here and found to be complex and nonlinear. Australopith body size varies enormously across both space and time. The pre-erectus early Homo fossil record from Africa is poor and dominated by relatively small-bodied individuals, implying that the emergence of the genus Homo is probably not linked to an increase in body size or unprecedented increases in size variation. Body size differences alone cannot explain the observed variation in hominin body shape, especially when examined in the context of small fossil hominins and pygmy modern humans.This article is part of the themed issue 'Major transitions in human evolution'.
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http://dx.doi.org/10.1098/rstb.2015.0247DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4920302PMC
July 2016

Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia.

Nature 2016 Apr 30;532(7599):366-9. Epub 2016 Mar 30.

Centre for Archaeological Science, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia.

Homo floresiensis, a primitive hominin species discovered in Late Pleistocene sediments at Liang Bua (Flores, Indonesia), has generated wide interest and scientific debate. A major reason this taxon is controversial is because the H. floresiensis-bearing deposits, which include associated stone artefacts and remains of other extinct endemic fauna, were dated to between about 95 and 12 thousand calendar years (kyr) ago. These ages suggested that H. floresiensis survived until long after modern humans reached Australia by ~50 kyr ago. Here we report new stratigraphic and chronological evidence from Liang Bua that does not support the ages inferred previously for the H. floresiensis holotype (LB1), ~18 thousand calibrated radiocarbon years before present (kyr cal. BP), or the time of last appearance of this species (about 17 or 13-11 kyr cal. BP). Instead, the skeletal remains of H. floresiensis and the deposits containing them are dated to between about 100 and 60 kyr ago, whereas stone artefacts attributable to this species range from about 190 to 50 kyr in age. Whether H. floresiensis survived after 50 kyr ago--potentially encountering modern humans on Flores or other hominins dispersing through southeast Asia, such as Denisovans--is an open question.
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http://dx.doi.org/10.1038/nature17179DOI Listing
April 2016

The evolutionary origin and population history of the grauer gorilla.

Am J Phys Anthropol 2016 Jan;159(Suppl 61):S4-S18

Department of Anatomical Sciences, Stony Brook University Medical Center, Stony Brook, NY, 11794, USA.

Gorillas living in western central Africa (Gorilla gorilla) are morphologically and genetically distinguishable from those living in eastern central Africa (Gorilla beringei). Genomic analyses show eastern gorillas experienced a significant reduction in population size during the Pleistocene subsequent to geographical isolation from their western counterparts. However, how these results relate more specifically to the recent biogeographical and evolutionary history of eastern gorillas remains poorly understood. Here we show that two rare morphological traits are present in the hands and feet of both eastern gorilla subspecies at strikingly high frequencies (>60% in G. b. graueri; ∼28% in G. b. beringei) in comparison with western gorillas (<1%). The intrageneric distribution of these rare traits suggests that they became common among eastern gorillas after diverging from their western relatives during the early to middle Pleistocene. The extremely high frequencies observed among grauer gorillas-which currently occupy a geographic range more than ten times the size of that of mountain gorillas-imply that grauers originated relatively recently from a small founding population of eastern gorillas. Current paleoenvironmental, geological, and biogeographical evidence supports the hypothesis that a small group of eastern gorillas likely dispersed westward from the Virungas into present-day grauer range in the highlands just north of Lake Kivu, either immediately before or directly after the Younger Dryas interval. We propose that as the lowland forests of central Africa expanded rapidly during the early Holocene, they became connected with the expanding highland forests along the Albertine Rift and enabled the descendants of this small group to widely disperse. The descendant populations significantly expanded their geographic range and population numbers relative to the gorillas of the Virunga Mountains and the Bwindi-Impenetrable Forest, ultimately resulting in the grauer gorilla subspecies recognized today. This founder-effect hypothesis offers some optimism for modern conservation efforts to save critically endangered eastern gorillas from extinction.
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http://dx.doi.org/10.1002/ajpa.22900DOI Listing
January 2016

Functional aspects of metatarsal head shape in humans, apes, and Old World monkeys.

J Hum Evol 2015 Sep 12;86:136-46. Epub 2015 Aug 12.

Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USA.

Modern human metatarsal heads are typically described as "dorsally domed," mediolaterally wide, and dorsally flat. Despite the apparent functional importance of these features in forefoot stability during bipedalism, the distinctiveness of this morphology has not been quantitatively evaluated within a broad comparative framework. In order to use these features to reconstruct fossil hominin locomotor behaviors with any confidence, their connection to human bipedalism should be validated through a comparative analysis of other primates with different locomotor behaviors and foot postures, including species with biomechanical demands potentially similar to those of bipedalism (e.g., terrestrial digitigrady). This study explores shape variation in the distal metatarsus among humans and other extant catarrhines using three-dimensional geometric morphometrics (3 DGM). Shape differences among species in metatarsal head morphology are well captured by the first two principal components of Procrustes shape coordinates, and these two components summarize most of the variance related to "dorsal doming" and "dorsal expansion." Multivariate statistical tests reveal significant differences among clades in overall shape, and humans are reliably distinguishable from other species by aspects of shape related to a greater degree of dorsal doming. Within quadrupeds, terrestrial species also trend toward more domed metatarsal heads, but not to the extent seen in humans. Certain aspects of distal metatarsus shape are likely related to habitual dorsiflexion of the metatarsophalangeal joints, but the total morphological pattern seen in humans is distinct. These comparative results indicate that this geometric morphometric approach is useful to characterize the complexity of metatarsal head morphology and will help clarify its relationship with function in fossil primates, including early hominins.
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http://dx.doi.org/10.1016/j.jhevol.2015.06.005DOI Listing
September 2015

The evolution of human and ape hand proportions.

Nat Commun 2015 Jul 14;6:7717. Epub 2015 Jul 14.

Department of Anatomical Sciences, Stony Brook University, Stony Brook, New York 11794, USA.

Human hands are distinguished from apes by possessing longer thumbs relative to fingers. However, this simple ape-human dichotomy fails to provide an adequate framework for testing competing hypotheses of human evolution and for reconstructing the morphology of the last common ancestor (LCA) of humans and chimpanzees. We inspect human and ape hand-length proportions using phylogenetically informed morphometric analyses and test alternative models of evolution along the anthropoid tree of life, including fossils like the plesiomorphic ape Proconsul heseloni and the hominins Ardipithecus ramidus and Australopithecus sediba. Our results reveal high levels of hand disparity among modern hominoids, which are explained by different evolutionary processes: autapomorphic evolution in hylobatids (extreme digital and thumb elongation), convergent adaptation between chimpanzees and orangutans (digital elongation) and comparatively little change in gorillas and hominins. The human (and australopith) high thumb-to-digits ratio required little change since the LCA, and was acquired convergently with other highly dexterous anthropoids.
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http://dx.doi.org/10.1038/ncomms8717DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4510966PMC
July 2015

Body mass estimates of hominin fossils and the evolution of human body size.

J Hum Evol 2015 Aug 17;85:75-93. Epub 2015 Jun 17.

Division of Anthropology, American Museum of Natural History, New York 10024, USA.

Body size directly influences an animal's place in the natural world, including its energy requirements, home range size, relative brain size, locomotion, diet, life history, and behavior. Thus, an understanding of the biology of extinct organisms, including species in our own lineage, requires accurate estimates of body size. Since the last major review of hominin body size based on postcranial morphology over 20 years ago, new fossils have been discovered, species attributions have been clarified, and methods improved. Here, we present the most comprehensive and thoroughly vetted set of individual fossil hominin body mass predictions to date, and estimation equations based on a large (n = 220) sample of modern humans of known body masses. We also present species averages based exclusively on fossils with reliable taxonomic attributions, estimates of species averages by sex, and a metric for levels of sexual dimorphism. Finally, we identify individual traits that appear to be the most reliable for mass estimation for each fossil species, for use when only one measurement is available for a fossil. Our results show that many early hominins were generally smaller-bodied than previously thought, an outcome likely due to larger estimates in previous studies resulting from the use of large-bodied modern human reference samples. Current evidence indicates that modern human-like large size first appeared by at least 3-3.5 Ma in some Australopithecus afarensis individuals. Our results challenge an evolutionary model arguing that body size increased from Australopithecus to early Homo. Instead, we show that there is no reliable evidence that the body size of non-erectus early Homo differed from that of australopiths, and confirm that Homo erectus evolved larger average body size than earlier hominins.
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http://dx.doi.org/10.1016/j.jhevol.2015.05.005DOI Listing
August 2015

Exploring phylogenetic and functional signals in complex morphologies: the hamate of extant anthropoids as a test-case study.

Anat Rec (Hoboken) 2015 Jan;298(1):212-29

Department of Anatomical Sciences, Stony Brook University School of Medicine, Stony Brook, New York; Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici Z (ICTA-ICP), campus de la UAB, c/ de les Columnes, s/n., 08193 Cerdanyola del Vallès, Barcelona, Spain; NYCEP Morphometrics Group.

Three-dimensional geometric morphometrics (3DGM) is a powerful tool for capturing and visualizing the "pure" shape of complex structures. However, these shape differences are sometimes difficult to interpret from a functional viewpoint, unless specific approaches (mostly based on biomechanical modeling) are employed. Here, we use 3DGM to explore the complex shape variation of the hamate, the disto-ulnar wrist bone, in anthropoid primates. Major trends of shape variation are explored using principal components analysis along with analyses of shape and size covariation. We also evaluate the phylogenetic patterning of hamate shape by plotting an anthropoid phylogenetic tree onto the shape space (i.e., phylomorphospace) and test against complete absence of phylogenetic signal using posterior permutation. Finally, the covariation of hamate shape and locomotor categories is explored by means of 2-block partial least squares (PLS) using shape coordinates and a matrix of data on arboreal locomotor behavior. Our results show that 3DGM is a valuable and versatile tool for characterizing the shape of complex structures such as wrist bones in anthropoids. For the hamate, a significant phylogenetic pattern is found in both hamate shape and size, indicating that closely related taxa are typically the most similar in hamate form. Our allometric analyses show that major differences in hamate shape among taxa are not a direct consequence of differences in hamate size. Finally, our PLS indicates a significant covariation of hamate shape and different types of arboreal locomotion, highlighting the relevance of this approach in future 3DGM studies seeking to capture a functional signal from complex biological structures.
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http://dx.doi.org/10.1002/ar.23079DOI Listing
January 2015

Geometric properties and comparative biomechanics of Homo floresiensis mandibles.

J Hum Evol 2014 Mar 21;68:36-46. Epub 2014 Feb 21.

Department of Anatomical Sciences, School of Medicine, Stony Brook University, Stony Brook, NY 11794, USA.

The hypodigm of Homo floresiensis from the cave of Liang Bua on Flores Island in the archipelago of Indonesia includes two mandibles (LB1/2 and LB6/1). The morphology of their symphyses and corpora has been described as sharing similarities with both australopiths and early Homo despite their Late Pleistocene age. Although detailed morphological comparisons of these mandibles with those of modern and fossil hominin taxa have been made, a functional analysis in the context of masticatory biomechanics has yet to be performed. Utilizing data on cortical bone geometry from computed tomography scans, we compare the mechanical attributes of the LB1 and LB6 mandibles with samples of modern Homo, Pan, Pongo, and Gorilla, as well as fossil samples of Paranthropus robustus, Australopithecus africanus and South African early Homo. Structural stiffness measures were derived from the geometric data to provide relative measures of mandibular corpus strength under hypothesized masticatory loading regimes. These mechanical variables were evaluated relative to bone area, mandibular length and estimates of body size to assess their functional affinities and to test the hypothesis that the Liang Bua mandibles can be described as scaled-down variants of either early hominins or modern humans. Relative to modern hominoids, the H. floresiensis material appears to be relatively strong in terms of rigidity in torsion and transverse bending, but is relatively weak under parasagittal bending. Thus, they are 'robust' relative to modern humans (and comparable with australopiths) under some loads but not others. Neither LB1 nor LB6 can be described simply as 'miniaturized' versions of modern human jaws since mandible length is more or less equivalent in Homo sapiens and H. floresiensis. The mechanical attributes of the Liang Bua mandibles are consistent with previous inferences that masticatory loads were reduced relative to australopiths but remained elevated relative to modern Homo.
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http://dx.doi.org/10.1016/j.jhevol.2014.01.001DOI Listing
March 2014

Phylogenetic, ecological, and allometric correlates of cranial shape in Malagasy lemuriforms.

Evolution 2014 May 4;68(5):1450-68. Epub 2014 Mar 4.

Department of Anthropology, Stony Brook University, Social and Behavioral Sciences Building, 5th Floor, Stony Brook, New York, 11794; Interdepartmental Program in Anthropological Sciences, Stony Brook University, Stony Brook, New York, 11794.

Adaptive radiations provide important insights into many aspects of evolution, including the relationship between ecology and morphological diversification as well as between ecology and speciation. Many such radiations include divergence along a dietary axis, although other ecological variables may also drive diversification, including differences in diel activity patterns. This study examines the role of two key ecological variables, diet and activity patterns, in shaping the radiation of a diverse clade of primates, the Malagasy lemurs. When phylogeny was ignored, activity pattern and several dietary variables predicted a significant proportion of cranial shape variation. However, when phylogeny was taken into account, only typical diet accounted for a significant proportion of shape variation. One possible explanation for this discrepancy is that this radiation was characterized by a relatively small number of dietary shifts (and possibly changes in body size) that occurred in conjunction with the divergence of major clades. This pattern may be difficult to detect with the phylogenetic comparative methods used here, but may characterize not just lemurs but other mammals.
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http://dx.doi.org/10.1111/evo.12361DOI Listing
May 2014

Ecological divergence and talar morphology in gorillas.

Am J Phys Anthropol 2014 Apr 21;153(4):526-41. Epub 2013 Dec 21.

Department of Anatomy, Des Moines University, Des Moines, IA, 50312.

Gorillas occupy a variety of habitats from the west coast to eastern central Africa. These habitats differ considerably in altitude, which has a pronounced effect on forest ecology. Although all gorillas are obligate terrestrial knuckle-walking quadrupeds, those that live in lowland habitats eat fruits and climb more often than do those living in highland habitats. Here we test the hypothesis that gorilla talus morphology falls along a morphocline that tracks locomotor function related to a more inverted or everted foot set. This proposed morphocline predicts that gorillas living in lowland habitats may have a talocrural joint configured to facilitate a more medially oriented foot during climbing, suggesting that they may be more adaptively committed to arboreality than gorillas living in highland habitats. To quantify the relative set of the foot in gorillas, we chose two three-dimensional measurements of the talocrural joint: mediolateral curvature of the trochlea and relative surface area of the lateral malleolus. Our results show that, in comparison to their eastern counterparts, western gorillas have talar features that reflect a more medially directed sole of the foot. This morphology likely facilitates foot placement in a wider range of positions and minimization of shearing stresses across the joint when the foot is loaded on more curved or vertically oriented substrates as occurs during climbing and other arboreal behaviors. In contrast, eastern gorilla talar morphology is consistent with habitual placement of the foot with the sole directed more inferiorly, suggesting more effective loading during plantigrade push-off on terrestrial substrates.
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http://dx.doi.org/10.1002/ajpa.22451DOI Listing
April 2014

The femur of Orrorin tugenensis exhibits morphometric affinities with both Miocene apes and later hominins.

Nat Commun 2013 ;4:2888

1] Department of Anatomical Sciences, Stony Brook University Medical Center, Stony Brook, New York 11794-8081, USA [2] Department of Vertebrate Paleontology, American Museum of Natural History and NYCEP, 79 St and Central Park West, New York, New York 10024, USA [3] Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona. Edifici ICP, Campus de la UAB s/n, 08193 Cerdanyola del Vallès, Barcelona, Spain.

Orrorin tugenensis (Kenya, ca. 6 Ma) is one of the earliest putative hominins. Its proximal femur, BAR 1002'00, was originally described as being very human-like, although later multivariate analyses showed an australopith pattern. However, some of its traits (for example, laterally protruding greater trochanter, medially oriented lesser trochanter and presence of third trochanter) are also present in earlier Miocene apes. Here, we use geometric morphometrics to reassess the morphological affinities of BAR 1002'00 within a large sample of anthropoids (including fossil apes and hominins) and reconstruct hominoid proximal femur evolution using squared-change parsimony. Our results indicate that both hominin and modern great ape femora evolved in different directions from a primitive morphology represented by some fossil apes. Orrorin appears intermediate between Miocene apes and australopiths in shape space. This evidence is consistent with femoral shape similarities in extant great apes being derived and homoplastic and has profound implications for understanding the origins of human bipedalism.
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http://dx.doi.org/10.1038/ncomms3888DOI Listing
July 2014

New wrist bones of Homo floresiensis from Liang Bua (Flores, Indonesia).

J Hum Evol 2013 Feb 4;64(2):109-29. Epub 2013 Jan 4.

Department of Anatomical Sciences, Stony Brook University, Health Sciences Center T-8 040, Stony Brook, NY 11794-8081, USA.

The carpals from the Homo floresiensis type specimen (LB1) lack features that compose the shared, derived complex of the radial side of the wrist in Neandertals and modern humans. This paper comprises a description and three-dimensional morphometric analysis of new carpals from at least one other individual at Liang Bua attributed to H. floresiensis: a right capitate and two hamates. The new capitate is smaller than that of LB1 but is nearly identical in morphology. As with capitates from extant apes, species of Australopithecus, and LB1, the newly described capitate displays a deeply-excavated nonarticular area along its radial aspect, a scaphoid facet that extends into a J-hook articulation on the neck, and a more radially-oriented second metacarpal facet; it also lacks an enlarged palmarly-positioned trapezoid facet. Because there is no accommodation for the derived, palmarly blocky trapezoid that characterizes Homo sapiens and Neandertals, this individual most likely had a plesiomorphically wedge-shaped trapezoid (like LB1). Morphometric analyses confirm the close similarity of the new capitate and that of LB1, and are consistent with previous findings of an overall primitive articular geometry. In general, hamate morphology is more conserved across hominins, and the H. floresiensis specimens fall at the far edge of the range of variation for H. sapiens in a number of metrics. However, the hamate of H. floresiensis is exceptionally small and exhibits a relatively long, stout hamulus lacking the oval-shaped cross-section characteristic of human and Neandertal hamuli (variably present in australopiths). Documentation of a second individual with primitive carpal anatomy from Liang Bua, along with further analysis of trapezoid scaling relative to the capitate in LB1, refutes claims that the wrist of the type specimen represents a modern human with pathology. In total, the carpal anatomy of H. floresiensis supports the hypothesis that the lineage leading to the evolution of this species originated prior to the cladogenetic event that gave rise to modern humans and Neandertals.
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http://dx.doi.org/10.1016/j.jhevol.2012.10.003DOI Listing
February 2013

On dental wear, dental work, and oral health in the type specimen (LB1) of Homo floresiensis.

Am J Phys Anthropol 2011 Jun 16;145(2):282-9. Epub 2011 Mar 16.

Department of Anatomical Sciences, Stony Brook University Medical Center, Stony Brook, NY 11794-8081, USA.

The claim that the lower left first mandibular molar of LB1, the type specimen of Homo floresiensis, displays endodontic work, and a filling is assessed by digital radiography and micro-CT scanning. The M(1) tooth crown is heavily worn and exhibits extensive dentine exposure that is stained white, but there is no trace of endodontic treatment or a dental filling in this Indonesian fossil dated to 17.1-19.0 kya. Dental calculus (commonly observed in foragers) is present on the teeth of LB1, but there are no observable caries. The pattern of dental attrition in the mandibles of both LB1/2 and LB6/1 (moderate to extensive flat wear across the entire arch) is consistent with that seen in Plio-Pleistocene Homo fossils and in modern hunter-gatherers, and is not typical of most agriculturalists. We conclude that the dental-work and farming hypotheses are falsified and therefore irrelevant to the debate over the taxonomy and phylogeny of H. floresiensis.
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http://dx.doi.org/10.1002/ajpa.21492DOI Listing
June 2011

Ecological divergence and medial cuneiform morphology in gorillas.

J Hum Evol 2011 Feb 19;60(2):171-84. Epub 2010 Nov 19.

Human Origins Program, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, 10th and Constitution Avenue NW, Washington, DC 20013-7012, USA.

Gorillas are more closely related to each other than to any other extant primate and are all terrestrial knuckle-walkers, but taxa differ along a gradient of dietary strategies and the frequency of arboreality in their behavioral repertoire. In this study, we test the hypothesis that medial cuneiform morphology falls on a morphocline in gorillas that tracks function related to hallucial abduction ability and relative frequency of arboreality. This morphocline predicts that western gorillas, being the most arboreal, should display a medial cuneiform anatomy that reflects the greatest hallucial abduction ability, followed by grauer gorillas, and then by mountain gorillas. Using a three-dimensional methodology to measure angles between articular surfaces, relative articular and nonarticular areas, and the curvatures of the hallucial articular surface, the functional predictions are partially confirmed in separating western gorillas from both eastern gorillas. Western gorillas are characterized by a more medially oriented, proportionately larger, and more mediolaterally curved hallucial facet than are eastern gorillas. These characteristics follow the predictions for a more prehensile hallux in western gorillas relative to a more stable, plantigrade hallux in eastern gorillas. The characteristics that distinguish eastern gorilla taxa from one another appear unrelated to hallucial abduction ability or frequency of arboreality. In total, this reexamination of medial cuneiform morphology suggests differentiation between eastern and western gorillas due to a longstanding ecological divergence and more recent and possibly non-adaptive differences between eastern taxa.
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http://dx.doi.org/10.1016/j.jhevol.2010.09.002DOI Listing
February 2011

Biomechanics: Barefoot running strikes back.

Nature 2010 Jan;463(7280):433-4

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http://dx.doi.org/10.1038/463433aDOI Listing
January 2010

"Life history space": a multivariate analysis of life history variation in extant and extinct Malagasy lemurs.

Am J Phys Anthropol 2010 Jul;142(3):391-404

School of Human Evolution and Social Change, Arizona State University, Tempe, AZ 85287, USA.

Studies of primate life history variation are constrained by the fact that all large-bodied extant primates are haplorhines. However, large-bodied strepsirrhines recently existed. If we can extract life history information from their skeletons, these species can contribute to our understanding of primate life history variation. This is particularly important in light of new critiques of the classic "fast-slow continuum" as a descriptor of variation in life history profiles across mammals in general. We use established dental histological methods to estimate gestation length and age at weaning for five extinct lemur species. On the basis of these estimates, we reconstruct minimum interbirth intervals and maximum reproductive rates. We utilize principal components analysis to create a multivariate "life history space" that captures the relationships among reproductive parameters and brain and body size in extinct and extant lemurs. Our data show that, whereas large-bodied extinct lemurs can be described as "slow" in some fashion, they also varied greatly in their life history profiles. Those with relatively large brains also weaned their offspring late and had long interbirth intervals. These were not the largest of extinct lemurs. Thus, we distinguish size-related life history variation from variation that linked more strongly to ecological factors. Because all lemur species larger than 10 kg, regardless of life history profile, succumbed to extinction after humans arrived in Madagascar, we argue that large body size increased the probability of extinction independently of reproductive rate. We also provide some evidence that, among lemurs, brain size predicts reproductive rate better than body size.
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http://dx.doi.org/10.1002/ajpa.21236DOI Listing
July 2010

Modern humans are not (quite) isometric.

Am J Phys Anthropol 2008 Dec;137(4):371-83

Department of Mechanical, Aerospace and Biomedical Engineering, The University of Tennessee, Knoxville, TN 37996, USA.

Allometric relationships are important sources of information for many types of anthropological and biological research. The baseline for all allometric relationships is isometry (or geometric similarity), the principal that shape is invariant of size. Here, we formally test for geometric similarity in modern humans, looking at the maximum lengths of four long bones (humerus, radius, femur, and tibia). We use Jolicoeur's multivariate allometry method to examine globally distributed samples of human populations, both collectively and individually. Results indicate that humans are not geometrically similar, although morphological deviations from isometry are small.
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http://dx.doi.org/10.1002/ajpa.20880DOI Listing
December 2008

Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism.

Science 2008 Mar;319(5870):1662-5

Center for the Advanced Study of Hominid Paleobiology, Department of Anthropology, George Washington University, 2110 G Street, NW, Washington, DC 20052, USA.

Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late Pliocene in early Homo.
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http://dx.doi.org/10.1126/science.1154197DOI Listing
March 2008

The bipedalism of the Dmanisi hominins: pigeon-toed early Homo?

Am J Phys Anthropol 2008 Aug;136(4):375-8

Interdepartmental Doctoral Program in Anthropological Sciences, Stony Brook University, Stony Brook, NY 11794, USA.

In the recent description of the hominin postcranial material from Dmanisi, Georgia, Lordkipanidze and colleagues (Lordkipanidze et al. [2007] Nature 449: 305-310) claim that the Dmanisi hominins walked with more medially oriented feet than do modern humans. They draw this functional inference from two postcranial features: a wide talar neck angle and a slight medial torsion of the tibia. However, we believe that the data provided by the authors fail to support their conclusions. Talar neck angle and tibial torsion values from the Dmanisi specimens fall comfortably within the range of modern human variation. We further submit that foot orientation cannot be reliably deduced from the tibia and talus alone.
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http://dx.doi.org/10.1002/ajpa.20827DOI Listing
August 2008

New hand bones of Hadropithecus stenognathus: implications for the paleobiology of the Archaeolemuridae.

J Hum Evol 2008 Mar;54(3):405-13

Division of Anatomy, Faculty of Medicine and Dentistry, University of Alberta, Edmonton, Alberta, Canada, T6G 2H7.

A partial, associated skeleton of Hadropithecus stenognathus (AHA-I) was discovered in 2003 at Andrahomana Cave in southeastern Madagascar. Among the postcranial elements found were the first hand bones (right scaphoid, right hamate, left first metacarpal, and right and left fifth metacarpals) attributed to this rare subfossil lemur. These hand bones were compared to those of extant strepsirrhines and catarrhines in order to infer the positional adaptations of Hadropithecus, and they were compared to those of Archaeolemur in order to assess variation in hand morphology among archaeolemurids. The scaphoid tubercle does not project palmarly as in suspensory and climbing taxa, and the hamate has no hook at all (just a small tubercle), which also points to a poorly developed carpal tunnel. There is a distinctive, radioulnarly directed "spiral" facet for articulation with the triquetrum that is most similar in orientation to that of more terrestrial primates (i.e., Lemur catta, Papio, and Gorilla). The first metacarpal is very reduced and represents only 48% of the length of metacarpal V, as in Archaeolemur, which suggests that pollical grasping of arboreal supports was not important. Compared to Archaeolemur, the shaft of metacarpal V is gracile, and the head has no dorsal ridge and lacks characteristics functionally associated with digitigrade, extended metacarpophalangeal joint postures. Proximally, the articular facet for the hamate is oriented more dorsally. Thus, the carpometacarpal joint V appears to have a distinctive hyperextended set, which has no analog among living or extinct primates. The carpals of Hadropithecus are diagnostic of a pronograde, arboreal and terrestrial (although not digitigrade) locomotor repertoire that typifies Lemur catta and some Old World monkeys. No clinging, suspensory, or climbing specializations that characterize indriids or lorises can be found in the hand of this subfossil lemur. The hand of Hadropithecus likely had similar ranges of movement at the radiocarpal and midcarpal joints as of those of pronograde primates, such as lemurids, for which the hand is held in a more extended, pronated, and neutral (i.e., showing less ulnar deviation) position during locomotion in comparison to that of vertical clingers or slow climbers. Although highly autapomorphic, the hand of Hadropithecus resembles that of its sister taxon, Archaeolemur, in having a very reduced pollex and an articular facet on the scaphoid for a sizeable prepollex. These unusual hand features reinforce the monophyly of the Archaeolemuridae.
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http://dx.doi.org/10.1016/j.jhevol.2007.09.010DOI Listing
March 2008

The primitive wrist of Homo floresiensis and its implications for hominin evolution.

Science 2007 Sep;317(5845):1743-5

Human Origins Program, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA.

Whether the Late Pleistocene hominin fossils from Flores, Indonesia, represent a new species, Homo floresiensis, or pathological modern humans has been debated. Analysis of three wrist bones from the holotype specimen (LB1) shows that it retains wrist morphology that is primitive for the African ape-human clade. In contrast, Neandertals and modern humans share derived wrist morphology that forms during embryogenesis, which diminishes the probability that pathology could result in the normal primitive state. This evidence indicates that LB1 is not a modern human with an undiagnosed pathology or growth defect; rather, it represents a species descended from a hominin ancestor that branched off before the origin of the clade that includes modern humans, Neandertals, and their last common ancestor.
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http://dx.doi.org/10.1126/science.1147143DOI Listing
September 2007

Homo floresiensis and the evolution of the hominin shoulder.

J Hum Evol 2007 Dec 13;53(6):718-31. Epub 2007 Aug 13.

Anatomical Sciences, Stony Brook University School of Medicine, Stony Brook, NY, USA.

The holotype of Homo floresiensis, diminutive hominins with tiny brains living until 12,000 years ago on the island of Flores, is a partial skeleton (LB1) that includes a partial clavicle (LB1/5) and a nearly complete right humerus (LB1/50). Although the humerus appears fairly modern in most regards, it is remarkable in displaying only 110 degrees of humeral torsion, well below modern human average values. Assuming a modern human shoulder configuration, such a low degree of humeral torsion would result in a lateral set to the elbow. Such an elbow joint would function more nearly in a frontal than in a sagittal plane, and this is certainly not what anyone would have predicted for a tool-making Pleistocene hominin. We argue that Homo floresiensis probably did not have a modern human shoulder configuration: the clavicle was relatively short, and we suggest that the scapula was more protracted, resulting in a glenoid fossa that faced anteriorly rather than laterally. A posteriorly directed humeral head was therefore appropriate for maintaining a normally functioning elbow joint. Similar morphology in the Homo erectus Nariokotome boy (KNM-WT 15000) suggests that this shoulder configuration may represent a transitional stage in pectoral girdle evolution in the human lineage.
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http://dx.doi.org/10.1016/j.jhevol.2007.06.003DOI Listing
December 2007